LEGUMES

OF THE
\VORLD
EDITED BY
GWILYM LEWIS
BRIAN SCHRIRE
BARBARA MACKINDER
MIKE LOCK

Published by The Royal Botanic Gardens, Kew

 Contents
PLANTS PEOPLE
POSSIBILITIES

© The Board of Trustees of the Royal Botanic Gardens, Kew 200'1 foreword·· ···· ·· ···· · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ······· vii

All righ1 s r's ·1-v •d . No pai1 c I' I his pub]j ·:11io11 may he re produced , stor •d in a rdrieval Preface · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ······· viii
system, or Lr:rnsmiued, in any ~ rm, or by any means, electroni -, mech~ink·tl ,
phot1)cupying, r ·cording or olh "rwise, w11hout writl •n permission of th · publis h r Acknowledgements · · · · . · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · ··· · ·ix
un less ill a ·ordnn ·e with Llie provbions or tlie Copyright Designs and Patents A -L1988. Artists and photographers ·ix
Copyright permission · · · · · · · · · ...... . . ..... .... . . . ...... . ..... . x
Great care has been taken to maintain the accuracy of the information contained in this
c;eneral acknowleclgements · · · · · · · · · . ........ ............. . · · · · · · · · · · · · · · · · · · · · ···· ·xi
work. However, neither the publisher, the editors nor authors can he held responsib le
for any consequences arising from use of the information contained here in.
About the book ...... .. ...... . .... . ......... ·· xii
First published 200') by
Introduction · · ...... ...... .. .. . · 1
Royal Botanic Gardens, Kew
Richmond , Surrey, TW9 3AB, UK Leguminosae or Fabaceae? · · · · · · · · · · · ....... . ...... .............. . . . . . . . . . . . . . . . . . . . 1
www.kew.org Cbssification of the Leguminosae · · · · · · · · · · · · · · · · . · · ......... . ............... ... . ... .. 3
Adv:mces in legume systematics · · · · · · · · · · · · · · . · ............. . . . .................... . . 4
ISBN 1 900347 80 6
Changes in generic alignment and tribal composition · · . .... . .............. ........ , ..... ... 4
L~conomi c importance of the legume family . ........... . , .. , ..... . .. , .. . . .. . . ..... 10
Production editor: Ruth Linklater
Typesetting and page layout: Christine Beard Complete synopsis of legume genera · · · · · · · · · · · .. ........................... .. .... . . .. 13
Cover design by Jeff Eden; cover photographs by Gwilym Lewis
Book design by Media lksources , Biogeography of the Leguminosae · '' •' •'' '''' o • o o' o o' o o'' I o o t o' o ' o' o o .. . . . •..... 21
Information Services Department, Introduction · · · · · · · · · · · · · · · · · · .......... . ........... . ......... ... ... .... ... . .... 21
!loyal Botanic Gardens, Kew
lliomes · · · · · · · · · · · · · · · · · ...... ' ... .. .... ............. . .. . ...................... 23
Printed in the United Kingdom hy The Bath Press (CPI Group) SysLematic biogeography of Leguminosae · · · · · · · · · · · · ....... . ... .. . ..... . .. ....... . .... 45

For information or to purchase all Kew titles please visit Tribes and Genera of Leguminosae ......... ......... . .. . ' .. .. ... . ..... . . .. .... . . 55
www.kewbooks.com or email publishing@kew.org Subfamily Caesalpinioideae
Tribe Cercic.leae o' ' o 0
. ... .. 57
o o 0 o o 0 o 0 0 o o 0 0 I 0 0 0 0 '0 o o o o o to 0 o

Tribe Detarieae · · · · · · · · · · · · ....... ............... .. ..... .. .... .. ... . . . . . . 69

Tribe Cassieae · . . . ....... . .. ................. ............... . 111
The costs of this publication have been generously supported
by a numbe r of individuals including
Tribe Caesalpinieae · · · · · · · · · · · · · · .. ............. . ... ' ... ' .' . ' .. . 127
Subfamily Mimosoideae
John Anton-Smith
P.D.S. Boardman Tribe Mimoseae · ..... ' ' ' ' ' ' ' ' ' ' o f o o •
0
I ' ' • ' ' o o • 0 0
'o o o 0 0
o o ' 0 0 0 0 • ' 0 o 0 ' 0 0 0 • o 0 o 'o o 163
Giles Coode-Adams Tribe Mimozygantheae ' ' ' ' ' ' • ' ' •' 0'' • • ' ' ' o. 184
o 0 0 0 o 0 o 0 o' 0 0 0 4 o 0' 0 o 0 o 0 • 0 o 0 o Io 0 o 0 o Io I

The Simpson Education & Conservation Trust Tribe Aca cieae . ....... ............... . ...... . .......... ..... 187
and three others who wish to remain anonymous. Tribe Ingeae .. . ...... ... ........ .... ..... .. .... . ............ 193
The editors warmly thank all the donors for their support which has helped to ensure Subfamily Papilionoideae
this book is accessible to those countries with the greatest legume diversity and highest
Tribe Swartzieae
- · · · · · · · · ..... . ........ . .. ... . ... . .............. . .......... ... 215
levels of endemism.
Tribe Sophoreae · · · · · · ... .. .. . ......... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227
Trihe Dipterygeae · · · · · · . .......... . ..... .. . . ...... . . . . . . . . . . . 250 .
Trihe Brongniartieae · · · · ............ ..... . ......... . . . . . . . . . . . . . . . . . . . . . . 253 .
Tribe Euchrcsteae · · · · · · · · · · · · . · .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
.. 260
Tribe Thermorsic.leae ...... . . .. . ..... . ..... .... ..... ... . . ... .. . ..
. . 26
3
:;:rihc Poclalyrieae · ....... .... . .. ... .. ...... . .. .... ......... .... 26 7
11 he Crotalarieae · · · · · · · · · · · · · · · · · · · · . · · . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273

CONTENTS v
 Subfamily Papilionoideae (contd.>
283
Tribe Genisteae · .. ........
Tribe Amorpheae •' +' o > o '
....... t

. . ....... ' .' ... .. . .. .......... .... .... . .. 299

Io •• o 'o''
•••••••••••••••••••••••••••••••••••••••••••

f oreword
• • • ' • • • • • • • • • • • • • ' ' • • • • • t • • 307
O 0 , O ' 0 ,, I'' O Ir '

Tribe Dalbergieae · · · · · · · · · · · · · · ·
0 0 0

................ ....... . ... .. ... .. 336

Tribe I-Iypocalypteae · · · · · · · · · · · · · · · · · · ·
' >
....... . ..... . ..... ' ..... ' 339
o o • •' • 4 0 • to I> o. I> t 0 o'

Tribe Mirbelieae · · · · · · · · ·
I 0 '' t
.. ..... . ' .................. 3c;c;
0 0 f > o 0' 0 o o 0 I Io 0 I 0'''

Tribe Bossiaeeae · · · · · · · · It is particularly appropriate that this fine book, the delimitation of tribes and subfamilies. The Editors
. . ... ... ' . ' . . ..... . . . ... . .. 361
Tribe Indigofereae ···················.·. . ............. . ... 367 covering all the genera of the Leguminosae, should - all from Kew - have drawn in 20 expert
Tribe Millettieae · · · · · · · · · · · · · · · · · · · ' · · · · · · · · · · · · · · · · · · · · ' · · · · c have heen written at an institute whose involvement in collaborators from all over the world. They have
.. .... ' . ' . .... . ........ .. ....... .. ..... 389 legume research is long and virtually continuous. skillfully woven together the traditional framework of
Tribe Abreae · · · · · · · · · · · · · · · · · · · · · · · · ·
....... ' ............. ' ... ... ... . .. .. . . .. ... ... ' ............. 393 Legume work at Kew began with George Bentham in the classification of legumes with the new system,
Tribe Phaseoleae ·
...... ' . ....... . ... . ....... 433 o I 0 • o o o o o o o'
the middle of the 19th century. He was the first to placing this within a novel biogeographical setting.
Tribe Desmodieae · · · · · · · · · · · · · · · · · · · · · ..... , .... . ........ 447 provide an account of the whole family as he knew it, They have made the information uniquely accessible
Tribe Psoraleeae · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · and much of his work on many legume groups is still through illustrating every one of the 727 genera,
. . . . . . . . . . . . . . . . . . . . . . . . .... . .. · · · · · · · 4S3
Tribe Sesbanieae · · · · · · · · · · · · · · · · · · ' · · · · . . . . .................. 455 valid today. He was followed by ].G. Baker and Daniel commissioning 10') new illustrations where an
Tribe Loteae · · · · · · · · · · · · · ·
> • 0 o o' • j I Io' I 0 o' 0' o o 0 o 0 I
Oliver, ancl by John Hutchinson, who in 1964 produced appropriate photograph, painting, or line drawing
... .... .. ... .. . ....... ' ............ 467 an account of all the legume genera known at that could not be found. They have consulted an enormous
Tribe H.obinieae · · · · · · · ' ' · · · · · · · · · · · · · · · · · ·
............. . ........... ... 47') dale. During the latter half of the 20th century, J.B. number of publications and incorporated much new
Tribe Galegeae · · · · · · · · · · · · · · · · · · ' · · · · · · · · · · · · · · ·
. . . . . . . . . . . . . . . . . . ' . .. . .. ' ...... 489
o • o • o '0 'o > 0 o o 0 o o I
Cilletl, J.P.M . Brenan, RM. Polhill and myself continued material during the preparation of the text .
Tribe I-Iedysareae , o t o 1 It I''''

.. . . . . ... . . . . . ' . . . . 496 this line, concentrating particularly on the floras of This work provides an encyclopaedic overview of
•••••••••••• t ' ••••••• - ' •• • ••

Tribe Cicereae ··········· ··· Tropical Africa. In 1978, Roger Polhill and others knowledge of the Leguminosae at the beginning of
Tribe Trifolieae ········ ·········· ·
............ . .. . .......... . .. . ... . ...... . 499
realised a longstanding aim of bringing together over the 21st century and is the first fully illustrated manual
............. . 50') '+ o o
400 scientists working on the legume family, to attend
t 0 0 0 lo t o o' Io I•

Tribe Fabeae · · · · · · · · · · · · · · · · · · · · · · · · · · · · · of all genera currently recognised in the family. While

. . . . . . . . . . . . . . . ... . . Sl l Lhe first International Legume Conference, held at Kew. the book is of specialist interest to the legume
Bibliography ·············· ··· ······· ·· ···· ···· ·· ·· ·· · ······ The L:onsequent first volume of Advances in Legume systematist, it has a much broader appeal in providing
S)ls/ematics (published in 1981) is effectively the a compendium of useful facts about genera for a whole
Indexes fmcrunner and seed of this book, containing as it does range of users of legume information. This includes
........ . . . . . .... .. ... . ... .... . .......... .. ............. . .. .. ... ' .. ' c;4c; a summary of all legume tribes and genera as then data on the number of species per genus, overall
Illustrations t. o' 0'
. ...... . ...... . ......... 5c;5
' o o 0 f Io 0 I 0 o Io o o 0 0 Io o o o 0' umlcrstood. Work published in the series (now 11 distribution and ecology, habit, etymology of the
Vernacular names · · · · · · · · · . . .. . . . . . . ......... 561 volumes) has been crucial in the preparation of scientific names, uses, reliable references for further
Scientific names (including phylogenetic and biogeographic terms) · · · · · · · · ·
l.eg11mes (!l the World. information and useful notes about their taxonomy.
This book is the culmination of more than ten years The book is also aimed at generalists wishing simply
of synthesising the massive advances in legume to visualise the plants they are interested in. I believe
systematics, charting the molecular phylogenetic this book will be a most worthy companion to the
revolution that has taken place between 1994 and 2005. current Advances in Legume Systematics series, leading
Phylogenetic analysis, based on multiple sources of legume research into the coming decades, as volume
data, has produced more objective hypotheses about one of the above series has done since 1981.
relationships between genera and species, and these
have led to new (and constantly developing) ideas on Bernard Verdcourt

FOREWORD VII
v1 LEGUMES OF THE WORLD

Preface
--mlJlflf,
Legumes hav been a major fo us of I< w ' r Jseas h
sin ·c Be ntham in 1855. The legum, colle ·tion in the
Herbarium comprises appr ximaLJ ly 725,000
sp dmens in ·!uding ·. 30,0 0 types a nd is o n f th
n1ost compr ·h1::ns iv · in he world hee:1us, il uniqt1c1y
contains gl h al re1 resen t~1tion at sp~ ·i s and genus
level. Kew holds at least one colle ·tio n of 726 of the
727 genera currently recognised (as yet we have no
material of th e monospecific Madagascan genus
Lemurodendron). The legume collections at Kew are
thus ideally suited to large-scale monographic and
floristic work and multi-disciplinary science. In terms
of existing expertise, Kew has a broad generic view
across the family, regional exp rtise in major tropical
areas, and taxonom i spe ia \ity at tribal and subtribal
level in select · I grou ps. Th r bas 'liso h en exteosiv ~
multi-di s ·tplinary work in rhe co.mparaLivc biology of
legumes b·1sed n wood anaromy poll n morphology,
mo le ·u !ar syst ma ti s, c mparativ ·h misLry and
us s. Th ·omb in ~cJ \at::i fr m th se lis ·ipUn1::s have
highlighted the need for a radi a l resu·u lltrin g f
higher level legume classificatio n, a proj ~ l in hich
the Kew le gume team is a ·live ly in vol ·<l in
collaboration with several int rn:ition al partners.
The idea for the book, Legu.m.f!S of the Wl'orld g< es
back to the first International Legum n nr , re n ·e, held
at Kew in 1978, during which num ~rous cof1f re n e
delegates commented that., while they worl e d on 1n
or a few legume genera, they Imel little ic.k:<1 al out th
hugJ m rphological div rsily of this vast pl:i nt fam il y.
A pau ·ityor illu. u-ations was highlight I a a gap that
n ecl ~d L I " fill e d so tha t this li v1t. r. ity oulcl b
3
visualised. The idea of one day proc.lu ·ing ~111 illustrated
guide to legume genera w . s born . Thro 1gh th e
VIII LEGUMES OFTHE WORLD
generosity of many ph tographers , the Kew legume
slide collection now numbers in excess of 12,500
photographs and has been tbe m·1in ·o ur · for th ·
illustrations in the book. As th • proje l got u nc.le1 ay,
the scope of the book evo lv <l to in ·!ud · Lb e
placement of the genera in a phyloge n etic context
based on all the recent molecular and combined
research being unde rwl en on th fam il y. Th g>ne1
are organised syst maLically so thot ac ·ess to facts
about each genus are most easily found by first us ing
the comprehensive scientific ind x on page. 561-577.
Legume· of tbe \Wo rld is a multi-~1utb ored
e n y ·I pa di a a nd the first authoritative illustrated
guid t: to Lh w rld 's 727 legume g n ··r:l . ll is an
important r·fer · n · work for l 'gl.ttne s1 i;tli:ts, bqt '
als a valuabl re >Lil' for a wi<l rang> or disciplines
that use legume informati n a.ncl wish to vlsurt!ise the
plants they are w rl<in g with a nd fine.I out more al o ut.
them and their r bt c.I gene rn. The ho I is a fa ·tual
compendium displaying Lhe wea lth of d iversity fmmd
in one of the world's mo. t economi all r imporLa nt
plant fom ll i ~ L g umin< sa , all . e l in :l m cl rn
g grap hi fram work . lt is a l o
d . ig n ed ·1 a su 1 pl m nt to th int rnatio nally
a l:tim ·d s ri s Adva nces in Lep,mn, . ystematics
(vo lu m '· 10 and 11 were published in 2003).
Numerous genera are il\u sLrat d f r the fir1'l time by
o tou r ' l photographs pa intings, line draw ings o r a
on1bina tion ol' th s >. Tb I o ok thus prov ides
d ingnosLi ' imag · s linked l bas line facts abouL all
·urr ·ntly r ·ognisecl legume genera.
Gwilym Lewis, Brian Schrire,
Barbara Mackinder and Mike Lock
Aclmowledgements
Artists and photographers
The c.litors and authors ;; ish to tha n! th many a rti rs
ho hav f r ' I ~ire l t:h !in drawing. used in th I > k.
he r · artis t's · nta ·1 a ldre ·s s wer known
specia ll y for Lh os • whc hav , ov •r rnany yea rs,
pr par -cl lin e \J-;twing. for f101·as, monogrnpbs
journ:tls published by Lh ~ R y· tl Bnwn ic.: .ard ·n. ~ ew
· i~Jn
tit · :1rt ls Ls we re 'tpproachec.I for p rm ls LC;
reprod I · their drawings. Wil11out cxceptio11 ·tll ~L gre d
to u::. using their Ogurcs in our bool a nd w ~~ire most
gral~{u l L > the m for ti~ ~ ir gen rosily . Many clr~iwing ·
WCJ' • prJparec.1 esp wily ~ r this volume (artists in
bold ~·::i. • in Lhe li s t b I w . Pa l H:Lllid ay wa •
m1111ss1o ned LO. pr? c.lu ., 10) c.lrn \ lngs of g n era
wlrre n altemauve 1mag ·oulc.11 e !' un<l . We thank
her for Lhis H rculea n achi ve me nl. P rm.lss lo n Lo
~· ' prt _lu ·' wo.rks I y oth r ::i 1t.ists h::ts I ecn given by \11
J urn.ti In whJch they we r first I uh lis hed se under
copyrigltt p rmis i n ). A b:rndful c f plar origina ll y
prepaie<l ~ r oth r worl s, but the n nev r publ ished
h:l e b !<!11 us d for th first time in Lhis b ok. Th~
name of t~e a:tist of each drawing or painting used in
the book is given next to the illustration. After some
se~rching we were unable to find the initials of a few
artists, for which we apologise.
Artists: Adam, M.-F.; Angell, B.; Asquith, A.; Barnard
A.; ~artusch, G.; Beaumont, A.].; Bergeron, B.;
Boutique, M. ; Bridson, D .; Catherine E . Chandlei·
J.''. Ch yp1e, . ' .,
G.; Coates Palgrave, O .H.; Crozier, F.; Curtis,
]., Dahlgren, R.; Davies, A.; Drake· Edwards S. Elk.ans
L . E" ' , 'I
., i asmus, D.; Farrer, A. (nee Davies} Fitch W H .
Foth · ·11 • • · ·•
V C .eigt , M.; Froeschne r, E. ; Goar. N.M.; Gordon,
.. ., Greenop, H.; Grey-Wilson , C.; Grey-Wilson CM.
GFW. nerson • M.,. H a11'e, N.; Halliday, . P.; Hart,].; Horne,' · ·'
'.mg., Ireland,
c H.E.; lwagu, T.; Jordan A· King B . I 'I
K ' .; Koorders; Lacour, J.E.· Ladete A: Laeren C
van & WI ' ' ,
].A. , . essendorp, J.; Lamourdedieu, H.; Langhorne,
R. ·· 1.aisen, G.A. & Ulibarri, E.A.; Lee, A.T. & Polhill
·•. Lemeux • J.., Lowe, J ·A. ( nee , Langhorne} Makar s.'
1 s· M'I • , .,
M
Och 11 ' i ne-Redhead, O.; Myers, ].; Natadipoera,
p. ]Joterena-Booth , H.., 0 s, J .H . van; Otero, M.C.; Owner
11., 0 I\ 'szym·k· s 1• · M.; n..C II1 1·1 1, H.M. ; R 'iche nbach '
R'.. •.; Rentz"\! · , r· vm1 &· Burka n · Riclat rd s M E .•
ier, L. van d .. Ri l , • ' ., ., . .
'.· R k ei P ey, L.M . Cn Mas >n); Ross-Cra ig
S · YP t:rna, H .; Saleem , M. Y.· '<1 ussolle- u ~r"'I I '.
, .111111 h r . 1· . ,
. A . . • · : ; • e 1ger, .J ,; Shaw, J S.; Smith , M.; Sobel,
1.M :·. ,0 ·la1 01an ltA: Speight, C.; Sr me, A. · Stone ,
·• Ian . ·1··•.; 'I' a nger111
· I' A. ·' ,lav, ·ra ' R .,· ·1.· b-b s,- M'!·
T'.n:l, K.; Trev~thick, W.E .; Turpin; Velmure, K.S.;
Villiers, J. -F.; Ynes, H. , de ; Warwick, M.; Webb, J.C. ;
Wessel, W.; Wessendorp , J.; Wickison, S.; Wise, R.;
Wood, H .; Zewald, I.
The editors and authors also thank the many
photographers who, over the years, have kindly
and donate.cl colour transparencies to the Kew legume slide
collect1on, and especially those who have responded
to ~ur requests for photographs to fill gaps for this
pro1ect . The name of the photographer is given
alongside each photograph.
Photog1·apbcrs: n Ir w ., .'.· Anton-Smith, J.;
very:inov, L. ; Bail y, D.; Barfo >l 1 .' .D: Beach 1 J H ·
Bradford , .J .C.· ~urgers, ~.;
.J.; But h 'r, Carr, G:D:;
C.~ .l.~.O .; Chap~1ll,].; Cheek, M.; Collenette, S.; Cope,
T., Cubb, P.; Cnsp, M.D.; Daly, D.; Davis, A.; De Wit,
H.D.C.; Downes, W.F.; Dransfield, J.; Du Puy, D.J.;
Fagg, M.; Ferguson, I.K.; Fortunato, R.H.; Foster, R.;
Fuhrer, B.; Gardner, M.F.; Gasson 1 P.· Geesink R.
G~ntry, G~·een, Gre;~
A.; George, A.S.; Gilbert, M.; P.;
Wilso.n , C.; Harley, R.M .; Harri s, D.J .; Hooper, J.;
H. pktns , Il.C.F.; llud{ 13,. Hughes, .. E.; Kirkup, D.;
l ltt gaa rcl H.13.; Lahar, .J .- : 1.aing, D. ; Lavin, M.; Mal:tn,
S.; Ma.a.' P.].M.; Mansano, . cl r .; Maslin , 8 .; McRobl ,
A.; Mil:Uk n, W.; ~ori, '. A..; Mu rata, .J .: Nich Ls G. R.;
~hash1, H.; Pennington, T.D.; Ratter, J.; Sanjappa, M.;
Simpson, B.B.; Sp lleob rg, R.; Stirton, C.H.; Svenning,
J. -C. ; Thomas, W.; Thompson, H.; Thulin, M.; U.S.D.A:
Van ~yk, A.E.; V:in Wyk, B .-E .; Van Wyk, P.;
I
Wagenmgen slide collection; Wie ringa , J.J.; Weston,
'
P.H.; Westra, L.Y.Th.; Wojciechowski M: Zabeau R .
Zhu , X. ' ' ' .,
We ~hank E. Robbrecht, Scripta Botanica Belgica 23
(National
f Botanic Garden • Be lgium) ' and DJ . . H arns
·
I 'I
or ~e.rmission to reproduce the photograph of
Berlinia hruneelii first used on the cover of Vascular
' .
Plants ~f Dzanga-Sangha Reserve, Central African
Republic, by DJ. Harris (2002).
R.; Four. p l.1otograp hs a r reprodu c •c.I with [b J kin I
per?1iSSIOL1 r the Australhn nlional Botanic Ga rdens
© : m tribe Brongniarlleae: I fovea /nmgens He n th .,
~t:~to: . M. D. ri 'Pi in tril , Mirheli 'a : Podolohium
th '!(~flu 11:1 Andrr..:ws r" p & P. H. w ston photo: M.
D. nsp ; m uib~ Bo ·siaeeae: JJos. i 1 ~t1 d~n.tC1ta It Br.
, .. ,
Benth., photo M. Fagg; Platypodium formosum Sm
photo M. Fagg. ·'
ACKNOWLEDGEMENTS IX
 111~~ m1111111ri,
.- . 'll'i I ._

Copyright permission
Th· >dl tors and autliurs wish I' > thank the institutions,
journa ls, ·ditors and arli~u; who ha e kindly given
permission lo reproduce the rollowlng I In , drawini;s .
Ph. lornt, und Puhlk:ition:-; cientifiqu •s du Mus •urn
n ~1Li nal d'l li. toie nmurdl , P~u·is ©: Flore du ahon
I'> Pl. l 8 : 9 1 ( J968): Loesener61 wulkeri a nd L.
p,aim1P11 ·i:; Plor • tlu Gab >11 l 5, Pl. 19: 9 ( l968l :
eocb ualierudendmn. stepbt111.ii· Flor du abon 15, Pl.
20: t 9 l 9t 8): f lyine nost ~ · ta j7v1·i/)ll JU/ct and fl.
nurnutn lii; Flor· dtt ·nhon 15, Pl. 39: l 73 ( 1968 :
, "iizduropsi:> Mesllli; I l1>re du :rabon 15, Pl. 2: 1 8.~
(1968 : Augo11ardle1 letc:stui; Flore du ~abon 15, l'I.
189 Cl' 8): A11thonotbo concbyliopbunun (originally
publish >d ri . l:>omacrolo!Ji'um cu11cbyliopbomm); Flore
lu G ~1hnn 15, Pl. 59: 2 5 ( 1908 : f,e0'/1f,/,/"d'nclro11
gab11·1pnse; and Flore du Gab >0 15, Pl. 71: 285 ( 1968):
Libre1Ji ll ~c1 kltlinei . Ph . Moral , :ind the Fl re du
amen un 9, Pl. 1 : 71 ( 197 ): G'll/ >llf demlron
pierrecm1t1n , C. mildbl'tPdii and ·. ki ant11e11se; Flore
du a111eroun 9, Pl. 17: 9 1 C 1970): Leonc1rc/oxa
a}i'fcantl ; r lore dL1 .am ' roun 9, I'!. l : 95 ('I<70):
Plci.gtnsipl cm longit11h1ts and P. muitfjugus; Flor· du
anProun 9, 1ll. 2 1: 1'11 (ll70 : l:.'z11-y/;efe1/um
1mij·11.p,um., Jo:. but >sii a nd JI lessm.a.nn'if; Plor du
·a m roun 9, PL 3 1: 151 197 )): 'J'c1.lbo1i'//ci. batesii and
7'. eketensis; Flor" dLI ..ameroun 9, Pl. 32: 153 Cl' 70);
Chi iia gabo1i>nsis ~1n I ·. J..~ tcti'nei; Fl o.r lu am roun
3
and Flore du ongo Beige 3: 331, Pl. 2 1952) :
fo/m.miod ' 1ttlro11 lef. tontb11w; Flor du ongo Uclg ~
3: 377, Pl. 28 1952): Cossureilemde1ldlw1 /}{1/scun {/emm;
Plor uu · ngo Belg ~ 3: 38'), Pl . 29 <195 . :
Ps1.mclonwcroluln11 m meng ' i ; Flore du Congo I~ "lg 3:
547, Pl. 40 ( I' 52): Al11/)bim c1s p1emccupoides; Flor du
C ng> Belg It: 287. Pl. J7 <1953): 1<0!~1msi11pbyto11
m1'1d ,1:11stii. F.H. Erteeh ~1n l Fl >1-a of Libya: 11/byltis
vu l neraria ·ubs p . mmtra Bilumfnarict /.Jit11mi110.'it1
published as P. · Jral rt /Jitumilt< s 1 , l :'hrm11s pinna/ct
a.nd H. am1ift1.i;wi, Hipp repis scabm, Lew ·11/i11aris,
M •di ago po~ymo11>hci , Ononis 1wtri.x, S/Jc1rli hum
srtbcmw, 'Frigon 4/a I.el/ala an I Trtpodion tulrtl/Jbyl/11.m
(pu l Ii. h cl :is Anthy!lis tetmpbyllc1) . Davi I H. Lor n ·.,
Anm.1 Stnn , ~ind · nvon 4 _ : I ·O ( J9 ): Ktt1tafoc1
ktiboolt:tw nsfs. A. McPherso n a n I N >v m 8 l l: 7
c1998): Hc11t1de11clmn acos/.a-solislc1111wt. A. M ·Pherson,
The lil'iSOL!ri BoLHILi ·al ra rd n Press, :inc.I wiLll the
aulh r's onsent: /famrdia cam._p yla tmlha (pt1 l li1ihed
as Pitb 'C •lluhitl/JI C(llllpyla 't /.'/lfl.nts ' in Annnts or the
issou ri 1301.anical Ga rd n 7. : 72!> ( 1986). l\ .H .
axwell and Th e Arrnals or Lh ~ MiliSOUri 13 Utnid tl
<1rd ·n ')7: 253 Jl70): G)1mho ·711w mseum. N. ilian,
and Willd · no> ia l8: 228 C1998): lf >rpyw 3rct1tcl(/7um·
WiUdcn wia 18: 227 ( l '>88): B1:v(../. 'IJ'->11/{S, . 13 B 1
13 rlin- D·1hl 111 . The m ri c n So · ie ly or J>l an.t
T:1xono111.ists an I Mclissn Lu kov r r p rmissi )11 to
use D sman1b11.s lJico rn11111 s first. puhli.<>h d ln
Syst , 111ati · 13 tany lo nogrnpbs 3 : i. Fig. 2.2 19 3) .
\'</. Burger, antl Fi ldi an~1 : T1ntlea leyer111c1rkif. T.
Rt1111ire:wl/a mi ·rcfl1/ba . T. Warra , and A 111:1nu:d c r
N ·w Guinea I gum s: .Agr1 nop , beptaµby lla and
Kingiorlendrnn allern((o//11m . W.J. Wiswall , and
on 1rihutions from the · .:. Nationa l Herharium :
'o/11p,e111 I 1. broussonetl fl ( pu blishcd as Cologrmicr
/e/11/.iS). Reim arutia: Ka!tlppiu c:ele/JiCCI. Bull . Jard . Bol.
n11itenzorg: !!11dertia spectcihf/.L'i. ln Lil ·a:,· if a small
nvml er >f illustraci ns , for whi h th e litors hav
s ught repro lu ·Lion permission on more th-in 11
<> ·c1sion , we h ' tV ·till r ·eivccl no response lo our
r •q u , srs. \Xie tru st that th ' edit.ors and artists involved
will not I .. displ ased 1hat we hav" used th ir lin
dr:iwings. The iJlustrnLions are: A1111nof1iplt1J1tb11
mm1j:!O/( ·11s, BmclJ.J'CJ /ix vageleri, Cb" ·1teya com posit
plate), DOl.1 ·11 ~>si p,erardi, 1uelde11stt1edtic1 bil'naluic:a,
. ~>011gio ·mj>ellc1 in/ ~rmedi t and Vavilovia formusc1 .
General acknowledgements
We most sincerely thank Chris Beard for typesetting the
text and designing the layout, and especially for so
p:lll ·ntly suf~ ting our many whim and sha1ing i.n our
pain; Ruth Lin.I Im r for m Liculously pr< of-readin the
t •xt and hr..dping to standardis it, :md al for her
wonderful supp01t throughout the editing process; Gina
Fullerlove, John Harris and Lloyd Kilton for assistance
with production, time-lining, and budgeting; Jeff Eden
for the cover design and John Stone for advice on design
being so willing to assist in fund-raising and Amy
Stockwell for liaising with donors, and Suzy Dickerson
and Ann Lucas for helping with editorial and copyright
issues. We particularly acknowledge, with gratitude, the
enthusiasm of Philippe de Spoelberch for all matters
dendrological, and Hiroyosbi Ohashi for encouragement
of this project from its inception . On the scientific side,
as editors we are much obliged to all the authors who
have contributed their expe1tise and time to provide a
truly global view of the legume family in this book. We
owe a large debt of gratitude to Roger Polhill for laying
the foundations on which this book is based, and for his
long-standing support and valuable advice throughout
the project; we also thank Alan Radcliffe-Smith for
assistance with the etymology of generic names, Dick
Brummitt for help with problems involving tribal
nomenclature, Neil Brummitt for providing additional
biogeographical data and taxon distributions; Justin Moat
for computer advice and generating figure 17; Sally
Dawson for preparing figure 16; Renee Fortunato for
providing information on Amburana, Anarthrophyllum,
Bergeronia, Collaea, Galactia, Ceoffroea, Holocalyx,
Lophocarpinia, Pterogyne, Ramorinoa, Rbynchosia,
Tipuana, and Zuccagnia; Rosaura Grether for the
distribution data of Mimosa; Keith Ferguson for his
persistence in tracking down some final biographical
details, and Susyn Andrews, Ruth Clark and Alex George
for proof-reading parts of the final manuscript. Out
thanks also to Alan Paton and Simon Owens for so
generously allowing us the time and resources to fully

Entwisl , and Th Roy:d B< tani · :ml ·ns , .'ydn y: and layout; Rachel Pedder-Smith for permission to use realise our objectives with this book. We are most
9, Pl. 3 : 175 1970): /emcmocole11s 111i ·rc111tb11s· Pl re
Aeuicl 11byton r , onclitum. Pfutylo/Ji11 m o/tern~/oll11 ·1n her stunning legume painting for the book end-papers; grateful to Bernard Verdcourt for providing the foreword
du am roun 9, Pl. 55: 24~ ll 7 : ddo'lliudenclro11
:.111tl P. ibt11sc111. •11lw11 , Templ •to11ia hi/ol;ct and T Marilyn Ward for help in tracing illustrations and liaising and much help besides with nomenclatural problems,
111icrantb11m; rl r du ameroun , Pl. 3: 267 ( 1970):
i11ca.1u:t original! pu! !ish d in thre dlf~ I' Ill iSS I of with several artists on our behalf; Simon Dickson for plant identification and for his continued friendship.
Mi ·1·0/pl'il11i&1 his1t!CL.tla and M. brt122 willensi ·; Flore
the j iurn a l M.ucll , ria are repro<lu e I with tile
du <1111 ·roun 9, Pl. 65: 275 ( 1970 : Tetwberlini.tl
p , rm is ·ion of th Hoyal 13oc~ini · ardens l >lbourn ,
h{(olioltt/cl ; ~ml Flore du mn ·roun c , Pl. 7 J: :H3
<Ul l.J.11. Ross. /lllcu..-ros,1111c11-u:a 11urrocc1~) 1Xi: l'JJ rinL d
I 70) : . '/a c/.l ·yutb )11'SN.S slcwdlii. Ph . Morar, and
/\uans nia Jl. : p .-3- 1 ( 1'89: Lemurodenclron with p ·rmission from I3ol bi /\ngell , IC '. Uarnel y an I
apuronii. Ph . M wat, an I Fl. 1 ~ la ouvell · .al ~doni :
J.W. rim •s, 'ilk r.r <.:, g u;ui.a ·ast , monkey's arring:
gen •ri · syst m for th · symm lrous Mimosaceae of
. cbleinllz ia insnlam.111 . J "<1n - u " I Labat, an I
the Am<::rica:,. Pttrl l. lwrenw , Alhiz ia, and allies,
Pu! ll ·aLi on ."cl ntifi 1u s tlu Mu se um nali >mll
M ' rnoirs of th .. , York Botanical G'1rdcn 7 I), c
d'llisLoir naturdle, Paris©: Peltiera nitidc1 in Laba t, J -
. & u Pu y, .J. - A revi Ion < I' Pelliem, a po rly
1c96, The Ne\. York l30lani ·al a rd ·n Pr ~ss . L.J. .
van c.le r Macse n , and LI W<tg "ningen gri ultt1rnl
known and pro! al ly cxlinct genu:-; o f L ·guminosa
niv ·rslty P:ip ·rs : Le11cmn;. balos CCl/JjJClrid us an I
C,Papilio noitle:.te - A ·s ·hynome n ac rrom Ma lag- ts ar.
Adanso nia ser. 3, 19<1 : S"i- c I, Pig. 1997). P. Baas, Bowringia discolor . .L.j .G . van d r Maesen :
Jiaplom1vsiu monopbylla ln Ubct·ian I ligh forest Tr es
an I Flora Maksi:ina s · r. ·1, Jl(I ): 1')2 Fig. -4 ( 1 92):
Falcc1/a.rirt mohtccartM (publish cl as Parc1serictnlbes
I y A. '· Voorl1o ~v » l. . ielscn , and p•rn Botani a:
;! palttth11 · 1,res sul s1 . t >res and A . fJ •rfolit1 /ll,~ subsp.
fal t /./arht suhsp. ff,//C:alaritt); Flor~1 Malcs iana • ~r. 1,
j>billif)Sii. . Ca Lr wicjo, and Flora fhcric1 : Eropbncci
L1( I): 16 1, Pig. _5 (1992): 1Jriw1tl.Ps gr,,mt!Ulom; PIOl'a
la ll'Si:10·1 s r. I I l( J : l 3, Fig . ~:> }C92): !?ti ' /fca © Flora fh eri ·:1 70 : 3 8 ( I 9);
Walla ·eotlc:11clron cal-/Jicnm: :md 11 rn Mal siana s r.
Ecbi:nospur/11111 algibic11r11 Flora ll>erin1 7( l : l27
1. , 12(2: 7 15, Pig. 52 (JC ): Uiltl nlu. 1nod >s/a . P. Raas,
(1999). ,. d " Tomasi , an I Fl >rl:I Patagonica : Acle:111 i ,1
an I l3lum ·a :~O< J): 84 (198 ): Mast rsiu bak >ri and 1.
grt1c ilis ~ind Ade mia guttu/if(;J/'CI. . C mnor, and J.
a ·scll'nica; Bluml'a . 6( 1):1 ·11. 9 1): Forciia c1lf1!f7orct;
Arnold Arbor.: ;v11111ocltul11s dioico . F. . Zuloaga, and
I nr iniana 2 l . 2 ): 02 U978} Si •11<Hll"jWn11m
Blum a 37 l : H ( J992 : Sfwtbolohus latistip11!11.;
/?ergU. M. Ederra, a nd Acme Ag ·n · S. A. : Do/icboj1sis
Blum.ea 8 2 : 82 ( 1994): K1mstleria sarnwal.ten ·ts. P.
parap,tlC1rie11sf,~. in L~1s Leguminos:is A.rgenlim1s. 11.M.
13aas, <tn<l Plora r the Guhnas: Hocotl prouacen.si.s. E.
RohhrJ •ht, Th NaLional Botani · i:ud · n of B ·lgium H ~ rn {1n I ·z and An ~tl . Tnsl. Biol. Nat Aul n . Mex .:

ACKNOWLEDGEMENTS XI
x LEGUMES OFTHE WORLD

About the bool<
,---·m111.,. ,. r==::::::==-
Editors (and authors): all from the Royal Botanic
Gardens, Kew
wilym L •wis: i. L \.Vis@I w .org
Uri.an Schrire: B.S ltrir @ K w.org
B~irhara Mackinder: B.Ma ·I ind r@ r<.e .org
Mile l.o :I : Roya l Botanic ar(h"U, I ·w (retired)
Authors
Juan Barh~11n : Roya l Botani · ·1rd ns, I w retir ~d )
j en ny Chappill : niv fl;ity of W<:!slern Ausu·alia
Mil<e Crisp : Australian Nationa l niv ' t-siry, a nhe rra
Rogi r cl I ok: R yal n t:rni ' ardens, f e
Feli Ii r •st: South African Na ti o na l Bi div e rsit y
Institute
Renee Fortunato: Instituto de Recursos Biol6gicos,
Bu nos Alr s
I-kl n Irdand: ardiff University, Wales
P te r .Jobson: Royal 13 tru1 ic arclens , . ydney
B nL Klltgaard : NaU.11'11 I-lisrory Museum , Lo ndo n
Mall La vi n: Montan~1 State Univer ity, B >z m:tn
Mdis$a Lu ckow: rnell ni v rsity, llhaca, 1::.w York
ig ~ 1 MaYt ·cl: 11iver:lly or Birmingham , J .
Hir yoshJ hashi: Toho! u Univ "'rsity , J'IJ an.
It Toby Pennington: Roya l Bot~tni · ~mien, Edin! urg h
Roge r P lhill : Roya l Bo ta nic a rd ns, K w r -·lired)
Lourdes Hico Ar ·e: Royal 13uwnic iard ns, K w
Jim Ross: Royal J3orn nj ,;miens, Victoria, Australia
Dmitry Soko loff: Mose< w Sla t ' Univ · rs ily
,harl ·s SLirton: Nat.ionaJ Botan ic Ga rd e n o f Wa l ~ ·
(retired)
Ben-Erik Van Wyk: Rand Afrikaans University, South
Africa
This v ilum • was urigin:illy dc:sign d Lo be :m illusuur >d
en 'Y fopaedia f legume genel'<I Whi •h WOU id Sc1ve as
a suppl ··m ' nl t() th lo ng-ru nning rtdvcm es in J.egwne
:ystemalics series PoU-1il! & Thwt:n, 1981; Stin o n, J 987·
Ile r ·ndee n o Dll ·h "r, 191 2; Sprt::nt & M Key, 1 9 ;
F ·rguson & Tuck r, 19 4; ,risp ' Doy I -. :I <)t '; ·
Pi ckersgill & Lo ·k !996; 1-leren 1 en & Bruneau, 20 ;
Crisp et al ., 2003; l litgmtrd & Bruneau, 2003). It Is also
a t'O r\lP ndium f fa ·ts abOLil <.'ti ·h ·urre ntly re ·o •nis d
legum• g ·nus ::111d a soun.:e b ok p >inting th e wny to
mo r • in~ nnati m. Th ' volume do s no t !ndude textual
d ~scripllons of Lht: g •nc ra and th ··rt:: ~' r no keys t
g ~ n •ra or trih •s: th I o k is thus not a full Genera
leg11.min1 sannn; this ls for a future dale . n of l11 •
'arly JLI •slio ns was wh Lh ·r Lo organi
a l p h~1hcll ·ally o r systemalicalJ . Th ~ ed itor o nclud d
tha t th ·hook sho uld represent the lat st 1111dernt~1ndlng
XII LEGUMESOFTHEWORLD
of legum · voluti n and g neri rel:ltionships, and
thal tl1 w·n ra . houkl be arr;:ing I in ·yslt!ITI<lllc r I r.
nee a . ystemali · a rrang menl had I cen ·hos ~n , Lb
scope or th ~ ulum . g rew l n ·ompass the la t l
phylogen lie information from legume r s 'art'b centres
around th w rid and rrom the burgeoning \iterarure.
A nalurnl ext m;ion. of thL as to invite a number of
legum xpen s from dif~ ~ r nt institutions to author a
·e le tion of th tril n l acco unls I res •ntc I h.erc in .
Fou rte~ n tril s bav I e ·n ritlen by s p ·ialists fr m
o utsi le I w, lifte n by Ke w I g ume rcsca r ·h ~rs
principa ll y th four editors of tl1e volu me and seven
w ·re co-auth re I by T " and non-I w ·ollnb< rators .
Th ~ fina l produ ·t ml in an lllu ·trat I guid to
I gum•. · l :n th fo r front of curr·n t leg um e
r hylog·n Li • r ~seM ·h ~111d tbis, in Lum is a signifi. , nl
step towa rd · a new lassifi ali n f the k:gume fUl1ily.
Th " Iner asi ng ly ·ompr h "ns iv, rhylogcnies now
avai lab le fo r L gumino. a prov! le ~' n< v I too l t
explur di:;trihulion pauerns that have hitherto b en
imp ssil le to visualis at fam ily, o nlin ntn l a n I g l bal
1 e ls. 'inc· 11i ~ I al l rm; f disu·ibution discu ·sc I in th
ch~1pter o n k:gum ~ biogeogr:Jph ' ,· ·hrir el et!., tlus
volum ) ar> und ·rpinn d I y these ph ylog · nies, we
have had LO Wk a s mewh·1t plu r.1listiC vie of iegum
re lali nshlps In th -• bo k. TlP syst -'marl ·ally arrang ·cl
u·ibal Lr :um •11ts that com prise lh , major pa1t o f thi
volume l:u g ·Jy fo ll ow more lr~1dili o n :t l !in •s of
cb ssiflcalion. This is simply I ·c:aus mol .. ·ular sa mpling
of many gen ra rema Lns s<> poor thar we wer ·· unal le
y ' t LO fo rmally ·ir urns -rib · n w higher taxa.
Th · 36 legume Lrihes arc d <Ill wil h in th ,
whi ·h they appcr1r in th · I ·gum · phyJog ·ny
Ea h tribal a ·ount in ludcs th .. aut ho r an I
publication Jf th tribal nam and th main tribal
synonyms . Th "S, cbw have I e n d1e ·I ·d using R v a l
(200 and by ref ·r ·n · · to th e origi nal pl ~t · of
publ ication of Lb m1111 >. Wh n ·onsid~ring m1 11Ps f
suprag · n ri · taxa in Legum in sae, those o f Ber ·hw ld
c' J.Pesl ( 182 ) have be n x ·lud ·c.l on th e advi ·e of
Marho lc.l , Kirsc hn ·r an<l , tepan ·k Turland, in
coJTe pon le n · w ith IU . Brummill al I cw, 5 April
000). Tur.l a nd (l. c. sw tes: "lsu ·11 m11n s l probably
h:1v to he tak 11 a: o rders "rncl") with s m . or them
divided into famili ("c,l , d "J. According to Marhold ,l
al., wh ile "rad" mea1 rde r 'c led " t:aJ1 o nl mean
family , hoth now ;,i n I in 1820". Tb s~ names thus have
no prio rity a t tribal level, .,g., fo r S I hor a ·', 'ytisene
(as Jitisea '), I alcn · a nd AsLrag:il '<le.
The l110Sl r '(.' '!'1 l class lfi ·ati )11S or the l.cguminos::i ~
a re thos of P lhlll "" Raven ll98ll a n IP >lhill 1994>.
Tit • intruduct o 1y t ~xr s L'i ea ·h tribe in its most current
phylogeneti · ·om xt and su mmarises the history of
rriba l cl ' limitation from 1981 hrough 1994 to 2004. The
11
u0ll 'f of gen ra ~111d sp ·i •s in each tribe is included.
Also incl uded Is a diagr;1mmatic representation of the
lat Sl Vi W of pltyJogeneti . relationships among genera
within th Lrib nncl w ith o ther related tribes. Groups
of gen 'nl : upported in phylogenetic analyses are
w;11ally referr d to a. clades; such aggregations that
ha v · r lalivcl p or or no s uppott at present are called
grc ups. T xtual I ox s ar ·olour coded. Blue boxes in
the li ~tgram intlicat taxa that are treated more fully
els Ii 're in d1e b ok and are cross-referenced to a
page m11nl ~r , w hile gr y boxes represent taxa fully
1r nt -'tl wi1hjn the trad itional circumscription of the
tribe u ncl r co n id ·ratio n. Buff-coloured boxes
represent other g roup ings of tax noml · ignifl a n e.
References to re · otly publish d phylug nies on whi ·h
the diagram is based are incJucl d in the figur ~ c~q tion
and, in detail, in the bibliography on pages 511-544.
The text accompanying each genus follows a
standard layout.
The generic name is followed by the author(s) and
date of publication of the name. Author's names are
abbrev iated according to Brummitt & Powell 0992).
The date of publication of the name has been
checked in Index Nominum Genericorum (Farr & Zilstra
(eds.), http://ravenel.si.edu/ botany/ ing//ingForm.cfm)
and, in many cases, by reference to the original place
of publication (especially where the date in ING
conflicted with that given in other published works).
Generic synonyms are given, but these are not
exhaustive, concentrating rather on those commonly
used in the legume literature during the past forty years
(e.g., those given in Hutchinson, 1964; Allen & Allen,
1981 , and Polhill & Raven, 1981), Fo r a more detailed
listing of synonyms see Gunn 0983).
Ne~t the number of species in the genus is presented.
~hts was based on a thorough re view of existing
literature, consultation with experts and new counts of
taxa made from the herbarium collections at Kew.
Where ranges of species numbers are given in the
gene ric treatments, spe cies totals for tribes are
represented with the lowe r and upper limits in
parentheses separated by the average number of
species , e.g. (lOO) - 120 - (140). This has resulted in a
freassess
. · ment o f t h e total number of species in the
amity, which has risen from c 18 000 in 1994 to c
19 325 111
, , ' I ,
th' this volume. This latter figure is the sum of
e average totals given for each genus.
Geographical d ·'tstrlb ution
. . ordered by continent
.
ts
Ca
grs normal!
. Y un Ic rs L o d) and then by country or
ff< ~ps of ·ciumri ~s , whi ·h ·v r is th · most informative
Ph) .Ow ing I lol li s & Brummitt, 1992 . For Africa,
Ytogltig rap hi ·a l r gions la rg ly f >llow the
phytoc horia of White (1983) . For the Americas
distinction is drawn between species numbers in the
Caribbean, Central America, Mexico and North America
from those in South America , as well as between
Ama zonia and circum- or extra-Amazonia. These
divisions were informative on more detailed analyses
of the biogeography of the family. Abbreviations used
in the text are N = notth; S = south; E = east; W = west;
C = central and sp. = species (spp. in plural).
Etymology is given next, where the derivation of the
gene ric name is explained. The main references used
when researching the origin o f names were: Wittstein
(1856), Exel! (1960), Allen & Allen 0981), Quattroccbi
(2000) and Stearn (2002), as we ll as the original place
of publication. Greek and Latin prefixes and suffixes
were checked in Radcliffe-Smith 0998).
Habit categories are simplified to trees, shrubs, lianas,
herbs , and occasionally suffrutices (where annual
shoots appear from an underground woody base,
White , 1977). Shrubs branch from the base, and lianas
are woody-stemmed climbers.
Ecology broadly combines the ecoclimatic zones of
Breckle (2002), Le., equatorial or aseasonal tropical,
seasonally dry tropical, subtropical, Mediterranean,
warm to cool temperate , and physiognomic vegetation
type following the definitions of White (1983), i.e.,
forest, woodland, bushland and thicket, shrubland and
grassland. Local terminology for particular vegetation
types, especially in South America, is sometimes
included in parentheses, e.g., semi-arid thorn scrub
(caatinga), is a subset of seasonally dry tropical
bushland and thicket.
References to each genus are a rranged in chrono-
logical order. They range from papers treating the
taxonomy of a few species to generic revisions and
monographs, and papers dealing with the phylogeny
of the genus. They also include , where appropriate,
refere nce to a series of regional checklists (Lock, 1989;
Lock & Simpson, 1991; Lock & Heald, 1994; Yakovlev
et al., 1996; Kumar & Sane, 2003; Lock & Ford, 2004)
published by Kew as part of its contribution to the
International Legume Database and Information Setvice
(ILDIS). All the references in the book are included in
the bibliography on pages 511 - 544, where the overall
arrangement is alphabetical by first author, then
chronological in multiple works by a single author.
Two-authored works follow single authors and are
arranged alphabetically by second author and
subsequently in chronological order if the same two
authors have published more than once. Three or
more-authored works are cited as 'First author et al.' in
the body of the text, and such works are arranged
chronologically in the bibliography (regardless of
number and order of names of the subsequent
authors), with dates of publication highlighted in bold
ABOUT THE BOOK XIII
to help readers locate the approptfot reference. Where
references cite an author or ci uU1 rs in the work of
another author(s) or editor(s), the reader should seek
the main publication in the bibliography (e.g., for
Verdcourt in Milne-Redhead & Polhill, the publication
is to be found in the bibliography under Milne-
Redhead & Polhill and not under Verdcourt). The
hi.h liog ra phy brings l >g Lh r for Lh e fi rst time an
c no rm us body of lite rnLure on legumes and it is a
res >urc Lh:it w ' trust wil l. be useful to th reader w l1<
seeks further information.
The notes section includes discussions of generic
relationship , phylogenetic cross-r fe r nces, alternative
taxonomic views, problems (such as taxa in need of
revision), and gaps in ur knowledge or Lhe genus or
gr up of species within it (e.g., geog raphica l regions
wher-> r·1xonomic d·1ta is lacking or weak). < meti.mes
the notes include comment on morphological
charn Le n; lhal clisringui h o ne genu · from its d se
relatives ( .g. in trib . Boss ia ae an d Mirbelic.:ae).
Thes .. cbta are espe ially pertin nt in genera for whid1
such details were not discussed in the tribal accounts
in Polhill & Raven (1981).
The uses entry aims to cover the major (and many
minor) modern and historical uses of the genus (use
categories largely follow those of Cook, 1995). Legumes
have an extraordinarily wide array of local and
commercial uses and a summary of these for the whole
fa mily is presented in the introduction (pages 10 -12).
Ln forma tion on uses has been gleaned from Burkill
0935), Watt & Breyer-Brandwijk 0962), Corkill 0979),
Allen & Allen 0981), Duk 1981), and Burkill 0995),
as well as from various monographs, revisions, floras
and other diverse sources, including the internet and
herbarlurn s p ~ ·im n Cield lab Is. ln lhe uses sectio n
nrnny locally used plani n:ini s, as w II ;1s names us cl
·0111 11·1 r ·ia lly (es pe ia lly in llJe Lirnhe r tr cl nre
in 'lud d. An index to vernacubr nam •sca n h found
on pages 5'i5 -559 :rnd th e re is also ::i s pa ra te
c mpre hen.sivc index to au sci ntific names, in lutling
informa l g rm1p and clad ' nam s, on pages 511 - 577.
Th e comp ilalion of ;1 ·olle ·Uo n o f' images of all
leg1-1111 genera has been a time· ·ons u1nl11g task. At
Lh begi nning of the projecL th " inte nLlon w<1s 1.0 us
o nly I l l l' photographs, bu t il soon I eca m , <tppar nl
tha t tbJs was 001 an a hi vable goal. M'tny gene ra bav
never been photographed, some are very rare, and a
f ~w are th ughl to b ·· extincl. 'om ~ genera com.prise
on ly tropica l trees with unkn ow n phcnol ogies, so
finding lhe 111 in flow r and rrnil is a clianc ve nt.
Some a r " mergent fore ·t g iants fro m whl h it is
cliffi ·u ll to ·o ll ·t fcnil e mate rial. WI thus opted f r a
mixture of illustrations, including paintings and line
drawing, . Ev "11 so, we were unable to find publishable
illustratio ns f ) r more than 100 genera. One artist, Pat
Halliday, was commissioned to prepare 105 new line
XIV LEGUMES OF THE WORLD
drawings of these g nera. Ea '11 of the 727 gen ra is
r presented by- at I "'~1 :> L o n · image, an I, wid1 a few
x ·eprions the maximum number of images per g nus
is tbre " Authorl'i of some Lril es provi le I 1mny of the ir
own images to illustrate their text, but the final
selection of all illustrative material lay with the editors
and Lil y t~1ke Full respo ns ibiliLy fo r any in.('o rre ·t
ldentifirnUons. Tmag s were ·e lecce I usln g the riteria
of p11 bl is bable qua I ity, aest beti app ea l, cle::t r
r presentation of dia •nostic F"'aturcs or rh genu s a
rang of' morphologica l dive rsity for s pt.:c.iose genera,
a nd a ba la nc of flowe rs and fruits throug hout tbe
book. Illustrations accompanying each genus are
re pro<lu ·eu al on qut1rrer A4 s iz a nd are Lhus, in
most c:1ses, mu h r luced fr m d1 orig inal. o as not
to clutter rh im·1 ~s us 1 in 1he bool , no sea l I ars
are included an I th e re ar no full I gends to rh
lrawings. While 1his will b fru Lrnting to om u rs,
·onomies r space dlclaled thl formal . J~~l ·h lmag r
illusLrnlion is labe lle d with the pl ant nam nn I the
name r Lb ph tographer or a 1tist. llor drawing: which
re pres nt a s ingle sp ci s, all numbe ring or lette ring
has b e n remov d , bu1 fo r illustratio ns wi th more lb.an
one species on a plate the lettering or numbering is
retained to show which parts represent which species.
A full list of a fo1owl edgeme nts to all lhose who have
l incll y given permiss ion to re produce anwo rk an d
pho1ogra phs is gi e n o n pages ix - xi. There is a lso an
ind x 10 ttll illustrati m n p~1ges 545 - 553.
The book contains three indexes, one to illustrations,
one to scientific names (including phylogenetic and
biogeographic terms) and one to vernacular names.
In the s ientiflc index, page numbers in bold refer to
the main citation of that entry in the text; all scientific
names are in roman and synonyms in italics. The
inde es, in a ldiLio n lo Lh contents pa ge and the
om ple Le ~ynopsis >f legum • g ncra ( pages 13- 19),
an.: designe I m lead Ill e read r ro all th info rmation
c nta ined in Lhi s vc lum . Tb , hook serv s different
audiences: it can be used simply as a pictorial
encyclopaedia to legume div · rs.iry; as a resource for
those with interests or rrof ssions that utilise legume
111forma1ion and who need to visua lise the pla nt. they
are dea ling with; as a ·omp ndium of facts ab i.H, ~ind
ref rences to, legum g n ra, m as a s ·ientifi c worl
placing the legume family in a modern phylogenetic
context for those seeking comment on legume
evolution, biogeography and relationship.
The editors intend to maintain Legumes of the World as
a current overview of the family. We will be adding
nnny- add itional imag f legum ca ·a as the pr je ·t
develops and we wo uld Iii to nc >urag read rs Lo
ont.dbute to this, parti \.llarly lf no I ur imag s a re
yet available for a genus.
Introduction
The t eguminosae c r I ea n and pea family is 1hc third
la rgest flow ering p.lant fam il y aft •r th e or ·bids
0 1 hl la ·eae) and dalsi s (Ast ra ·ea r mp sltae).
Legumes vary in habit from ·phemeral her! s l shrub ',
vine ·, wo cly ·limbers li::mas and giant e m ~ rgem
f r st u·ecs; B w a re aq uati s. They are to be found
ns major t:ornp ne.nls of most f th world 's vegeration
ryp •s and many hav the abili ty to ·o lo ni. ma rgin al
or I arren lands I e ·aus o f th ' ir ·apa ·ity L fix
atmosphe.ric niu·ogen thr ugh ro >l no clul ,
200 1). In this book we re ognls 727 legunp g n ra
and c. 19,325 species. The last overview of the
om pl ' l ~ family is th at of Polhill 099 )
r •ogni:;ed al ta! of 671 ge11c1y1, 56 less than pre ·e ntcd
her . Thii:. in · as in the number of genera in rh past
ten years is large ly du to the recognition at g n ric
rank of several segrega tes of larger and previous ly
pa ra phyl ll (unnatural) genera. fn ther w rd s, om e
g n ·r:1 hav bee n sp lit into two or mor s mall e r
g n ·r~1. Tn 1his boo k we have largely re ogni d
gen ri · Sl:g r gates, where th se ar ' suppo ned in the
r '<.:enl lit ~ ran.1r~, I e ·a use this will encourag · "' ider
sampling within large and problematic taxa.
As noted by Polhill 0994) the principal unifying
feature of the family Leguminosae is the legume. This
(with a small number of exceptions) comprises a single
superior carpel (a few species in tribe Ingeae have
several free carpels per flower) with one locule (some
species of Astragalus and Oxytropis have a septum
intruding from one of the sutures rendering the carpel
essentially bilocular), parietal placentation along the
aclaxial suture, and ovules 2 - many, in two alternating
ro':'s .on a single placenta. The most common type of
frLut is a pod with two valves that separate and twist
to expel the seeds, but this has been modified in many
ways to facilitate dispersal by animals, wind and water.
Sometimes the seeds are in hardened seed-chambers
which do not open and are variously nut-like, winged
flesl1Y or lx1oyant. In other taxa the seeds are brightly'
coloured and exposed on fruit valves of contrasting
colour, or p 10v1 · 'de d wit. h fl es h y stalks which
n1nmm·11 • ' "incl I . ·d . I ·11 tJ
11 s. n st1 o 1e r gro 1ps f gen ra Lh "
whole f1 11' t 1·s <-1·1v1.<- · IecI mlu
· s .mgl - edec.I ch ambers
. ith di~''·1 1·l· 1· cu l·ale , '- 11· j 1 d ·is p e rsa
Wh '
l by
anrma.ls
I Lll'tl\ ·r <.!ri h
an cI I1y I ns1 · les or hookeu hairs o n the
I10mb e r ~ T l . . .
·· 1c: g nu s !Jn.tada 111
Mimosoid . a <".1n proc1u ' f ru1ts . we ll >V ' r n • metre
1On g from
. . flow rs Lhal meas ure I s. Lhan fiv
n11l111nctre 5. Tl t
•. · i · egume "e I- m il ls also unique· the
Pit 1c r111Js 0 1111 . . . 1. . .. . . .
,~, . s a c istm ·r1ve palisad e
•ra 11sa nd lh 11 I . .
fh Yl oc enn1s 1s almost always comprised
ourglass-s ha pe<.l cells Po lhill , 19 4 .
Leguminosae or Fabaceae?
Two questions are frequently asked about the legume
family: what is the correct name for the family?, and
should legumes be treated as one family or three?
(Lewis & Schrire, 2003a).
There is an increasing body of evidence in support
of the legumes being one monophyletic family (Doyle
et al., 2000; Wojciechowski, 2003; Wojciechowski et
prenL al., 2004) and this seems likely to be reinforced over
time given the degree of interrelatedness between
taxonomic elements within the legumes compared to
h that between legumes and 'nearest neighbour' families
(Polygalaceae, Quillajaceae and Surianaceae) in the
Fabales. The legume supertree (Fig. 1) is based on the
chloroplast matK analysis of Wojciechowski et al.
(2004) which found 100% bootstrap support for a
single monophyletic family.
Several recently published floristic accounts still refer
the legumes to three separate families (e.g., Nielsen,
1992 and Hou et al., 1996 in Flora Malesiana and Beny
et al., 1998 and 1999 in the Flora qf the Venezuelan
Guayana), but the most recent regional account of the
legumes, The Leguminosae qf Madagascar (Du Puy et
al., 2002), follows the latest systematic evidence and
recognises the legumes as one family with three
subfamilies: Caesalpinioideae, Mimosoideae, and
Papilionoideae.
The three-family argument for treating legumes
must now be considered untenable on two counts.
Firstly, the Mimosoideae and Papilionoideae are
apparently unique and distinct lineages which arose
independently within the basally branching, non-
monophyletic caesalpinioid alliance and they are not
comparable to it on the same taxonomic level.
Secondly, the Caesalpinioideae, as traditionally
circumscribed, is currently under detailed scrutiny
(Bnmeau et al., 2000, 2001; Herendeen et al., 2003a)
and division of the subfamily into several more clearly
definable groups comparable in status to the other two
. attract
subfamilies seems inevitable once further detailed
studies have been concluded.
Having established that the legumes are one family
·
oflen there remains the problem of what name to give it,
and on this issue legume specialists differ on whether
subhmiJy to use the family name Leguminosae or Fabaceae.
Both names are acceptable following Articles 18.5 and
18.6 of the International Code of Botanical
Nomenclature (Greuter et al., 2000). Fabaceae, based
.'
w ith L
w 1ste d on the genus Faha (now considered as a generic
synonym of Vicia), was first used by]. Lindley 0836) .
For those who prefer that the suffix 'aceae' be added
INTRODUCTION 1
in all cases to the stem of a legitimate name of an The options, therefore, ;ire :
TABLE 2 Size classes of legume genera by species number
in cl ud ed gen u s , Fabacea e s tand s as the o nl y 1) Legum inosae (at family level): Papilionoideae,
acceptable name fo r the family derived from genus Mimosoideae and Caesalpinioicleae (at subfamily level). Number (and% of total) genera
~ tal legume genera 727 Number (and % of total) species
Paha (Stafleu , 1978; Isely & Polhill , 1980). Use of the 2) Fahaceae ( = Leguminosae, at family level): Faboideae T~tal legume species 19,32 7
term Fabaceae is, however, ambiguou s because it may (= Papilionoideae), Mimosoideae and Caesalpinioideae 192 (26%); 130 of which are range
Monospecific genera 192 (1%)
refer eith er to the legume family as a whole (thus (at subfamily level). The legumes are here treated as restricted
including all three subfamilies, as in Zarucchi, 1993), one family.
or to subfamily Papilionoideae (which contains genus Genera with 2-10 spp. 304 (42%) c. 1,400 (7%)
3) Fabaceae (= Papilionaceae, at family level). This is
Faha) w h en this is considered under its alternative Genera with 11- 99 spp. 190 (26%) c. 5,305 (28%)
treated here as one of three families with Mimosaceae
taxonomic sta tu s as one of three sepa rate families , the Genera with 100 spp. or more 41 (6%) c. 12,430 (64%)
and Caesalpiniaceae as the other two.
Papilionaceae (as in eill et al., 1999).
The u se of the family name Leguminosae has There remains a need to avoid the ambiguity presented
existed as an :.1ccepted alternative name for Fabaceae, in options two and three, both of which are illu strated
internation:d community into agreement about this Adesmia (c. 240 spp.); Rhynchosia (c. 230 spp.); Lupin11s
since the 1978 International Code of Botanical in the various examples given above. The preferred
" l"" 'll1d avoid in future the often confusing citation
u s.,.,~ .
(c. 225 spp.); Swartz ia (c. 180 spp.); Aeschynomene (c.
Nomenclature (ICBN) permits the use of alternative option is, therefore, the first one, where the legumes
of F:i haceae in the lite rature. 180 spp.) and Dalea (c. 165 spp.). With a current family
names for certain families "sanctioned by long usage" are recognised as a single family, the Leguminosae.
tota l of 19,327 species (averages are taken for genera
(Stafle u, 1978). The most recent ICBN (Greuter et al., This is the nomenclatural standard set in volume 10 of
where a range of species numbers is given), the above
Advances in Legume Systematics by the editors
2000) also states (Article 18.5) that Leguminosae is a
Klitgaard & Bruneau (2003). It is also the standard
Classification of the Leguminosae figures show that over a qua1ter of the species in the
validly published name and it is thus an acceptable family occur in just the four largest genera .
alternative to Fabaceae. adopted here in the hope that we may e ncourage the The family is currently divided into three subfamilies The next twenty one largest gen era (i.e. all the
(Tables 1 & 4) and 36 tribes. Subfami ly Caesalpinioicleae o th er genera containing over 100 species) are:
comprises four tribes and c. 2,250 species, subfamily Hedysarum (c. 160 spp.); Vicia (c. 160 spp.); Lathyrus
Mimosoideae four tribes and c. 3,270 species, and (c . 160 spp .); Bauhinia sens. strict. (c. 155 spp .);
TABLE 1 Main diagnostic characteristics of the three legume subfamilies
subfamily Papilionoideae 28 tribes and c. 13,800 species. Millettia (c. 150 spp.); Lotononis (c. 150 spp.); Eriosema
The top twenty largest legume genera are: Astragalus (c. 150 spp .); Calliandra (c. 135 spp.); Daviesia (c.
(AE SALPI NI OID EAE MIM OSOIDEAE PAPI LIONOI DEAE
(c. 2,400 sppJ; Acacia sens. lat. (c. 1,450 spp.); Indigq/em 135 spp.); Alhizia (c. 130 spp.); Ormosia (c. 130 spp.);
Trees, shrubs, lianas, Herbs, shrubs, trees, liana s, twiners (c. 700 spp.); Crotalaria (c. 690 spp.); Mimosa (c. 500 Machaerium (c. 130 spp.); Onohrychis (c. 130 spp.);
Trees, shrubs, lianas
rarely aquatic herbs spp. ); 0.\)1tropis (c. 350 spp.); Tephrosia (c. 350 spp.); Phanera (c. 125 spp.); Lotus sens. strict. (c. 125 spp.);
Ownwecrista (c. 330 spp.); Inga (c. 300 spp.); Senna (c. Lonchocarpus (c. 120 spp.); Erythrina (c. 120 spp.);
Flowers relatively large Small regular flowers aggregated Pea flowers 300 spp. l; A.~palathus (c. 278 spp.); Desmodium (c. 275 Gastrolobium (c. 109 spp.); Mucuna (c. 105 spp. );
into heads or spikes spp. l; Dalhergia (c. 250 spp.); Trijblium (c. 250 spp.); Vigna (c. 104 spp.) and Pultenaea (c. 104 spp.).
Flowers generally zygomorphic Flowers actinomorphic, radially Flowers zygomorpic
symmetrical TABLE 3 The main characters distinguishing Leguminosae subfamily Papilionoideae from Polygalaceae (the milkwort
family)
Petals imbricate in bud Petals valvate in bud Petals imbricate in bud
LEGU MIN OS AE (Papilionoideae) POLYGALACEAE
Median petal overlapped by others Median petal not overlapped by Median petal (standard, banner or
(when these present) others, similar in shape and size vexillum) overlaps others (these Stipules generally present Stipules usually lacking (sometimes represented by glands)
occasionally absent)
Leaves mostly compound (rarely 1-foliolate), mostly Leaves simple, usually alternate, sometimes reduced to
Sepals united at base into a calyx tube alternate (one genus opposite), with pulvini for "sleep scales, without pulvini
Sepals generally free Sepals (and petals) generally movements"
united at the base
Sepals 5 (upper 2 occasionally fused), rarely 2 sepals Sepals 5, the two lowermost united or the two laterals
Seeds generally without pleurogram Seeds usually with open Seeds (if hard) with complex hilar petaloid (3 genera) enlarged and petaloid
(if present this closed); also without pleurogram valve (beans and peas); pleurogram
Corolla with (0-) 5 petals, usually a standard, 2 free Corolla usually reduced to 3 petals, the lowermost concave
a hilar groove absent
wings and 2 joined keel petals, no fringed crest (boat-shaped) and sometimes with a fringed crest
Embryo radicle usually straight Embryo radicle usually straight Embryo radicle usually curved Stamens usually 9 or 10 (sometimes fewer or more) Stamens (5-) usually 8

Leaves bipinnate or pinnate (rarely
simple or 1-foliolate)
Stamens (1-) 10 (-many); sometimes
dimorphic or heteromorphic
Petals most showy part
Compound pollen (polyads) rare
Root nodules uncommon, but many
associations with fungi
Leaves mainly bipinnate and often
with specialised glands; Australian
acacias have phyllodes
Stamens (3-) 10-many (sometimes
over 100); all the same
Stamens most showy part
Compound pollen (polyads) common
Root nodules generally present
Leaves 1-foliolate to once pinnate
(a few palmate); some with tendrils;
only one rare species bi pinnate
Stamens (9-) 10-many
(sometimes dimorphic)
Petals most showy part
Pollen in single grains
Root nodules generally present
Anthers dehisce by longitudinal slits
?vary generally with a single carpel with 2 - many ovules
in two alternating rows on a single parietal placenta
Style never bi lobed, although stigma sometimes complex
and hairy
Fru it a legume dehlscing along upper and lower sutures,
or a lornent breaking into single units, or drupa ceous,
Se\lerat va riations on these gene ral typ es, some
lnctehlscent
Seeds not hairy
Anthers usually dehisce by 1-2 apical pores or a
v-shaped slit
Ovary of at least 2 united carpels with a single pendulous
ovule on an axile placenta in each of the 2 locules
Style bilobed with a receptive lobe and a hairy, sterile lobe
Fruit usually a loculicidal capsule (but several exceptions)
splitting into two 1-seeded valves
Seeds often with stiff rigid hairs

2 LEGUMESOFTHEWORLD INTRODUCTION 3
 Nearly 500 legume genera (68%) are small, being
either monospecific or containing up to ten species. In
total these ~1ccount for c. 1,592 (or 8%) of legume
species. This compares with c. 231 larger genera
(containing 11 species or more) which account for c.
17,735 (92%) of species in the family. These statistics
for the family are presented in Table 2 on page 3.
It is now generally accepted that the Leguminosae
is most closely related to Polygalaceae (Table 3),
Surianaceae and Quillajaceae, which together form the
order Fabales (Angiosperm Phylogeny Group, 2003).
This revises the long-standing view that the family was
considered to be more closely related to the
Connaraceae and Sa pindaceae (Dickison, 1981 ;
Gottlieb et al., 1994).
Advances in legume systematics
.ln Lhe last jO y ·~1rs, legume sy ·temaLics ha. I nefoe I
fro m mnjor a Jvanc s in our un l e rs tanding of th
morphology an I d:1ssiflcati<>n of th family (Polhill ~
Rav Jn, 1981; Polhill 199 ), as w ll as fr m more than
t<::n yt::Lrs of inL ~ n. iv· molecu lar phylog n tic research
by severa l grmq1s or r ,~~1r b rs around Lh ·world. By
r vealing that severa l or the \nforn ally m11n d g n ric
grt upin gs, l c.:gum~ rrib s, and v " 11 sul rainily
,a s;tlpi ni i .lt.:He, as t:racHtionally cir: ·t1rn. cril "U, ·ire
n ) I mtluraJ groups, Lhc analy s hw ~ oflen re ulled
in radl ·1tl r <.I finlti nor g ncri . an I tribal limits, and
to the discovery of previously overlooked relationships
amongst taxa. Although it is too premature to recognise
all of these new groupings formally, the analyses do
indicate a largely robust pattern of overall relationships
in the family. For this introduction, we have therefore
constructed a family-wide supertree (Fig. 1) derived
from the latest analyses (cited in the legend to Fig. 1
and in the tribal accounts of this volume), with terminal
taxa representing putatively monophyletic groups of
genera. The supertree also repr s nl. an important
facet of this volume - it is a 'snapshol' of our current
state of knowledge about relationships within these
putative monophyletic groups, set within the traditional
tribal context of the remaining chapters for ease of
cross-reference between the old and new placements
of genera.
The order of the tribes presented in Legumes of the
World follows this supertree of the family, starting
with the more basally branching tribes ::incl working up
Lbrough th • le gume phylogeny by pr s nLing eac.b
sl, t r g r up in turn. Where a tribe , as traditiona lly
circumscril d, is now kn wn to be no n-monoph.yleLic
( >.g., th oplior ~1 "; l~e nnington et al., Lhis volume ,
it w.lll app ~:ir on th • su1 rt re mor Lilan o nce. The
Suphcm:ae is thus repres nL cl on lhe tre hy some
e ight differenl lineages Fig . ll , rn b ing Lh ·
Soph o r a • sens. strict. and the ot h r s v n being
km.~ ms of S >phorc..:ae sens. fut . LL is thu. •vidcm that
th · tri l ·, allh ugh d all with in its traditional all-
embracing sense in this hook, will ultimately be
4 LEGUMES OF THE WORLD
segregated into several distinct ent1t1es, only one of
which will continue to carry the tribal name Quillajaceae
Sophoreae . We do not provide new tribal names for
~---------------- Polygalaceae
these entities in the book, although the authors of the
1 - - - - - - - - - - - -- - - - - - - Surianaceae
tribe do use a series of informal clade names to denote
Fabales
groups of genera. \Xlhere st1pporl for Lriba1 segrga tion . - - - - - - - - - - - - - - - - - Cercideae (page 57)
is already robust tb e n > e bave followed this, e.g., the Detarieae sens. lat. (page 69)
Sesbanieae is reinstalecl as ·1 Lribe distin ·1 from the
Cassieae sens. lat. (Duparquetia; Dialiinae), page 111
Robinieae (Lavin & Schrire, pages 452-453, this
Legumino sae Caesalpinieae sens. lat. (Umtiza clade), page 127
volume) . Similarly, where there exists strong support
for lumping tribes wg ' Lh ' I' w have folio' ed this, ! - - -- - - - - - - - Cassieae sens. strict. (Chamaecrista, Senna, Cassia), page 112
e.g. , Dall ergi :\ "' · e 11.'i . lat . here includes the Caesalpinieae p.p., incl. sens. strict. (page 129)
Adesmiea , the A s liynom ·ne;i and subtribe B1yinae Dinizio, Pentaclethra (page 165)
J
of the Desmodieae (Klitgaard & Lavin, pages 307-335, Mimoseae sens. lat. (page 163) incl. Mimozygantheae (page 184)
this volume). Mimoseae sens. strict.
Caesalpinioideae - r----- Acacieae sens. strict. (page 187)
Mimosoideae Acacieae p.p. (Senega/ia)
Changes in generic alignment and Papilionoideae lngeae (page 193); Acacia p.p. (subgen. Phyl/odineae)
j
tribal composition in the legumes \
=weakly suppor ted clade in
Wojciechowski et al., 2004
Swartzieae sens. strict. (page 215)
Sophoreae p.p. (Alexa, Angy/ocalyx, etc.), page 228
l
since Polhill (1994) 1 - - - - - - - - - - - Sophoreae p.p. (Myrospermum, etc.); Swartzieae p.p. [=Aldinoidclade]
The following is an update of the tribal and generic Dipterygeae (page 250)
classification of the family since 1994. The reader is also Sophoreae p.p. (Cladrastis, Styphnolobium, Pickeringia) , page 228
referred to the complete updated generic synopsis on r - - - - - - - - - - Sophoreae p.p. (e.g. Calio); Swartzieae p.p. (Lecointeoid clade), page 216
pages 13 - 19 and Ll1e sysLenK1t\ · biogcogrnphy s ·ctioll !---------- Dalbergieae p.p. (Vataireoid clade), page 309; Sophoreae p.p. (Sweetia,
of l.cguminosa in th second rx11t of tJ1 , I iog ography Luetze/burgia), page 228
·hapter (.' fo'ir ,, al., pages 1-5 , thb v lum . Each Sophoreae p.p. (Ormosia, Acosmium, Dip/otropis, etc.), page 228
trib -· i.n l he I ol has its own intr It 1cwry · 'cli n and r - - - - - - -- Brongniartieae (page 253)
50 Kb Inversion
more detailed phylogeny relevant to that tribe - - - Sophoreae sens. strict. (page 228)
(represent I in an abbreviated form as a diagram), but Euchresteae (page 260)
Genistoid clade
here we s ummarise the major generic realignments Thermopsideae (page 263)
and tribal updates that have taken place since Polhill's Podalyrieae (page 267)
(1994) classification and also indicate where major Crotalarieae (page 273)
realignments are to be expected in the future. Genisteae (page 283)
Dalbergieae p.p. (Hymeno/obium, Andira), page 3o 9
Tribes in subfamily Caesalpinioideae Amorpheae (page 299)
Dalbergieae sens. lat. (incl. Adesmia, Da/bergia & Pterocarpus clades),
As in Polhill (1994), the Caesalpinioideae are divided Dalbergioid sens lat. clade page 307
into four tribes , the Cercicleae, Detarieae, Cassieae and Sophoreae p.p. (Baphioid clade), page 228
Dalbe rgioid clade
a sa lpinJ ac. Th ea rliest I ranching d . de i.n the ~--- Hypocalypteae (page 336)
I gum phylogeny (Fig. 1) is trib r ·id 'a in hich f--- - - Mirbelieae (page 339)
Polhill ( ·1 re ognised five gen ~ .i Jn two :-;ul trib . : Bossiaeeae (page 355)
Old World clade
Cercidinae and Bauhiniinae. In the account of the ~--- lndigofereae (page 361)
Cerci<leae by Lewis & Forest (pages 57-66, thi:;
Millettieae (page 367); Abreae (page 389); Phaseoleae p.p.
volume) t'> elve gener::1 are re ·ogn ised. Seven n·1111es,
._____ Phaseoleae sens. strict. (page 394); Desmodleae (page 433);
pr · iou s ly con.s ider' d as gen ri · synonyms of
Psoraleeae (page 447)
Htmbfnia , are reinstated at g "n ' ric rank ba ·ed on
Sesbanieae (page 452); Loteae (page 455)
molecular and morphological data.
~-- Robinieae (page 467)
The Detarieae continues to comprise 82 genera
(Normcmdiodendron, described by ].Leonard in 199.'1 IR LC millettioids (Callerya, Wisteria, page 369), Galegeae sens. /at. (Glycyrrhiza)
was included in Polhill's synopsis as genus 1.4.8a l, Hologalegina Galegeae sens. lat. (Astragaleae), page 475
although four genera have been excluded since Polhill Inverted Repeat Lacking Clade (I RLC)
Hedysareae (page 489)
(1994) and four recently described genera added Cicereae (page 496); Galegeae sens. strict. (Ga/ego)
(Mackinder, pages 69-108, this volume). The genus Trifolieae (page 499)
Umtiza has been moved out of the Detarieae to the Fabeae (page 505)
Umtiza clade of tribe Caesalpinieae (Herendeen et al., FIG 1 Ph
penntngton
· Ylogeny
et al. (of Le~m I~osae corn piled as a supertree, based on analyses by Doyle et al. (2000); Crisp et al. (20oo); Wojciechowski et al. (20oo; 200 )'
2003a & b; Lewis, page 131, this volume).
are In bo(d type Pa20 oi),/<a11ta et al. (2001); Herendeen et al. (2003a); Luckow et al. (2003); Wojciechowski (2003). The 36 tribes dealt with in this volu~;
Pseudosindora is considered to be a synonym of · ge re erences are either to the beginning of the tribe or to the diagram of relationships following the introduction to that tribe
INTRODUCTION 5
 Copaifera, and Thy/acanthus, originally thought to be
a monospecific genus in Brazil, has proved to be based
on African material of julbernardia (Breteler, pers.
comm . in Mackinder, page 105, this volume).
Monopetalanthus has been disbanded (Wieringa, 1999)
with species being redistributed to other genera in the
tribe, including the recently described Bikinia
(Wieringa, 1999). !curia was also described as new by
Wieringa 0999), as was Ecuadendron by Neill 0998).
Micklethwaitia (Lewis & Schrire, 2004) is another
generic addition to the tribe. The genus is based on
Cynometra carualhoi Harms, which was first given
separate generic status as Brenaniodendron by
Leonard 0999a). Brenaniodendron is a later homonym
of Brenandendron in the Asteraceae and thus required
a new name. The correct name for Bathiaea Drake is
Brandzeia Bail!. (Du Puy & Rabevohitra in Du Puy et
al., 2002) . Phyllocarpus has changed name to
Barnebydendron (Kirkbride, 1999b). The Detarieae is
well-supported as a monophyletic group once Umtiza
is removed (Bruneau et al., 2000), but none of the
informal generic groupings of Polhill 0994) are
supported as strictly monophyletic and these are not
retained by Mackinder (pages 69-108, this volume).
In 1994 the Cassieae comprised 20 genera in five
subtribes. The tribe has long been known to contain an
artificial grouping of genera which, pending further
molecular analysis, should be segregated into probably
two to three separate tribes . It is dealt with largely in
its traditional sense by Lewis (pages 111-124, this
volume) , and now comprises 21 genera. The
phylogenetic position of the enigmatic genus
Duparquetia remains equivocal and monogeneric tribal
status would seem appropriate. The Cassieae sens.
strict. now contains only three genera: Cassia , Senna,
and Chamaecrista, and this group is more closely allied
to the Caesalpinieae than to the other elements of
Cassieae sens. lat. The largest group of genera within
the Cassieae (in its broad sense) falls under the
umbrella of subtribe Dialiinae which is expanded to
include subtribe Labicheinae, and now also includes
the genus Poeppigia, transferred from the
· Caesalpinieae. Uittienia, reinstated as a genus separate
from Dialium (Rojo, 1982), is also included in this
generic grouping. Ceratonia is transferred from the
Cassieae to the Umtiza clade of the Caesalpinieae.
Tribe Caesalpinieae has increased from 47 (Polhill,
1994) to 56 genera (Lewis, pages 127-161, this volume).
Poeppigia is now excluded (see above) . Sclerolobium
is considered to be a synonym of Tachigali, although
far from aU the necessary new combinations at species
level have been published in the latter. Cordeauxia is
reinstated as a genus separate from Stuhlmannia , and
Ceratonia and Umtiza are transferred into the tribe
from the Cassieae and Detarieae respectively. A major
change since 1994 is the recognition at generic rank of
eight names resurrected from synonymy under
Caesalpinia, a genus which is now reduced from
approximately 140 to 25 species. The reinstated genera
6 LEGUMESOFTHEWORLD
are: Coulteria, Erythrostemon, Guilandina, Lihidibia,
Mezoneuron , Poincianella, Pomaria, and Tara. The
tribe is currently paraphyletic with respe ct to the
Cassieae sens. strict. and subfamily Mimosoide ae.
Within the tribe only one of the informal groups of
Polhill 0994) remains unchange d with respect to
generic content, the monogeneric Pterogyne group.
Several new groups are referred to following Haston et
al. (2003, submitted) . Of considerable biogeographical
interest is the Umtiza clade (Herendeen et al. , 2003b;
Lewis , page 129; Fig. 23 , this volume) which
amalgamates seven genera previously positioned in
three different tribes .
Tribes in subfamily Mimosoideae
The number of tribes in the Mimosoideae has been
reduced from five (Polhill, 1994) to four, with tribe
Parkieae h ing subsum >d inl the Mimoscac (J.u ckow,
pag s 1 - J83, this o lum . Th n on ogi:neri · trib
Mim ozyga nth e::i Cr o nuna t , p ag ··. IR - 1 'i, thJ.
volume) will a lso be lnduded in th Umosca in th
near future (Luckow, page 163, this volume). Dinizia
and Pentaclethra are among the most basally
br:.111 hing I me nls of th subfa mily and might w ell I e
b •tter pJacecJ in Lh ~ Caesa lpirtio ideae, as first sugg sled
fo r Pentacl>thro I y Be nthRm 1875). Po lhill 199 )
placed 36 genera in the Mimoseae. This has now
increased to 40 due to the addition of the two genera
from the Parkie ae , and two newly described genera:
Kanaloa 0994) and Alantsilodendron 0994). The
twelve informal generic groupings proposed by Lewis
& Elias 0981) , and followed by Polhill 0994), remain
largely unchange d , although the Xylia group is
disbanded and its two genera added to the
Ad nanth ' ra g r 1p. The Pentad · thra , IJ ipta I nhstrum
und yli o llscus groups are ne w a klitio ns sin ·c 1994.
Trih Mlmos ae ls paraphyle ti c ith rl.:!spe l 10 the
Acacieae and Ingeae and will need to undergo supra-
generic restrU<.:turlng in the fULure.
With th g ·nus Fa itlberf;ta havi ng I een tran fe rred
to t.he lnge~l e ( Po lhill, 1 94; Lu ·k w c>/ al., 200. , tribe
·a ieae is c urr ' nil y mo n g Jn •ri ·, alth ug h the
taxonomic status of the single genus Acacia is, as yet,
unresolved. The genu s, in its traditi o na l
certainly not monoph ylcti · and it is l b · ·p ·t I
that at least five gen - ra wiU be resurr 'Clc d o r new I
described from within it. The speciose Acacia
su bg •nus Pb,J llodi11.eae · mstILut s n wc ll s upported
grour nested within tribe fn~t'a >,as discusse I In mor •
de ta il by l.c v is C raw J87, tlii s volum 1 and in rhe
ch·q ler o n biog ·og r::t phy ( · ·hri1·e et. ed ., this vo l um >J:
A ctc.: ia s ubgenus Acacia is mo re I I re bl ti to
tribe Mimoseae .
The Ing a · has in r "asccl in numb ·r of gen ~r..1 fr 111
25 to 36 s in ·e 199 . Thi s brg ly restd ts fro m th
treatme nt of Barn I> ~ rrirn s 19< 1) w hich d scribed
as n ew reinswtecl c r ~ l eva t d l( gen ·ri · staru • a row I
of nine genera: Blanchetiodendron, Ebenopsis,
Frt~ -, 1wri,t, /·tes/ K!rctlbizlct, Hydro b CJPll l e11cocblunm ,
J ii iJ1fc •ria, P ·<?11dosm11em e t, and . 1j>bi11ga. In additio n,
iwo oL11cr ge ne ra have b ·e n tl s Ti! ' d sin ce 1994:
1tfllC'lill L. Rico & M ., o usa in Rk:o et ct!., 1 <t ) a n$1
V(~i tierm 11/:ms Villie rs in I u Pu y el al., 2002) a nd
17; illc:11tc1dupsis has I l.:!n r instat.ed :1s :i dislin ·1 genus
( Lewis ~ Sc:brir _, 2003bl. bofinga, in ·lud "d ill th ~
tribe hy Politi II ( 1( ), is now cons idered a syno nym
f C<!jo/Jo foll wing B;i rncl y & ·rim s ( I9 ;)7). Po UlilJ
19911) rccogni s I no gen ·ri · grou in ' S wilhin the
Ing ·1 and the phylogen >f Ll1 ' IJ'i l e r main poorl y
r .-.olv ·<l d ue to a pau •ity or mo lct:UhLr data. L '\ is &
Hi ·o (p;1ge 19'l, Lliis volume) pla ·em SL o f the gt:nc ra
into si. informal alliances foJlo win,' B;um:I y & rim s
L9lJ6, I' 7) an<l Lu cko el a.I. {2003).
Tribes in subfamily Papilionoideae
In the Papilionoideae numerous changes in generic
gro11p1ngs have t:i1 en place s in · 1994. Wh r
e:illgnm •nts are stro ngl suppone I by molecu b r and
t mhme I datasets aut.ho rs have •ffected these d1~ui ge
in their tribal treatments, and many genera have been
moved from one tribe to another. Within the subfamily,
radical shifts in inter and intra-tribal relationships are
indicated. Nevertheless, the majority of the tribes are still
defined in the more traditional sense of Polhill 0994)
with respect to the genera contained within them,
pending more robustly supported phylogenies. The
most striking example of a tribe which is polyphyletic
in its traditional sense is the Sophoreae (Fig. 1),
elements of which are scattered throughout the
papilionoid phylogeny, from the basally branching
groups which lack the 50 Kb inversion, i.e.: a) Alexa,
Castanospermum, Angylocalyx and Xanthocercis, and
b) Dussia and the group of Myrospermum, Myroxylon
and Myrocarpus associated with the Aldinoid clade of
the Swartzieae; to elements within the 50 Kb inversion
clade, i.e.: c) a basally branching Cladrastis clade, cl) a
Calia-Uribea clade associated with the Lecointeoid clade
of the Swartzieae, e) a Sweetia-Luetzelburgia clade
associated with the Vataireoid clade, f) a basally
branching element within the Genistoid clade, g) a
more derived group within the genistoids comprising
Sophoreae sens. strict., and h) the Baphioid clade, sister
to a remaining clade of predominantly Old World tribes.
The subfamily can be broadly di vid ed into the
following ma111 · groups ( Fig. . 1): the , wa r1 z1eae
. sens.
Slr/ ·~· · :in a$, ortcd group c f I a sa l papiUo n ids, the
enistold l ttd~. th An lira gr >Li p th e /\111 rpbe·1
an . I Lht.: D·1• I l1 ' 1g
. 1. •t
, I Ia d " (.r I1e las
' t two toge t.h••r
r •l ened l<> ,·1·s II 1e D a Ji. Jerg 101d . . se ll . fCJI . ·la d e
f (Jl!owing w · ,· h . was dissuaded from following their conclusions because
1
0 1c: 1 c ows ' ' e l a l. 200 and a n
essenU·1II • •
·I· ' . Y Id Wo rld d adc (i n lu<ling th Bapltioicl
1
1 ;:~e, .Mtrl e li >id sens. lat. lad , Mill tti i I ·en '. la t.
.ind I f loga legi11·1
'I'I 1 • • .
In cn.~' l'i clw~1nzieac,
r 01 as traditionall y ' ir ' lll11 · 'fil ed Ins
• I
'lcldi '. r 15 to 17 g ne ra s in ·e 1994 lu ·· Lo L11 e
• l ton of n I . '
<> ?l-1 1t11111c1 seg regm d fr >m • wa r1ztrr
(Kirkbride & Wiersema, 1997) and Trischidium,
reinstated as a genus distinct from Bocoa (Ireland, this
volume). Molecular data strongly suggest, however,
that Swartzieae sens. strict. should be redefined to
include only the genera Swartzia, Bobgunnia, Bocoa,
Candolleodendron, Trischidium, Cyathostegia, and
Ateleia, with the other ten swartzioid genera requiring
transfer to other tribes (Ireland, pages 215-224, this
volume).
Polhill 0994) included 47 genera (Alexa was allotted
number 3.2.12a) in his synopsis of the Sophoreae.
Pennington et al. (this volume) reduce the number to
45. Etaballia, lnocarpus and Riedeliella are removed
from the tribe and replaced in the Dalbergieae,
following Polhill Cl981d, but not 1994), Lavin et al.
(2001a), and Klitgaard & Lavin (pages 307-335, this
volume). Ammothamnus is reinstated as a genus
distinct from Sophora. Pennington et al. continue to
recognise the genera Bowringia and Baphiastrum as
separate from Leucomphalos, although Breteler 0994b)
considered the three to be congeneric. Further analysis
is needed to resolve this issue . Baphiopsis in the
Swartzieae has also been shown to belong in the
Baphioid clade of Sophoreae sens. lat. (Ireland et al.,
2000; Pennington et al., 2001). Polhill's 0981b, 1994)
six informal generic groups are largely abandoned and
replaced with a series of informal clade names (e.g., the
Baphioid, Genistoid, Vataireoid, Aldinoid and
Swartzieae sens. strict. clades) which, as indicated
earlier, shows the affinity of groups of Sophoreae sens.
lat. genera to several different areas of the papilionoid
phylogeny (Fig. 1; Ireland, page 216, this volume;
Pennington et al., page 228, this volume).
The Dipterygeae (Barham, pages 250-251, this
volume) are unchanged from Polhill 0994), although
Lima (pers. comm.) suggests that the tribe might include
Monopteryx from the Sophoreae.
Genistoid clade
The most basally branching group of the Genistoid
clade (Fig. 1) comprises several genera of the
Sophoreae sens. lat. Sister to this group is a clacle
containing the next six tribes in the Papilionoideae,
together with the Sophoreae sens. strict. (discussed in
its broader sense above). The Brongniartieae has
increased in genera from two to ten since 1994 when
it only housed Brongniartia and Harpalyce. Polhill
0994) alluded to the work of Crisp & Weston 0987)
which proposed moving the Australian Templetonia
group of tribe Bossiaeeae to the Brongniartieae, but he
of the analysis of Chappill 0995). Ross & Crisp (pages
253-258, this volume) include the Templetonia group
in the Brongniartieae, adding two genera recently
segregated from the genus Templetonia: Cristonia and
<
Thinicola. Neotropical Poecilanthe and Cyclolobium
are also transferred into the tribe from the Millettieae.
The monogeneric Euchresteae remains unchanged
INTRODUCTION 7
since 1994. It5 affinity lies with the Sophora group of the
core genistoids, most closely allied to Sophora (Kajita et
al., 2001). Euchresteae will thus have to be included
within a recircumscribed Sophoreae sens. strict., as
suggested by Ohashi (pages 260-261, this volume).
The Thermopsideae, with six genera, also remains
unchanged since 1994, although Wojciechowski et al.
(2004) find 100% bootstrap support for including
Pickeringia within the Cladrastis-Styphnolobium clade.
This transfer, however, has yet to be formally made.
The Podalyrieae (Van Wyk, pages 267-271, this
volume) now includes tribe Liparieae (in large part) in
synonymy and the number or g n ra has grm n from
four to eight since 1994. Priesli ~)ltl Is clisl :ind •cl , Its
species being redistribut ' U b e t w ~ n ll/Jtll'ia a nd
Xiphotheca. Coelidium is placed in synonymy under
Amphithalea. Stirlonanthus (Van Wyk & Schutte, 1995b)
is added as a new member of the tribe. Hypocalyptus
is removed from this assemblage of genera and placed
in its own monogeneric tribe in an, as yet, weakly
supported clade together with tribes Mirbelieae and
Bossiaeeae. The Crotalarieae retains the same eleven
genera as listed for the tribe in 1994, although Van Wyk
et al. (1989), on the basis of biochemical evidence,
suggest that Spartidium may be better placed in the
Genisteae. The Genisteae, sister to the Crotalarieae,
comprises the same 25 genera, but there is better
understanding of inter-generic relationships within the
tribe (Polhill & Van Wyk, pages 283-296, this volume).
The recent exciting rediscovery of the previously only
once-collected genus Sellocharis (Miotto, pers. comm.,
2004), confirms the existence of this morphologically
rather unusual taxon in Brazil. Further study will reveal
whether it is correctly placed in the Genisteae.
Dalber9ioid sens. lat. clade (following Wojciechowski
et al., 2004, Fig. 1)
The weakly suppo1ted lineage sister to the above clade,
which includes tribes Amurpheae and the Dalbergioid
clade (sensu Lavin et al., 2001a), comprises two genera,
. Andira and Hymenolobium. Pending further analysis,
these, together with Vatairea and Vataireopsis, are
dealt with by Klitgaard & Lavin (page 309, this volume)
under a greatly enlarged concept of tribe Dalbergieae
sens. lat., although the authors clearly discount this
group of four genera from their cryptic Dalbergioid
clade diagnosed by the synapomorphy of
aeschynomenoid root nodules. Sister to the Dalbergioid
clacle is tribe Amorpheae which comprises the same
eight genera as in 1994. These are grouped into two
clades, the Amorphoid clade and the Daleoid clade,
following McMahon & Hufford (2004), who also
indicate that Psorodendron should be resurrected as a
ninth genus in the tribe. The Dalbergieae sensu Polhill
(1994) comprised 17 genera divided between two
groups, the Andira and Dalbergia groups. The Andira
group contained the same four genera (discussed
above) now partly referred to the Vataireoid clade. The
8 LEGUMES OF THE WORLD
Dalhergieae (sensu Klitgaard & Lavin, pages 307- 335,
this volume) is greatly increased in size to contain 49
genera, indllding the monogeneric tribe Adesmieae,
tribe Aeschynorn •neae, and subtribe Bryinae of the
Desmodieae in synonymy, as well as the three genera
Riedeliella, Etaballia and Jnocaipus returned to the
tribe from the Myroxylon group of tribe Sophoreae.
The genus Belairia is treated as a synonym of Pictetia
following Beyra Matos & Lavin (1999), and
Arthrocarpum and Pachecoa are subsumed into
Chapmannia in line with the work of Thulin 0999).
Three new genera, Peltiera , Zygocarpum , and
Maraniona, described as new since 1994, are added to
the tribe . In Fig. 40 (page 309) of this volume the
Dalbergioid clade is further divided into three informal
groups: the Adesmia, Pterocarpus and Dalbergia clades.
Old World clade
The remaining 17 papilionoid tribes group together in
a large, predominantly Old World clacle which has the
Baphioid clade (comprising seven genera of tribe
Sophoreae sens. lat.) as its most basally branching
element. The 17 tribes cluster into three groups: the
Mirbelioid sens. lat. clade, the Millettioid sens. lat. clade,
and the Hologalegina alliance. The latter is further
divided into the Robinioid clade (with a large
neotropical element) and the Inverted Repeat Lacking
Clade (IRLC) .
Mirbelioid sens. lat. clade
The Mirbelioid sens. lat. clade comprises the
Australasian Bossiaeeae and Mirbelieae together with
the South African Hypocalypteae which is weakly
supported as a member of this clade in the matK
analysis of Wojciechowski et al. (2004). Hypocalyptus
was misplaced in the genistoicl tribes and has been
removed from the Liparieae (this is now part of an
expanded Podalyrieae in this volume) and placed in its
own tribe Hypocalypteae (Schutte & Van Wyk, 1998b;
Van Wyk, pages 336-337, this volume). Polhill 0994)
recognised 26 genera in the Mirbelieae, now reduced
to 25 (Crisp et al., pages 339-352, this volume). Three
genera, Nemcia, Bracbysema anc\jansonia are placed
in synonymy under Gastrolobium; Stonesiella is new
since 1994 and Otion, dealt with as a member of the
tribe, remains to be validly published. The !30 ·sia eae
is reduced from ten genera to six due to th • Lr~tn · ~ r of
all four genera of the Templetonia group to the
Brongniartieae.
Millettioid sens. lat. clade
Basally branching in the Millettioid sens. lat. clade is
tribe Indigofereae that retains the same seven genera
as in 1994, although current evidence indicates
Vaughania must be subsumed within Indigo/era
(Schrire et al., 2003). The concept of what comprises
Millettieae sens. strict. is changing rapidly based on
•vid , 11 ·~ from mol c:ular J hylogenies. A traditional
t·lr ·umscr lplion o f th Millettieae is thus followed
hrire, p:1ge:; j67- 387, tMs volume) with 45 genera
in cluJed in ch • Lrihe Can increase of two on the
Lre:1 101e nt:o of G >csi nk 11 :>84) and Polhill [1994]) . A
sJgnifi ·ant advan · l. th:1t rhe genera of the Millettieae
:ire 110 kmg<::r nrr:rng ·ti alphabetically as they were in
ihe sy n )psis of Po lhJll t1994), but are placed in a
number ol' infonn·tl supr~1generic groupings. The
gener<t G)1 ·/n/obiulll and Poecilanthe are transferred
from the Mill ·tti ae to tribe Brongniartieae. Six genera
fr rn Q 1/! 'l~)ltl l( \ isteria w ill have to be transferred to
::i ha ·:tl l I ran -bing r> >silion in the IRLC clade (Schrire,
ra " h
%9 • thi. olurn ·: Mi llettieae). Neodunnia and
•
p 011 gomia ·ire 110\· c nsidered as synonyms of
Mllle1tic1, and \ i!/ardia is placed as a synonym of
Lonchocarpus. The genera Paraderris, Paratephrosia,
Philenoptera, Ptycholobium and Requienia are
reinstated. Pyranthus and Sylvichadsia, two genera
endemic to Madagascar, have been described as new
since 1994.
The monogeneric tribe Abreae is retained
unchanged since 1994. Its phylogenetic position lies
with the Millettieae sens. strict., together with two
subtribes of Phaseoleae now excluded from tribe
Phaseoleae sens. lat.
As for the Millettieae, the Phaseoleae (Schrire, pages
393-430, this volume) is also treated largely according
to the traditionally held view of the tribe, as
recircumscribing Phaseoleae sens. strict. is currently
very problematic. The Phaseoleae sens. lat. clade (Fig.
47) includes two traditionally independent tribes, the
Desmodieae and Psoraleeae (both dealt with separately
in this volume). Polhill (1994) recognised 83 genera in
eight suhtribes in the Phaseoleae. The number of
genera is increased to 89 here. Ramirezella is reinstated
as distinct from Vigna, and Barbieria as distinct from
Clitoria. Baukea is now treated as a synonym of
Rhynchosia. Based on morphological, palynological
and molecular evidence, Phylacium and Neocolletia are
transferred from subtribe Lespedezinae of trib e
Desmodieae to subtribe Glycininae of the Phaseoleae .
Three genera, Lackeya, Mysanthus and Oryxis have
been described as n ew since 1994. Schrire (this
volume, Table 9, Fig. 47) places the genera of the
Pl:aseoleae into two groups: Phaseoleae subtribes
Diocleinae and Ophrestiinae allied to Millettieae sens.
strict., and the Phaseoleae sens. lat. clade, comprising
subtribe Clitoriinae, tribes Desmodieae and Psoraleeae,
ancl the core-Phaseoleae.
The Desmodieae now excludes subtribe Bryinae,
~~nsfrrred to the Dalbergieae sens. lat., and
ylacium and Neocolletia moved to the Phaseoleae.
Nevertheless, the tribe has increased in size since
l 9~q · fro m 26 to 30 g n r~1. Th· gen ra Apby ll > 1111111
WILh 0 icerma
· ·
111 synun my) , D >·111 0,liastr11m ,
Hun~t I((
· , negnera,
u
Mo11arthmcc11p11s ~ind Oug , ini&I
are reinstated. Akschindlium Hylodesmum and
Ohwia a1e
. new genera, added since
' 1994. The tribe
Desmodieae is most likely sister to Mucuna within the
Phaseoleae sens. lat. (Fig. 47 of tribe Phaseoleae).
Ohashi (pages 433- 445, this volume) recognises
three groups of genera within the tribe: the
Desmodium and Phyllodium groups (corresponding
to subtribe Desmodiinae in the synopsis of Polhill,
1994) and the Lespedeza group (subtribe
Lespedezinae, excluding Phylacium and Neocolletia).
The Psoraleeae retain the same nine genera as in
1994. The tribe is nested within the Phaseoleae sens.
lat., sister to Phaseoleae subtribe Glycininae.
Holo9ale9ina
This alliance of mainly Old World tribes contains two
main clades: the Robinioid clade sister to the Inverted
Repeat Lacking Clade.
Robinioid clade
The Sesbanieae (Lavin & Schrire, pages 452- 453, this
volume) is resurrected as a separate monogeneric tribe,
distinct from the Robinieae. Sister to the Sesbanieae is
tribe Loteae that has increased in number of genera
from 17 to 22 since 1994. Acmispon, Hosackia,
Syrmatium and Tetragonolobus have all been
reinstated as segregate genera, although indications
are that Tetragonolobus should be subsumed, once
again, into Lotus. Kebirita and Ottleya are new since
1994. Verm~(rux is now treated as a synonym of
Dorycnopsis. Sesbanieae and Loteae are together sister
to the Robinieae. With Sesbania transferred to its own
tribe, Poissonia reinstated as a genus distinct from
Coursetia, and Hybosema placed as a synonym of
Gliricidia, the Robinieae now comprises 11 genera, a
decrease of one since 1994.
Inverted Repeat Lacking Clade (IRLC)
The basally branching lineage of the IRLC (Fig. 1)
comprises a group of millettioid genera (including
Callerya and Wisteria) sister to Glycyrrhiza of the
Galegeae (as traditionally circumscribed). Polhill 0994)
distinguished five subtribes in Galegeae (the Alhagiinae
DC. being added since his earlier classification [Polhill,
1981h]) and recognised Galegeae and Carmichaelieae
as distinct tribes. More recent studies suggest that
Carmichaelieae should be treated as an additional
subtribe of Galegeae sens. lat., and this is followed
here. In 1994 Galegeae and Carmichaelieae together
comprised 27 genera. In their account of Galegeae,
including Carmichaelieae (Lock & Schrire, pages
475-487, this volume), the tribe contains 24 genera.
Alhagi, Caragana, Calophaca, and Halimodendron
are transferred to tribe Hedysareae. Astracantha and
Neodielsia are returned to Astragalus, although,
conversely, Biserrula is reinstated as a distinct genus.
Chordospartium, Corallospartium, and Notospartium,
considered distinct genera in 1994, are all placed as
synonyms of an expanded Carmichaelia. Erophaca
INTRODUCTION 9
is reinstated as a separate genus, and Tihetia is
accepted as a distinct genus following Tsui Cl979l.
Ophiocarpus 0977) and Barnehyella 0 994) are
tentatively re co gnised, although both might be
returned to Astragalus in the future. Montigena
(1998) is also recognised, although indications are
that it could be returnee\ to Swainsona. Recent
molecular studies point strongly to the polyphyly of
the Galegeae and we can expect disintegration into
a number of smaller tribes in the future. In addition
to the isolated position of G~YCJirrhiza mentioned
above, the genus Galega is also isolated from the rest
of the 'Galegeae', and is possibly more closely related
to the Cicereae. The remaining bulk of the genera in
Galegeae sens. lat. are thus likely to become more
narrowly defined as tribe Astragaleae.
Tribe Hedysareae (Lock, pages 489-495, this volume)
comprises 12 genera, five more than in 1994. Four genera
are transferred from tribe Galegeae, as discussed above.
In addition, Corethrodendron and Sulla are reinstated,
while Stracheya is placed under Hedysarum.
Tribe Cicereae is monogeneric, and unchanged
since 1994. The Trifolieae is retained in its traditional
sense (Lock, pages 499-503, this volume) and
comprises the same six genera as in 1994. The tribe is
not monophyletic and some restructuring seems
inevitable, pending further sampling and analysis. The
Fabeae (Lock & Maxted, pages 505-508, this volume)
is the correct name for tribe Vicieae. The tribe houses
the same five genera as it did (as tribe Vicieae) in the
treatments of Kupicha Cl981a) and Polhill 0994).
Economic importance of the
legume family
Legumes have been gathered, cultivated, e::iten and
used in a multitude of other ways by humans for
millennia and are arguably as important as grasses in
global terms. Certainly the range of uses of legumes is
broader than that of the grass family (Doyle & Luckow,
2003). Legume products contribute enormously to the
world's economy through food (for animals and
humans) and drink, pharmaceuticals and medicine,
biodiesel fuel, biotechnology (as industrial enzymes),
building and construction, textiles, furniture and crafts,
paper and pulp, mining, manufacturing processes,
chemicals and fertilisers, waste recycling, horticulture,
pest control, and ecotourism. Ceratonia siliqua L. (the
carob tree, locust bean or St. John 's bread) is
remarkable for its wide range of uses over the past two
thousand years. Carob seeds, said to be the original
carat used as a standard weight by jewellers, are used
to make a chocolate substitute, and, blended with
chicory, a coffee substitute. Carob seed gum is used in
foods and cosmetics, adhesives, inks, paints, polishes
and as a photographic film emulsion. Across the globe
legumes have been used as substitutes for coffee, tea,
10 LEGUMES OF THE WORLD
tobacco , betel-nut, hops, garlic, ebony woocl, cocaine,
soap and liquorice (true liquorice, Gf:ycyrrbiza glahra
L. is also a legume).
Ancient cultures were aware of the ability of many
legumes to improve the soil, even if they did not then
appreciate that this results from symbiotic nitrogen
fixation (Van den Bosch & Stacey, 2003). Some 40 to
60 million metric tons of nitrogen are fixed annually by
agriculturally important legumes and a further 3 to 5
million metric tons by legumes in natural ecosystems
(Graham & Vance, 2003). Many species are used as soil
improvers, green manures and stabilise rs and in
reforestation programmes. Natural accumulation of
nitrogen has also resulted in predation of legumes by
a wide range of animals and insects. To combat this,
the family has evolved a wide repertoire of chemical
defences based on secondaiy compounds, especially
alkaloids. Humans have exploited the chemistry of
legumes by utilising many species as medicines,
insecticides, molluscicides, abortifacients, purgatives,
fish, arrow and ordeal poisons, anti-fungal agents,
aphrodisiacs and hallucinogens. Some legumes have
been used as antidotes to poisons, as anti-
inflammatories and antiseptics . The senna pod is a
well known laxative. Several legumes are rich in gums
used as glues and food thickeners (e.g., Acacia,
Astragalus), resins used in paints, polishes and
varnishes (e.g., Hymenaea , Copa(fera, Prioria) and
oils used in lubricants and cosmetics. Important dyes,
such as brasil, indigo and dyer's greenweed all come
from legumes, and several species are used as inks, and
for tanning leather.
Grain and forage legumes are grown on
approximately 180 million hectares (12 to 15%) of the
Earth's arable surface and account for 27% of the world's
primary crop production with grain legumes alone
contributing 33% of the dieta1y protein nitrogen needs
of humans (Graham & Vance, 2003). The main cUeta1y
legumes (the pulses) include several species of bean
(Pbaseolus species), the pea (Pisum sativu.m L.) ,
chickpea (Cicer arietinum L.), broad bean (Viciafaha
L.) , pigeon pea (Cajanus cajan (L.) Huth) , cowpea
(Vigna unguiculata (L.) Walp.) and lentil (Lens culinaris
Medik.). Legumes (mainly soybean, Gf:ycine max (L.)
Merr., and peanut, Amchis bypogaea L.) also contribute
more than 35% of the world's processed vegetable oil
(Graham & Vance, 2003). Forage legumes provide the
protein, fibre and energy that have underpinned daily
and meat production for centuries. In temperate regions,
alfalfa (Medicago sativa L.) is an important forage crop
for cattle. Other important temperate pastures used for
forage, hay, silage and green manure, include clovers
(Trifolium species), trefoil (Lotus corniculatus L.)
sweetclovers (Melilotus species) and vetches ( Vicia
species). In the tropics, species of Aeschynom.ene,
Arachis, Centrosema, Desmodium, Macroptilium, and
Stylosantbes are all being used to improve tropical
pasture systems, with Stylosanthes the most
geographically widespread (Graham & Vance, 2003).
--
FIG. 2 A sm 0 II .
selection of useful products derived from legumes Photograph by B. o. Schrire
INTRODUCTION 11
 In northern Europe species of G'leditsict, Lahurnu.m I er · nt;1gl' )r e ·situ. onserv::1 ti on prngr:1mme:;. complete synopsis of legume genera
and Robinia have been widely grown as street, p;.i rk Spe · ie:; of crtcitt, Ainule1zantbera , Da/IP rg la ,
J:,'1yt bri11 1, Prc1:ofJIS an I l'l ero e1rp11s arc al l importan t Legume genera, with page reference, arra nged systematica lly within tribes and subfamilies
and garden trees for centuries, and exotics, such as the
wi nter-flowering Australian acacias, are becoming very woody tr' , I g umi;:s i.n foreslly . L~gurn ' timber and
popular. A handful of species are sole.I as cut llo rs wood from many species has long been put to a
CAESALPINIOIDEAE 1.2.38 Maniltoa .......... ..... . ... 88
('florist's mimosa ' being, confusingly, the c )mmnn multitude of uses, ranging from he avy construction
1.2.39 Cynometra .. ...... .. ........ 89
name of a small number of Australian Acacia species). (house and boat building, railway sleepers and can CERCIDEAE
1.1 1.2.40 Tamarindus . ....... . .... , ... 90
Sw ' t p eas Latby ru.s s pec ies), lupins (l 11j i111ts wh eels), to paper and plywood m:mufacture, and fine
1.2.41 Intsia ....... . ..... ......... 90
sp · ' i s) :m I broo ms G°(mistct and y lisus species furnitu re production, ca rpentry, marquetry and veneer 1.1.1 Cercis .... ................. 59
u.2 Adenolobus .......... . ...... 59 1.2.42 Afzelia ........ , ... ...... . .. 91
have a ll h;td p ak:o; or rorn.tl ~1rity large ly based 011 work. Wooc.ls high in silica (e.g. , Dicorynia guianensis
Brodriguesia .. . . .. . . . ....... 92
rr
mu ltiple new ·ol Lil' fonns I' ·suiting )J1'l intcn iv Amshoff) are of particubr val u e in ma rine 1.1.3 Griffonia .. ..... ....... ..... 60 1.2.43
i.H !3renierea . . . . . . . . . .. .. ..... 60 1.2.44 Loesenera .......... ........ 92
hybridi z::it ion . All · >nlim1 Lo he~ p< pul ar gard n construction. Some species (e .g ., Kalappia celehica
u ."i Bauhinia . , ................. 61 1.2.45 Neocbevalierodendron ... .. .... 93
plnnts. In the tr pies, avenu ~sa nd pa 1·ks ~II"' In aria I ly Kosterm ., Dalhergia nigm (Veil.) Allem:lo ex Benth.)
lJ/1 Gigasipbon . ...... . ......... 62 1.2.46 Normandioc.lenclron .. ........ _ 93
adorn -· I wi th a represe ntation of /fJi.zia , as fa , are now considered rare and endangered because of
u .7 Tylosema ... ....... , ........ 63 1. 2.47 Zenkerella . . ...... . ....... _ . 93
Delonix, Senna, Castan o,\permum. and Tipu.ana. over-exp! italion due to their commercially valuable
1.1.8 Barklya ... ................. 64 1.2.48 Humboldtia . .. ..... ......... 94
Spectacular, showy-flowered species of Caesalpinia, timbers. Caesetlpinia echinata Lam. (brasil wooc.I or
1.1.9 Lysiphyllum ... . .......... . .. 64 1.2.49 Hymenostegia . ..... ...... _ .. 94
Calliandra, Mucu.na and Strongylodon have also pau brasil , the tree l'rom which the country Brazil took
l.},JI) Phanera ... .... . ........ . ... 65 1.2.50 Leonardoxa . . . . . . . . . . . . . . ... 95
becom ~ pc pu lar in tropi. ·al gard ~ ns. In th sul trop ics its name) , once a sour· of an important red dye and
u. n Ltsiobema . . . . . . . . . ......... 66 1.2.51 Amherstia . . . . . . . . . . . . . . .... 95
an l!ven grl!a L r vari ' l y f g n ra and sp i s hav • still the preferred wood for violin bows, has been
1.1.12 Piliostigma . ....... .......... 66 1.2.52 Ecuadendron ... . .. ...... .... 95
been introduc d into horli ulLur . ."orne f Lhes ',S U ·h reduced , by major habitat destruction, to a few
1. 2.53 Paloue . . . . . . . . . . . . . . . . . . . . . 96
as Acacia, Dichrostacbys, Leucaena , Mimosa an d p pularlorn; along the Atlanti · ·cast of Rt. zil .
1.2 DETARIEAE 1.2.54 Paloveopsis . . .. . ... ..... . , .. 96
Sesbania, have become weedy and invasive, having Th p •a, M n lei 's study organism b ·:im th· Cir:-;t
Neoapaloxylon ... ......... .. 72 1.2.55 Brachycylix . ....... . _ .... . .. 97
escaped th ·ir na tural pew; and diseases. lullipurpo. e m< d I s st ·m for quantirativ -• ge ne ti · and 1 gunie:-; 1.2. l
1.2.2 Schotia ......... .. , ........ 73 1.2.56 I-Ieterostemon . . . . . . . _ . . . . . . . 97
trees and slirul s have long b en selected and r"fined ha ve I · -n 111 del organisms for many biol gi al studi •s
1. 2.3 Barnebyclendron ... _ ...... , .. 73 1.2.57 Elizabetba ........ ......... . 97
by loca l communlli s for s hade, ornament, l'orngc, throughout the 20th century (Van den Bosch & Stacey,
1.2.4 Goniorrhachis .... ......... . . 73 1.2.58 13rownea .. .. .... .......... . 98
fodder, fuelw<. o I, i)ee for:ig f r honey produ tiun , 2003) . Recently, new genetic and genomic tools have
Branclzeia ... .... ........... 74 1.2.59 13rowneopsis . . . . . . . . . . . . . . . . 98
and soi l en ri ·hme nL. Reg i nal favourites includ e permitted an a ·celerated adva nc -' in our und •rst~nc.li ng 1.2."i
1.2.6 Oxystigma . . ..... . ........ .. 74 1.2.60 Macrolobium ..... ........... 99
Butea, Dalhergia ancl Millettia in India; Callicmdra, of legume biolof.'Y· Major \ o rk on positional doning
1.2.7 Kingioclendron . . . , .......... 75 1.2.61 Paramacrolobium . .. ......... 100
Gliricidia, Inga and Lettc&Wl'/Ct in Central America of I gu mc genes is being und rtaken >n Metlicago
1.2.8 Gossweileroclenclron ..... _ ... , 75 1.2.62 Cryptosepalum ..... ......... 100
(Polhill, 1997) and Faidher/Jia and Acacia in Africa. tn111cau t!o. <1e1t11. , Lo/us jc1pon.ic11s (Reg I) l .Lars n
1.2.9 Prioria .......... ... __ ...... 75 1.2.63 Dicymbe ..... ... . ..... . _ .. 101
Some are grown as impenetrable spiny hedges, living and Glycine max.
1.2.10 Colophospermurn .......... .. 76 1.2.64 Polystemonanthus .. . , .... . _ . 101
fence- lines and windbreaks. In the 1980s and 1990s a Further details of current and potential uses of
1.2.1 l Hardwickia ...... . ....... ... 76 1.2.65 Pseudomacrolobium . ......... 102
number o f seed banks ta rgeted multipurpose I ~gum ~s can be found in the economic notes which
1.2.12 Daniellia ... ... .. .. . . . ...... 77 1.2.66 Englerodendron . , .. .. .. .. ... 102
leguminous trees fo r introduction as trial species in for m part of the textural block of each genus in this
u.n Eurypetalum .... . ... . ....... 77 1.2.67 Anthonotha ....... ......... 102
agrofo rest ry syste ms and legumes form a large volume.
1.2.14 Eperua ... . .. ... .. . ........ 78 1.2.68 13erlinia . . ........ ......... 103
1.2.15 Augouarc.lia ...... ..•........ 79 1. 2.69 Librevillea . ... .. .. .. .... .. . 103
1.2. 16 Stemonocole us .. . .. ... ... . .. 79 1.2.70 Didelotia ......... .. _ . ... .. 103
1.2.17 Peltogyne ....... ........... 79 1.2.71 Pellegriniodenclron , . . . ....•.. 104
1.2.18 Hymenaea ....... ........... 80 1.2.72 Gilbertiodendron ... ......... 104
1.2.19 Guibourtia . ...... ..... , . , ... 81 1.2.73 Isoberlinia ... ... .. .. . ...... 104
1.2.20 Hylodendron . . .. . ........... 82 1.2.74 Ockloniodenclron .. . . .. . , ... . 105
1.2.2 l Gilletiodenclron .... ...... .... 82 1. 2.75 Microherlinia . ..... ......... 105
1.2.22 Baikiaea . ....... ... ........ 82 1. 2.76 Julbernarclia .... . . . .. .. ..... 105
l.2.2j Tessma nnia .............. . , . 83 1.2.77 Brachystegia , . . ... ......... 106
1.2.2 1! Sindora ....... ...... _ ...... 83 1.2.78 Tetraherlinia ... . .... , ...... 106
1.2.25 Sincloropsis .. , . .. ..... , , , . .. 83 1.2.79 Bikinia ... .. .. ..... ....... _ 107
1.2.26 Copaifera . . . , . . . . .......... 84 1.2.80 Icuria ..... . ...... ......... 107
1.2.27 Detarium ...... . . ........... 84 1.2.81 Aphanocalyx ...... ......... 108
1.2.28 Enclertia . . . . . . . . ........... 85 1.2.82 Michelsonia .... .. .. ..... 108
1.2.29 Lysidice ... . .. ... ........... 85
1.2.30 Saraca .. ..... . .. ........... 85 1. 3 CASSIEAE
1.2.31 Leucostegane .. ... ...... . ... . 86 1.3. l Duparquetia ... ............. 11 3
l.2.j2 Talbotiella . ...... . .... _ ... _ . 86 1.3.2 Poeppigia .... ..... . ....... 11 3
1.2,53 Scoroclophloeus . . . . . . . . . . . . .. 86 1.3.3 Baudouin0 ... ..... _ .. . .. , . 114
1.2,34 Cruclia .. .. ...... ........... 87 1.3.4 Eligmocarpus . .. _ ... ........ 11 4
1.2,35 Lebrunioclendron .. ... ........ 87 1.3.5 Mendoravia . . ... ... . . ... _ .. 115
1.2.36 Plagiosiphon ..... . . .. ....... 88 1. 3.6 Distemonanthus ............. 115
1.2.37 Micklethwaitia . ... .. . _ . .... . . 88 1.3.7 Apuleia .. . .. .............. 115

INTRODUCTION 13
12 LEGUMESOFTHEWORLD
 In northern Europe species of Gleclitsia, Letlntrnwn p erce ntage of ex situ conservation programmes. complete synopsis of legume genera
ancl Robinia have been widely grown as street, park Species of Acacia, Anade1zanthera , Dalbergia,
ancl garden trees for centuries, ancl exotics, such as the Eiythrina, Prosopis and Pterocmpus are all important Legume genera, with page refe rence, arranged systematically within tribes ancl subfamilies
winter-flowering Australian acacias, are becoming very woody tree legumes in forestry. Legume timber ancl
porular. A handful of species are sold as cut flowers wood from many species bas long been put to a
multitude of uses , ranging from heavy construction CAESALPINIOIDEAE 1.2.38 Maniltoa ......... ... . ...... 88
('florist's mimosa' being, confusingly, the common 1.
(ho use and boat building, railway sleepers and earl 1.2.39 Cynometra ..... . .. .......... 89
name of a small number of Australian Acacia species) .
1.1 CERCIDEAE 1.2.40 Tamarindus . . . . . . . . . . . . . . . . . O
Sweet peas (Latbyrus species), lupins (Lupinus wheels), to paper and plywoocl manufacture , and fine
furniture production, carpentry, marquet1y and veneer 1.1.l Cercis ... ......... ... . ..... 59 1.2.41 Intsia .......... . . .......... t O
species) and brooms (Genista and Cytis11s species)
work. Woods high in silica (e .g., Dicorynia guianensis 1.1.2 Adenolohus .. . . . ... ......... 59 1.2.42 Afzelia . . . . . . . . . . . . . . . . . . . . J
ha ve all had peaks of popularity largely basecl on
Amshoff) are of particular value in marine 1.lJ Griffonia . . .... .... ......... 60 1.2.43 Broclriguesia . . . . . . . . . . . .... 9
multiple new colour forms resulting from intensive
construction. Some species (e.g., Kalappia celehiw 1.1.4 Brenierea ...... . .. ..... . . .. 60 1.2.44 Loesenera . . . . . . . . . ..•..... 92
hybridization. All continue to be popular garden
Kosterm., Dalhergia nigra (Veil.) Allemao ex Benth.l 1.1.'i Bauhinia .......... ......... 61 1.2.45 Neochevalierodenclron ..... ... . 9.
plants. In the tropics, avenues and parks are invariably
are now considered rare and endangered because of 1.1.6 Gigasiphon ........ . . ... .. . . 62 1.2.46 Normandiodenc.lron .. .. . •.. . .. 93
adorned with a representation of Alhizia, Cassia,
over-exploitation clue to their commercially valuable 1.1.7 Tylosema .... .. .... . . .•..... 63 1.2.47 Zenkerella ......... .. . ...... 93
Delonix, Senna, Castanospermum and 1'ipuana.
timbers . Caesa!pinia ecbinata Lam. (brasil wood or 1.1.8 Barklya ........... ... .. .. . . 64 1.2.48 Humboldtia ...... . . .....• . .. 9
Srectacular, showy-flowered species of Caesalpinia,
Calhandra, Muczma and Strongylodon have also pau brasil, the tree from which the count1y Brazil took 1.1.9 Lysiphyllum .... . . . . ...... ... 64 1.2.49 Hymenostegia .... . . .....•... 94
its name), once a source of an important red dye ancl 1.1.10 Phanera ........... ......... 65 1. 2.50 Leonardoxa . .... . .. .... .. ... 95
become popular in tropical gardens. In the subtropics
still the preferred wood for violin bows , has been 1.1.1 1 Lasiobema .. ... .. .. ......... 66 1.2.51 Amherstia ..... . . .. ......... 9
an even greater variety of genera and species have
redu ced, by major habit::it d est ru ct ion, to a few 1.1.12 Piliostigma .... . . .. . ......... 66 1.2.52 Ecuaclendron ....... . ..... ... 95
been introduced into horticulture . Some of these, such
populations along the Atlantic coast of Brazil. 1.2.53 Paloue ............ .... . •. . . 96
as Acacia, Dic/Jrostacbys, Leucaena , Mimosa and
The pea, Mendel's study organism, became the first 1.2 DETARIEAE 1.2.54 Paloveopsis ........ . . .. . . . .. 96
Sesbania, have become weedy and invasive, having
model system for quantitative genetics and legumes 1.2. l Neoapaloxylon ... ...... . . . .. 72 1.2.55 Brachycylix ...... . . ..... . ... 97
escaped their natural pests and diseases. Multipurpose
have been model organisms for many biological studies 1.2.2 Schotia ......... ........ . .. 73 1.2.56 Heterostemon ...... . . ....... 97
trees and shrubs have long been selected and refined
throughout the 20th century (Van den Bosch & Stacey, 1.2J Barnebydendron .. . . ....... . . 73 1.2.57 Elizabetha ....... . . .... .. ... 97
by local communities for shade, ornament, forage,
2003). Recently, new genetic and genomic tools have 1.2.lt Goniorrhachis .... ........... 73 1.2.58 Brownea .......... ....... . . 9
fodder, fuelwood, bee forage for honey production,
permitted an accelerated advance in our understanding 1.2.'i Brandzeia ..... ............. 74 1.2.59 Browneopsis ....... ...... . .. ;>8
and soil enrichment. Regional favourites include
of legume biology. Major work on positional cloning 1.2.6 Oxystigma ...... ............ 74 1.2.60 Macrolobium .... . . . ........ . 99
Butea, Dalbergia and Millettia in India ; Calliandra,
of legume genes is being und e rtaken on Medicago 1.2.7 Kingiodendron ... .... . ...... 75 1.2.61 Paramacrolobium . . . ..... . . . 10
Gliricidia, Inga and Leucaena in Centra l America
tntncatula Gaertn., Lotus japonicus (Regel) K.Larsen 1.2.8 Gossweileroclenc.lron .......... 75 1.2.62 C1yptosepalu111 ..... ....... . . 10
(Polhill, 1997) and Faidberhia ancl Acacia in Africa.
and Glycine max. 1.2.9 Prioria ....... . .. ........... 75 1.2.63 Dicymbe ......... .. . .... . . IOI
Some are grown as impenetrabl e spiny heclges, living
Further details of current and potential uses of 1.2.11) Colophospennum . . . .. . .. ... . 76 1.2.64 Polystemonanthus . . ......... 10 1
fence-lines ancl windbreaks. In the 1980s ancl 1990s a
legumes can be found in the economic notes which J .2.1 L Hardwickia ...... ........... 76 1.2.65 Pseudomacrolobium . ....... . . 1 2
numb e r of seed banks targeted multipurpose
form part of the textural block of each genus in this 1. 2.12 Daniellia . . . . . . . . . . . ..... . .. 77 1.2.66 Englerodenclron .... .. . ... . . . 102
leguminous trees for introduction as trial species in
volume. 1.2.13 Eu1ypetalum .... ............ 77 1.2.67 Anthonotha ........ .. . ..... 102
agroforestry systems and legum es form a large
1.2.1 4 Eperua ......... ...... . .... 78 1.2.68 Berlinia .... . .... . ... . ..... I 03
1.2 .1 'i Augouardia ...... ........... 79 1.2.69 Librevillea ........ ......... l03
1.2. H> Stemonocoleus ... ........... 79 1. 2.70 Diclelotia ......... ......... lO
1.2.17 Peltogyne . . . . . . . . .......... 79 1.2.71 Pellegriniodenclron . . ......... 10'1
1.2. 18 Hymenaea ....... ........... 80 1.2.72 Gilbertioc.lenc.lron . . . ..... . .. . JO
1.2. l 'J Guibourtia ....... ........... 81 1.2.73 Isoberlinia ........ ... . ..... 10
1.2.20 Hylodenclron . . . ........ . .. 82 1.2.74 Oddonioclendron . . . .... . . . . 105
1.2.2 L Gilletiodendron ... ... . .... . . . 82 1.2.75 Microberlinia ...... .. , ... ... 105
1.2.22 Baikiaea . . . . . . . . . . . . . . . . . .. 82 1.2.76 Julbernardia ....... . .. .. . . .. I O"i
1.2.23 Tessmannia ..... . ... .. . . .. .. 83 1.2.77 Brachystegia . . ... ..... .. .. l.06
1.2.24 Sinclora ......... ........... 83 1.2.78 Tetraberlinia .. .. . . . . .. .. .. . 106
l.2.2'i Sincloropsis . . . . . . .... . . . ... . 83 1.2.79 Bikinia .. .. .. ... .. ......... 107
1.2.26 Copaifera ........ .......... 84 1.2.80 lcuria ........ . .. .... . .. , .. LC 7
1.2.27 Detarium . . . . . . . . . .... . ..... 84 1.2.81 Aphanocalyx .. .. .. .. , . , .... 108
1.2.28 Endertia . ... ... . ........... 85 1.2.82 Michelsonia . ...... ..... . ... 108
1.2.2') Lysiclice .. . .. . . . . ........... 85
1.2.3() Saraca .......... ........... 8'i 1. 3 CASSIEAE
1.2.3 L Leucostegane . . . . . . . . . . . . .... 86 1.3.1 Duparquetia .. .............. 113
1.2.32 Talbotiella . . . . . . . . , . . . ...... 86 1. 3.2 Poeppigia ... .............. 113
1.2,33 Scorodophloeus ... ........... 86 1.3.3 Baudouinia .. ......... , . ... 114
1.2.:'>4 Cruclia .......... ........... 87 1.3.4 Eligmocarpus . ........... . .. 114
1.2.3c; Lebruniodendron . . ......... . . 87 1. 3. 5 Mencloravia .. .............. 11 'i
1.2.36 Plagiosiphon . . . ....... . ... 88 1. 3.6 Distemonanthus ......... . ... 11 'i
1.2.37 Micklethwaitia . . . . . .. . ....... 88 1.3.7 Apuleia ..... .............. 115

INTRODUCTION 13
12 LEGUMESOFTHEWORLD
 1.3.8 Storckiella . . . ....... . ...... 116 1.4.44 Lemuropisum ... ...... . ..... 156 MIMOZYGANTHEAE 3.1.11 Aldina ..... .. ..... . ....... 221
Labichea .... ....... . ... ... 117 Pachyelasma . .. ... ......... 156 2.2.
1.3.9 1.4.45 Mimozyganthus .... ......... 185 3.1.12 Zollernia .. .. . ..... . . . ..... 222
1.3.10 Petalostylis . .. . . . . ....... ... 117 1.4.46 Erythrophleum . ....... . ... .. 157 2.2.1
3.1.13 Holocalyx .... .. . .... , .. ... 222
1.3.11 Koompassia . .. .......... ... 117 1. 4.47 Dimorphandra . .. .. . . . ...... 157 3.1.14 Lecointea .. . ... ............ 223
2.3 ACACIEAE
1.3.12 Martiodendron ...... .. ..... . 118 1.4.48 Mora . .. . .. .. .. ........... 158 3.1.15 Harleyodenclron .. .. ......... 223
1.3.13 Androcalymma ....... . .... . . 118 1.4.49 Burkea ... . . .. ............ 158 2.3.l Acacia .. .... .. .. . . . . . .... . 188
3.1.16 Exostyles ... . . ..... ..... . .. 224
1.3.14 Kalappia . . . . .......... . . .. 119 1.4.50 Stachyothyrsus .. ....... . . ... 159 3.1.17 Baphiopsis .. .. .. .... , ..... _ 224
1.3.15 Zenia .... . .. ............ . . 119 1.4.51 Sympetalandra . .. .. ... ..... . 159 2.4 INGEAE
1.3.16 Uittienia . . . . . ........... .. . 119 1.4.52 Campsiandra .. .... . ... . .... 159 2.4.l Faidherbia . . . . . ... ......... 196 3.2 SOPHOREAE
1.3.17 Dialium .. ... ...... ... . .. . . 120 1.4.53 Chidlowia . ... .. . .......... 160 2.4.2 Zapoteca ... . . ... .......... 196
Guinetia .. .. . . . .......... . 197 3.2.1 Alexa . . ..... . . . . .. ........ 229
1.3.18 Dicorynia ... . ....... . .... . . 121 1.4.54 Diptychandra .. .. .. .... .. ... 160 2.4.3
Calliandra . .. . . . . ... .. . .... 197 3.2.2 Castanospermum .. . ..... . ... 229
1.3. 19 Chamaecrista . . . . . . ..... . ... 122 1.4.55 Orphanodendron .......... .. 161 2.4.4
Viguieranthus . . . . . ...... . ... 199 3.2.3 Angylocalyx . . . . . . . . . . . . . . . . 230
1.3.20 Senna . ... . . ... ...... . .. . . 123 1.4.56 Vouacapoua . .. .... .. ....... 161 2.4.5
Macrosamanea . .. .... . ...... 199 3.2.4 Xanthocercis . . . . . . ... . .... . 230
1.3.21 Cassia ..... ........ . ... . .. 124 2.4.6
Cojoba ... . ... ... ... . .. .... 199 3.2.5 Dussia . . . . . . . . . . . . . . . . . . . . 231
2.4.7
Inga .. . ... . . . . .... . ...... 200 3.2.6 Myrocarpus ... . . .. .. ..... .. 231
1.4 CAESALPINIEAE 2. MIMOSOIDEAE 2.4.8
Cedrelinga . . .... ......... . . 201 3.2.7 Myroxylon . . . . . . . . . .. ...... 232
1.4.1 Gymnocladus . . . .......... . . 130 2.4.9
2.1 MIMOSEAE Zygia .. . . .... .. ........... 201 3.2.8 Myrospermum . .......... . .. 232
1.4.2 Gleditsia . . . .. . .... . ..... . . 130 2.4.10
Marmaroxylon .. .. .......... 201 3.2.9 Monopteryx . .. ........ .. ... 232
1.4.3 Umtiza .. . . .. . .. . ... . ...... 131 2.1.1 Dinizia . . . . . . . . . . . . 166 2.4.11
Archiclendron . . . . . . . . . . . . ... 202 3.2.10 Claclrastis . .. . . ...... .. ... . . 233
1.4.4 Tetrapterocarpon .......... . . 132 2.1.2 Pentaclethra ... .. ...... . .. 166 2.4.12
Falcataria . . . . . . . . . . ...... . . 202 3.2.11 Styphnolobium . . . . . . . . . . . . . 233
1.4.5 Arcoa .... . .. .. .. . . .. . . . .. 132 2.1.3 Aubrevillea . . . . ........ .. . 166 2.4.13
Serianthes . . . . . . . . . . . .. .... 202 3.2.12 Calia . .. . .... ... ...... .... 233
1.4.6 Acrocarpus ...... . . . .... .. . 133 2.1.4 Adenanthera . . . .. ..... ... 167 2.4.14
Paraserianthes . . . . . . . ... .... 203 3.2.13 Uribea .. . ... . ...... . . .. . . . 234
1.4.7 Ceratonia ... . . ..... ... ... .. 133 2.1.5 Tetrapleura . . ............ 167 2.4.15
Archidendropsis .. .... , ..... . 203 3.2.14 Sweetia . . . . . . . . . . . . . . . . . . . 234
1.4.8 Pterogyne . . .. ........... . . 134 2.1.6 Amblygonocarpus . . . . .. . ... 167 2.4.16
Wallaceodendron . . .......... 204 3.2.15 Luetzelburgia . . . . . . . .. , . . . . . 234
1.4.9 Haematoxylum ..... . ....... 135 2.1.7 Pseudoprosopis ..... . . . . ... 168 2.4.17
2.4.18 Pararchidendron . . . ........ . 204 3.2.16 Ormosia ... . . .. ... .... ... . 235
1.4.10 Cordeauxia . .. .... . ... ... .. 135 2.1.8 Calpocalyx .. . . . . , .......... 168
2.4.19 Hyclrochorea . .. . . . ......... 204 3.2.17 Haplormosia ... ............ 235
1.4. 11 Stuhlmannia . . . . . . . . . . . ..... 136 2.1.9 Xylia . . . . . ... . .. .. . . . . 168
2.4.20 Abarema . . . . . . . . . . . . ...... 205 3.2.18 Pericopsis . . .. . , .... .... ... 236
1.4.12 Mezoneuron .. ... , , ... .. . .. 136 2.1.10 Piptadeniastrum ..... ..... .. 169
2.4.21 Blanchetioclendron .. . ........ 205 3.2.19 Acosmium . . . . . . . . . . ....... 236
1.4.13 Pterolobium . .. ........ , . . .. 137 2.1.11 Entada . . . . . . . . . . . . . . . ... 169
2.4.22 Leucochloron . . . . . ... . ...... 206 3.2.20 Bowdichia . .. . . . ........... 237
1.4.14 Tara . .. . . ... . .. .......... . 138 2.1.12 Elephantorrhiza . . . . . . . . . . . 170
2.4.23 Chloroleucon .. . . ......... .. 206 3.2.21 Diplotropis ... ... . ... .. ... . 237
1.4.15 Coulteria ... . ... ...... .... . 139 2.1.13 Plathymenia . . . . . . . . ...... . . 170
2.4.24 Cathormion . . . . . . . . . ..... .. 207 3.2.22 Clathrotropis ... ... . ........ 238
1.4.16 Caesalpinia ... .... . ........ 140 2.1.14 Indopiptadenia . . . . . . . . . . . 170
2.4.25 Thailentadopsis . . ........... 207 3.2 .23 Petaladenium . . . . . . . . . . . . . . . 238
1.4.17 Pomaria . . . . . . . . . . . . . . . . . . . 141 2.1.15 Lemurodendron . . . . . . , .. .... 171
2.4.26 Sphinga . . .. . ... ........... 207 3.2.24 Sakoana ~ .. . . . . ........... 238
1.4.18 Erythrostemon . ......... . . . . 141 2.1.16 Newtonia . . .... . ..... 171
2.4.27 Havardia . . . . . . . . . ........ . 208 3.2.25 Neoharmsia ... ... .. .. ... . .. 239
1.4.19 Poincianella . . . . . . . ......... 142 2.1.17 Fillaeopsis . . ..... . ...... 171
2.4.28 Ebenopsis . . . . . . . . . . . ...... 208 3.2.26 Bolusanthus .. . ....... . .... . 239
1.4.20 Cenostigma ... .. , .... .... .. 142 2.1.18 Cylicodiscus ... . ........ 172
2.4.29 Painteria . . . . . . . . . . ........ 208 3.2.27 Platycelyphium ............. 240
1.4.21 Guilandina ... . ............. 143 2.1.19 Prosopis . . . . . . . . ...... . 172
2.4.30 Pithecellobium . . . ........... 209 3.2.28 Dicraeopetalum ....... ... . .. 240
1.4.22 Libidibia . . . . . . . . . . . . . . .... 144 2.1.20 Xerocladia ......... ... 173
2.4.31 Hesperalbizia . ... ........... 210 3.2.29 Cadia .. .. .. . . .... ... .. . . .. 241
1.4.23 Stahlia .. .. .. . . ............ 145 2.1.21 Prosopidastrum . . . . ........ 173
2.4.32 Pseudosamanea .. .... . ...... 210 3.2.30 Ammodendron . .......... , .. 242
1.4.24 Hoffmannseggia . . . . . . . . . . . . . 145 2.1.22 Piptadeniopsis . .......... .. 173
2.4.33 Samanea . . . . . . . . . ... ..... . 210 3.2.31 Ammothamnus . ....... . .. ... 242
1.4.25 Stenodrepanum . . . .. ... ..... 146 2.1.23 Neptunia . . . . . .. ..... . ... 174
2.4.34 Albizia .. .. .. . .. ........... 211 3.2.32 Maackia .. . . .. ........ .. . .. 242
1.4.26 Zuccagnia . . . . . . ........... 146 2.1.24 Leucaena . . . ............ 174
2.4.35 Enterolobium .. . . ........... 212 3. 2.33 Sophora . . . . . . . . , . . . . . . . . . . 243
. 1.4.27 Lophocarpinia . . .. .. . . ...... 146 2.1.25 Schleinitzia .. . .. ........ ... 175
2.4.36 Lysiloma . ... . . . ........... 212 3.2.34 Salweenia . . . . . . . . . . . . . .... 244
1.4.28 Balsamocarpon ... . ......... 147 2.1.26 Desmanthus . . . . . . , . . . .. 175
3.2 .35 Camoensia . .. . .. .. . ...... .. 245
1.4.29 Moullava .. . .. . . ..... ... . .. 147 2.1.27 Kanaloa . ... . . . ...... 176
3.2.36 Dalhousiea . . . ......•...... 245
1.4.30 Batesia . . . . . . . . . . . . . . . . . . . 148 2.1.28 Calliandropsis . . . . . . . . . . .. . 176
2.1.29 Gagnebina . .... ... .... . 176 3. PAPILIONOIDEAE 3.2.37 Airyantha ... . . ...... . . . .... 245
1.4.31 Recordoxylon . . . . ... . ...... 148 3. 2.38 Leucomphalos . ............. 246
1.4.32 Melanoxylon . . . . . . . . . . . . .. . 148 2.1.30 Dichrostachys . . . . . . . . . . . . . 177 3.1. SWARTZIEAE 3.2.39 Bowringia . . . . . . . . . . . ...... 246
1.4.33 Moldenhawera . ..... .. . . .. .. 149 2.1.31 Alantsilodendron . . . . . , ... .. 177
3.1.1 Bobgunnia . ... . .. .......... 217 3.2.40 Baphia .... . .. ......... .. .. 246
1.4.34 Tachigali . . . . . . . . . . . . . . . . . . 150 2.1.32 Parkia . . . . . . . . . . . . . . . . . 178
3.1 .3 Bocoa ......... . ... . . ..... 217 3.2.41 Baphiastrum .. ............ . 247
1.4.35 Arapatiella . . . . .. . .. .. ... . .. 151 2.1 .33 Anadenanthera . . .. . .... . . . 179
3.1.2 Swartzia . . . .. .. .. .......... 218 3.2 .42 Amphimas ... . .. , ......... . 247
1.4.36 Jacqueshuberia .. . ..... . ... . 151 2.1.34 Pseudopiptadenia .......... . 179
3.1.4 Candolleodendron . ..... ..... 219 3.2 .43 Panurea . . .... ... .. . . . ... . . 248
1.4.37 Schizolobium . .. .... ....... . 151 2.1.35 Piptadenia ... ......... , . 180
3.1.5 Trischidium . .. . . . ..... . . . . . 219 3.2.44 Spirotropis . . . . ....... .... .. 248
1.4.38 Bussea . .. . .. .... ... , . .. . . . 152 2.1.36 Parapiptadenia . . . . . . . . . . . 181
3.1.6 Cyathostegia .... .. ......... 219 3.2.45 Uleanthus .... . . .. ... , .. .. . 248
1.4.39 Pelto phorum . . . . . . . . . . . . . . . 152 2.1.37 Microlobius . . . ...... . ..... 181
3.1,7 Ateleia . .. ..... . . . .. ....... 220
1.4.40 Parkinsonia .. . ..... .... ... . 153 2.1.38 Stryphnodendron . . . . . ... . . 182
3.Ls Amburana . . . . . . . . . . . . . . . . . 220 3.3 DIPTERYGEAE
1.4.41 Conzatt~ .. .. . . .... ..... .. . 154 2.1.39 Adenopodia . . .. .. . . ... . . . 182
3.1.9 Mildbraediodendron . . . . . . . . . . 220 3.3.l Taralea ... ... .......... . .. 251
1.4.42 Delo nix . . . . . . . . . . . . . . . . . . . 154 2.1.40 Mimosa ... . . ......... . 183
3.1.10 Cordyla . .. . .. . .. . . ... . .... 221 3.3.2 Pteroclon .. . . ..... . .. . ... .. 251
1.4.43 Colvillea .. ... ..... . . . .. .. . 155
3.3.3 Dipteryx .... ........ . . .... 251

14 LEGUMESOFTHEWORLD INTRODUCTION 15
 BRONGNIARTIEAE 3.9.11 Petteria ....... ............ 290 ;_ Grazieln Ien iron ............ 326 3.15 INDIGOFEREAE
3.4 3 11 ...
3.9.12 Laburnum ..... .......... . . 291 · 1. 33 L)a lb rgi~1 .. . ... , ........... 327 3.15.1 362
3.4.1
3.4.2
Cyclolobium .... ........ . .. 254
Poecilanthe . . . . . . . . . . . ..... 254 3.9.13
3.9.14
Podocytisus .... .. . ...... ... 291
Hesperolaburnum .. . .... .... 292
;:u.3 '
1
Macha riu m . .... ........... 328
?i.ll .35 A ;-;d1yn mene .... ......... 329
3.15.2
3.15.3
Phylloxylon ...
Cyamopsis ....
Indigastrum ...
.............
.............
.............
362
362
3.4.3 Harpalyce ...... . . . . . . ..... 255
Brongniartia . . . . . . . . . . . . . . .. 256 3.9.15 Cytisus ........ ......... . . . 292 3.11.3 > y fo ~ r ~<l . .' . . . . . . . . . . . . . . . 330 3.15.4 Microcharis ... ............. 363
3.4.4
Plagiocarpus . . . . . . . . . . . . . .. 256
3.9.16 Lembotropis .... ...... . . . ... 293 .11.37 So~ m'.nenngrn .............. 330 3.15.5 Rhynchotropis . ............. 363
3.4.5 3
Templetonia . . . . . . . ......... 256
3.9.17 Calicotome .... ... ... ...... 293 3.11 .38 ."1rnLhW · ·. · · · · · · · · · · ...... 330 3.15.6 Vaughania .... ...... . , .. ... 363
3.4.6
Hovea ....... . . ........... 257
3.9.18 Echinospartum .. . ....... .... 293 3. U .39 K~ ts ·hya · . . · .. ............ 331 3.15 .7 Indigofera .... ............. 364
3.4.7
Cristonia ....... ... , ....... 258
3.9.19 Erinacea ...... ............ 294 . (1. . ) Ht1mula ria ................. 332
3.4.8 3
Thinicola . . . . . . . . .......... 258
3.9.20 Retama ....... . .. . .. ...... 294 .11.41 Bryaspis ........ . .. ... ..... 332 3.16 MILLETTIEAE
3.4.9 3
3.9.21 Gonocytisus .... ........ . ... 294 .11.42 Geissaspis ... . .. . . ...... ... 332 3.16.1 Callerya ..... .............. 370
3.4.10 Lamprolobium . . . ........... 258 3
3.9.22 Genista ....... ..... .. .... . 295 .11.43 Pictetia ........ ... . .. .. . .. 333 3.16.2 Antheroporum .............. 370
3
3.9.23 Spartium ...... ...... ... ... 296 3J1.44 Diphysa ........ .. . ........ 333 3.16.3 Endosamara .. ... . . ......... 370
3.5 EUCHRESTEAE
3.5.1 Euchresta .... . ... . . , . , , . .. . 261
3.9.24 Stauracanthus ... . . .......... 296 3.11.45 Zygocarpum .... ........... 334 3.16.4 Sarcodum ... .............. 371
3.9.25 Ulex ......... . . .... .. , ... 296 3J1.46 Ormocarpum .... ........... 334 3.16.5 Afgekia ..... ........ , .. . .. 371
3.11.47 Onnocarpopsis .. ........... 335 3.16.6 Wisteria ..... ............. , 372
3.6 THERMOPSIDEAE 3.10 AMORPHEAE 3J 1.48 Peltiera ........ ... ........ 335 3.16.7 Austrosteenisia .... . ..... .. .. 372
3.6.1 Pickeringia . . . . . . . . . . . . . . . . . 264 3.10.1 Apoplanesia . ..... . ......... 301 3.11.49 Weberbauerella .. ...... . ... . 335 3.16.8 Leptoderris .. . ............. 372
3.6.2 Ammopiptanthus . . . . . . . . . . .. 264 3.10.2 Parryella ... .. . ... .... ... . . 301 3.16.9 Dalbergiella .. .............. 373
3.6.3 Anagyris . . . . . . . . . . . ....... 264 3.10.3 Amorpha ... ... .. . ... ...... 302 3.12 HYPOCALYPTEAE 3.16.10 Aganope . . .... . .... . ... . . . 373
3.6.4 Piptanthus . . . . . . . ........ .. 265 3.10.4 Errazurizia .. . . .... . . ....... 302 3.12.1 Hypocalyptus .... . . . . . ...... 337 3.16.11 Ostryocarpus ... ............ 373
3.6.5 Thermopsis . . . . . . . . ... . .. . . 265 3.10.5 Eysenhardtia ............... 303 3.16.12 Xeroderris ..... ............ 374
3.6.6 Baptisia ....... .. .. . .... .. . 265 3. 10.6 Psorothamnus ........ .. .... 303 3.13 MIRBELIEAE 3.16.13 Fordia ........ ......... ... 374
3.10.7 Marina ..... ......... ....... 303 3.13.1 Gompholobium ....... , ..... 340 3.16.14 Dewevrea ..... ............ 375
3.7 PODALYRIEAE 3.10.8 Dalea ..... ............... 304 3.13.2 Sphaerolobium ............. 340 3.16.15 Platysepalum ... ............ 375
3.7.1 Cyclopia . . . . . . . . . . . . . . . . . . 268 3.13.3 Daviesia .... ... ........... 341 3.16.16 Sylvichadsia .... ........ , ... 375
3.7.2 Xiphotheca . . . .......... .. . 268 3.11 DALBERGIEAE 3.13.4 Erichsenia ... .............. 342 3.16.17 Schefflerodendron ........ . .. 376
3.7.3 Amphithalea . . . . . . . . . . ..... 269 3.11.l Vatairea ...... ......... ... . 310 3.13.5 Viminaria .... .............. 342 3.16.18 Craibia ........ . .. .. . ... . .. 376
3.7.4 Stirtonanthus . ... .... ....... 269 3.11.2 Vataireopsis ... ............. 310 3. 13.6 Isotropis .... ............. . 342 3.16.19 Disynstemon ... ............ 376
3.7.5 Podalyria . . . . . , , .. . . , , .... . 269 3.11.3 Hymenolobium . .. ....... . . . 310 3.13.7 Jacksonia .... .............. 343 3.16.20 Platycyamus .... ......... . .. 377
3.7.6 Liparia ...... .............. 270 3.11.4 Andira ....... ........ ... .. 311 3.13.8 Leptosema ... .............. 344 3.16.21 Kunstleria ..... .... . ... .... 377
3.7.7 Virgilia . . . . . . . . . . . . . . . . . . . . 271 3.11.5 Adesmia ..... ... .. . ....... 312 3.13.9 Latrobea .... ....... . ...... 344 3.16.22 Burkilliodendron .. . .. . ... . .. 377
3.7.8 Calpurnia .... ... . .......... 271 3.11.6 Amicia ....... . ... . ........ 312 3.13.10 Euchilopsis .. .... . ......... 344 3.16.23 Craspedolobium . ............ 378
3.11.7 Zornia ....... . . ...... , .... 313 3. 13.11 Phyllota ..... .............. 345 3.16.24 Philenoptera ... .. . ......... 378
3.8 CROTALARIEAE 3.11.8 Poiretia ...... . . ... . ...... . 313 3.13.12 Otion ....... .............. 345 3.16.25 Hesperothamnus ........ , .. , 378
3.8.1 Spartidium . . . . . . .... ...... . 275 3.11.9 Nissolia ...... .. .... .... . .. 314 3.13.13 Aotus ....... ............ .. 346 3.16.26 Piscidia ....... .. . ......... 379
3.8.2 Lebeckia ..... ............. 275 3.11.10 Chaetocalyx ... .. ... .... . . . . 314 3.13.14 Urodon ..... .............. 346 3.16.27 Dahlstedtia . . .. . . .. . ...... . 379
3.8.3 Wiborgia . . ... ............. 276 3.11.11 Riedeliella .... ............. 315 3.13.15 Stonesiella ... . ....... . ..... 346 3.16.28 Deguelia ...... ............ 380
3.8.4 Rafnia ....... ..... ... . .... 276 3.11.12 Discolobium .. . . ........... 315 3. 13.16 Almaleea .... ..... .. . ...... 347 3.16.29 Lonchocarpus .. .. .... ...... 380
3.8.5 Aspalathus . . . . . . . . .... . .... 277 3.11.13 Cranocarpus .. . ............ 316 3. 13.17 Eutaxia ..... .............. 347 3.16.30 Behaimia ...... ......... ... 381
3.8.6 Lotononis . . . . . . . . . . . . . . ... 278 3.11.14 Brya ........ ............. 316 3. 13.18 Dillwynia .... . ........ . .... 347 3.16.31 Bergeronia ..... .. . ......... 381
3.8.7 Bolusia . . . . . . . . . . . . . . . . ... 278 3.11.15 Platymiscium .. ........ . .... 317 3. 13. 19 Pultenaea .... ..... .... ..... 348 3.16.32 Margaritolobium . ............ 382
3.8.8 Crotalaria . . . . . . . . . . . . . . . . . . 279 3.11.16 Platypodium .. ..... . . . . . ... 318 3.13.20 Mirbelia ..... .............. 349 3.16.33 Muellera ...... . .. ..... .. .. 382
3.8.9 Pearsonia ..... ..... . ... . . . . 281 3.11.17 Inocarpus .... ............ , 318 3.13.21 Chorizema .. . . . ............ 350 3.16.34 Derris ........ .... . . . ..... 382
3.8.10 Rothia ....... ... . . .. .... . . 281 3.11.18 Maraniona ... . . ...... . . . . . 319 3.13.22 Oxylobium .. .............. 350 3.16.35 Paraderris ..... ............ 383
3.8.11 Robynsiophyton ............. 281 3.11.19 Tipuana ...... ............ . 320 3.13.23 Podolobium .. . ... .. . .. . . .. . 351 3.16.36 Millettia ....... . . . .. . ... .. . 383
3.11.20 Ramorinoa .... ..... ....... . 320 3.13.24 Callistachys .. .............. 351 3. 16.37 Pongamiopsis .. .. . .. . ... . .. 384
3.9 GENISTEAE 3.11.21 Centrolobium .. ... .. ....... . 321 3.13.25 Gastrolobium . .. ........ . ... 351 3.16.38 Pyranthus .... . . ..... .. . ... 384
3.9.1 Melolobium ... ....... .. .... 285 3.11.22 Paramachaerium . . .. .. . .... . 321 3.16.39 Chadsia ....... ............ 385
3.14 BOSSIAEEAE 3.16.40 Mundulea ..... ............ 385
3.9.2 Dichilus . . . . . . ...... .... ... 285 3.11.23 Etaballia ..... .. . . . ..... .. . 321
Polhillia ...... .. , .......... 286 3. 11.24 Pterocarpus ... ..... . ...... . 322 3.14.1 Goodia .... ............... 356 3.16.41 Tephrosia ..... ..... . . ... , . 386
3.9.3
Argyrolobium . . ............. 287 3.11.25 Cascaronia .... .. . .. . ...... . 323 3.14.2 Bossiaea ... ............... 356 3. 16.42 Apurimacia .... ......... .. . 386
3.9.4
Lupinus ...... .. . ........ .. 287 3.11.26 Geoffroea .... .... .. . ... .. . 323 3.14.3 Platylobium . ............... 357 3.16.43 Paratephrosia ... ... . . . ...... 387
3.9.5 3.14.4
3.9.6 Anarthrophyllum . . . ...... , . . 289 3.11.27 Fissicalyx ..... . . . ..... . . .. . 324 Muelleranthus .............. 358 3.16.44 Requienia ..... ............ 387
3. 11. 28 Fiebrigiella . . . . . . .......... . 324 3.14.S Ptychosema . . . . . . . . . ....... 358 3.16.45 Ptycholobium .. .... ........ 387
3.9.7 Sellocharis . . . . . . . . . . . . . ... . 289
3.11.29 Chapmannia .. ............ . 324 3.14.6 Aenictophyton . ........ ..... 358
3.9.8 Adenocarpus . . . . . . ........ . 289
3.9.9 Cytisophyllum . . ......... . . . 290 3.11.30 Stylosanthes ... ..... . . ..... . 325 3.17 ABREAE
3.9.10 Argyrocytisus . . . . . . . . . . . . . . . 290 3.11.31 Arachis ...... . , ..... . .... . 326 3.17.1 Abrus . ................... 390

INTRODUCTION 17
16 LEGUMES OF THE WORLD

3.18 PHASEOLEAE
3.18.1 Dioclea .. ..... ..... , ...... 395
3.18.2 Luzania ...... . . . . ......... 395
3.18.3 Macropsychanthus ........... 395
3.18.4 Canavalia . .. . . .. ... ... ..... 396
3.18.5 Cymbosema .... ..... .. ..... 397
3.18.6 Cleobulia .. ... . ... . . . ...... 397
3.18.7 Camptosema .... . . . ........ 397
3.18.8 Cratylia ....... ......... . .. 398
3.18.9 Galactia ....... ...... .. . .. . 398
3.18.10 Collaea ....... ............ 398
3.18.11 Lackeya ..... . . ........... . 399
3.18.12 Rhodopis ....... ..... ... . . . 399
3.18.13 Neorudolphia .. . .... . .... . .. 399
3.18.14 Cruddasia ..... ... ......... 400
3.18.15 Ophrestia .. . . .. ........... . 400
3.18.16 Pseudoeriosema . ......... ... 400
3.18.17 Clitoria . .. .. . . ... ... ...... 401
3.18.18 Barbieria .... . . .. .. .. ... ... 401
3.18.19 Centrosema .... .... .... . , . . 402
3.18.20 Periandra . . . . .. ............ 402
3.18.21 Clitoriopsis .... ... ...... ... 403
3.18.22 Apios ........ . ........ . .. . 403
3.18.23 Cochlianthus ... ... ...... . . . 403
3.18.24 Shuteria . .... . . .... .... .... 404
3.18.25 Mastersia .. ... ............. 404
3.18.26 Diphyllarium ... . . .......... 404
3.18.27 Mucuna .. .... . ............ 405
3.18.28 Kennedia ..... ..... ... ..... 406
3.18.29 Hardenbergia . .. ........ . ... 406
3.18.30 Vandasina ..... ............ 407
3.18.31 Spatholobus .... ....... .. ... 407
3.18.32 Butea ........ ......... . .. 407
3.18.33 Meizotropis .. .. . . .. . .. , ... . 408
3.18.34 Adenodolichos .. ....... . .... 408
3.18.35 Paracalyx ..... .. ........... 408
3.18.36 Bolusafra . . .. . . ....... .. ... 409
3.18.37 Carrissoa . . . . . . ............ 409
3.18.38 Chrysoscias .... .. . ... . . .... 409
3.18.39 Rhynchosia . ... .. .. ........ 410
3.18.40 Eriosema . . .. . . . . • ... .. . ... 411
3.18.41 Dunbaria . . . . . . ............ 411
3.18.42 Cajanus ....... . . ..... . .. .. 412
3.18.43 Flemingia ...... ... ......... 413
3.18.44 Erythrina .. ... .. ........... 413
3.18.45 Psophocarpus .. ............ 414
3.18.46 Dysolobium .... .... ..... ... 414
3.18.47 Otoptera ...... . . .......... 414
3.18.48 Decorsea . . . ... ........... . 415
3.18.49 Strongylodon ... .... . . . ..... 415
3.18.50 Calopogonium . . ... .... ..... 416
3.18.51 Cologania .. ... ............ 416
3.18.52 Pachyrhizus ... . ............ 416
3.18.53 Herpyza .. . .... .. , .... ..... 417
3.18.54 Neorautanenia .. .... ... ..... 417
3.18.55 Neonotonia .... ............ 417
3.18.56 Teyleria ....... ............ 418
3.18.57 Dumasia . ... .. ......... . . . 418
18 LEGUMESOFTHEWORLD
3.18.58 Pueraria ..... ...... . ....... 418 ,19.25
3
3.18.59 Nogra ... ... . .. .. . .... .... 419 .19.26 l ys1ca r~u s . .. ... .......... 444
3
3.18.60 Eminia .. . ... .............. 419 fp9.27 D sin d1as1. rum . ... .. . . . . . .. 444
3.18.61 Sinodolichos . .... ... ....... 419 3 ..19.29
1 9.2~ l llLniella .. . ......... , .... 445
3.18.62 Pseudeminia . .............. 420
. l9.:i0 TI!eiotls ............. .. .... 445
3.18.63 Pseudovigna . ....... . ...... 420 3
3.18.64 Amphicarpaea .......... ... . 420
po
3.18.65 Teramnus . .. ..... . ........ 421
3.20.1
3.18.66 Glycine . .. .. .. . . .. ....... . 421
3.20.2
3.18.67 Phylacium ... . .. ... ........ 421
3.20.3
3.18.68 Neocollettia .. .............. 422
3.20.4
3.18.69 Wajira .. . ... .............. 422
3.20.5
3.18.70 Sphenostylis . . .. ............ 422
3.20.6
3.18.71 Nesphostylis . .. ......... ... 423
3.20.7
3.18.72 Alistilus ..... ...... .. ...... 423
3.20.8
3.18.73 Austrodolichos .. ........... . 423
3.20.9
3.18.74 Dolichos .... . ............. 424
3.18.75 Macrotyloma . ............ .. 425
3.18.76 Dipogon .... ..... .... .... . 425 3.21 SESBANIEAE
3.18.77 Lablab ...... .............. 425 3.21.l Sesbania ... ............... 453
3.18.78 Spathionema . .. ..... ... ... . 426
3.18.79 Vatovaea .... ...... .. ..... . 426 3.22
3.18.80 Physostigma .. . . ..... . ...... 426 3.22.1
3.18.81 Vigna ....... ........... ... 427 3.22.2
3.18.82 Oxyrhynchus .. ........ . .... 427 3.22.3
3.18.83 Phaseolus ... .............. 428 3.22.4
3.18.84 Ramirezella .. ... .. ..•.... . . 429 3.22.5
3.18.85 Strophostyles ... . ........... 429 3.22.6
3.18.86 Dolichopsis . ...... . . . . ..... 429 3.22.7
3.18.87 Macroptilium .......... . . . .. 430 3.22.8
3.18.88 Mysanthus .. ........ . ...... 430 3.22.9
3.18.89 Oryxis .. . . . . .. . . .......... 430 3.22.10
3.22.11
3.19 DESMODIEAE 3.22.12
3.19.1 Campylotropis . ............. 434 3.22.13
3.19.2 Kummerowia .. ... ...... .... 434 3.22.14
3.19.3 Lespedeza . . .. .......... , .. 435 3.22.15
3.19.4 Dendrolobium . ........ . .... 436 3.22.16
3.19.5 Phyllodium .. . ....... . ..... 436 3.22.17
3.19.6 Ougeinia . .... ... . . ....... . 436 3.22.18
3.19.7 Aphyllodium .. ............. 437 3.22.19
3.19.8 Ohwia .. ..... ... . ......... 437 3.22.20
3.19.9 Hanslia ...... ............. 437 3.22.21
3.19.10 Arthroclianthus .... . ........ 438 3.22.22
3.19.11 Nephrodesmus . ............. 438
3.19.12 Tadehagi ..... .. .. .... .. .. . 438 3.23
3.19.13 Akschindlium .. .......... .. . 439 3.23.1
3.19.14 Droogmansia .. ..... . .. .. . . . 439 3.23.2
3.19.15 Monarthrocarpus .. . ......... 439 3.23.3
3.19.16 Trifidacanthus . ............. 440 3.23.4
3.19.17 Desmodium ... ........... .. 440 3.23.5
3.19.18 Codariocalyx .. ............. 441 3.23.6
3.19.19 Hylodesmum .. ..... .... .... 441 3.23.7
3.19.20 Hegnera ..... .... . ........ 441 3.23.8
3.19.21 Pseudarthria ... .... .. .... ... 442 3.23.9
3.19.22 Pycnospora ... ............. 442 ;·23.10
3.19.23 Mecopus .. . .. ............. 442 .23. 11
3.19.24 Uraria . ...... .... .. .. . . . .. 443
hri~t.fa . .................. 443 3.24 GALEGEAE
3.24.1 Glycyrrhiza .. .............. 478
3.24.2 Chesneya .... .... .. .. ...... 478
l.epl clesmia .... .. ........ . 445
3.24.3 Spongiocarpella .... .. ....... 478
3.24.4 Gueldenstaedtia ...... . ...... 479
3.24.5 Tibetia ...... ...... .... .... 479
3.24.6 Erophaca .... .. . .......... . 480
PSORALEEAE 3.24.7 Oxytropis . .. ........ .. .... 480
Otholobium . . . ............. 448 3.24.8 Biserrula .... . . ............ 480
Psoralea . . .. . .. . ...... , .... 448 3.24.9 Astragalus ... .... . . ..... ... 481
Orbexilum . . .... . .......... 448 3.24.10 Ophiocarpus . .... .... .. . ... 482
Hoita ... . .. . .. . ........... 449 3.24.11 Barnebyella ... ............. 482
Rupertia .. . . ............... 449 3.24.12 Colutea ..... •..... ... ... .. 482
Psoralidium .. ....... ....... 449 3.24.13 Oreophysa . . .. ............. 483
Pediomelum ..... . ......... 450 3.24.14 Smirnowia .... ..... .... .... 483
Bituminaria ..... .. ......... 450 3.24.15 Eremosparton . ..... ........ 483
Cullen ... ... ... ....... . ... 450 3.24.16 Sphaerophysa . ... .. .. . ..... 484
3. 24 .17 Lessertia . . . . . . . . . . . . . . . . . . 484
3.24.18 Sutherlanclia ... ............. 485
3.24.19 Swainsona .... ............. 485
3.24.20 Montigena .... . . ........... 486
3.24.21 Clianthus . .. .. ............. 486
LOTEAE
3.24.22 Carmichaelia .. ............. 486
Hippocrepis ... ... ... . ...... 457
3.24.23 Streblorrhiza .. ..... .. ...... 487
Scorpiurus .. .. .. ........... 457
3.24.24 Galega .... . .. .. .... .... . .. 487
Securigera .... . . ..... ... ... 458
Coronilla .. . .. ............. 458
3.25 HEDYSAREAE
Podolotus .... .. . .......... 458
Anthyllis .. .... ............ 459 3.25.1 Calophaca . . .. . . . . ......... 491
Hymenocarpos . .. . ......... . 459 3.25.2 Caragana . .... ... .. ... ..... 491
Pseudolotus ... ........ . . . .. 460 3.25.3 Halimodendron .. .. ... .... . . 491
Antopetitia .. .. .......... . .. 460 3.25.4 Alhagi ....... ............. 492
Hosackia ..... ............. 460 3.25.5 Eversmannia .. ........ ..... 492
Ornithopus .. . . . ........... 461 3.25.6 Hedysarum . . . ............. 493
Dorycnopsis .. .. . ... .. . . ... 461 3.25.7 Corethrodenclron ..... . ...... 493
Kebirita . ... . . ......... . .. . 461 3.25.8 Sulla ... .... . ........... .. 493
Ottleya ...... ...... , ...... 462 3.25.9 Taverniera .... .. . ...... ... . 494
Acmispon ... . ............. 462 3.25.10 Onobrychis ... ........ ..... 494
Syrmatium . . . . .. . . .. ....... 462 3.25.11 Sartoria . . .... ........ . .... 495
Lotus ...... . .. .... ..... . .. 463 3.25.12 Ebenus ... ... .... . ... ..... 495
Dotycnium ... ............. 464
Tetragonolobus ........... .. 464 3.26 CICEREAE
Tripodion .. . . . . ........... 465 3.26.1 Cicer . . . .... .............. 497
Hammatolobium .... . . .. ... . 465
Cytisopsis ... . .... . . ....... 465 3.27 TRIFOLIEAE
3.27.1 Parochetus ... , ..... . ....... 501
ROBINIEAE 3.27.2 Trifolium . . ... . .......... .. 501
Hebestigma .... . .. . . .. ..... 468 3.27.3 Ononis ... . . ..... .. .. . .... 502
Lennea ....... .... . ....... 468 3.27.4 Melilotus .. . ............... 502
Gliricidia ...... ..... .. .. ... 468 3.27.5 Trigonella . . ... ......... . .. 503
Poitea ........ ............ 469 3.27.6 Medicago .. ............ ... . 503
Olneya .. . .... ............ 469
Robinia ....... .... ... . .... 470 3.28 FABEAE
p 01ssoma
. ' . . . . . . . . . . . . . . . . . . 470
3.28.1 Vicia .. , .... . ...... . ... ... 506
Coursetia ...... .... .. .. . . . . 471 3.28.2 Lens . ... ............... .. 506
Peteria ... .. . . . ........ .... 472 3.28.3 Lathyrus .............. . . . . . 507
Genistidium .... ............ 472 3.28.4 Pisum . ... , ........... . ... 508
Sphinctospermum ...... . .... 472 3.28.5 Vavilovia ... . ..... . ........ 508
INTRODUCTION 19
 Biogeography of the Leguminosae
by B. o. schrire, G. P. Lewis and M. Lavin

Ba/samocarpon brevifolium - Tribe Caesalpinieae
Introduction
M< rley (2 ()0 implies <t 'moist 'quat rial m g:ithermal'
o.rlgin of I •gumes, c. 8 - 7 Mn In lhe :impanhn. c>r
ta:i:-.tn ·htian of the Upp ·r r lac ·<)US t11u. s upi rung
a e:L >ondw<1na (or ausLrotropical) o rigin >r lh •
fatnll (Raven & Axdr d , 1974; Raven & Polhill 1981).
'l'hl::. Jo ng-h Id Wes t ondwana hyp the. is fo r rh
origin >f I 'gu m s requir·s dP f::tmiJy diversifl ~ttion
i.t: ., lhat or Lh ' I gu m ·rown ·la I ; Pig. 3) to b' at
t ast 100- O Ma in ag ~( Lavin er al. _Q ; Davis "' ul. ,
2002b), when Africa and South America were last in
near contact, although Raven & Axelrod 0974) and
Morley (2000; 2003) suggest that dispersal routes
existed over islands and ridges between these
continents until c. 65 Ma. In addition, it is often stated
(e.g., Morley, 2000) that the tropical angiosperm fossil
record is biased to Laurasian collection localities, and
that if more fossils were available from South America
and Africa, the stratigraphic record for many groups
would be older.
Although it is true that legumes are now highly
diverse in both Africa and South America, fossil data
alone indicate that neither a moist megathermal origin,
nor a Mesozoic age of legume diversification is likely.
The clear message derived from fossil legume studies
such as those of Herendeen et al. 0992), Herendeen
& Dilcher 0992), Herendeen (2001) and Jacobs (2003)
is that all three subfamilies of legumes are well
represented in the fossil record in North America,
Europe, Africa, and Asia by at least fruits and leaves
from recent times back to the Palaeocene. Putative
earlier legume fossils include only pollen and wood
specimens that lack any specific legume
synapomorphies, and even then such fossils go back
only to the latest Cretaceous. Given the temporal and
sp. Lial ·on Linuity >f Ii er · ' legume ma ro foss il :
lhmughou1 Lhe C nozoi ·, and th • ·1hrupl abs nee f
d · ·Iclu uus l ·gume lcafl t and pods prlor to rhe Late
htl<t o · ·ne, the origin of I gu rnes is unlikely to b
much bdore 60 Ma . lkmark abl y, the r:1 pld
cli·v'rsin ·:nion of rite fami ly must h:.tvc >c ·urr ~d oon
alt ~r. B the mid 11 • Eo en· ( ·. 'ill Ma) n arl y all or lh ·
~1 HJOr lincag s h:1V1: a fossil recor<.l in orth America,
lt'r pe, fril:a, and Ash .g., /\ xdrod , 1992; La in.,
00
9 ; II l" nc.I en ,, al. l 992; H er •ndccn, 2001). Tn
:tc.ldllion
e . • •' I ·111·0 n gram o r· Ll 1 • 111a/.1(, . p I1yIogeny (I,avm
1
. nl., lrl rress shows minlrna I r ·st lu1 ion at ilw base
O Ii , Ir ...... 1~"t
itlJ lh .
'"ween_niaior T< w n l ' Ia I · c11ve · rl': ii ..1•a lions,
al~ ~· 1 llng the abntpl cm rg ' 11 .,. ,r :tll hut 0 111:
>f l e~'Clm s b ·rw ·en O an I 50 Ma .
Photograph by P. Baxter
The legume supertree (Fig. 1) is derived from a family-
wide phylogeny based on DNA sequences of the
chloroplast matK region (Wojciechowski et al., 2004)
and complemented by chloroplast trnl region analyses
of the Caesalpinioideae (e.g., Bruneau et al., 2000;
2001; Fougere-Danezan et al., 2003; Herendeen et al.,
2003a; Forest (unpubl. data) and Mimosoideae
(Luckow et al., 2000; 2003; Miller & Bayer (2003), as
well as chloroplast trnl and rbcl region analyses of
Papilionoideae (Doyle et al., 2000; Pennington et al.,
2001; Kajita et al., 2001). The emerging supertree is
largely stable, although the circumscriptions of
terminal monophyletic groups will continue to be
refined as more taxa are sampled; it is, however,
already requiring us to modify our traditional concepts
of legume biogeography.
Data from these large-scale molecular phylogenies
have recently been compiled by Schrire et al. (2005),
to identify the major subgroups of legumes and the
biogeographical inter-relationships among these
groups. The chloroplast matK phylogeny representing
a comprehensive sampling of all major legume groups
(Wojciechowski et al., 2004) served as the backbone for
the supertree (summarised with modifications in Fig.
3). In addition, major legume subclades ·detected in
the phylogenetic analyses of chloroplast trnl and rbcl
sequences noted above, were used in supertree
construction because they represented a more
exhaustive sampling of various local subclades. A strict
supertree (sensu Sanderson et al., 1998) was readily
constrncted manually because of the high compatibility
of all the component trees. Essentially, the large-scale
matK phylogeny (Wojciechowski et al., 2004)
represented all major clades of legumes, and the other
molecular phylogenetic studies identified the broader
constituents of these monophyletic matK subclades.
The local subclades were then scrutinised for global
distribution patterns. Four generalised areas of
endemism predictive for legume distributions were
identified at the biome level , including a newly
recognised amphi-Atlantic Succulent Biome. Schrire et
al. (2005) subjected the resulting taxon-biome
supertree (Figs. 4 -14) to cladistic vicariance analyses,
to detect a generalised pattern of biome relationships
. for legumes. Chronograms derived from rate-smoothed
bayesian consensus trees of the matK phylogeny (Lavin
et al., 2004; in press) provided comparative clade ages
major within the family, based on thirteen time constraints
derived from fossil evidence.

BIOGEOGRAPHY OF THE LEGUMINOSAE 21

20 LEGUMESOFTHEWORLD
 predictable occurrences, e .g., S-biome taxa frequently
cCll .c
Cll
... "'
Cll
... .., Biomes have secondary G - (but rarely R-) centres while the
... 'C'
..,
i
Cll ..
u reverse is rarely the case in predominantly G - and R-
.........
.5
r l •rminal taxa usetl in Lhe legum
"'..,."" M -r of tht: nm i
.... "'Cll
Cll Cll Cll ...
v.....
7.. ·~ ~ biome taxa . Taxa in the latter two biomes associate
!!: ..,. ";ij ~ ~ e-
SU I ertre , ro.::pr s nt ·d well suppo rt d dadcs llrnt ·ould
much more closely with each other, and many genera
"' v g.., be I , 5 nib •d in g n •ral ~co l g i <I t rm s in vo l ing
..,"'
:§ "'
-0
·s
..,.
..,..,..... ... .~
.:!:.
...c
N C.,.
'-'
........
....: ra c
. .., Cll

!11 is lllre '" · t to dry l te mp ' ratur , (L r pi ni l Lo

w ith a predominance of species in one biome also

.."'
0
Cll
·;:
:s l"I
"g
-.::: QI
.... ""...
·;:; t
II..
E
Cll
c
~Cll
...
Qj

; ~·a
cii-
c .. re mp ,,~ ,l~ )1 antl Iistu rlx 1nc (llre- l1isLOry to no fir _ have substantial centres in the other. Many legume
terminals assigned to R-areas, therefore, had to be
<(
......
..,..... .!!!
.c ~ 0-g
"'c:, hit 1y) g1,1Jicnts. The g ·m.:ralisctl combination of1h /;)'
..,·c;"' II.. 3:..!!
"g l,J
~
.c
t:
0
·;:: 1l
.~ N ,._
41
c
0
~ 0
0
.b
hMl resu lt I in recognising the four I io rne. which a r•
designated as also inhabiting G-areas (i.e., in Figs.
"'E :::E...
=
~ '°"'
0
..:E... ~
~
-:;;
::I
~
~c £::I
<(
--~ ...
Q ......
cit E-;
ie, c
....
Cll"'

::i "-
d •s ril ·d h I x . Thes in lud l l a semi-arid, fl re-
4-14 , R/ G or G/ R depending on which biome
contained more taxa) . Clades in the T-biome are
i V)
int I ' ran t, . uc.: ·ul 111-ri h a nd grass-poor, dry tropi ·a l
LI.I ...~
Cll

~
<( ~ v"'
0
Ill w
::I ::I Cll
.c ~
::I

~ >r •st, 1liid el , ncl bushlan I hiom ahhr ~via ted h •1 l()
subdivided into no1thern TN- and southern TS-regions,
• •••
0
c:s: ,.... ......
•• .., >
Cll
.!! ,.... based on their distinctive patterns of diversification
LI.I e, 'M ;
g
fti
c...~
succulent or . -), 2 a fir -lo lernnt, sue ul nt-poor and
:a across the phylogeny. Although each colour-coded
-0
Q
.E
~:§
~~ l"I
'°,....
N

0
O"&?~
Cll 0
0 CL grass-rich, sea on· dl y dry tropical forest, woodland and
savanna bi m ~ (Grass r G-), 3) a tropical wet forest box in the taxon-biome supertree (Figs. 4 -14) denotes
z .;1 N
0
N .~ Z
'-:'i_~
.... Cll

biome (Rainforest or R-), and 4) a temperate biome the predominant biome for that clade, the species
0 " ....
-.....
II)
l"I
~i ... E,1:-:::
ti ... ... including both the Northern and Southern Hemispheres numbers given are for the entire clade and not for the
..,
·cs"' ao :z: +
~ 2 ·=] (Temperate or T-). T hese areas reflect major
designated biome (u nl ess boxes are divided to
Q: :c.... "',.... o bn 0E II)
..:.= ...
::1
represent this split where the data for species numbers
~ .2.. ~ .! zonobiomes (Breckle, 2002): tropical wet (Zonobiome
~ ...
1111 ..,... 3: ........ a:!!!
~=~~ I), two areas of tropical d1y (in Zonobiomes II and III) , per biome are known). It is likely, therefore, that some
~z g ~~ E and temperate (Zonobiomes IV -X). species in single boxes may be centred in secondary
~ g--ro ~ (or tertiary) neighbouring biomes.
..,. ::::~ ti t:c.e Assignment of major subclades to one or more
Of the four fundamental biomes identified here
-0"' "'3: '° ... "' >-
'1~·-Q1

!9~4;~
biomes initially involved ascertaining the distributions
the Succulent biome is perhaps most novel and need~
::l 0
~c :~Cl.I 't ~ of c. 730 legume genera world-wide. Within each
.
l.:J
~~
,....
~
"'~.cc

~.!
0 ......

i :2
genus, patterns were especia lly sought among the to be distinguished from the similar, largely tropical
Southern Hemisphere (and Asian) Grass biome. These
~3: g ~:;;j range restricted species, e .g., the genus Caesalpinia
-=tZ ~ 11'1 GI sens. strict. has a worldwide distribution, but most biomes (Fig. 15) are described in detail below.
-='06:5!
.
~ ~~ ~CU ~ ~';
RI ._ .C
species are essentially limited to dry tropical areas in
A Succulent (S-) biome, comprising a semi-arid, non-
1111
°' c ......
0
~el
Cl.I .....
c ......
... .e
North America (twelve species in the Caribbean and
fire-adapted (or non-resistant and intolerant to fire),
·= three species in Mexico to Central America) with two
":I cc Iii~ ~z- succu lent-rich and grass-poor, dry tropical forest,
LI.I
c:s:
..."'g:, -.c Gt
t........£ :90 ~btl
species in South America , and in Africa (four species
thicket and bushland biome (Zonoecotone II/ III and
LI.I .E
in north-east Africa and Arabia, five species in southern
s~:t; Zonobiome III [Breckle, 2002) and highlighted as the
-
c
0
U')
ci.
ci.
.
Cll
Cll
·;:;
"'3:
~o
~t~;
() ·-
.c z .><
g~-g~
....
Africa, one species in Madagascar and one species in
central Africa).
Many legume clades have distinctive geographical
red area in Fig. 15). Pennington et al. (2000b; 2004)
refer to Neotropical Succulent regions as seasonally
0 ... ~"' ta .. dry tropical forests (SDTF's). The S-biome is the
~ -:-si~
:e ... o:i: and ecological phylogenetic structure, where stmcture
-:e
Cll
!;Oz ~~ Cll 0 ~
·-- smallest and most fragmented of the four biomes (Fig.
Cll
::I°:~ ~ refers to geography or ecology predicting phylogenetic
"'E
0
:;;$ "'~ .... Cl.I
rel~tedness among terminal taxa. Caesalpinia sens.
17) and is prone to bimodal or erratic rainfall patterns,
i ,.... ~:;:; ~ z
2 ::I .., .c
.... Cll and legume subclades inhabiting it have a northern
a:ri c - .,, strict. was thus attributed to a global distribution
i ..J 0 bn range - but with predictable Southern Hemisphere
.
•• ::I

."'
.,; ,....-..J a> c
0.W.c·-
8!~g
pattern suggestive of a Su ccu lent biome which
occurrences (as seen below) - characterised by a
I LI.I
<
.!!
·=
.... Ev
"g
Cllt'O
QI
~&f·;;;Q,I
-:

~11~
... a>~
occupies the red areas of the map (Fig. 15) and may
represent the remnants of a persistent dry vegetation
predominance of amphi-Atlantic disjunct taxa. Legume
I LI.I ~ ci. S:,c pe that was Lin k •c.i , in th· T "rtia1y, from ·irn 1n1-
genera in Succulent biome clades are listed in Table 5,
c ci. ~o~e

-
I 0-z " ...
Cll
Cll
'iii
"'3:
,...o
co~
-12
QI 3:
...
.!! .!:!, .g
Q
~.
1; - c
.&J
11)'1:1
m a~.om.m . ·s UL I1 Am •rk:-1 through .emni l meri a,
Me~ko and th Ca ril h >an, along lh , Telhys eaw:1y Lo
which identifies c. 26 amphi-Atlantic, c. 12 pantropical
and nine Africa -Madagascar -Asia disjunctions occurring
:Q ... ~ ... 0 ...
c ClJ •n;og

I -.....
0..
...
Cll
·;::
!!
v
,.... 3:
~o
~:ii! E ~
cu"'G>.C
::: ~ £
... .c ....
·i
fn :i, rahi ~1 an I Indi a (F.ig. L ). In Africa Lilis
~x t·cn cI ·c.If10 · 111 11! · H >J'll sou rhw:.i r ls lo southern AJJica
among and within genera. No tropical New World-
Asian disjunctions are found among S-biome taxa,
N~ u 0 uf althou gh seven pantemperate or temperate North
• . •Sll •h •s - I) 10
Q.I

~ Q
Cll
co~ ~ ·~ -s g alld
• Ma hga •se1r · 111 " t:ix a ar " o fre n sister lo
American-Asian disjunctions occur among and within
LI.I ."'
Cll
"'3:
~o
g ·om~
·-E :it•• ·-c ·-c
lades• conv.ttnmg
Bn Zmol)tome
1 ·
· · n ombinrn io n f -hiom · taxa (i.e.
LI , hut •xdu ling Z noec.:ownc IT/ ITT·'
the S- and TN -biome genera Cercis, Gymnocladus-
~ -0
·e ...
N~
~
Sl,~"'""
"""' E
~8~8 r · ·I' 1 • .. OO~) and H-b10 . 11 .. taxa (Figs. - J ). • Gleditsia and Cladrastis-Styphnolobium.
v"' 'O ~ ~ ~
"' >--"' ::I 0 f All g 'n ·ra cm1I 11 a. s1 ·gne <l to al I ·ast n >I' these The present-day S-biome encompasses regions in:
~;~o
"'~... bn 0 0 ...
pourl glo . lY::i I I11 · 0111 ·s on lb I a:-. is or lia ing a a) the Neotropics: semi-arid tropical to subtropical
.
c 000
3: o.c u; .5
>i: GI.,,
.C·-"O GI ofr t. nm man ·c.· 0 S(· ec~·1 . o curnng
nurse l .1
· the re. M·1ny g n rn Mexico , Central America and the Caribbean
• " es 'CC nc..b 1 (and ::;ome, te rtb1y ccnlr s,'
.!! LI\ '{fl.
CL .~_!!.~
s l"I
"Cl 0 u -
Cll :it ... gi
~ . (particularly the Greater Antilles; Wolfe, 1975; Lavin et
Ill
!E ~ ~ .!!! Pl ar ·ntly · · ' ..tal e ti wit. 11 subseque nt diversifi ·~ui o ns
.el
Cll '° Q..ru ... >
E g' .2. ..?'
into . ·is.-;( al., 2001b), linked frequently to circum-Amazonian
~ ~ :0
iii '-'
suppn ighhouring biom "s. This hyrmh ·:;is is al l a.'H 'Pleistocenic Arc' dry forest elements in South America
... nrtcd wile·, . :
i
;;;:.
(V'\....:
0 a,._
.! !GI l ~ll l in , . re gen n · P11ylog •n1es Mr ·· avail:,1hl for
g. St1 h s · ·o n la r Ii ·rs in ·a ti o ns hav
including the inter-Andean valleys of Ecuador and
Peru , the Piedmont area of north western Argentina
0
II..
w:l'.i~~

BIOGEOGRAPHY OF THE LEGUMINOSAE 23

22 LEGUMES OF THE WORLD
and central Bolivia, the Misiones region of north
eastern Argentina and adjacent Paraguay, and the
Caatinga of eastern Brazil (Prado & Gibbs, 1993;
Pennington et al., 2000b; 2004). The Neotropical centre
of S-biome clade distributions is thus mainly North and
Central America and the Caribbean with distinctive and
predictable South American occurrences. Neotropical
centres are linked (with intervening fossil evidence
from Tertiary tropical North America and Europe
[Herendeen et al., 1992]) across to:
b) the Old World, in the S-biome Somalia-Masai
regional centre of endemism (White, 1983) of the
Horn of Africa, with various dry forest and thicket
'arid corridor' disjunctions and extensions through to
the Nama-Karoo, Succulent Karoo, Desert and Thicket
biomes of southern and south western Africa
(Verdcourt, 1969; De Winter, 1971; Rutherford &
Westfall, 1986; Thulin, 1994; Low & Rebelo, 1996;
J\.irgens, 1997; Thulin & Lavin, 2001; Van Wyk &
These genera were thus previously placed in three
separate caesalpinioid tribes with no suggestion of CAESALPINIOIDEAE 1
any close relationships between them. The legume
phylogeny has thus not only revealed relationships FIG.4
between genera that were previously widely separated
taxonomically, but has elucidated new associations CERCIDEAE
supported by previously unforeseen morphological
synapomorphies. In this case, dioecy was found to
occur in all but two genera in the clade, and monoecy
is considered to be a reversal in Arcoa and Umtiza
(Herendeen et al., 2003b). In addition, these novel
geographical patterns of association of genera are
frequently repeated across the phylogeny, e.g., the
Umtiza clade pattern is largely repeated in the Cercis,
Schotia and Poeppigia clades, each sister to large
Rainforest and Grass biome diversifications comprising
the rest of tribes Cercideae, Detarieae and Cassieae
DETARIEAE
subtribe Dialiinae respectively (Fig. 3). Most of the (CAESALPINIOIDEAE 2;
Umtiza clade genera are narrowly distributed and Fig. 5)

Smith, 2001); western Madagascar (Leroy, 1978; comprise one to a few species inhabiting S-biome
Schatz, 1996; Jansa et at., 1999; Lavin et at., 2000; Du vegetation, such as the monotypic Arcoa from the
Puy et al., 2002; Schrire et al., 2003) and Arabia to Dominican Republic, and its sister genus
West Asia and north-west India (Quezel, 1978; White Tetrapterocarpon, with two species from Madagascar
& Leonard, 1991; Miller & Cope, 1996; Klirschner, (Fig. 18). The reiterated amphi-Atlantic disjunctions s TN GR
1998; Conti et al., 2002; Cortes-Burns et al., 2004). within the Umtiza clade are consistent with an original
The Nubo-Sindian local centre of endemism, or distribution of this lineage along the margins of the S TN (G R)
subzone 3 of the Saharo-Sindian Regional Zone Tethys Seaway (Fig. 16). Similar north-south
(White & Leonard, 1991) is a critical part of this disjunctions to those between Gleditsia and
distribution as it is a floristic continuation of the Gymnocladus and the semi-arid southern African
Somalia-Masai regional centre of endemism into Asia genus Umtiza are also repeated in Cercis and
(Fig. 15). The remaining two subzon s (Sahara and Adenolohus (Cercicleae), Prosopis and Xerocladia
Arabia) comprise mixed S- and T-1io m ~ elements and (Mimoseae) and outside the legumes, Ephedra and CAESALPINIOIDEAE 2
are thus coloured yellow (Fig. 15), but their main Welwitschia in the Gnetopsicla. Ickert-Bond &
affinity is to the S-biome. Wojciechowski (2004) resolve a w 11 s upported FIG. S
The Umtiza clade (Herendeen et al., 2003b), sister Welwitschia-Gnetum clade sister to E'pbedrn. A further
and basally branching in the large Caesalpinieae sens. north-south disjunction is noted between mainly
lat. plus Mimosoideae clade (Fig. 3) exemplifies a northern temperate Gleditsia and its outlying southern
Succulent biome distribution (Fig. 18). The previous Brazilian and Argentinian species, Gleditsia
tribal affinities of the various elements of the Umtiza amorphoides Taub. The ITS analysis of Schnabel et al.
clade (Polhill & Raven , 1981; Polhill, 1994) were : (2003) reveals this species as sister to the rest of the
Gleditsia-Gymnocladus (from eastern Asia and North extant northern temperate lineages of Gleditsia. The
America), and Acrocarpus (south east Asia) placed affinity of this species is to the S-biome Misiones and
basally in two separate groups in tribe Caesalpinieae; Piedmont nuclei of 'Pleistocenic Arc' vegetation
-----1 DETAR I EA E1-----1
Arcoa (Caribbean) and Tetrapterocarpon (Madagascar) (Prado, pers. comm .). ··--···········-·····-·····················-··-··-········-···········--·-····-··1
in the more derived Dimorphandra group of tribe Disjunctions in the S-biome occur in both species-
Caesalpinieae; Ceratonia (north-east Africa and poor basally branching lineages such as the Cercis,
Mediterranean) in its own subtribe in tribe Cassieae Schotia, Poeppigia and Umtiza clades, and Basal
and Umtiza (South Africa) in the Cynometra group of Papilionoideae (including Cladrastis), or in speciose
tribe Detarieae. clades nested high in the tree. Speciose sister clacles
I
FIGS 4 - 14 Taxon· blome supertree. with major legume subclades representing a more exhaustive sampling of various local subclades than were used
=
In the matK phylogeny. Colour-coded biomes (as in flg. 15) were superimposed on the supertree: S = Succulent; G Grass; R "' Rainforest; TN =
Temperate Northern Hemisphere; TS= Temperate Southern Hemisphere. At= Africa ; Mad = Madagascar; As= Asia; Aus = Australia; Pac= Pacific: C/N [R)
"' Central or North America & Caribbean; S Am = South America; Eur/Med= Eurasia & Mediterranean; Pan= Pantropltal. The ratio's above each group
~the proportion of Rainforest (k-) to Grass (G·) blome species In the group if R/G, or vice versa If G/R. Clades are named, e.g. DET 1 = Detarieae sens.
strict. clade 1, according to their position within the major subclades. Many nodes are also Identified by the biomes (placed In boxes with arrows
pointing at the critical nodes to which they refer) as optimised In the DIVA analysis of Schrlre et al. (2005). The blomes are colour coded as In Rg. 15,
and are listed In decreasing order of occurrence among possible optimisations In the analysis, I.e. the first area Is either optimised most frequently or
at least no less so than the others. Biomes In square brackets= the only optimisation found; biomes underlined= present in all optimisations, and biomes
in round brackets = present in only one of a number of optimisations

24 LEGUMES OF THE WORLD

BIOGEOGRAPHY OF THE LEGUMINOSAE 25
CAESALPINIOIDEAE 3 MIMOSOIDEAE 1
FIG. 6 FIG.8
Parkia +
Piptadenia grp.
CASSIEAE p.p. (DIALllNAE) (1 SAm+Af+
Mad+ As+ Pac;
5 S Am; 1C/N + IM11 J
SAm)
MIMOSEAE
(c. 16: 4)
Gf R

Labichea,
Storckiella
grp. (1 Pac+ (19)
Aus; 2 Aus)
G
IE:l (31: 7) Neptunia,
Prosopidastrum, Leucaena
DETARIEAE G/ R
iI CAESALPI NI EAE s. /.
Piptadenlopsis,
Mimozyganthus
grp. (2 C/N
+ S Am; 1 As
Entada + Pipta· (25Am; 1C/N+ +Pac; 1 Pac)
CASSI I NAE
! MIMOSOIDEAE
deniastrum grps.
(2 Af; 1 Af + Mad
+As (1 sp. C/N +
SAm; 1 C/N +S
Am +As+ Aus)
PAPILIONOIDEAE SAm)
The position of Dlnizia,
Duparquetia is
equivocal I Pentaclethra,
Aubrevillea (1
Af; 1S Am;1
l-·- - -·-····- ···-··· -,·- -················-···· Af+C/N+S I~
i Arn)

[S]

CAESALPINIOIDEAE 4 MIMOSOlDEAE MIMOSOIDEAE 2 (1050)

FIG. 7 G
FIG.9

Abarem~ Chloroleu· Pithecellob· Acacia

!~~~~=~
(2
ium Alliance subgen.
Alliance (1 S con Alliance
CAESALPINIEAE s./. (25Am; 1 (3 C/N; 2 Phyllodineae

__
Am; 1 C/N +
Core Peltophorum (1 Aus+As
SAm;lAs+ C/N+SAm; C/N+SAm; C/N; 2 C/N +
grp. (2 AF+ Mad; 2 1 As) SAm) +Mad) _,
,__
Aus) 1 As+ Aus)
Mad; 15 Am; 2 C/N
(1 unpubl.); 1 C/N + I~ l 1NG10 J l tNGll l
SAm+Af; 1As+Af
+SAm)
PAPILIONOIDEAE genera.
· ~-···-----~ ----·- -----------~---....__--.-----.,
Possibly in
(See Fig. 10) Caes 4 to 7
(4 SAm)
INGEAEj t "'" I
i

(48: 13) ~ ACACIEAE i j

G/ R i
!!
Umtiza, Gleditsia,
Arcoa , Ceratonia
Poincianella,
Caesalpinia s.s. grp. (1
C/N ; 2 C/N + 5 Am; 2
Mezoneuron,
Pterolobium grp.
(1 5 Am; 1 C/ N +
Hoffmannseggia,
Libidibia grp. (1
C/N; 2 C/N + 5
Molden·
hawera grp.
I
C/N +S Am+ AF; 1 C/N SAm; lAs+Aus Am; 4 5 Am; 1 As) (1 SAm)
grp. (2 C/N temp.
+ 5 Am+ Af·Mad +As; +Pac +AF+ Mad;
+ W + E As (1
with 1 spin 5
2Af) 1 Pan; 1 As-AF) l CAES 51
Am); 1 C/N; I
Mad; 1 Af; 1 Af-
Eur-Med; 1 As)

! CAES t l

G IMOSEAE I

BIOGEOGRAPHY OF THE LEGUMINOSAE 27

26 LEGUMESOFTHEWORLD
PAPILIONOIDEAE 1 pAPILIONOIDEAE 3

(S9) flG.12
TH
(1 0)
(10) G

G
Thermopsid- Genisteae (18 Eu"Med; 3
Af; 1 Af +Mad + Eur-Med;
~ MIRBELIOIDS INDIGO- MILLETTIOIDS s.s.
eae (1 C/N
sens. lat. FEREAE
~--
l·---·············---·-···-··--·-·-1
Sophora s.s. grp. & 1 Eur-Med+ temp. C/N +
temp.; 1 C/N
Eu chresteae (4 Eurasia; 5Am;25Am)
temp.+ E Asi
1 As+ temp. As; 1 As+
3 Eur-Me d)
Aus+ Pac +temp. S Am
+ Af (coastal)

(21)
G
Diocleinae (2 C/N;
1 C/N temp.; 1
SWARTZIEAE C/N + SAm; 45 Ophrest·
Am; 1 SAm +As; iinae (1 As;
2 As; 1 C/N + S 1Af;1Af+
Am+Af+As+ As)
Aus; 1 C/N + S Am
+ Pac+ As+ Mad)

PHASEOLOIDS

IHALO GA LEG INA I

IDALBERGIOIDS I
GENISTOI DS/DALBERG IOI DS

PAPILIONOIDEAE 4
FIG.13 (80: J)
PAPI LIONOI DEAE 2 G/ R
Glycininae (4 Af; 7 As; 2 Af +As; 1 Af +As Phaseolinae (7 Af; 1 Af +Mad;
FIG.11 + C/N + S Am; 1 C/N temp. + E As; 1 As+ 1 Af +Mad+ As; 2 Af +As; 1
Aus; 1 As+ Mad; 2 C/N + S Am; 1 C/N Aus; 1 Pan; 2 C/N + S Am; 1
~ '---~-------' C/N +As; 3 S Am; 2 C/N)

. . .~-----------11
. DALBERGIOI DS s.I. 1-1- - -- --------•.- ..__..,,..___~~~--------~~~--' ~
'-------r-------.1~
Cajaninae . .- - - , - . . . l [S G]
-..~---------"[ DALBERGIOID CLADE li------------~)lo~ L--···- -··· · -·-·---- --·-----· ~- .. --... ········-·· .. ······-·· ·--------·-·1
PHASEOLOIDS I I l
!
ADESMIA
CLADE ~
PTEROCARPUS CLADE -----1 DALBERGIA CLADE 1--- ---1•
)lo

!I
(61:
44) i
G/ R Desmodieae
(197: 185) Desmodium grp. (7 As;
(91) 1 C/N temp.+ E As; 1 j
G/R
As +Aus; 3As+Af;1
Chapmannia,
G As+ Mad; 1Af;1 Pan;
I
(246) (114) Styiosanthes Daibergia, 1 Af +As+ Aus) iI
clade (1 C/N + S Machaerium grp. Ae schynomcne,
1S 6(15) Am+Af+ Mad+ (1SAm;1 C/N + S Humularia grp. (1 S Am Diphysa clade Clitorilnae (1 !
As; 1 C/N + Af; Am; 1 C/N + S Am 1 C/N + SAm +Af+ Mad (2 C/N; 1 Af; 2 S Am; 2 C/N + I
Adesmia clade (3 S Am; 1C/N;4 SAm) + Af +Mad+ As) +As; 2As+Af;1 Af+
Mad; 2 Af; 1 As)
Mad; 1 Af+
Mad +As)
S Am; 1Af;1
C/N + S Am+
i
1C/N;1 C/N + SAm; 1
IP181 ~ i
~ ~
Af +Mad+ As
~
Amorpheae C/N +SAm +Af+ Mad
+Aus)
(1 C/N temp.; +As+Aus)
5 C/N; 2 C/N
+SAm)

~ ILLETTIOIDS S.S. I
lGENISTOIDS/VATAIREOIDS I BAPHIOIDS
HOLOGALEGINA

BIOGEOGRAPHY OF THE LEGUMINOSAE 29

28 LEGUMESOFTHEWORLD
 PAPILIONOIDEAE 5
FIG.14
Wet
Boreotropical

IRLC
ROBINIOIDS

(27)
TN

(16:14)
G/ R

Callerya grp. Galegeae (8 As; 5 Hedysareae (5 Cicereae, Wet

(4 As; 1 As+
Aus; 1 C/N
Eur-Med; 1 Eur-
Med + C/N temp.;
As ; 3 Eur-Med ;
2 Eur-Med+ As;
Parochetus,
Galega (2
Austrotropical
temp. +EAs; 1 Eur-Med + C/N 1 Eur-Med +As Eur-Med+
1 Eur-Med+ temp.+ S Am; 2 + C/N temp; As+Af; lAs
Aus+ C/N Af; 5 Aus) lAf+As) +Af))
temp.+ S Am
temp.)

s (, R TNI
lndigofereae; Millettioids/Phaseoloids

flG. 16 Palaeocene map reconstrud.ing the Tethys Seaway, based on Scotese (2001). A seasonally dry to arid tropical belt apparently became Tethyan-
wlde only in the Tertiary, and covered much of the S-blome regions in CAmerica, N Africa and Arabia to W Asia. This belt Is sandwiched between two ba.nds
of wetter tropical climate (Scotese, l.c.), the boreotroplcs to the north and the austrotropics of the Southern Hemisphere and SE Asia to the south. This
arid belt is noted by Scotese (I.e.) to expand northwards into N America, the Mediterranean region and W to C As ia in the Neogene and then it ls
squeezed southwards again in the late Neogene by expanding T-biome dlmates

with ± equivalently sized amphi-Atlantic diversifications
(where each is well supported as well as the two
together being well supported), reflects a pattern of
reciprocal monophyly (Cunningham & Collins, 1998;
Lavin et al., 2001b). Examples of genera (or sections
within them) showing this pattern of continentally
disjunct and genetically disparate sister clades include
Baubinia sens. strict.; Caesalpinia sens . strict.;
Chamaecrista and Senna; Mimosa and Acacia subgenus
Acacia; the Ormocarpum clade, Chapmannia and
~yloscm1b 'Si Sesba nia , Lh L' /11 rltgvfera :ml clades,
1t:/Jbi-ost 1, IJ'1y1hrl11a , and tJ1e \l(r1,t1c1-Pl t1s >o/us clad ·
lrable ')). R •lativ I few amp l1i- tJanli · di ·jun ·rions
c ur in Pf'l'dc min:intJy R- and G-biome clacles (e.g.,
th e 1Iym ·m1~a- >u ibo urtia-Peltogyne clacle in the
D 'tari •a ·ee bc lowl).
J [
th. Vicarian e. nalysi. (Sci 1rir el al. , 200')) um1ers ·o re ·
b'e pred minance or su b 1~1 des assigned lO rhi.! ._
1 111
t thm are s islcr (and f)LIL:.Hivdy h:1~a lly bra n hing)
,. ·~o · ub hcl ' ·s ,.1ss · 1·gn , l 1 to Lu
1.
' R- anLI - I11
·0 111.cs.
c.Xni11J)I ·s 0 1· S I I
Ln , _· • _ ., 0~ 1 chides ·howi ng Lh is r 1 ~11i nsllip
FIG. 15 World map with four generalised legume distribution patterns at the biome level: Red =the Succulent (S-) biome; Brown= the Grass (G-) biome; s 11 c1 · lh e r ·1s.-Ad no lo l us grou1 (Fig. ·i ) th
Green= the Rainforest (R-) biome and Blue= the Temperate (TN-) and (TS-) blomes · )Olia -13arn ,, d ci '
Y ' n run group (Fig. "\ , the Po pplgi:1-
Baudouinia group (Fig. 6), the Umtiza clade and
Pe llo j h< ru m group fi'ig. 7 , rJ1 Faic.lhe rb la-Z·1pot ·a
g roup Fig. 9), Lil ' Leld.1 Styph no lobium lad rnsLi ·,
a nd Lh J Acosm i11 m- D I ·r:-1 or ·talum grou ps ( Fig . .LO),
tri h Am rph a J Pig. I l ) and l ri h · Tn ligofer ac (fi g.
12J. Tiu· 11 g h ut the fa 111il , run 1:1111 •ma l :wi.tches in
biome pr •f ·ren , • occur bet\ e n 'arly hra n ·I tin g
li.n · :1g 'S an I th ir siste r grou r .' a rpea ring l() drlv,
diversification across the phylogeny.
A Grass (G-) biome, comprising a fire-adapted (i.e.
prone to, a nd t >lerant of or reslst;.i nt to nre), succulent-
! >r and gnss-iich, seaso na lly dry tro p ical rorcst,
w clland and savanna (w ><Jed grasslan d ) biom
(Zonobiome II excluding Zonoecotone II/ III [Breckle,
2002], and highlighted in brown in Fig. 15). The G-
biome is the second largest of the four biomes (Fig. 17)
and generally has a unimodal rainfall pattern. Genera
often also occupy th e R-biome (Zonobiome I),
sugge sting that dispersal or habitat s w it ·!ting is
common between th ese two, whose I o u n J aries
fluctuate constantly in response to climatic change.
Distribution centres a re confined largely to the

30 LEGUMESOFTHEWORLD B\OGEOGRAPHY OF THE LEGUMINOSAE 31


GRASS Biome
18%
TEMPERATE
Bio me
41%
SUCCULENT Biome
6%
FIG. 17 Relative sizes of the four biomes (and areas falling outside these) as a percentage of the total land surface of Earth. Biomes colour-
coded as l n Fig. 15
seasonally dty tropics of the southern continents and
Asia. The Neotropical G-biome is referred to as
savannas by Pennington et al. (2000b; 2004).
The main area of G-biome clade distributions in the
Neotropics is in South America, in seasonally dry forest
(but not the 'Pleistocenic Arc' SDTF's sensu Pennington
et al., 2000b; 2004), woodland (cerrado), savanna and
grassland environments. In the Old World similar
habitats predominate in the Sudanian and Zambezian
regional centres of endemism (Savanna and Grassland
biomes sensu Rutherford & Westfall, 1986) in Africa, to
the monsoon forest and more open vegetation types
RAINFOREST Biome
Miscellaneous
7%
in Madagascar and tropical Asia to Australia . Clades are
distributed more commonly right across the tropics in
G/R-biomes and Table 5 lists c. 26 pantropical, c. 17
New World -Asia (i.e. excluding Africa), c. 37 Africa-
Madagascar-Asia, and c. 15 amphi-Atlantic disjunctions
occurring within or between genera in G- and R-biome
clad es.
Data for generic disjunctions is combined here for
G- and R-biomes since G- and R-biome clades are
closely associated. Rainforest biome taxa often occupy
gallery or riverine forests that are part of a mosaic
within the G-biome vegetation (Figs. 4-14).
'fbe re ·en! ages of 1nany R- div rsific:it:ions (Jlarris
_ooo: Richardson el 1/., -< 0 I ; Pcnn.ington >I ol.
Ill (/ I., . ) 11 .d
A· uivin ct al. 111 pr s · , as we as ev1 enc
.ZO 11• , r·, 111 ., analys s t..hat su ·h wcl ·la I s re cl ri e I
1
rhC: '1 '
. VIdi·y d ·11.J..:s Figs. /i-J. 4 ), ::.ugg scs th:it the R-hl >m,
from
, • •en co• l nis d more r ' · ' ntJ I I 'gum s than th ·
11
, .,, 0111 "'.S. Su rpn• Ing Iy re ·em 'Pl
b,ts .
~ e1stoc ' ne) ages 1f
1
:.~, 11 Uvcr. ill .1tions hav I 'en proposed for R-biome
ta ·a ·uch as N.uprecbllll Cl'olyg na cac; !' ndr!' 1 . 20~J4
~ind f//,W1 (Hic h~u:cl· on et cil., 2001~· Th 1 nterd1g1t~n n
f IZ- ~ind r-lor SL blom . fov urs a ' refuge '
Inc ·rpret:11ion of a liopatric di "rg n " CPra n ·e, ·1973;
p 'nnington t!I ctl. 1 200 ) ·aus cl I y Lhl! expansion ~ind
·ontl~l Ci 11 r \.Vet and dry-adapt " I veg l:Jlion. An lire/
tliermls (W.WrighL) I . sul sp. inermi'i o curs in rain
for SL anti woo I.eel gni:;sland in both , ·o uth Ameri ·a
nnc.l /\fricn (Penn ington, 200. ), and sp ·i s pain; in
Lop/JircJ (~apota ne :ind An ·/w111 aue · (Araceae) oi1
th· W s L J\fri "al1 R/ -biom " b undary ar bar ly
dilT 1" nliat J, and sp :imen · ·a n h • c.Liffi ·ult ro id ntify
iLhOlll h,1bit:it data Lo k pers. on1111 .). Ta:xl-l ac.lapt ~ I
to both G- and R-biomes may often be pioneer species
which can take advantage most effectively of post-
disturbance conditions, i.e. in competing for light gaps
in wet forests and resprouting or reseeding rapidly
after fire in savanna environments.
The S-biome lacks the R-biome affinity of the G-
biome (Figs. 4-14). For example, the S-biome does not
include terminal taxa like Lonchocarpus and allies,
which comprise many constituent species and close
relatives from the G-biome forests of South America.
Many of the North American constituents , however,
are R-biome species and indeed , North American rain
forests differ from thos e in South America by a
preponderance of Lonchocarpus species (Sousa &
Delgado, 1993; Wendt, 1993). Sousa & Delgado (1993)
noted that the wet for es ts in Mesoamerica are
floristically different from those in South America, and
Gentiy (1982) emphasised the fundamental difference
between the Tertia1y floras of South America and North
America, thus underscoring the demarcation of the
biomes circumscribed here .
A Rainforest (R-) biome (Zo nobiome 1), with
distribution centres confined to the equatorial tropic5
(wet forests) worldwide, is the second smallest of the
four biomes (Fig. 17). Many legume taxa occurring
within the R-biome, however, are found in edaphically
dry substrates, e.g., Amazonian South America is a
complex mosaic including drier savannas and Campina
forests on white sands; the Atlantic forests of eastern
Brazil comprise large areas of dty Tabuleiro forests ; the
Guineo-Congolian and Swahelian wet forest regions of
Africa also include extensive areas of drier forest as do
much of Inda-China and north-eastern India. Recent
molecular analyses show that most R- and G-biome
legume clades, in contrast to those in the S-biome
comprise elements that are widely scattered across the
tropics, without the relatively narrow and predictable
areas of distribution of the S-biome clades, except in
a general sense of being restricted to one or other
continental region.

Gymnocladus Acrocarpus
r ASll\ N AM EIWA li'!lll;\, SE AS IA
6 spp. 1 sp.

Gleditsia Tetrapterocarpon Ceratonia

Umtiza E /\SIA N AMf l<. lt A MADAGASCAR NE AFR ICA Tropical AS IA
S AMERICA 2 spp. M l ll l ll· l~ I U\Nl /\ 1 1
S AFRICA
Arco a 2 spp.
1 sp. c. 13 spp.
CAR IBBEAN
1 sp.

S AMERICA AFRICA
FIG 19 p
FIG . 18 Umtiza clade after Herendeen et al. (2003b) with clade ages superimposed after Lavin et al. (2004; in press). Distributions colour-coded to biome · alaeogene se d"
e •ng of Leguminosae from a putatively Tethys Seaway (Lauraslan) origin
as in Fig.15

BIOGEOGRAPHY OF THE LEGUM INOSAE 33

32 LEGUMES OF THE WORLD
 In the R- and G-biome clades, the predominant
pattern is that elements within higher level taxa are
more closely related to each other across the entire
equatorial tropics. A very brge percentage of lower-
level taxa, however, are more closely related to
elements restricted to only one of three continental
r -•gio ns. For • ample, 1> upra -g •ncric grc up: ~ u d1 ~JS
Lill! Copaif rn-:u1 I m1 -T ssmannia clacle in. D ·taJ:ieae
(Fig. 5) or th, Dialiurn dad· in Di aliinae (fiig. 6) or th ~
rrnosi:i lad e in Ba s~d P:-ipilionold ae (Fig. 10),
contain genera that are sister to each other across the
Lro pics, however, c 2 percent or g fl ra J11 !fo~ Derarieae,
3 1995). An original Tethyan-wide distribution is thu~
89 p r · nL Ln I i::tlli nae, a n I 95 per enL iJ Basal
Paril ionoi I ·ae ar relit ricLcd Ln elLh ·r Lh NeoLr pi ·s,
Afrlc<1-Madagascar or Asia-Pa ·ific-Austnil ia (Sdui.r ~ el
ttl. , 005; T:.ihle ). Th e D "tarl ae , Dialiinae and Ba. ·d
3 between-genus (i.e. intergeneric) - and consequently
Papilionoicle<t · main lh high 'St pr p rtions of R-
hiOl'ne species ti. ·. , ~7% >n av rage) ·ompar · d with ,
endemism arising through subsequent diversification
into the R/G-biomes of the individual southern
continents and tropical Asia . When fossil evidence is
available for taxa in the R-biome, this too supports a
Tethyan (i.e. Northern Hemisphere) link existing
between them in the Tertia1y (despite there being no
extant northern taxa). Examples include the amphi-
Atlantic Swartzia-Bobgunnia clade in Papilionoideae
and the reciprocally monophyletic amphi-Atlanti~
Hymenaea-Guibourtia clade in Detarieae (Herendeen
1992; Graham, 1992; Axelrod, 1992; Lavin & Sousa'
also hypothesis e d to account for the current
distributions of R- and G-biome taxa.
The R/G-clades thus have high percentages of
low proportions of within-genus (intrageneric) _ g e'N g g g c'N
disjunctions. The largely G/R- clades show lower .. .
... ... ...c
c
N

"'
c ......
"'
N

.g., aesalpin i e~1 ~ and .M imosea (having t11~my more
-bio m ~ s p ··i ·s ith o nl y 12% <tn I l % H-hiom
·1pdes r i1p ·c[ivdy. The l::i tter two Lrihes 1!~1ve 7'5%
rn1 I 76% or gen ra reslri t d tu un ly o ne o f th ab v ··
continental regions respectively (Table 4). Clades in R-
and G-biome areas, therefore, show a high degree of
interrelatedness across the southern continents and
tropical Asia, despite the high levels of generic
endemism within each continental region and the long
percentages of genera restricted to any one
continental region and this is correlated to higher
numbers of intrageneric disjunctions. Examples of
pantropical R- and G-biome intrageneric disjunctions
(Table 5) include Copaifera, Cynometra, Parkia,
OcJ//Je r.~lr1, Pterocarpus and D • modium, while
inle rgc n .. ric pantropical disjunclions o ·cur between,
for exa mrle, JJr/Of'l..Gl ( ~entral Ail1 rica), lh ' 0 .. )!Stig ma
group (Africa and f iugiode 11.dro11
1-Jroch'lg, uesia CSouL b i\me ri a), A.fz e/ir;1 AJri 'a -Asia)
si a ; -I

I
I I f • o
.............. -... ....... .._
• • o ' '
........
periods during which these landmasses have been
isolated from each other. This pattern is reflected in the ~1 nd l n.lsin ( A:> ia); or Platycyamus ( outh i\n1 erica) $ ~ -:;- 8

predominance of pantropical or New Worl I- Asian

(rather than biased to amphi-Atlantic) c.lisjun '. Lion.
and the Old World millettioids. There are also many
examples of Neotropical-south-east Asian disjunctions,
.!:>
~ --- - s
"'
.-<
~

DO
N
...
"
/'-.'

within these clades. such as Androca~ymma (South America) and the

Early-diverging legume cl a des (c. 60 - 55 Ma)
apparently became distributed across the Tethys
Seaway which was subject, at least in part, to a
seasonally dry tropical climate throughout much of the
Tertiary (Fig. 16; Scotese, 2001). Such elemc.:nts are
K11 ompc1ss ic1 - lr1./Jl chea groups (Asia-Australia);
Leucaena (Neolropics) and Schleinitzia (Asia-Pacific);
G'oj 1fu1-Zyg ia (N mro pi s) and the A 1·cb/dB11dro n
group ( Asil1-/\usrra li a); w ithin Ormosio (N olroi !cs· ...
,:;;,
A s i ~t & Au stralia) ; an d Lhc Tipuana g r up South "'"'
N

thus considered likely to have seeded norrh wards into
the wetter boreotropics and southwards into seasonally
dry vegetation types emerging in South America,
eastern to southern Africa and Madagascar, and Asia
(Fig. 19). This h poLh esis d •rives from th h.igh d 'gre '
of intergeneric rel al ~dn ess foun I across th e 1.ropi ·s ,
and also account for Lh ~ high lev ~ l s of ge r ri ·
America) and lnocarpus (Asia). Amphi-Atlantic
lis jun('Lions oc ' Ur h ' l We n , ror ins tance FjJe/'11.C/
(. ' outh Ameri ·a ~ind '/. nwnocof •11s-Eurypetulum
(A fri ca l; Di y tnfJe Sout h Arn ·rl ·al and che
JJo fyst monan.tb11s group ( Africa) ; D/11 izit1 (Smith
Arn ·ri ·:1) and Aubr,villoc1 (Afr i «t) , anti withi n
Pentaclethra (Neotropics-Africa).

TABLE 4 (RIGHT). Statistics of genera and species in Leguminosae. No. (%) =number and percentages of genera and species, or 0
of inter-continental disjunctions. An additional four genera to the main treatment of the book are recognised here (i.e., 731 vs. ...... ,:;;,

727 genera), representing the putative segregates of Acacia. Numbers in bold highlight the highest percentages and numbers <t
"'00
N

of genera and species across the table. Gen/spec= genera/ species; the four continental regions used are Af-Mad =Africa·
Madagascar; As/Pac/ Aus= (tropical) Asia-Pacific-Australia; New World= Neotropics and temperate North and South America;
Eur/Med= (temperate) Eurasia-Mediterranean (Including Africa north of the Sahara)-Macaronesla. lntergeneric disjunctions
are noted when sister genera, within phylogenetically supported clades, are distributed exclusively In different inter-continental
regions (i.e., one genus does not have endemic species in the same region as the other). lntragenerlc disjunctions are noted
when genera are distributed in more than one Inter-continental region (I.e. they have endemic species in two or more continental
=
regions). [SJ = Succulent biome; [G) Grass biome; (R] = Rainforest blome, and (T) = Temperate biome. * ·= this figu re is
artificially high with S-blome genera being scored as though all species occurred in the S-biome, thus further researd1 Is
needed to derive more accurate figures; in general those removed from the 5-biome will be transferred to the G-blome. For the
total number of tropical/subtropical dryland species add figures (percentages) in the S· and G-biome columns. Crown age refers
to the crown clade of a higher taxon, i.e. the age when the diversification of extant species occurred, as seen on a tree where
a single lineage begins to branch

BIOGEOGRAPHY OF THE LEGUMINOSAE 35

34 LEGUMES OF THE WORLD
A Temperate <T-l hiome, with distribution centres
co nfin e d to th e Mediterranean, warm and cold
temp e rate reg ions of the Northern and Southern
Hemisphe res, linked directly through mo ntane tropical
regio ns , cove rs the largest area of the four biomes (Fig.
17). Temperate clades are principally either Northern
TN- or Southern TS- Hemisph e re in distribution.
mirbe li o ids to the hulk of Papiliono ideae l. Temper~ile
TS- cbd es have thus largely reoccupie d tropical tir
montane tropica l areas , where~ts this is rare in TN-
diversifica li o ns .
I
The TS-biorne, South African mediterranean lrihe z
,_
HypoGdypteae, is moderately supported in a clacle Vl
with the warm te mperate to subtropical sderopliyll
~

Northe rn Hemisphere TN- tax a gene rally comprise commu nity, TS-biome , Australian tribes Mirheli eae :tncl 'CJ
<:: ·~
e
elements derived directly from S- or G-biome clades Bossiaeeae <Figs. 1 & 3). This group of tribes, referred i::"
13
<:: E' ~e .sc:
Q ;g 2Q -<: ~.,Q

(Figs. 4 -1 4). Southern Hemisphe re TS- taxa , e.g., the to he re as the Mirhelioids se11s. lat., have a crown cbde -<:
_, -~ <:; "' ~Q e-Q §..0..
'-"
Aclesmia, Hypocalyptus, Mirbelieae- Boss iaeea e, age of c. 55 ± 1.1 Ma (la In l!I al. in pr ss), indicating ..:
u
"'0 <::
<:: - E .~
.~
a surprisingly old <lisjun ·Lion l tw e n so uthe rn Africa a:0 <»
"'"' ;t cs ":'
Podalyrieae and p oss ibly Pso raleeae clades, however, 8
~2 8
~~E

are themselves principally sister fo nd often basally and Au:-;Lralia for this grou p of I 3gum .. A 111.inimal bO ,_
z"'
!:; -!:!2
c: '-'
b c:c: "'
Q; <»
..: ·- <»u b .!:: ·- Q.

bran c hing) to the rest of their clacle components,

although the phylogeny indicates that they all have S-
Ma age (based on fossil evide nce) is reported hy Linder
et al. (2003) for the crown clade of African and
a.

----
~

\.9
Vl
l9
-.
I

Vl-<:
NU
,!::
Q

Q
E
I I
I
l9
N
l!)
-.
Vl -.
.-<
I.!)

""C:-
-
Q
..E
..-< ""
or G-biorne outgroups. Australian Restionaceae . The last opportunity for dirl:ct
Legume clades with a tempe rate distribution are ove rland dispersal b lWe n these regions was c. 160-
3

130 Ma . In the early TC!lt(a1y, however, the south Indian

"'..:u
lis te d in Table 5. Five Asian-New World , nine Vl
..:
pantemperate Cmontane tropical) , two Africa-Asia and O c an was fragmen L cJ by rnd o -Matl a gas ·ar th e
3
;I! ~
0 Vl
three :unphi-Atlantic disjunctions occur in temperate Nin ry-East Rid •e and th Kergu Jle n Plateau, providing ..: <l:
st pping-st me route for subseque nt disp rsal b tw ' •n ~I
biome clades (excluding the S- and TN-biome ..:
~
Africa and Australia.
disjunctio ns noted under the S-biome). Tropical S- and
G-biome clades have e ach dive rsified inclep e nckntly An S-biome area invo lvin g the Sa haro-Sinclbn-
w est-A ·ian reg io ns t WhiL & L ' ornml, 199 .L) is o l"tcn
"'
u..
<l: --
\.9
~

into temperate regions resulting in distinctive TN- and

TS- centres, partic ularly in papilionoicl legumes (Figs. sisL r .i nd puwliv ly I asa ll y I ran ·hlng) to major TN-
0

4-14). Predominantly TS- clades more often have Euraslan ·rncl M ' cl it rran<.:}111 clades, which may h:1ve
secondary diversifications back into the tro pical G- di s jun c t' :rnb ·lacl · s ln th l mp ra t N w World.
0

biome (e.g., the Adesmia clacle in the dalbergioids and

Crotalarieae) but this is rare ly the case in
Exarnpl · lncluue Lote<.1e of the robinloi<.l l · gumcs 11 11cl
Lh IRL Ce.g., Lrib s al egeae, H d ysa r a , Ci r<.>: te,
--
..: ..:
v;
<l: ~
:J
z
----
I-
\.9
Vl
TriJi iii ·•1e <In I 1"~11> a . A group of J e rlved , Jnis t<Jicls
f--
I Vl
predominantly TN-biome cbdes (e.g., a few species of 0
_, <l:
::>

Sopbora sens. strict.). Schrire et al. (200')) resolved the includ ing the l'oclalyriea , rotalarieae anJ Gen isl :ie °'--
Ou
two temperate areas in a basal polytomy , revea ling have th ir h ttsal ly bran ·hJng Jame n Ls in w a rm 5~
3: u
that tempe rate-in ha biting clad es are often re late d to temperar s uLhcrn Afri ·a, alth ough th S-hl me z rf_
each other, but they are inconsistently related to the A ·osmium-Di ·rne petalun dade Is a [>UtaLiv ' outgro up
or this a ll ian ·e. M mb1.;rs of th ·s g •nistoi I groups
other three biomes.
Temperate distribution patterns include a subtropical, ::ipp 'ar to hav ~ eithe r migrate I northwards through
mo nwn ' trop l :a l Africa to Lh Mee.lit " rran a n ,111 d
--
(.!)
~

rnecliterranean o r temper:lte North American to northern

Mexican distribution, some times linked to the Andes
and warm tem perate South America (e.g., some
Amorpheae and Adesmieae in the clalbergioic.ls ancl
various tribes in the Inverted Re peat Lacking Clade
[mLC], e.g., Galegeae and Faheae). Warm tempe rate
South African distributions are often linked through
m o ntane tropic al Africa to th e Me dite rranean and
M:.icaron •siHn regi o n (e .g ., l.o l o n onis ;1n d
Argyrolol;iu.m and ·uhs qucntly to the ew 'oriel
(e.g. , Lupf.uus), o r t hnv sc · n laril.y invad ·d tl1
[ropics (e.g. (.'rolctlttria). Th , 13rongniarti ··ae hav , ii
N >OLropic:al disLribuLl n, disjunct to Lr pi al a nd warm
t ·mpcrate Australia . Within Ps ra l · a , 1h '1tl/eu -
Bil11mi11ctrht a nd ~w \1 o rld Psora l ~ ·a div rsifl 'itcions
3

Ma caron es ia , for example , th e ge nisto id tribes clc>s ~l y trH ·k Lh · d.istril utional palt rn ob:»rved in 1lte ~
Vl
vi
Crotalarieae and Genisteae and the phaseoloic.I tribe L ) t ~1 , a llh< ugh Cullen has a n xte n s iv cJ fi eel
J
3
.!;:!
.s
Psoraleeae. Southern, central and eastern European Austra lian liver ifi nLion .rim s, Jl 7). The puw1j e ly -<:

distributions are commonly linked to montane western basnlly branching so uLh rn Afri ·an tbo/11bimn :i nd ~Vi's"'
c:"'
Psora/ea lim.ag s, howev r, rei t rat ·1 -g ~
to eastern Asia, or cold temperate central and north- z - c:
t: g
in LI ge nJsLo ld tril e · rotalari :1
--
I
eastern Asia (e.g., tribes in the IRLC). Australasian I-
V'I ~ U)

di stributio ns ma y be la rg ely e nd em ic (e.g., th e ( i rim s, 1 · 0) . Th ~ · uri an ~1 ·ea :ire a I utati v~ Vl .-< <O: .-<

mirb e lioic.l s) o r linked to north ern patte rns (e .g ., ( utgrm1p or Lhe I gumes (see page 45), and ha ' a
Psora leeae) . All TN-hiome clacles are derived from S- disjunct listril l!Lion in Centra l a n I orth Ame rict1 ::i nd
or G-hi ome clades <Pigs. 4 -1 4 ), e.g., the Sophora Aus tralla ( myn et 611., 19 5 , · hich is similar to 1h:1t f _,
u.J
<(
ci.
Oii
"'
"'
c:"'
:~ :~
0.
Lhe Brongnia niea ~ in Lb gen ist lei subdade .
Vl
group including The rmopsid eae, and the mLC tribes ,
best illustrate this pattern. Most TS- dades, however, Id ·r d ive rsWcalio ns in th · TN- r g io ns bas cJ o n
"'
z
w
z
u.J
0
0
"'
·~
..c: -
l9 c:
Vl -
l9 c:
Vl
u"'
- Vl
ro -

Cl> Cl>

~~
u
z ~ ~.g -. 0
"'"'y
th rate:'> ;u1a l ys~s or Lavin. '/ u./., 2004 ; in press) are
0::: ·~ Cl>

are sister or basally bran ching to other clac.les (e.g ., 0

u
f--
U c::
_, Cl> Cl>"' -ro
:~ ~
,z "' ~~vi %~Vl >
Adesmia, sist e r to the rest of the clalbergioids; ~1s o ·iat d wi th bas;dly I. ran i ing ·la I s in th ·' fo1nil · "' ::> :z <( ,"! ·c c: ro0. bJ) .--
w -
f-- Vl
0
x
VJ
u.J lg -Q)
"' ~
Cl>
~ Qj "'
Cl> 0~
E
Vl.
e-
Poda ly ri eae to the d er ived ge nistoid s, and the Th s · in ·lutle taxa now restride I Lo warm L ·mpcrate ~a ~
<(
u
OJ
Cl Cl 'O
.2::
0 :0"' > u
ro
Q.. M

36 LEGUMES OF THE WORLD BIOGEOGRAPHY OF THE LEGUMINOSAE 37

IN
00 MIMOSOIDEAE
r-
rn 1 R/G - inter 2 R/G - intra 1 R/G - intra (Entada)
Gl Mimoseae [Pentaclethra,
c (Pentaclethra-Aubrevil/ea) (fldenanthera; Xylia)
Adenanthera, Entada,
srn Newtonia grps.] 1 R/G - intra (Pentaclethra)
<.fl
1 G - inter (Leucaena- 1 S/G - intra
.,,
0 Mimoseae [Plathymenia, 3 5 - inter (Plathymenia-
Schleinitzia) (Dichrostachys)
-I Prosopis, Neptunia, lndopiptadenia;
I 1 G- intra (Neptunia)
rn Leucaena, Dichrostachys Prosopis-Xerocladia;
~ grps.] Calliandropsis-
0 Dichrostachys grp.)
Al
r- 1 S - intra (Prosopis)
0

1 S - inter (Mimosa- 1 R/G intra (Parkia)

Mimoseae [Piptadenia
& Mimosa grps.] Adenopodia)
2 S - intra (Mimosa;
Adenopodia)

1 S - intra (Acacia 1 G - intra (Acacia subgen .

Acacieae
subgen. Acacia) Aculeiferum)

lngeae [Faidherbia-Zapoteca 2 5 - inter (Faidherbia-

-Calliandra-Viguieranthus Zapoteca; Calliandra-
grps.] Viguieranthus)

3 R/G - inter (Cojoba 1 G - inter (Lysiloma 1 G - intra (Acacia 1 R/G - intra (Albizia)
lngeae [Inga - Old World
lngeae -Albizia- grp. -Archidendron; -Acacia subgen. subgen. Phyllodineae)
Enterolobium-Abarema grps.] Abarema grp.- Phyllodineae)
Pararchidendron;
Chloroleucon grp.-
Cathormion)

lngeae [Pithecellobium- 1 B/S- inter

Samanea-Acacia subgen. (Pithecellobium grp.
Phyllodineae grps.] -Thailentadopsis)

PAPILIONOIDEAE

Basal Papilionoideae (Pl - PS) 1 S - inter (Cordy/a- 3 R/G - inter (Bobgunnia- 1 R/G - inter (Alexa- 2 S/TN - intra
Aldina grp.) Swartzia; Amphimas- Castanospermum) (Styphno/obium;
Myroxylon grp.; Cladrastis)
Mildbraediodendron-
Amburana grp.)
1 R/G - intraspecific (flndira)

Genistoids [Basal Genistoids 1 R/G - inter (Ormosia- 1 G/R- intra

(P6 & P7)] Haplormasia [Africa]) (Pericopsis)
1 R/G - intra (Ormosia)

Genistoids [±tropical groups 1 S - inter (flcosmium- 1 S/G-inter

(PS & P9)] Dicraeopetalum grp.) (Brongniartia-Hovea)

1 TN - intra (Lupin us) 1 G/T - intra (Sophora) 1 G - intra (Rothia) 1 G - intra (Crotalaria)
Genistoids [±temperate
groups (P10- P14)] 1 TN- intra
(Thermapsis)

Dalbeigioids{Adesmia clade (P16)] 1 S/G- intra (lomia)

Dalbergioids [Pterocarpus 2 S - intra (Chapmannia; 1 R/G-inter 1 R/G - intra (Pterocarpus)

clade (Pl 7 - Pl 9) J Stylosanthes) (fnocarpus-Tipuana)

Dalbergioids [Dalbergia 1 S - intra (Aeschynomene) 1 G - inter 3 G- intra (Smithia;

clade (P20 - P22)] 1 S - inter (Diphysa- (Soemmeringia- Cyclocarpa; Ormocarpum)
Zygocarpum) Cyc/ocarpa) 1 G - inter (Geissaspis-
Bryaspis)

Baphioids (P23) 3 R/G - intra (Dalhousiea;

Airyantha; Bowringia)

Millettioids 1 [lndigofereae (P25)] 1 S - intra (Cyamopsis) 2 S/G - intra (fndigofera)

Millettioids 2 [Basal millettioids 1 R/G - intra (Aganope)

& phaseoloids (P26) J

Millettioids 2 [Abreae (P27)] 1 S/G - intra (Abrus)

Millettioids 2 [Dioclein ae (P28)] 1 R/G - intra (Dioclea) 2 G - intra (Canava/ia; Ga/actia)

Millettioids 2 [Ophrestii nae (P29)] I

1 G - intra (Ophrestia)

Millettioids 2 [Core-Millettieae
(P30 - 34)] I
1 G- inter (Piscidia-
Philenoptera) I 2 R/G - intra (Derris;
Millettia)
1 S/G- intra (Tephrosia)

Millettioids 3 [Phaseoloids - I 1 G/R- intra (Clitoria)

Clitoriinae (P35)] I
Millettioids 3 [Phaseoloids - 3 TN - intra (Apios; 5 G - intra (Pseudarthria; 2 intra; 1 G/R- (Mucuna)
Apios - Desmodiea e grps. Lespedeza; Hylodesmum) Uraria; Pycnospora; 1 G - (Desmodium)
(P36-41)] A/ysicarpus;
I Leptodesmia)

I ~
Millettioids 3 [Phaseoloids - 4 intra; 3 G- 3 S/G - intra (Erythrina;
basal core-Phaseoleae (Psophocarpus; Flemingia; Rhynchosia; Eriosema)
r
(P42 - P44)] Cajanus) 1 S - (Paraca/yx)

Millettioids 3 [Phaseoloids - 1 TS- intra (? 1 G/R- intra 9 intra; 5 G- (Neonotonia; 2 intra; 1 G - (Teramnus) 1 TN- intra
Glycininae-Phaseolinae Otho/obium) (Oxyrhynchus) Dumasia; Strongylodon; 1 S/G- (Vigna) (/lmphicarpaea)
(P45 - 47)] Nesphostylis; Lab/ab)
4 S/G - (Cullen; Wajira;
I
I
I Macrotyloma; Dolichos) I
Robinioids [Sesbanieae (P48)] I 1 S/G - intra (Sesbania)

Robinioids [Loteae (P49)]

I
1 TN/TS- intra (Ornithopus)

I
I I
1 S/G/T - intra (Lotus) 1 TN - inter (New World
Loteae genera -
Eurasian/Med. genera)

IRLC [Millettioid IRLC (PSl)] 1 TN - intra (Wisteria) I 1 TN - intra (Glycyrrhiza)

IRLC [Galegeae (P52)]

IRLC [Hedysareae (P53)]

I
I
I I
2 T - intra (flstraga/us;
Oxytropis)

1 TN - intra (Hedysarum)

I RLC [Trifolieae-Fabeae
(P54- P56)] L L L L 1 TN - intra (Parochetus) 3 T - intra (Trifolium;
Lathyrus; Vicia)
or subtropical North America and eastern Asia, linked
often to mesophytic forest refugia such as those in
west Asia (i.e. the Caucasus), the Himalayas in southern
Asia, and China. Examples of taxa with this distribution
type are Cercis (Cercideae; Davis et al., 2002b),
Gleditsia and Gymnocladus (Caesalpinieae) and
Styphnolohium and Cladrastis (Basal Papilionoideae).
Such older no1thern temperate diversifications probably
reflect the northward expansions of the Tethys Seaway
taxa into the boreotropics (Fig. 16), and their relictual
distributions now occur within the TN-biome, which
became superimposed over these areas in the late
Neogene (and are thus discussed under the S-biome).
More recent genistoid and IRLC temperate legume
diversifications (again based on the rates analyses of
Lavin et al., 2004; in press), however, appear to have
occurred within co-existing temperate conditions.
Legume Biogeography
Many of the amphi-Atlantic disjunctions in the legume
family lie within the S-biome of the Neotropics and
Africa to adjacent Asia, a pattern consistent with a
historical legacy of a once widespread Tethyan
distribution. The Arcto-Tertiary Geoflora hypothesis
(Chaney, 1947) and the Boreotropics hypothesis per se
(as developed by Wolfe, 1975 and Tiffney, 1985a & b)
are rejected here in favour of a more narrowly defined
Tethys Seaway explanation, despite attempts to
broaden the original concept (e.g., Lavin & Luckow,
1993). A once widespread Tethyan distribution
involving seasonally dry tropical vegetation is different
from the Boreotropics hypothesis, which was erected
to explain Tertiary similarities in wetter boreotropical
floras of the mid-latitude Northern Hemisphere (Fig.
16). The Madrean-Tethyan hypothesis (Axelrod, 1975;
Fritsch, 1996; Liston, 1997; Hileman et al., 2001) is
also rejected here because it refers specifically to
subtropical Tertiary sclerophyllous vegetation types in
North America and Eurasia giving rise to similarities in
the vegetation now seen between the western North
American ('Madrean') and the Mediterranean basin
('Tethys') regions. Axelrod (1975) places this band of
laurophyllous, sclerophyllous vegetation to the north
of the Tertiary tropical vegetation, which is the focus
of our seasonally dry tropical Tethys Seaway. The
Madrean-Tethyan areas also fall within our TN-biome
and are marginal to (and probably derived from) the
S-biome areas distinctive of the Te thys Seaway.
Present-day disjunctions between the California and
Mediterranean sclerophyllous regions , and possible
linking taxa used to exemplify the hypothesis, are
often either rejecte d as being similar by convergence
(e.g., Wolfe, 1975) or to have crown ages too young
to be explained by such vicariant events (e.g., Fritsch,
1996; Lavin et al. , 2004; in press). The Tethys Seaway
flora thus evolved within a seasonally dry tropical
climatic belt (Scotese, 2001), sandwiched between the
40 LEGUMESOFTHEWORLD
wet tropics of the southern continents and south-east
Asia (austrotropics), and the boreotropics to the north
(Fig. 16).
Good evidence of the long association of legumes
with seasonally dry areas comes from fossil sites rich
in legumes (e.g. , Herendeen et al., 1992; Herendeen
2001), as well as the presence of a tropical seasonal
climate (Scotese, 2001) and deciduous forests in and
around the margins of the Tethys Seaway at the same
period that legumes were initially diversifying (e.g.,
Upchurch & Wolfe , 1987; Cavagnetto & Anad<>n
1996). Many of the adaptations that distinguish
legumes are to a seasonally dry warm climate. These
include compound leaves, which photosynthesise
rapidly in favourable periods (and exhibit leaf
nyctinasty to promote its efficiency) while avoiding
excessive water loss through leaflet shed in
unfavourable periods (Runde!, 1989). The high
nitrogen metabolism of all legumes (Sprent, 2001)
may confer a competitive edge in colonising
seasonally dry environments because leaves can be
produced economically and opportunistically in
unpredictable climates (McKey, 1994) . Deciduous
leaves with a short life span influence interactions
with herbivores, because rapid leaf turnover allows
phenological escape from herbivores and pathogens
(McKey, 1994 and references therein). Diverse and
mobile chemical defences in legumes (Janzen, 1981),
such as alkaloids and other small molecules, can be
shifted from senescing leaves to other organs and
from seeds to seedlings, unlike non-mobile defences
such as tannins and fibres. Seed adaptations (Gunn,
1981; Van Staden et al., 1989) include hard testas,
long dormancy, long viability and the ability to store
nitrogen in seeds, permitting rapid seedling
development. Ant associations such as extrafloral
nectaries, pearl bodies, beltian bodies (in Acacia),
hollow stems and thorns, have arisen numerous times
and confer considerable competitive advantage in
legumes (J. nzen , 1981; McKey, 1989), although such
ant associalions and mobile chemical defences would
be advantageous within a range of habitat types . The
widespread occurrence of wind dispersal by winged
fruits and seeds, especially samaras (Augspurger,
1989), attest to the long association between legumes
and open environments.
Trends toward increasing humidity started in the
early Palaeocene and continued into the Eocene,
rea hing maxi m~t in rh · Early Eo e n 55 - 50 NW·
. <.w in, I 77; Wo lrc, 1( 85; Jani s, 1993). Much of
sou1h ' rn J..aurasl<i w as cov r cl by mega the rm;:d 20"C
+ l I o reolr pi a tl •vergr n rn lnfor ·ts w ith humid
su btropica l to w;1rm tem per~le c.lt::dtluoLJS mes )the1111al
1 " - 20" forests e xtending to 65u (Axelro I, I 92 ·
Southwa rds, a belt of seasonally diy tropical climate
sti· ~ ngLh e ned between the wetter tropical areas in the
r the Soutllern
n rrh anti the austrorr pi 'S
H misphe re and SE Asia (Fig. ·1 >) , ~u1 I th is cu rred
consiste nUy across the latlrud s or much of the: TetJ 1ys
rom U1 La te Pala oce ne w Lh e lioc ne
'
· 0 awa Yf
_ ?O()j) Jn ti
. .
Mrdcll e Eocene c. 50- . Ma),
( · ·01es . .. ·
0
. .
h mc;gathL!rma l T thyan llor:.1 I e um 111creas111gly
1
, anally dr (WoJfe & p ·hur ·h, 1987; avagn tc &
st:.ts , 19 > ;:in I Ieg~rn • I'rv rge(I rart'd iy d unng . lineages for the other biomes. This pattern is fractal
M d6 1 •
I I . p r!od. Ax Ir tl Cl. . nor d th t.1 l n :w r
1
~ 1;pcr'tll.lll'eS flu tu::necJ 'OQSLU ra J ly, Wil h tn:Jj r shifts
·cun·ing hetv e n seasonall y dry ·.in I humid d l~1aL ·s
0
und ·u h dlnl,~lli · puJs · resulted rn lhe pans10~1 of
lry tr pia d for~ · 1 and _wo dland by 49- 8 Ma , 1TIJ.Xed
de ·ic.Juou ha rcl \l oo I loreslS by 7 - 6 Ma, scleropbyll
wo c.llnn I hy 3 - 35 Ma :inc.I thorn fo rest by 0 - 29
M.a. rr as' e propose tb _. w SL Gondwana byp Lhes ls
is cU · (> 1nled an I th e r · is n< ··vid en e o f an
all er 1.roplc.:a l origin for the fami ly, LI e n ciisp n;al of
I menls s n11.Liwnrds in to I olh the dri ~r and w ll r
l!~p!cs of 1he outhcrn cominents is ax iom:ni (Fig.
I • Th rn >Sl likely inrerprc tntion , th ~refo r , of ea rly
I ran hing c la I s In th R-1 io m of th outh rn
continents and Asia, is that they are derived from
seasonally d1y tropical Tethyan elements that dispersed
there during the Palaeogene (65 -24 Ma).
During the Eocene (c. 56- 34 Ma), Tethyan legume
dispersal to South America and Africa became
increasingly possible as 'stepping stone' links appeared,
particularly across the Turkey -west Asia landmass
between Africa and Europe, which split the Tethys
Seaway (c. 50 Ma; Osborne & Benton, 1996). Global
cooling at the Eocene-Oligocene boundary (c. 34-33
Ma), lowered sea levels thus enhancing these links
Oanis, 1993). The Middle Miocene 05-14 Ma) saw a
rapid spread of drier climates in the tropics with the
widespread development of savannas and grasslands in
Africa and South America, a process which accelerated
at 7 - 5 Ma, the driest part of the Tertiary, and by 2.5 - 2
Ma, another major expansion of savannas and open
grasslands caused a large-scale retreat of forests and
woodlands (Axelrod, 1992; Janis, 1993; Maley, 1996).
Grass biome savannas and open environments thus
nppear to hav .. <l "v loped more recently than the
LI 'lrl .nt hiom · dq fo rest and thicket formations in
fri ·a and Soulh Ametica.
The persistence of the S-biome indicates that
remnants of early Tethyan legume floras should be
~ SL vident wi 1hln lhi s biome. Tn addition Lh
Linda.mental drough t tol 1~mc of the -l i me remained
un. •hano cl cJ ·
, . r; un ng 1a l N • gen ·' and u arre rnary
lrmmrc: ch:rnge <Axelrod , 1975· u ' zel, 1c78; Maley,
1980· M arr~· / I l
' .' tO e a·· · 8). Lineag - · o · ·upying •-a nd
l,. 1)tomes I 10 I
f : ' w v ·r iav I e n l ss p "rsistc:nl, · 1 re:ulL
O the higher W"l, . . •
af~ , " e r req utr 111 nl I Ihe. hiorn ·s being
J 992. I
ct cl mu h mor , l)y •!'
. c u11a ll . l,J L~iau.o n s !Lv:. Iroe.I ,
, 11
20() ): a;;~ ~'. :; ![.~ti Y, :9 ); Pern:t~gl m et al., 20 Ob·
1 et al., 2004; in press). The notable pattern detected in
biorne. . g c.tt 1 P rs rst n e of Im.cages in the S-
1 disperse back to the S-biome from the R- and G-
et ctl support ed by Lh .. vicariance tmalyses (Schrir
2
·• ..005JwhJr· tl1i··'t · · ·'aJ
lhc ha 1· ·
< ea is optums ' u · o ng many of
11
lh la )~anch ·s o[ the l •gum phylogeny, e ithe r a ·
bit1q1 U .
s e b!l me < J1
>r t e m s-r consist ' ntly o pllrnis I
' tgs. - I ). The.'>e a nalys s also : how 1hat the
S-biome is optimised in an intermediate position relative
to the other two pairs of more highly connected
biomes, i.e. G/ R- and TN/ TS-biomes, emphasising the
underpinning role of the S-biome as a source area of
.
and occurs repeatedly, suggesting it has been constantly
generated throughout the evolution of legumes, as
indicated across the legume phylogeny (Figs. 4-14).
The cladistic vicariance analyses all suggest that tropical
d1y areas have been occupied by legumes since their
inception, which counters Raven & Axelrod's 0974)
and Morley's (2000) assertions of a wet biome
austrotropical origin of legumes.
A biogeographic metacommunity
alternative
The four fundamental biomes identified here could
equally be viewed as the result of dispersal assembly,
where taxa with similar ecological preferences can, in
the given timeframe, disperse to similar ecological
settings worldwide. If closely related species tend to be
similar geographically and ecologically, as revealed by
the study of community phylogenetic structure (e.g.,
Webb, 2000; Webb et al., 2002), and if dispersal has
such enormous consequences over large spatial and
temporal scales (e.g., Hubbell, 2001), then geographical
or ecological structure observed in phylogenies of
globally distributed legume taxa may be the result not
of continental history but rather the product of
ecological drift playing out in immigration and resident
speciation (Hubbell, 2001). Ecological drift, similar to
genetic drift, is a random walk of individuals of any
taxon in a community through time; low immigration
rates into a biome result in endemic diversifications
suffering lower extinction rates and thus higher levels
of resident speciation (Lavin et al., 2004; in press).
Legume clades inhabiting the S-biome are
distinguished from those in the G-biome by a different
set of phylogenetically linked centres of endemism;
the former have range-restricted taxa mainly in the red
areas shown in Fig. 15, whereas centres of the latter
occur in the brown areas. The S-biome is the smallest
and most fragmented of the four biomes (covering c.
6% of the earth's surface compared to 12%, 18% and
41 % for the R-, G- and T-biomes respectively, Fig. 17),
and since there is a predominance of geographical and
ecological phylogenetic structure in clades endemic to
the S-biome, this suggests a connection between
phylogenetic structure and low immigration rates (Lavin
this analysis is that legume taxa are much less likely to
· biomes. This may be due in part to the relative
abundance of many R/G- habitats compared to the
relative scarcity of S-biome habitats, and the greater
aridity (i.e. drought tolerance) and often bimodal to
BIOGEOGRAPHY OF THE LEGUMINOSAE 41
erratic p;:i1tem of rnl nfall o f mi1ny ext:1nt S-biom ar as
as opposed to the mu ·h m r r 0 ul:lr and pr di ·La l I
unimod a l pattern in the R/G-hi o mes (\XlhiL ~. 1983;
Garcillan et al. , 2003). The great age of plant taxa
occupying the S- habitats (e.g., Pennington et al., 2004;
Lavin et al., in press), therefore, may not be clue just to
persistence of v ·•g1::tati >n and .omp nent lineages that
are adapted to unpre di table rainfall , but also to the
reduced rate of recent dispersal (i.e. immigration) into
this dry biome.
l is1 rsa l and pers iste nce of tax a ha ving a
predil e ·ti n for the S-biome is well •x mplified by
the ·o nUn nwl structur in the phyl oge ny of the
Umtiza clacle (Herendeen et al., 2003b) . Disjunct
Northern Hemisphere species of the once supposed
s iste r ge nera Cleclilsia and 1y mJ1 0 ·tadu · have b e n
xplain cl by Te rtiary interconllne nw l land I ridges
( ~ . g ., S ·hna be l et al. 2003 . N w Cl ,dl/. ·icl o f the
Northern Hemisphere is known to be sister to Umtiza
from an S-biome local region in South Africa. This
ecological pattern is matched by the one disjunct
South American species, Gleditsia amorphoides, which
inhabits S-biome local regions in the Piedmont and
Misiones centres of 'Pleistocenic Arc' vegetation
(Prado, pers. comm.). The Umtiza clade provides
another example with the Dominican Republic
endemic Arcoa, which is sister to the Madagascan
Tetrapt.e rocarpon. Historical migrations among
sue ule m-rl.ch, seasonally dry tropical forest, bushland
and thicket worldwide provide a more parsimonious
explanation of the global distril uLion of the Umtiza
clade, than invoking the extinctio ns of certain lineages
in puli ·ular regio ns of online nts , om or whi ch
w r pu ta tiv "ly o nnecte d by now submerged la nd
I ridg ··. , a ·h o r,, hi ·h has a di ff rent ag1:: l .avln et ell.,
2004; in press).
The Umtiza clade is also typical of clades confined
to, or centred in the S-biome in that it shows two
common biogeographic patterns that are otherwise
uncommon in the other biomes. One is that trans-
Atlantic clades predominate and often show reciprocal
monophyly, or a deep phylogenetic split se pa rating
Old and New World sister clacles . This ma y manifest
itself as continentally confined clades each showing a
large degree of phenotypic divergence. The second is
a lack of species that are distributed on both sides of
the Atlantic within the S-biome.
The pattern of reciprocal monophyly is also well
illustrated by the Ormocarpum clade and
Chapmannia (Lavin et al., 2000), and a clade including
Old World Vigna sens . strict. and New World
al., 2000). The pattern of reciprocal monophyly is also
reveal e d in clades occupying the mainland and
continental islands, e.g., the Greater Antillean robinioid
genera Pictetia (sister to Diphysa from Mesoamerica)
and Poitea (sister to Gliricidia from Mesoamerica;
Lavin et al. 200lb; 2003). This pattern is not evident
among clades occupying the mainland and oceanic
islands (e.g., Hawaiian Silverswords [Asteraceae];
Baldwin, 1997).
Regular dispersal of new taxa into local regions of
a global metacommunity (biome) means that a few
immigrants will establish over time , some will become
more common through ecological drift, and more or
less equivalent numbers will go extinct as a result of
this process (Hubbell, 2001). The lack of detected
reciprocal monophyly in most G- and R-biome clades
can be explained by high levels of immigration into
local regions of these global metacommunities
increasing resident extinction rates through ecological
drift, thus reducing species accumulation within trans-
oceanic sister clades. In S-biome clades, however, the
predilection of legume. to persist in this biome has not
resulted in an x Lin tion rate that eliminated the
pattern of reciprocal monopbyly. The build-up of
more or less equal sized trans-oceanic clades with
time, therefore, is likely to be the result of restricted
immigration (i.e., <lis p rsa l into local r g ions of this
biome and standing dlversily being gene rnLcx l mostly
by endemic speciation.
A manifestation of reduced immigration rates into
the fragmented local regions of th e global S-
metacommunity is that lineages of species separated
from sister clades for sufficient time come to occupy
continentally confined and widely disjunct (often
amphl- At la ntic trans-continental) distributions. The
Umtiza ·lad pattern is widely re peated in other
endemic taxa of the S-biome. For example, in the
Ormo ::i rpum cl a de, Diphysa is confined to
M iw a me rica, Pictetia to the Greater Antilles,
Zygocarpum to the Horn of Africa, Orm.ocarpopsis
and Peltiera to Madagascar, and Ormocarpum to
e a stern and central Africa , Madagascar with one
species widespread in Asi.H (Lavin et al. , 2000; Thulin
& Lavin, 2001). In ·o nLrast in R/ G- clades, although
intrageneric species diversity is to a gr ·a t r or lesser
extent continentally confined, rel a1Io ns between
g ·11 ra an d ofl en within larg g ~ n ra ) are
"ha ra t risec.I by e ithe r pa ntro pi ·al c.listrib1.1tio ns ( .g..
within Cyn o met ra, o r I cw n P r foritt [ entral
Am · ri .il, 0 o:tig nu1 [Afri ·al a n I K i11giod>ndron [ i;J I,
in l e wriea ), o r by Afri ·a-As ia d istribuLions (e.
( . e I 'ss c ngru •nl, na rr wly define<.! a nd
srrucwr · 1· · · . . . 1 ]' ·I · "'
. 11 ·tal le dlstrib11l1 ns) o mfM I <. L~ t 1~ • -
J
'?m .
P' t 'thin Lh , H/ li -bio m ·s, ·xp · n n ·ing lugh r
en •r:i 1 •
, 1i 11 r'll 'S thus · ;:1n o nl y b ass ign ~ c.I lo
I
imi n gr.1 . ' . ~ ... .
ti n •nL::i l-w1d R- OJ C1-.u e<1s.
t)~l'IW secon I · 111111 n bi ogcogrr1pLtic patte rn - the
l:i k , f widcl ulstri blll. d :- p ' i 'S '.n th~ '_-I iome ~
y LS with in t rco11t11P nta l sp ·c1cs d1s1nbutlo ns m
on11 .l~ . .
th , TN -, H- ::ind r-l iom •s. l' ''. m pl he re mclud ?
t ·/l'af~ci!ll · rilpimts L. , A . a m >ru.:am1 I fc ok.) M.E.
J oes, ;inti !1. a/Jorl~it~11111. L~i · h ·1 ~~s . ~ ~x .'p1"ng.
(B~i rnt::by , 19 , :Y.yt1 opl.~ a~ m~; s fn~ ( I.. ) D .,
iodo ar/ a A. ra 1 , and . u1s 1dt1 utt. Ba rneby,
1 52 , ~ u wl le pr ad in the T -bi me, and A ndin:i
1 1
lntJrm is, Lori bocmpus s>ri ell , and iVJa c;hue rtum
lnrtflll.tllZ whid1 va rimLo; ly inhabi t R/ -bio mc.:s in the
>btr pies ancl N "rl ·:1 (e.g., Thom , l l 72; Penningto n,
2003). Lists of i.n te r o nlin •nl'dly lisLrilx m.:cl sp c ·ies
uld in ludc o ·r l '.'i R- o r -hi >111 1 ·gum ' r.axa
dl tril \H ,d throug hou t the I ro pi ·al wd o · a ni · isla nds
of lb world ( Lavi.11 , u npubl ished l<1t<1 ); th s oft ' 11
being linked to the presence of resistant, lo ng-noa ting
seeds. Notably, many intercontinentally dis nibuted
species have infra-spe cific taxa confined to one
continent (or oceanic island) , suggesting tha t
immigration is ::in ongoing process that over time
manifests itself as phenotypically divergent
populations. The presence of widespread species in the
RIG- and TN-biomes which also comprise relatively
more recent diversifications than in the S-biome, again
points to the more current role of dispersal assembly
(i.e. species assembly through immigration) in the
former biomes.
The lack of species dis tribute d among distinct
regions of the S-biome can also be found intra-
continentally in South America (Prado & Gibbs, 1993;
Murphy & Lugo, 1995; Pennington et al., 2000b; 2003;
2004; Lewis et al., 2003a, Linares-Palomino et al., 2003;
Prado, 2003; Wood, 2003) and Africa (Verdcourt, 1969;
De Winter, 1971; Thulin, 1994; Jurgens, 1997). Jurgens
0997; his Figs. 2 and 4) illustrates two of the most
common Africa-wide disjunctions between the arid
north and south of Africa. His Fig. 2 represents an
t!ss ' utia lly S-hi<me Horn of fri -:.1 disjun t i .n w ith the
sou Lh-w st Africa n Ka roo- a mih regio nal e ntre o f
•nd rni m <White, l 98 a nd :ilso id nti.fi ed by Le brun
(lt '7) :ind Quezel 0978 > ·1s rh u·op i ·~ii r mi • nora
or 'R<111d nor::i '. Disjun ·Li on. h r ar comm n al th
slst r ·S() ~,... i cs, , s · ti.c n ~Hl d g ·nu s le e l, but f w
ed' ·nmp l · l1·1v" • .. I) en 1'd n 11·r·1 , d o 1· 111
. rra. p e ' 1·t··1 •
1
0 • IU n 'lions ·urring b twee n Lhes · regio ns (Th ulin
·•
1994 11 . •
e lement (e.g. , Nordenstam, 1974), is thus characterised
by disjunct taxa being found predominantly above the
species level. Widespread taxa have thus probably
de creased in species abundance and the resulting
refugia have been isolated from o ne another for at
least one to two million years , more than sufficient
time for speciation to occur (Cowling & Hilton Taylor,
1997; Hubbell, 2001).
The predominance of reciprocal monophyly, the
abundance of genera confine d to the same single
regions within continents , and the lack of
intercontinentally distributed species among legume
taxa centered within the S-biome regions of the world
are probably the result of one process: the contractio n
in size of the S-biome during recent geological times.
The S-biome has apparently been restricted in size as
a target area for sufficient time to reduce successful
immigration into local regions of this metacommunity.
Well delimited species or clades of species are thus
largely limited to old lineages. The drought tolerance
of the S-biome is probably the most important factor
limiting immigration , espe cially from other biome
elements that have a high water requirement.
While monophyletic clad es in legumes are often
confined to distinct continental regions, cladistic
vicariance studies show no consistent pattern of
continental relationships (e.g., Lavin & Sousa, 1995).
These authors considered that the phylogenetic and
biogeographical patterns of the tribe Robinieae are
seemingly as well explained by chance dispersal as
by the Boreotropics or Geoflora hypotheses. To
determine if vicariance or dispersal is more likely to
ex plain legume distributions across the Tethys Seaway,
age distributions of a range of trans-continental crown
clades were established using a Penalised Likelihood
rate-smoothed Bayesian likelihood analysis of cpDNA
m.atK or nrDNA ITS sequences (Lavin et al., 2004). Of
59 trans-oceanic clades of taxa ranked at the species
level and above, 51 have ages between 1 and 22 Ma
with the rest scattered through 30 to 50 Ma. The
majority of continentally confined clades have crowns
less than 10-15 Ma. Estimated ages of these clacles,
therefore, show a strong Neogene to Quarterna1y bias,
not older modes as would be predicted by continental
hi s tory. As the dates of these major peaks of
disjunctions correspond to when dry environments
were rapidly expanding, but post-elate large-scale
continental movements and the existence of putative
stepping-stone land-bridges, metacommunity processes
(sensu Hubbell, 2001) are most likely to explain these
disjunctions in legumes, rather than processes invoking
I

Phaseolinae (Lavin et al., unpublished data) . Even the many mille LLi ids a nd pins o lo i Is), o r N ·w orld- · ev r. Fig. of Jurgens C1 97) describes r1 singular historical continental events. Fossil and
m , <-r• ·n ~ r'J l nortJ)t:fll :ind souLhe
th
.
rn African d isjunclion
genus Wajira , largely confined to the Somalia-Masai As in distri butions e.g., w ithin Ormosia lg nisl ic.lsl, r mol e cular evidence sugg e sts that trans-oceanic
r gio n · ntc.:recl in the Horn f Africa, r ·pr s nls a b e tw ·n 'J 'i/mana [South Am •ri a l a nd J11 oc;c11pas . HI has . • 1<1 u1v rs1 1·a u. ns int
suhst·1nt·· I . 1· 'f' . LI1C . - I 1
.omc disjunctions in legume taxa arose neither at the same
ll(.1:.t n1an in tJ ' · 1r • •
[Asi a l in th dalb rgio lcls ), o r h y a n1phi -Atl an1ic l" 'fl' 1C 110111 :.t ncl Za ml 7..1:111 ( 111 lh SOUlh ) time nor under the same set of geophysical and
mo nophyl •lie group separated fro m ils loscsl isl r by -...('t .1onal · o 1• nd rnism (
·•rt' res
clistrihuli <> ns ( e.g., I c rw Jn Hy m ew.Jet1 a nd . climatic conditions. Rather, such disjunctions have
>Vt.: r 10 Ma Thulin et al., 200 ) . Th c.I · p : pJii Or tn frn .-;n " f 1
J
..
separating sister clades, resulting in the pattern Guiho11rlla ID •tariea I o r u •ttrtz ia and l3obg 11 m1ia $Lli... •1·t!U c <l isJlincl i >ns r·1nke d a l vari e Ly a n I multiple geographic origins, dispersion pathways and
I) , p ' ·1' I I a r . 1111110 11 h IW et.! 11 th e: · lW O
referred to as reciprocal monophyly, is a signature of [Swartzieae]) . Such a mphj-A1lantic distributions are . . q, CV divergence times.
.1 .as. Th SI .
long isolation of each of the sister lineages (Lavin et relatively few, and sho w I ss ontinental phylogenetic ' - 1 me Afric:.ui 'a ri I ·o rrid r' r Afro-arl I

BIOGEOGRAPHY OF THE LEGUMINOSAE 43

42 LEGUMES OF THE WORLD
Summarising discussion
To test the hypotheses presented above we framed
the following questions: what does the evidence of
legume distributions and the phylogeny of the family
say about a wet versus dry megathermal origin of
Leguminosae, and can the crown clade ages of a range
of trans-oceanic disjunctions between sister legume
taxa inform on whether vicariance (continental history)
or dispersal, is largely responsible for the existing
pattern of legume distributions? Four global biomes
were delimited (Schrire et al., 2005) which at the
broadest level have greatly increased understanding
of biogeographical patterns in the Leguminosae. A
northern (Laurasian) rather than austrotropical (West
Gondwana) hypothesis for the origin of Legumes is
supported both phylogenetically and by fossil
evidence, and this is consistent with a Tethys Seaway-
wide ancestral distribution.
The West Gondwana 'equatorial megathermal'
hypothesis for the origin of legumes (e.g., Raven &
Axelrod, 1974; Raven & Polhill, 1981; Morley, 2000) is
rejected here for the following reasons. The vicariance
analyses of Schrire et al. (2005) firmly place the semi-
arid S-biome as the only biome diagnosing the critical
nodes of diversification in the legume phylogeny. A dry
origin for legumes is in accordance with key
morphological synapomorphies and a high nitrogen
metabolism in the family. Lineages confined to the S-
biome gave rise to sublineages occupying all other
biomes and the Rainforest (R-) biome taxa are primarily
derived from dry Succulent (S-) and Grass ( G- biome
taxa . The essentially Tethys Seaway, S-biome (i.e.,
excluding the extensions into the Southern
Hemisphere) largely overlaps with the belt of
seasonally dry to arid tropical climate illustrated by
Scotese (2001), which links the Caribbean and Central
America with North Africa (including the Horn) and
extends to Asia. Scotese (I.e.) shows this palaeoclimatic
belt largely emerging to its full extent only at the
beginning of the Tertiary, when seasonally dry forest
is known to have occurred along the Tethys Seaway
(e.g., Upchurch & Wolfe, 1987). This also coincides
with the known spatial and temporal distribution of
legume fossils (Herendeen et al., 1992).
The rapid divergence of the family, as noted by the
chronogram data of Lavin et al. (in press), and the near
instantaneous Tethyan-wide representation of fossils of
almost all major taxa, is in accordance with a putatively
'sudden' opening up of a novel dry-adapted flora along
the Tethys Seaway in a post-Cretaceous extinction
environment. There is increasing evidence that many
major angiosperm families diversified during the
Tertia1y and do not extend back into the Cretaceous
(Tiffney, 1985b; Magallon & Sanderson, 2001; Davis et
al., 2002a). In the Tertiary, many extant families and
genera first appeared as part of a modern flora, spurred
on by coevolution with pollinators and dispersal agents.
This was a period of major change in global climates,
44 LEGUMES OFTHE WORLD
of significant mountain building, and a shift from warn1
and moist-equable regimes to seasonally dry and cool
climates (Wolfe, 1990; Axelrod, 1992; Brown &:
Lomolino, 1998). Distributions of largely amphi-Atlantic
disjunct S-hiome genera, together with super-generically
akin but panu· pi ·ally disjunct R- and G-biome genera
emphasise the Tethyan-wide nature of the early
diversification of legumes. This has been followed by
subsequent diversification occurring to a greater or
lesser extent within each of the three major continental
tropical regions. A number of R-biome diversifications
are now being shown to have remarkably recent origins
(e.g., Richardson et al., 2001; Pendry, 2004), supporting
the hypothesis of wet derived from dry being the
primary trend in legume evolution. Current data thus
point to a Tertiary origin of the Leguminosae along the
Tethys Seaway, with taxa probably originating in
restricted, edaphically dry sites and in terrains of diverse
relief (Axelrod, 1992).
The second aim was to establish if the existence of
closely related taxa in disjunct areas is largely clue to
separation of biota by the formation of natural barriers
(vicariance) or to the movement of organisms from
one area to another independently of other organisms
and continental history (dispersal). An argument can be
supported for vicariance as an explanation for the
present disjunct distributions of a number of
Palaeocene to Oligocene-aged clade diversifications in
legumes, e.g., most tribal and many super-generic-level
clades. Geological, physiographic and palaeoclimatic
changes over this period almost certainly resulted in
altering and breaking up the continuity of the Tethys
Seaway, as well as fragmenting the boreotropical and
Southern Hemisphere extensions of the S- and other
biomes. At the biome (metacommunity)-level,
however, the processes of ecological drift, immigration,
extinction, and resident speciation have played out
over such enormous spatial and temporal scales that
any historical signal of the Palaeogene is likely to have
been largely swamped if not obliterated altogether. We
have seen too that the majority of continentally
confined clades and trans-oceanic crown clades in
legumes are Miocene to Holocene in origin, thus much
of the present diversity in th · fam ily is Neogen ' in
age. This is loo recent for clisjun ·tions to be ex p lained
by o nlinenlal history, e.g. , l.a rge-s ale con lin •ntal
movements and the existence of putative stepping-
stone land-bridges. The present pattern of legume
distribution thus appears to be overwhelmingly the
result of global dispersal within and between
neigh! uring biomes . The historical legacy of these
melacommunities is one of dispersal assembly within
similar ecological settings world-wide, thus the biorne
concept as used here ta kes on extra significance for the
unde r.::rancl ing :ind im rpre tation of plant distribuLions.
Phyl og -•oles can b used in hist 1i -a l blog ogrrq hY
for cir ·ums ·ii ing gl< bal biom s, unde rsta nding 1h
relationships between them and estimating relative
rates of immigration within and among them.
systematic biogeography of
Legummosae
'I foJlo~ ing flC ·ount is a J Lailed :rnalysis o f UK:
~1: 11u uper1r e (Figs. ~14 , :1ssessing ~1. h major
d;de for crow n ag · L:ivm et al., 2004· in press),
iorne iifl1nity (S ir!r ,, ii. , 2 05 and the c.lii;Lrlbution
1
nern ~in d sp des :1 ·ross ·omin nLal rt:gi ns,
>r ·ng
g f l I . I.
l<lkI 11'·1rti ·u!ar n. le o JOl, incer- an. 111l'ragen · nc
disjun ·tions o ·currin.g :.im.<>1:g Lh ·. e reg1 ns (Ta ble 5).
·th s, lala rs1in1man ed 111 f<1ble 4 , br a k !own th
l1Llfllb r ·ind J r entages of •en ra pres ~m in - a11d
g n eru nntl sp >de~ e nd e mi ~o - rh
Mad agasc~1r, Asla-Pa ·di -/\u su-:-das1a, New World an I
Europc-MecliLeJ·r•1n ,a n regions c s how th majo r
li stril u tion patlerns w ithin leg um e ciadcs. Th
per 11rng of g n rn l e r clad r ·tri 'l ' d ro nly one
f ;ll1y ()f I h se fo ur regions is , pr c.I i 'Liv Sl ali, Li f
b.ion1 "' ·1ffinity, I ecause fi ures f 89% and above :11' •
all correlated to predominantly R-biome taxa, while
figures of 83% and below apply only to taxa from the
drier (S, G and T) biomes. Numbers and percentages
of inter- and intrageneric disjunctions per clade are
also correlated to biome. If 53% or more of all generic
disjunctions occur between genera (i.e. are
intergeneric) then taxa belong predominantly to the R-
biome, and if 63% or more of all disjunctions occur
within genera (i.e. are intrageneric) then taxa are
largely confined to the other three biomes. The
numbers and percentages of species with a
predilection for each of the four biomes are also given
as an indication of the biome affinity of each clade,
along with the biome optimisations noted at critical
nodes in the vicariance analyses of Schrire et al. (2005;
Figs. 4-14). The data in Table 4 are further
summarised for each of the three subfamilies and then
for the Leguminosae as a whole.
Current evidence (e.g. , Soltis et al., 2000) favours
the Surianaceae as the immediate outgroup of the
L gurninosac, a ll11 >ugh th r anu l s s sugg ·t
P Jygalacea :tnd Quill, ja ea ~.g., Cr~1yn et Cl!. 199'i;
J (
P rs .on 20(1 1; r r Sl, unpubl.). In ~ uda na e:-1e
1
lc~ ica~ g ~us R 'cCb ici is disj un t Lo a large ly
Aus1ral1 . an l1 \' e 1·s1·r·c·I <I t I' Orl, a ( 1·LS[rJ·1
) Ll[t. On f)artcrn
mn11nisc •nt of rl1 u_· . I .
·ri , ' : · pap 1 no1t tnbe Brongn ia rtieae .
· l Sunan·1 · ...... 1 •
. • ..... \sensu rayn •f r-1 / ., 19< 5 , ·o mprises
l I11 e spc ·le of I'
. \eccl.11&. 1 .10 Mexko, o n , in L11 R- and
two tn Lile I1 ·1 h pantropi al liLLCral sh rub
• ome., l
U l'/cllJtl II/Cl 11 ··t · ·
peur. . rni a, :inti In Au s tralia , Ccf. ch:l!lt1
/
ti ty is (a ti"e In th e G-1 lom seas nally dry
woodk1 ncl 'ind LI1. ·k . ·
oJ·s '. K et, lrom ue·nsl a n cD , two s p ~ ies
1vtobasm11 1 c·I 1 . 1
th' ,1· s lf1.I >. m cry Gir-biome s rubland to
le L on san I 1:.1
A\ . .
h
, om art ern Terril ')' to \J' stern
11.ilra 1ia) ·1nd G ·1r. / ·
I ie m , f. • lll'JO.Y za mo1wstylis a tree in th e H-
. )rt;i ~·
1 111
Quee nsland ). Crayn " f c1l. <19 5) nnc.J
Ce1 ~/el! · unpu l I. ~\lp po n R " cb ic1 b in' . ister to
· ta , .ind boll · only one of any of the three major inter-continental
llria11 . l ar S IS l e r LO rb e r S l o f th
a .ie. In Fig. • Re 'bict i. ·oded a:; an ·-1 !om
taxon, with one wet R- species, while the remaining
genera are largely restricted to the G-biome, including
one tropical wet R- species. In Surianaceae, the largely
disjunct S- and G-biome taxa, if similar in age to the
Brongniartia-Hovea clade in Brongniartieae (Lavin et
al., 2004; in press), could be no older than c. 10 Ma,
too recent for hypotheses other than long distance
dispersal to explain this disjunction.
·
Cercideae
The crown clade of the Leguminosae is c. 59.5 ± 0.2 Ma
(Lavin et al., in press; Table 4) and the Cercideae is
fri ca- putatively the first-branching tribe sister to the rest of
the family (Figs. 1 & 3), with Cercis the most basally
branching genus in the tribe (Forest, unpubl.) . In a
pattern repeated across the family among basally
branching groups, Cercis (Cerc 1; Fig. 4) has an extant
TN-biome distribution, i.e. North America to east Asia
closely associated with relictual S-biome areas, e.g.:
Adenolobus (Nama-Karoo biome, southern Africa),
Tylosema (Horn of Africa to southern Africa),
Brenierea (south-west Madagascar), Bauhinia sens.
strict. (Neotropics, Africa-Madagascar, south Asia).
Species in the latter three genera extend into the G-
biome. S-biome taxa account for 30% of the Cercideae.
It remains equivocal whether Grif(onia (Cerc 2; Fig. 4),
with three species in R- and one in G/R-biome tropical
west African centres, is allied to this S-biome alliance
or to the large, predominantly Asian and Soutl~
American R/G-biome diversification centred around
the genus Phanera (Cerc 3; Fig. 4). Gigasipbon with
two disjunct R-biome species in west and east Africa,
one in north-east Madagascar and three species, one R-
and two G -biome, in Malesia to Papua New Guinea,
may be more closely allied to the Asian Lysiphyllum
and hence to the Phanera alliance. New World-Old
World disjunctions are shown in Table 5.
The S-biome is the only area to occur in all four
optimisations at the basal node of Leguminosae (Schrire
et al., 2005), diagnosing the clade comprising Cercideae
sister to the rest of the family, with the TN-biome
' Lh , (reflecting the scoring of Cercis) also in three of the four
optimisations (Fig. 4). The G- and R-biomes are each
optimised only once at this node. The node diagnosing
tribe Cercicleae, however, is optimised by all four areas
more or less equally. The age of the Cercideae stem
clade is c. 56.5 ± 1.7 Ma, based on penalised likelihood
(PL) rate smoothing in the rates analyses of Lavin et al.
(in press); i.e., this is the earliest putative age of the
Cercideae lineage. The Cercideae crown clade,
however, marks the age of the diversification of extant
species (as seen where the lineage begins to branch)
and the PL age here is c. 23.4 ± 3 Ma. Since all genera
have not yet been sampled, the crown may be older but
probably not significantly so.
In the Cercideae, 58% of genera are restricted to
tropical regions, with 67% of genera present in the
BIOGEOGRAPHY OF THE LEGUMINOSAE 45
Asian-Australian region and 25% of genera (38% of
species) endemic to this region. This compares (Table
4) with levels of 33% generic (14% specific) endemism
in Africa-Madagascar (with 58% genera present in the
region) and no generic but 45% specific endemism in
the Neotropics (with 25% of genera present here). The
lack of generic endemism in the New World indicates
an Old World centre of diversity of the tribe. Four New
World-Old World and two Africa-Madagascar-Asia
disjunctions occur among 12 genera (Table 5) with
one (i.e., 17% of all disjunctions) being intergeneric and
five intrageneric (Cercis comprising both a single inter-
and intrageneric disjunction) . The number of
intergeneric New World-Old World disjunctions may
increase as relationships become better known in
Cercideae . Such relatively low percentages of
intergeneric (and consequently higher intrageneric)
relatedness across the tropics, compared to e.g.,
Detarieae and Dialiinae, is suggestive of widespread
ancestral taxa and substantial levels of dispersal , given
the comparatively large number (58%) of drylancl
species in the tribe.
Detarieae sens. lat.
In tribe Detarieae the basally branching S-biome
Schotia group (Det 1; Fig. 5) is sister to the rest of the
tribe (Det 2 -Am 10) which essentially comprises R-
biome diversifications with varying numbers of species
occurring in the G-biome. The Schotia group contains
Schotia in the S-Nama-Karoo and Thicket biomes and
G-southern Zambezian biome of southern Africa (Low
& Rebelo, 1996); Neoapaloxylon and Brandzeia from
S/ G-biome north, west and south Madagascar;
Barnebydendron from S-biome areas in Central
America disjunct to circum-Amazonian South America
(here also in G/ R-biome forest), and Goniorrhachis in
G-biome forest in South America. Two sister G-biome
genera, Colophospermum from south tropical Africa
and Hardwickia from India (Det 2; Fig. 5) are in turn
sister to the R-biome diversification comprising the
Prioria clade, itself sister to the rest of Detarieae
(Fougere-Danezan et al. , 2003; Bruneau et al., unpubl.
data). This link, together with the rather relictual
distributions of both monospecific genera, is perhaps
indicative of a vicariant origin with both genera being
now restricted to G-biome refugia in Zambezian Africa
and peninsular India respectively.
Rainforest biome taxa are clearly derived in Detarieae
(Schrire et ell., 2005), with the S-biome unequivocally
diagnosing the clade comprising Detarieae sister to the
rest of Leguminosae (Fig. 4), and the next four nodes of
the supe1tree which are basal to the diversification of all
the other major clades in the family (Figs. 6, 7, 9 & 10).
The (S, R) biomes diagnose the Detarieae clade in
Schrire et al. (2005). The crown clade age of Detarieae
is c. 18.6 ± 2.4 Ma (Lavin et al., in press).
With the exception of the neotropical Brownea,
and African 'Macrolobieae' clades (Bruneau et al.,
2000), phylogenies show that most Detarieae clades are
46 LEGUMES OF THE WORLD
distributed fully across the tropics rather than being
dusre red into generic groups restricted to any one
continental region. At least 14 New World-Old World
(and three Africa-Madagascar-Asia) disjunctions occur
among 82 genera (Table 5) with nine being intergeneric
(53%) and eight intrageneric. This high degree of
intergeneric relatedness across the tropics contrasts
markedly with the high percentages of genera endemic
to any one of the major continental regions. Detarieae
has 92% of genera restricted to only one of three
continental regions (Table 4), with none being endemic
to Australia and the Pacific area . Such high levels of
continental generic endemism (thus lowe r levels of
intrageneric disjunctions) probably resulted from
subsequent diversifications which then largely became
restricted to each of the southern continents and Asia,
particularly Africa with 60% generic and 48% species-
level endemism and South America with 20% generic
and 34% species-level endemism (Table 4). When
relationships are better known the number of amphi-
Atlantic disjunctions may decrease in favour of
pantropical distributions.
Dialiinae (= Cassieae pro parte, excl. Cassiinae)
Placement of the Duparquetiinae (containing the
monospecific Duparquetia; Fig. 6) is uncertain; it is
treated as sister to a combined Dialiinae and
Caesalpinieae sens. lat. plus Papilionoideae plus
Mimosoideae alliance by Herendeen et al. (2003a), or
sister to the rest of the family above Cercideae by
Bruneau et al. (unpubl. data). Another S-biome group
is indicated as being sister and putatively basally
branching to the Dialiinae but its monophyly is yet to
be ascertained (Fig. 6). Herendeen et al. (2003a)
grouped Poeppigia (Dl; Fig. 6) from S-biome Central
America, the Caribbean and circum-Amazonian South
America with Baudouinia from S- and G/R-biorne
deciduous to evergreen (often coastal) forest in
Madagascar. Forest (unpubl.) finds support to group
the S/ G-biome Eligmocarpus from deciduous forest in
south-east Madagascar with Baudouinia, and it seems
likely that the S/ G-biome Mendoravia, also from south-
east Madagascar, belongs here. The rest of the Dialiinae
(D2 - D5; Fig. 6) again comprises largely R-biome
diversifications with varying proportions of species
found in the G-biome (Table 4). The S-biome is again
unequivocally optimised as the only endemic area
diagnosing the node Dialiinae sister to the rest of the
family (Fig. 6), and the (S, R) biomes diagnose the
Dialiinae which has a crown clade age of c. 28 ± 3.3
Ma (Lavin et al., in press).
In the Dialiinae, 89% of genera are restricted to
only one of the three major inter-continental regions
but here some 33% of genera ( 44% of species) are
•ntlemic l<.> ch Asian-Au.'> U'<.I Lian regio n (Ta b! · ). Th i.
·o mpm s with lev Is or 22% g n ri · ( 0% s p~dlk)
e n le111ism Jn Afri ·a-Mad~igascar an I 28% w·n ·ri · (16%
specific) c:.: ndemism in the Ni::otr pk ·. Th I I Wo rld
I ia.liinne chus conta ins 61% ge ne ri an I 83% s p ·inc
1,...., Five Ne::w Wo rld - 01 I Wo rld
e ndem s,." •. r- 'ia disjun ·lio ns o • ·ur am ng .l 8 ge n , ,~t
•A~id nga sc,1 ·
iv,. ' four 80% b ing l nte rg ' 11 r! n 1 on ·
\J !Lh ·. (1., I I 5). Hi oh p r ·e nw e · o f 1 level of intrageneric disjunctions in Caesalpinieae sens.
. ·ag •ner1c a) "
int~; neric n.::lar , dn ss :1 ·ross the tropics uggest a
Int g •ra )h ica1 s ·e m:trio irnilar to Derari e'te e xce pt
blog g. eI mphas is >f tiI e . . t'·1c'1t10ns
G - d 1ver:;1 . .
111
thHt ~ I1 · · J.•
. .. ·s in rh Asian - ustr':'dian r g1on, not /\ n ·a-
l 1a111n11 1• •
Mada ,:i sca r and the eotrop1 ·s.
Caesalpinieae sens. lat.
Th . rown clad omprising tl:e P~q~ilionoideae s i~~er l'o
ae5<llplnieae S "llS. /{ff. plus Munos ttle:.1 clades Fig'. 7
· · 't 1• to the Dia liimtt: ( .g ., T-kre nd n t rll. , 2003a·
is Sl
Woj I ·howski, _o . a' alpinieae ·>11 •. k/i/. broadly
c:ooiprls s t.l1J·ee major bi ge~>gra phi ~ti e l rn . ~t. ; a
basally I ,1n ·hing -blame Umuza :hde ( .aes 1; h g. 7
siste r to a ·om bin cl a ssiinae-Pterog yn c/
caesalpinia group alliance (Cass-Caes 3; Fig. 7) plus
an R/G-biome Batesia-Tachigali-Dimorphandra- and
an S-biome Peltophorum group- alliance (Caes 4-Caes
8), which is sister to the Mimosoideae. The Umtiza
cJade comprises the S- and TN-biome Gleditsia-
Gymnocladus which have an extant temperate New
World to east-Asian endemic area distribution,
although Gleditsia has a single S-biome species in
South America . Gleditsia is sister to Umtiza
(Herendeen et al., 2003b), which has a relictual S-
Thicket biorne (Albany Centre) area of endemism in
South Africa. Sister to this group are the monospecific
S-biome Caribbean endemic Arcoa; Tetrapterocarpon
(two S/G-biome species from deciduous forest in the
north, west and south of Madagascar); Ceratonia (S-
and TN-biome) with two species from north-east
Africa, Arabia and Mediterranean; and the
monospecific Acrocarpus in G/ R-biome forest in the
Western Ghats of the Indian peninsula disjunct to
north-east India, Indo-China and Malesia (Herendeen
el al, 2003b). Th .'-bl oru , is un quivo ·aUy ptirnisccJ
a:; th~ only ·nd mi · a r •a di ag n in g t h e no d
Papilion i<leae sist r to the r st of lh f~ mily (rig. 7 ,
which h::t.<; ;1 r w n cbde age of c. 59.2 ± 0.2 Ma Lavin
et al., in pr ·ss). Th lJmtiza r wn clade ;tge is c.
± 3. M;i (Lavin et at., 20 ).
Th IV 1 -hiome Bnt s ia-Ta ·hfgali-Dimorphandra
allian . ntnin s 8 1% o f R-hiom e taxa . Tn ·o m
;~~lyrs \ ~.g. , ~ !er n I 11 el Cl/. 200. a· Wi J 'ed10wsl .i ,
th1. ::tll 1an <.: appe ar to be s is te r and not
Pb?raph yl tic as in l laslo n et al. 2003· Fig 7) ro the ·-
rom p I • ·' · '
,, ... • tophoru m grc up ( aes 8, rig. 7 . The C',
_ " , nnac-Pt
. rogyn I ..1 sa Ipin1a . . group a lLia
. ncc C~t
. s·
~ s 3; !1lg 7) c Illa ins 9 % of :-b!om • and 6:~ % of
1
~)I m Laxa. Togerh r wiLh t.l1e '50/o of -bio me taxa
Occurring in ti I-' l
m. ie e to1 he rum aroup , a nd the -I iome
17 lLZa la<l - · lJ es LI1rc gro ups accoum fo r J3 of
C;p 5.11cl . Wo r l d - Id Wo rld di sjunc tions in
, ' p 111 ,\e_ ·~eu · t-ti· 1· ('!'a ble 5 , ·omparcd with th
f ~1r ln h
g n ri L . ~VG~a llia n ·~ . Tw nty int r- ·ontin enw l
111
ud ing three Af ri ca -Ma d aga ar- . fa )
disjunctions occur among 60 genera (Table 4), six
being intergeneric (30%) and 14 intrageneric. When
compared to the Detarieae and Dialiinae, the higher
lat. indicates that it is the c. 87% dry land (i.e., S- and
G-) biome taxa (Table 4) which have diversified
significantly across continental barriers. S-biome taxa
account for 72% of Caesalpinieae sens. lat. (although
see Table 4 for caveat) . Only 75% of genera in
Caesalpinieae sens. lat. are restricted to only one of the
three major inter-continental regions. The
Papilionoideae, and Caesalpinieae sens. lat. plus
Mimosoideae crown clade marks a major neotropical
node in the diversification of legumes which also is the
oldest within the family at c. 58 Ma. In Caesalpinieae
sens. lat., 50% of genera (71 % of species) and in the
basal Papilionoideae, 55% of genera (72% of species)
are restricted to the New World (Table 4), while Africa-
Madagascar has 20% generic (14% specific) and 18%
generic (12% specific) levels of enclemism respectively
in the above two taxa.
Mimoseae
The Mimoseae are sister to the S-biome Peltophorum
group and/ or the Dimorphandra group of
Caesalpinieae sens. lat. (e.g., Herendeen et al., 2003a;
Wojciechowski et al., 2004), and one of the two R/G-
biome diversifications in Mimoseae occurs in the
basally branching groups closely allied to the
Dimorphandra group. Vicariance analysis optimises
the R/ G-biomes at the node (Fig . 8) diagnosing
subfamily Mimosoideae (Schrire et al., 2005). Groups
Ml to M4 (Fig. 8) comprise a basally branching R/G-
biome grade while a large S- and G-biome alliance
occurs from groups MS to M9 (Fig. 8). A third element
is the derived R/G-biome Piptadenia group, sister to
a S-biome Mimosa group, which in turn is sister to the
Acacieae-Ingeae. The Mimosoideae stem clade age is
c. 56 Ma, while that of the crown clade is 38.4 ± 3.8
Ma (Lavin et al., in press).
The basally branching R/ G-biome Pentaclethra-
Adenanthera-Entada-Newtonia alliance accounts for
47% of Mimosoid R-biome taxa and 52% occur in the
derived R/ G-biome Piptadenia group. In both
groupings S-biome taxa are absent. The S- and G-
biome Plathymenia-Prosopis-Neptunia-Leucaena-
Dichrostachys alliance has 30% of the S-biome taxa in
Mimoseae and the S-biome Mimosa group 70%, while
both have 20% of the G- and a small number of the R-
biome taxa. S-biome taxa account for c. 73% of
Mimoseae (although see Table 4 for caveat). A total of
at least 16 inter-continental generic (including three
Africa-Madagascar-Asia) disjunctions occur among 41
genera, six being intergeneric (38%) and ten
intrageneric (Table 5). Some 76% of genera in
Mimoseae are restricted to only one of the three major
continental regions; 34% of genera (and 76% of
species) are endemic to the New World while the tribe
also has 34% generic (but only 18% specific) endemism
BIOGEOGRAPHY OF THE LEGUMINOSAE 47

in Africa-Madagascar (Table 4). The predominanrly
New World Mimosa largely accounts for the diff rence
in levels of species endemism between the Neotropics
and Africa-Madagascar.
Th ' patterns f di ers ifi ca lion in Mimosea · :ire
c.:ompar:1hl LO thos . ccn above in ac..:salrini ae suns.
lot ., altiloLJgl1 in Lile la(ter more gen n ar ·enc.Jemie LO
the N ·oLropl ··.Noia hie in th' pr 's ~ nt cir umscription
of Mimosea · is the la ·I of ~I putativ ·ly basal bra nching
S-biome group, sister to the otherwise R/ G-biome
Pentaclethra-Adenanthera-Entada-Newtonia alliance.
This may indeed be an exception to the rule , but given
the current uncertainty of where the bounda1y lies
between the Mimoseae and the Dimorphandra group
of Caesalpinieae sens. lat. , an alternative might be to
consider the S-biome Peltophorum group as sister to an
expanded Mimusea ~ (in ')udlng ::ii least part uf th"
Dimorphandrn group) . Tribal d lirniwtions in th large
,n "sa lplni a · · us. lat . plus Mirnosoi lea · nlll.ancc a re
ompli «tl1.: I, It wcver, hy the extended phylog •n •ti ·
gr:u..l e K' ·u1Ting hctw en traditionally · ir~· ums ·ril • I
tribes in this group.
Acacieae-lngeae
Th Ac:tcie:ie plus Ing a · allianc' is poorly r ·sc lved
phylogen ·'lil'llll y w ith tJ1 Ing ae l'(:111aining on " Lbor
lea ·t s:1mp l d Lrib ·s in mol • ·ul a r ::ina lyses . Th ~
foUm ing : ssesi-.rn m ls 1m 1<le ith th · unde rsland lng
ilrnL data wi ll h:ivc to b ~ r ~ view· I when r ·lation hips
are h ucr und rstood . Tb , -I io m is unequlv .ill y
optimis ·d :ts l'he on.ly araa c.li:ign ising th a Acad a'
plus lngea datle (crown lade agl.: = " 4.7 ± .3 Nh.1;
u 1vin el (i/., in pr ·ss) as \ ell as die Tng ~~H:: dad .. ( ·r< wn
age = ·. 19. 1 ± 3.8 M~d indi ·aLing thnt R-biomc ingoi<ls
~ir • deri ec.I Vchrirc ,, rt!., 2) 5 .
The l{/G-bJOml.: Inga-Old World lngeae-Albizia-
Enterolobiu m-AbaM11a alliance (Ing 3-Ing 8; Fig. 9)
accounts for 91 % of R-biome taxa in lngeae, with S-
I i >me tax:t b ing x)mpletcly abs ~ m in this gr >ur. Th •
• / -I iom<;; ca ·i:t gr up and basal ing ids (A I - lng - ;
.Pig . 9) , ;,1n I tli ·· Pith '.•ll oh iu111 -S·1manea-Aca ·i;i
subwn. Ph yll o Jin ·a -· groups Ing - lng l.L ; rig. 9.l,
contai n all th• . -I lom ', nnd 88% or -1 iom ~ wxa . A
total of two inter-continental, intrageneric disjunctions
are found in Acacieae (i.e., 100% of the disjunctions in
the tribe; Table 5) and at least nine inter-continental
dlsjqn ·tions < • ·ur amo ng 37 ln gea " g ·'11 · ta, ' • n
b ·Ing int •rg •neric 78% and tw o lnLrag n •ric
(in 'iuding ne in Ma lagas -:.1.r- si:l, In t lc{l(;fct s ul gen.
Pbyllodi11ec1e . Some ')0%1 >f genera In Ac~1ciea :1n l
t 2;.){, in loge::ie a rc r ~~ tri ·t· d t only on · of a ny of th e
rm1jor continl:nta l r , gi n s (T<d I 4), with 50% of genera
(and 7% sp cies) in Aca i -;1 ·and 65% f g ·n ra (and
::i ~1i, >I' sped s) in Lngea · e nd ml · to the ~ \Xlorld.
fric:i -Madagas ·ar has no ge n e ric hut 3') 0;., sp' ·Ifie
nc.Jemislll i.11 AC<l 'i<:!ac, whil the Ingeae has 2 % o f'
0
gen ~ ra endem ic in Lhc A.->ia-/\us trali a-Pacifi · regi )11
(a nd 61% f p "d s lu e mainly to the inclusion o f
the largely Austra lian G-hiom ' flea · itl subge nu s
48 LEGUMES OF THE WORLD
PbvttmfiJLette) . Most of the generic diversily in lngeae
is ~i Lhin Lh<.: Neotropics and Asia with onl one genus
endemic to Africa (Table 4). The high percentage of
interge n ~ ri · n.:'lat edn ss e s pe ia lly b ·tween th e
Ncot:ropl s and A!'lia) , t >g tiler w ith high levels of
•en ~ra ende mi · ro onJy one< r th, ·onlin •nwl region:;,
~ ch e. Lill! pauerns di . cust>C: d fo r D ~1 : 1ri a and
es['> ·ktl ly th Di.nliinae (-.;: ilh its imilar high I ·v · I <>r
sp ··cies en km ism in Lh · Asin-/\u str<.dia r gion .
In summarising GI salpiniold ·u1tl mime so i I
patterns (Tai ks & 5), Lil • three higher t<Lxa
Detarieae, I i:tliinne and Ingeac ( th e lntl er in Lhe
Mim >soi I ae h::i e high I er ·enLage. f genera
restricted to only one of the three major continental
regions (i.e., the Neotropics, Africa-Madagascar or
Asia-Australia-Pacific). They also have the three
hig h ·st 1P rtenrnges or ·l ntcrgem;ri (an I thus j( w :-,[
perc ntages r in trngen l"ic disj unctions I 'LWe 11
thes . regions as we ll as Lil · higlP:t per ·enlag<;.:.s of R-
hi m • ta, n ( I ngeae is skew "d by th larg numbers or
-I lorn ' tax:l in A ·a ·ic1 :u h g ·n us Pby//odl11eoe) .
Ing "a • (with ut 11 ti ia subg ~nu:-, Phyllotli1Pt1e has ·.
9'i l s p •c i s, . 5' % of w hi ch ar R- biom • lax a
miparal I · Lo l v ·Is in ialiinn . All tb r ~ e;; groups
have b;1sally bran hlng S-hiu111e I ~ ments sister I<> tile
H-hiome diversifi ·a tions ~ hi ·h nr putativ ly derived
f'rom Te rh y.in- id e · I ·m nt s (hen ·e the high
per ' ntag ·s of in terge n er i ' cli ·jun(LionS , Wil li
su l s •quent diverslficarion~ bei ng li m it ti b rg ·ly to
inc.Ii idu:tl ·omin ' JHal r gions in lV ,-hi0111 s (he m: ·
IO\ p ·1-r ntag . f intragen , ric disjunctions) . Tn
J e tari eae th gr a t "S t diversity o ·curs in AJr l ·a-
MaJagas ar, in Dlalilna ~ Ln rropical Asia , and in
Acaci ac-Ingl!ae in the N o trori · ~ a lth ugh tb e larg ·
ustra li an div ·rsJrl ·nLion f Acn ·ic1 subgenu s
Phyllodineae has skewed s p • ·j •s level end •mism to
the Asian-Australian-Pacific region . The Ca salp inieae
sens . !ttt . and Mi mo:> ·a , h owev r, h av , low r
perccmagc..:. ( f gene .t restricted 10 o nl one of th<.:
inc r- con tin nt~tl regions low r p rcenlag s of
imerg ·neri (Ll1ll great r nurnh rs of intr::tg ·n ricl
c.li sj unctio ns o cu rri11 g bctw ·n Lh s regions and
mu h higher pcrc ntages of S- ;in l ~ - bl me , and
cons (jll ntl y Ii ·wcr R~ bi m wx~1. Many ·- and
hiom • chides ha e s u e essful ly di s 1 rsc I a ·ro:ii
continental barriers. A critical node in the phylogeny
l.inl ing Lil .. n ·sa lpi r1i •ae sens. lat. nnJ .Mim >s ~t t'
r mains p >rl r s< lv d < n I r s ·nt lata . Thi:! lat ·st
ana lyst:s ( .g. 1-JasLOn et ti!. 2003; J-1 ~ rend · ·n el rt!.
2003a; Lu ckow ' I al., 003 ; Woj ·Iechowski, 200 ~·
Woj .i · ·howski et ct!., 200 .) ~ii I sugg , st that the S-
I iom · P It< pli rum group is Iii dy 10 b • s is t r ' iLh ' r
LO two ·epar:1t " H/ ,-1 iome diversifi «1Lions comprising
til e Dim rplrnn Im group of a ·salpini ae .'1:'.llS. ft //.
and the Mimllsl'a or to a sing le grncle fr< m th e
former to the latter. Such S-biome diversifications have
been shown elsewhere in the phylogeny to form
critical sister groups at the base of major new
diversifications of higher level taxa in legumes.
Basal Papilionoideae
'fht! b:isal papili< noi ls eo111 1 rise ·. 0 g •ner;.1 CgroLJps
Pl -P'i, Pig. IOl, lo a basa l gr::1d ·of <uious le 111ents
• Lhe Ira litionnlly dr ·umscrih d tri b · s S arl7.i ae ,
() f . .
S ph ur>a :1nci D'.1lb.crgLea ~ Penrungton I al., 200 1).
.R •l:ll i< ns h ips w1th 111 th ·se groups ar " sc m w h:1t
ol s ·ur> ,,11 huugh a 'i .I' b ··hl oroplast DNA in "'rsion
(Ooyl, <JI c1/., I c >) di. tingui.-;hes groups P4 an I I 5
(Fig. JO) al I ' \ ilh LIP r SI or th Parill moi J ae,
fr in groups Pl to P3 (Fig. 10), whic h t g thc r' ilh the
aes:il1 inioideae and Mlmos id •ae Ln k Lhis inv rsion.
The S- hiorn ~ is une 1ui ucaJ ly ip imi.sed ::ts Lb· only
ar :1 dl :1gnosing th ·base of the P~1pilionoi lt:a (, ·llrire
el al., 2005 whi ·h hris a st 'Ill l::t I age or c. 59.2 ± 0.-
MH a nd c.ro n dade of " ';8 ± 0. Ma (Lavin et a l., in
pr ssl. F :-.si l cvl 1· nc · p la · ·s Swartz /a Pl and
G'/cidrct tis-Sl.J~Jl.1J1olo/Jtmn (P ) In arly TerLiwy Norrh
Am •rit·a ( I I rend n al a l. 1992; \Xlh ~ 1 ' I' ~ Baas,
J 92; La in & Sousa , .I )95 . l'Ollps PI - P •a I h~1v'
, -bi o111 ~ l ' m ·ncs, e.g., Aleleitt- yatbost •gia [PI] ;
Cordylct [P2] and Cladrastis-Styphnolohium [P3], either
sister to , or embedded within large R-biome
diversifications in Pl and P2, and sister to TN-biome
diversifications in P3. Wojciechowski et al. (2004) and
Lavin et al. (in press) have clades Pl and P2 grouping
together w i1h moderate supp xt, and basally branching
to a well :.up1 orted Cla lrastis clade (P3) which is sister
to the rest of the Papilionoicleae. The crown age of the
swartzioid-sophoroid sens. lat. (Pl- P2) clade is c. 56.3
± 0.7 Ma and the Cladrastis clade c. 47.4 ± 2.6 Ma
(Lavin et al., 2004). The S-biome Calia-Holocalyx group
(P4J is in turn sister to (or in Wojciechowski, 2003, in
a polytomy with) the large R-biome Lecointeoicl and
Vataireoitl clacle diversifications in P5 . Kite &
Pennington (2003) identified quinolizidine alkaloids in
Calia that suggest there may be a close link with the
Genistoid clade. With improved sampling, elements of
the vataireoids may group together with the
dalbergioids (e.g., Wojciechowski, 2003).
At least eight New World-Old World (and no Africa-
Madagascar-Asia) inter-continental generic disjunctions
occur in basal papilionoids, five being intergeneric
(63%) and three intragene ric (Table 5), although this
total is likely to rise as relationships become better
understood. Some 95% of genera are restricted to only
one of the three major contine ntal regions, with 78% of
genera and 92% of species endemic to the Neotropics
(Table 4). Only 2% each of genera and species are
endemic to the Asian-Australian-Pacific region. In basal
papilionoids, 75% of taxa occur in the R-biome with
only c. 9% occurring in the S-biome. The basal
Papilionoideae thus show a similar pattern to e.g.,
Detarieae but with a Neotropical centre of diversity.
Genistoid clade
The basal genist i I dad s P6 and P7 (Fig. 10) comprise
RIG· biome diversifications (with 71% of the R-biome
species), showing similar patterns of distribution to
the basal papilionoids with 70% of genera and 60% of
species restricted to the Neotropics. The crown clade
age of the genistoids is c. 57.4 ± 0.16 Ma (Lavin et al.,
in press) , and fossil evidence places Ormosia (P6) and
Diplotropis (P7) in early Tertiary North America
(Herendeen et al., 1992; Wheeler & Baas, 1992; Lavin
& Sousa, 1995). As noted above, the S-biome P4 clade
(including Calia) may b e basally branching to the
genistoids. All four biomes are optimised at the base of
the genistoicls (Schrire et al., 2005; Fig. 10). Current
phylogenetic evidence (e.g., Pennington et al., 2001)
suggests that Clathrotropis (P7) is a putative outgroup
of Brongniartieae (P8) and tha t the South American
Cyclolobium and Poecilanthe are basally branching
with a G/R-biome distribution (Fig. 10). Clades P8 and
P9 contain the bulk of S-biome genistoid taxa (55% of
species in these two clades), with 28% of genera and
62% of species restricted to the New World. The crown
clade age of Brongniartieae is c. 32.8 ± 3.2 Ma (Lavin
et al., in press), while that of the Brongniartia (New
World)-Hovea (Australia) disjunction is c. 9 .9 ± 1.8 Ma,
too recent a date for continental history (i.e. vicariant
events) to account for this highly disjunct North and
South American and Australian (excluding Africa-
Madagascar and Asia) distribution. The African-
Madagascan S-biome Cadia group (P9) is putatively
allied to Acosmium, with a crown clade age of c. 44.3
± 2.7 Ma (Lavin et al., in press). The third, predominantly
temperate grollp of genistoids (with 65% of species in
clades Pl0-Pl4 occurring in the T-biome), has 33% of
genera and 62% of species restricted to the Africa-
Madagascar region. The stem clacle of the Sophora
sens. strict. group (PIO), including Euchresta (and
hence tribe Euchresteae) is c. 38.8 ± 2.8 Ma (Lavin et
al., in press), with the (P11) Thermopsideae (crown
clade c. 25.6 ± 3.4 Ma) also likely to be an inclusive
subclade. The group of genera around Sophora mark
a major papilionoid div e rsification into montane
tropical to warm temperate T-biomes, although the
distribution of Sophora is unusual. Besides having a
largely G- and T-biome west, central and east Asia
distribution (with two, tropical, coastal vegetation
species in Africa), Sophora also occurs in montane
tropical south and south east Asia crossing over to
Australasia, the Pacific and temperate South America (in
Section Edwardsia). It should be noted that South
American Sophora appears closely allied to Pacific
island and Australasian taxa.
The almost exclusively mediterranean and warm
temperate TS-biome tribe Podalyrieae (Pl2) is centred
in the Cape region of South Africa. Putative outgroups
include tl1e S-biome distributed Cadia (from the Horn
of Africa-Arabia and Madagascar regions) . The basally
branching element Cyclopia is restricted to the Cape,
often in montane regions. Only the more derived genus
Calpurnia extends beyond the Cape into subtropical
south east Africa, and bas one species ranging through
montane tropical Africa to India. The crown clacle
(Pl2 -P1 4) is c. 43 ± 2.7 Ma (Lavin et al., in press),
BIOGEOGRAPHY OF THE LEGUMINOSAE 49
while that of P13-Pl4 is c. 40.S ± 2.7 Ma and
Genisteae (P14), c. 20.5 ± 2.6 Ma (although as indicated
below, when putatively basal branching taxa are
sampled in these clades, the tribes may be older). Such
taxa, i.e., Lebeckia, Wiborgia, Rafnia and Aspalathus in
Crotalarieae (P13) and Melolobium, Dichilus, Polhillia
and Argyrolobium in Genisteae (P14) occur in the TS-
biome warm temperate and mediterranean Cape region
of southern Africa, and a probable outgroup is the Cape
Podalyrieae. Since both tribes are sister to Podalyrieae,
similar trends are seen with migration northwards into
south-east subtropical and montane tropical Africa (i.e.,
Pearsonia [with one species in Madagascar], Rothia [to
India and Australia] and Robynsiophyton in Crotalarieae,
and Argyrolobium in Genisteae). Both tribes have
secondary diversifications in the T-biome Mediterranean
and Macaronesian regions (i.e., Lotononis in
Crotalarieae, Lupinus, Argyrolobium and the major part
of another 18 genera in Genisteae). Lavin et al. (in
press), indicate a crown age of c. 20.S ± 2.6 Ma for the
clade comprising Anarthrophyllum (one of the endemic
South American genera) and derived elements of
Genisteae. An Old World-New World disjunction in
Lupinus, however, is dated at c. 6.8 ± 1.3 Ma,
suppo1ting the phylogenetic distancing of Lupinus from
Anarthrophyllum (Lavin et al., in press). Polhill (1976)
remarked on the overall similarity of Spartidium to
genistoid genera such as Retama (while placing the
genus in Crotalarieae because it is almost
indistinguishable from Lebeckia). Van Wyk (this volume)
notes recent chemical findings that might place
Spartidium in the Genisteae, where there are many
Mediterranean elements, rather than in Crotalarieae
where Spartidium would have a highly anomalous
distribution. Grata/aria (with the segregate south central
African genus Bolusia) has a pantropical G-biome
distribution with at least one New World disjunction.
While Crotalarieae diverged significantly in the tropics,
Genisteae remained a T-biome warm temperate tribe
with massive divergence in Mediterranean,
Macaronesian and west to central Asian regions .
At least ten (including one Africa-Madagascar-Asia)
inter-continental generic disjunctions occur in the
genistoids, seven being intrageneric (Table S) and three
intergeneric (c. 30%), although the latter is likely to rise
as relationships are better resolved. Some 79% of genera
are restricted to only one of any of the four major
continental regions (i.e., the T-biome Mediterranean-
Macaronesian and Europe region is here considered
distinct) , with 32% of genera and S4% of species
endemic to the African-Madagascan region (Table 4)
and 18% genera (23% species) to the New World. In the
genistoids, SS% of taxa are in the T-biome with only c.
S% each occurring in the S- and R-biomes.
Dalbergioid sens. lat. clade
Outgroup genera to the dalbergioids sens. lat. are
likely to be in clade PS (vataireoids), although with
further sampling genera in PS may join the Dalbergioid
50 LEGUMES OF THE WORLD
sens. lat. clade (Fig. 11). Sister to the Dalbergioid sens.
lat. clacle sensu Wojciechowski et al. (2004) is Andira
and Hymenolobium, and Lavin et al. (in press) date
this Dalbergioid sens. lat. crown clade at c. S7.4 ± 0.3
Ma. The Dalbergioid sens. lat. clade is optimised by
the S-, G- and T-biomes (Schrire et al., in press),
indicating that the c. 1S% of R-biome dalbergioid taxa
are derived in the Dalbergia and Pterocarpus clades.
The largely shrubby to herbaceous Amorpheae (PIS)
is an entirely New World group (stem clade age c.
SS.7 ± 0.7 Ma and crown clade c. 37.7 ± 2.9 Ma), with
91 % of species in the S-biome and Dalea having a
secondary centre of diversity in the TS-biome of
Andean South America . Amorpheae is sister to the
Dalbergioid sens. strict. clade (i.e., Dalbergioid clade
sensu Lavin et al., 2001a) and Amorpha is a TN-biome
genus in warm temperate North America. The Adesmia
clade (P16), sister to the combined Pterocarpus and
Dalbergia clades (Pl 7- P19 and P20- P22; Fig. 11)
together comprise the Dalbergioid clade sens. strict.
(Lavin et al., 2001a), with a crown clade age of c. 49
± 1.2 Ma (Lavin et al., in press). The TS-biome warm
temperate to montane tropical Adesmia is basally
branching in the predominantly herbaceous Adesmia
(P16) clade (crown age c. 34.S ± 2.4 Ma), which has
83% of genera and 92% of species endemic to the
New World. The Dalbergia and Pterocarpus clades
have a stem age of c. 46.S ± 1.2 Ma (Lavin et al., in
press), with the Pterocarpus crown (Pl 7 -P19) c. 44.7
± l.S Ma, and the Dalbergia crown (P20-P22) , c. 40.9
± 1.2 Ma in age. The Pterocarpus clade has 82% of
genera and species endemic to the New World, with
the S-biome Riedeliella and Discolobium (crown c.
32 .S ± 3.4 Ma) sister to a largely G/ R-biome
diversification comprising the rest of the clade except
for the derived S-biome Chapmannia-Stylosanthes
subclade (Lavin et al., 2001a; crown c. 16.3 ± 1.6 Ma).
The intergeneric amphi-Atlantic disjunction in
Chapmannia is c. 14.2 ± 1.7 Ma in age, and the
disjunction between Platypodium-Tipuana (South
America) and Inocarpus (south-east Asia) is c. 20.3 ±
3.6 Ma (Lavin et al., 2004). A pantropical disjunction
also occurs in Pterocarpus with European fossil
evidence of Pterocarpus reported by Buzek (1992)
from the lower Miocene. The Dalbergia clade has c.
43% endemic species in each of two regions, the New
World (with 29% of genera) and Africa-Madagascar
(with 3S% of genera). Based on current analyses (e.g.,
Lavin et al., 2001a), the Dalbergia clade comprises
four subclades forming an unresolved polytomy: the
G-biome Weberbauerella; the G/ R-biome Dalbergia-
Machaerium group (crown c. 34 Ma; Lavin et al., in
press); the S/G-biome Aeschynomene-Bryaspis group
(crown c. 32.6 ± 1.7 Ma) and the S-biome Diphysa-
Ormocarpum group (crown c. 13.4 ± 2.S Ma) .
At least 13 (including four Africa-Madagascar-Asia)
inc ~ r- o n.llnental generic disjunctions occur in
da ll e rgioids, nine being intrageneric (Table S) and
four intergeneric ( c. 31 %) . Some 83% of genera are
restricted to only one of the major continental regions,
~ ith 6R% of g nl!ra <i nd 69%l of SJ ecies endemic lo Lh e
NeW World and 11 % genera C~ % s pecies ) lo Africa-
JVfadnga. ·a r. T11e S-bio me taxa account for 38% of'
lalbcrgio icls witJJ c. 18 % a nd 15% res p ~c r ivc l y
c urring in Lhe T- and R-biomes.
Baphioid clade and Mirbelioid sens. lat. clade
The predominantly Old World clade (Figs. 1 & 3)
mpri fog the remaining pa pilion oic.b has a row n
ag or " 56.2 ± 0.5 Ma (Lavin et al., in press and is
01 timls d hy all fo ur bio mes excluding TN) I y 'chrir "
et 6it. (2001). Th basa ll y bra n lung Ba phio id cl ade
(P23; Fig. 12) with a stem age as for the Old World
clade and a crown age of at least c. 22 ± 2.8 Ma
(perhaps older when more gen ra ~1r sampled), have
S7% of genera and 93% of s pecies e ndemi · in Africa-
Madagascar. The R-biome taxa account for 63% of
species while S-biome taxa are absent. Putative
relationships between the mediterranean TS-biome
South African Cape tribe Hypocalypteae and the warm
temperate to subtropical sclerophyll community, TS-
biome, Australian tribes Mirbelieae and Bossiaeeae
(P24; Fig. 12) have been discussed earlier. This group
of Southern Hemisphere tribes is here referred to as
the Mirbelioid sens. lat. clade, with a crown clade of
c. SS± 1.1 Ma (Lavin et al., in press), which is almost
the same age as that of the Old World clade itself,
indicating the surprising age of this disjunction
between southern Africa and Australia (i.e., without
evidence of intervening taxa occurring in Asia,
Madagascar or the Indian Ocean islands). The
Australian Mirbelioid sens. strict. clade (i.e., mirbelioids
sensu Wojciechowski et al., 2004) have a crown age of
48.4 ± 1.3 Ma (Lavin et al., 2004). The phylogeny (Fig.
12) suggests that the outgroups of these shrubby to
herbaceous mirbelioids sens. lat. are likely to be
among the R/G-biome baphioids (P21), which
comprises a number of Africa-Madagascar-Asia
disjunctions, again supporting the possibility of
Tertiary Tethys (and Indian Ocean) Seaway links
underpinning these relationships. As in many other
legume clades, the extant 'signal' of these links is
s nmy, an d ·o nn ~ ti ns Lo tb '·e o ld , now t ~ 111pe ra 1 ·
· outh rn II misph r ·nLre. h a ve likely i1he r b e n
obs ured, r ear ly dispersal c f mirb lio id · to Austra lia
Was followed by a long period of isolation there.
. . nly lhre • Africa -Mada g as ·a r-Asia ge ne ri c
~ 1 s! unt:1io~s occ~1r in rJ1 hasnl Id Wo rld Lrib s, , U
ing U'lli ag n n (T~1bl 5). om e 57% of ba pW Id
11 . ra ru·e re:Lri 'L •c.I L o nl y o n o f Lh c tw major
lrop1. ·al Id Wo rld continenta l regi n , w iLh 17% of
g · i e ra a nd 93% o f sp e it:s e n d e mi tO Afri a-
Maclagasca r, while all mlrbe lioid genera are restricted
l either Australia (with a minor exte nsion Lo Asia or
S ll th fri ·a (Tab! ~ o taxa o cur in 1h ew
Wor[(l, In I aphioids % e>f t·1~ a rc in Lh ' H-biom . and
nearly
.
·111 "" ' I 1· . I l
"cLl'J ·• 101( :-; sens. al. ' u·e nd mi · lo LIP T -
b 10111e <Ta ble ).
Millettioid sens. lat. clade
The clade from P2S to P47 (Figs. 12 & 13), including
tribes Indigofereae and the millettioid-phaseoloid
alliance has a crown age of c. S3.6 ± 1.0 Ma (Lavin et al.,
in press), and is here referred to as the Millettioid sens.
lat. clade. The predominantly S/G-biome, herbaceous
to shrubby Indigofereae (P2S) are sister to the large
millettioid-phaseoloid diversifications, and have a
crown age of c. 30.8 ± 3.4 Ma (Lavin et al., in press).
The S- and G-biomes are optimised at the base of the
Millettioid sens. lat. clade suggesting that R-biome
millettioids-phaseoloids are derived from dryland taxa
(Schrire et al., 200S). Outgroup relationships were
tested with the basally branching Old World tribes,
particulary Hypocalypteae (Schrire et al., 2003), and
Indigofereae shares an obscure origin along with the
mirbelioids (sensu Wojciechowski et al., 2004) and
baphioids. Representing the earliest branching lineage
of Indigofereae , the woody genus Phylloxylon,
endemic in Madagascar, has a strong resemblance to
the baphioids, and Pennington et al. (2001) have a
Baphioid clade in a polytomy with Indigofereae.
Indigo/era diversified c. 26.3 ± 2.5 Ma (Schrire et al.,
2003) and comprises four distinct clades with at least
two Old World-New World diversifications dated at c.
16.9 ± 2.6 Ma and c. 8 ± 2.2 Ma. The Indigofereae
(Table S) have three inter-continental intrageneric
disjunctions (one Africa-Madagascar-Asia); 71 % of
genera are restricted to only one of the three tropical
continental regions (Table 4) and 71% of genera (74%
species) are endemic to Africa-Madagascar. Some 66%
of species are in the S-, and 34% in the G-biomes.
The crown age of the combined millettioid-
phaseoloid alliance (millettioids sensu Wojciechowski et
al., 2004) is c. 4S.9 ± 1.8 Ma (Lavin et al., in press), and
this is optimised by the G-biome (Schrire et al., 200S).
The basal millettioid and phaseoloid group (P26)
comprises woody elements of both groups since
relationships are too poorly resolved currently to assign
them to either single group. The predominantly R/G-
biome clade P26 (69% of species in the R-biome) has
S9% of genera and 62% of species endemic to Africa-
Madagascar, with only 2% of species in the New World:
in the genus Platycyamus which is probably basally
branching to the phaseoloids (e.g., Wojciechowski,
2003). In current analyses (e.g., Wojciechowski, 2003),
the S-biome Abrus clade (P27) marks the first
dichotomy in the Millettioid sens. strict. group
(P27-P34; Fig. 12), with a crown of c. 37.5 ± 2.4 Ma.
The predominantly R/G-biome Diocleinae (P28) , shown
to belong here rather than in the phaseoloids (e.g.,
Kajita et al., 2001), has 8S% of genera (79% of species)
endemic to the New World and no genera (and only c.
1% of species) endemic to Africa-Madagascar. Rhodopis
and Neorudolphia are S-biome Caribbean endemics
and the TN- genus Lackeya is probably sister to
Galactia. The Old World Ophrestiinae (P29), similarly
shown by e.g., Kajita et al. (2001) to be better placed
here than in the phaseoloids has 76% of species in
BIOGEOGRAPHY OF THE LEGUMINOSAE 51
Africa-Madagascar and all species occur in the G-biome.
The crown age of core-Millettieae (Hu et al., 2000) is c.
26. 5 ± 2.1 Ma (Lavin et al., in press) and 39% of genera
(26% of species) are endemic to the New World and
30% genera (41 % species) to Africa-Madagascar. The
core-Millettieae (P30- P34) is a predominantly G/R-
biome diversification with 61% (G), and 22% R- species,
although the Tephrosia clade (P32) represents a derived
S-biome diversification with 17% of species. The Old
World G-biome Philenoptera and a canavanine-
accumulating part of Millettia sens. lat. (P30) are sister
to the remaining clades (P31-P34), which have a
basally branching New World S-biome element in
Hesperothamnus and Piscidia (e.g., Hu, 2000; Kajita et
al. 2001). Related to the latter is clade P31 containing
the G/R-biome distributed Lonchocarpus group. P33
represents the R/G-biome distributed Derris and
Paraderris diversification in the Old World. P34
comprises the Old World R/G-biome non-canavanine-
accumulating Millettia sens. strict. and allied genera.
Of 11 inter-continental generic disjunctions in the
millettioids sens. strict., only one is intergeneric (Table
5), although more may become evident as relationships
are better resolved. Some 82% of genera are restricted
to only one of the major continental regions, with 32%
of genera and 37% of species endemic to Africa-
Madagascar (Table 4) and 32% genera (32% species) to
the New World. The Asia-Pacific-Australia region has
19% of genera and 30% of species; species diversity in
this group is more or less evenly spread across the
three continental regions. The G-biome taxa account
for 66% of the millettioids sens. strict. with c. 26%
occurring in the R-, and 13% in the S-biomes.
The crown age of the Phaseoleae sens. lat. clade
(P35-P47) is c. 40.3 ± 2.2 Ma (Lavin et al., in press),
and this is optimised by the G-biome (Schrire et al.,
2005) with the S-biome only becoming optimised
(together with the G/R-biomes) at the base of the
core-Phaseoleae clade (P42- P47; Fig. 13). The
Clitoriinae clade (P35) is related to genera in the basal
millettioid and phaseoloid group (P26), e.g., to the
only neotropical genus Platycyamus (Wojciechowski,
2003), or to Austrosteenisia (Kajita et al., 2001). The
G/ R-biome Clitoriinae comprises largely shrubby to
herbaceous elements with 60% of genera (86% of
species) endemic to the New World, and 96% of
species occurring in the G-biome. The phaseoloids
above P35 (Fig. 13) form two sister clades with a
crown age of c. 28.1 ± 1.9 Ma: P36-P41 including
the Apios group, subtribe Kennediinae, Mucuna and
tribe Desmodieae, and P42 - P47 comprising the core-
Phaseoleae inclusive of tribe Psoraleeae (Kajita et al.,
2001). Basally branching elements of the Apios-
Desmodieae clade (e.g., the Apios group and the
desmodioid Lespedeza group) are montane tropical to
warm temperate east Asia-North America in
distribution. Some 95% of genera in this clade are
Asian-Australian, with 69% genera (85% species)
endemic here, and no genera but 28% of species
(occurring in the single genus Desmodium) are
52 LEGUMES OF THE WORLD
endemic to the New World. The G-biome accounts for
88% of species and 13% occur in the TN-biome. The
crown age of the Desmodieae is c. 14.5 ± 1.5 Ma
(Lavin et al., in press), although further sampling may
show this to be older. Evidence supports the R/G-
biome distributed Mucuna (P38) as the sister genus to
Desmodieae (Kajita et al., 2001). More than one
diversification occurs in the New World in
Desmodium, representing separate dispersal events
from the Old World (Kajita, pers. comm.). The basally
branching core-Phaseoleae (P42 - 44), including the
R/ G-biome Spatholobus group, the S/ G-biome
Erythrina-Psophocarpus group and the S/ G-biome
subtribe Cajaninae, have 33% genera (50% species)
endemic in Africa-Madagascar and 28% genera (25%
species) in Asia-Australia; no genera are restricted to
the New World but 25% of species are endemic here
in the genera Erythrina, Rhynchosia and Eriosema.
Apparently basally branching elements of Erythrina
are South American (Bruneau, 1996) but the centre of
diversity is in the S-biome in North and Central
America, with one or more disjunctions to the Old
World. Clades P42-P44 have 66% of species in the G-,
and 31 % in the S-biomes and on present evidence, the
basally branching element of the Cajaninae (P43) is the
G-biome African Adenodolichos. If similar patterns of
distribution noted in the Phaseolinae hold true,
however, the S-biome Paracalyx may prove to be
basally branching after more comprehensive sampling
of the Cajaninae. The crown clade age of the
Glycininae-Phaseolinae clade (P45-P47; Fig. 13) is c.
19.4 ± 1.5 Ma, and contains tribe Psoraleeae (P46)
sister to elements within the Glycininae (e.g. , Kajita et
al., 2001; Wojciechowski, 2003). The montane tropical
Asian Pueraria (at least in part) and Dumasia (with
one species extending to Africa) are possibly basally
branching in the Glycininae (P45), which has 35%
genera (61% species) endemic in Asia-Australia with
16% species in the S-biome. A near basal disjunction
is found with the New World element comprising
Cologania, Calopogonium, Pachyrhizus and Herpyza
(e.g. , Lee & Hymowitz, 2001) . Compared to
Glycininae, the Phaseolinae have most taxa endemic
in Africa-Madagascar (29% genera and 53% species)
and the New World (33% genera, 36% species) with c.
52% of species in the S-biome. This subtribe has a
basally branching S-biome element, Wajira (and
Vatovaea) with a major disjunction occurring between
Old World Vigna and the S/TN-biome New World
Phaseolinae (including elements of Vigna). The
Psoraleeae (crown clade age c . 6.3 ± 0.9 Ma) is
predominantly temperate to montane tropical in
distribution, with 60% of species in the TS- and 17%
in the TN-biomes. Putative basally branching elements
are the S/G-biome Cullen (e.g., Wojciechowski et al. ,
2004) although Grimes (1990) favours the southern
African Otholobium and Psoralea in this position
(Stirton, this volume). Current evidence suggests that
outgroups for Psoraleeae are perhaps near Glycine
and the TN-biome distributed Amphicarpaea, in
~ t ydnlna ~ . Africa has U % of g 11er1 (57 Vti of pc f(::S
'i6% o f g nera (a nd - 5% of sp d es) are e nd mic
~111ll~he Nt!W Worl I. Pmther res arch will establish if the
uth m.erl an sp ci s f Otbolobium. belong i11 V'lis
th 1w ise soutll rn to .1stern Afri can gem1s or if their
~ffiniLI ··s Ii ' rath r with the No rth American g ne ra.
~rh r , e nt ag of c:he la rge ly T-bi me Psora l e~1
llv rsifi -.:Ilio n a ppt!ars t pr dud Te tbyao vicarianc ,
in fa v ur f dispe rsa l xplanati ns for th - clistril ution
of this clade.
No clear examples of intergeneric disjunctions occur
in the phaseoloids, although a few may become evident
when relationships are better resolved, thus the c. 32
intercontinental disjunctions recorded are all
intrageneric (Table 5). Some 71% of genera are
restricted to only one of any of the major continental
r~glons, with 36% of g · nera. and species endemic to the
in-l'acific-Australia region (Table 4), 18% genera (33%
species) to Afri a-Madagascar and 17% genera (30%
species) to th ~ N w W<>dd ; thus species div rsily (as in
the millettioids sens. strict.) is more or less evenly spread
across the three tropical continental regions. In the
phaseoloids, 63% of taxa are in the G-biome with c.
21% occurring in the S-, and 12% in the T-biomes.
Hologalegina
The Hologalegina clade (Wojciechowski et al., 2000),
sister to the Millettioid sens. lat. clade, comprises the
final two major papilionoid subclades (Fig. 14), the
robinioids (P48 - P50) and the Inverted Repeat Lacking
Clade (IRLC [P51- P56D. Hologalegina has a crown
age of c. 51.1 ± 1.1 Ma (Lavin et al., in press) and is
optimised with the S-, G-, R- and TN-biomes (Schrire
et al., 2005).
Robinioid clade
The Robinioid crown clade is c. 48.9 ± 1.2 Ma (Lavin et
al., in press) and is optimised only by the S-biome
(Schrire et al., 2005). The Robinieae (P50), distributed
entirely in the New World, is centred in the Neotropics
and subtropics of North and Central America and tl1e
Caribbean. The crown age of the Robinioid clade is c.
38.7 ± 1.3 Ma (Lavin et al., 2003; in press), with c. 63%
of species restricted to the S-biome and 30% to the G-
biome, and Robinia in the TN-biome. Lavin et al. (2003)
found conflicting evidence between ITS and matK
sequence data as to whether the Sesbanieae-Loteae are
sister to Robin i ae (e.g., as Lndkate<l by the matK dat:.1
set , >r if Sesbanieae is sister ro Robinieac e.g., as in
th fl'S data set . Neither optimisation is su p po rt d by
parsimony bootstrap values. The I -bi >m . eslx 111ieae
~'48), ' ith a crown f c. 19 ± 1.8 Ma, cornprls · th
sing) ' pa nLrop i ·al g ~nu s Sesbanitl w ith . 8% o f
p ~c:ic!'i endemi L Afric:a-Mad·1 1ase 1r and 15% LO th
New Wort I. Th P49) Loteae d ad " CJ wn . 14.6 ± 1. 5
Ma hat; 23% of genem and 7 % of sp ies ndemic to
th Eurc1. ·ia-Mcditernne an region with c. 88% of sp cies
re!itriued Lo tl1e T -I io me; the1wi:e 18% of g(::nera
and 16% of species are endemic in the New World.
This largely temperate Eurasian clade (with extensions
into the montane tropics of Africa and South America)
has putative basally branching elements including
Hippocrepis, Scorpiurus, Coronilla, Securigera and
Podolotus (this volume), which are biased towards a
south-west Asia, Horn of Africa, Arabia to north-west
India S-biome distribution. Current phylogenies place all
four New World taxa in a subclade including e.g., the
Old World Kebirita, Dorycnopsis and Ornithopus
(although this has one New World species). The clade
ages reported here, however, are considered too recent
to be the result of vicariant events and are probably best
explained by dispersal. One putative, pantemperate,
intergeneric disjunction between the New World genera
(Hosackia, Ottleya, Acmispon and Syrmatium) and Old
World Eurasian-Mediterranean genera, requires further
sampling to clarify relationships. Three other
intrageneric, intercontinental disjunctions occur in the
robinioids (Table 5). Some 68% of genera are restricted
to only one of the major continental regions (Table 4),
with 44% of genera (31% of species) endemic to the
New World and 15% genera (50% species) to Eurasia
and the Mediterranean. In robinioids, 61% of taxa are
in the TN-biome with c. 24% occurring in the S-, and
15% in the G-biomes.
IRLC
The IRLC crown is c. 39.l ± 2.4 Ma (Lavin et al., in
press) and is optimised by the TN-, R- and G-biomes
in Schrire et al. (2005), and by only the TN-biome at all
the other nodes (Fig. 14). The basally branching
millettioid IRLC clade (P51) contains nearly all the
tropical taxa in the IRLC, with 67% of genera and 51 %
of species endemic to Asia-Australia, and c. 43% of
species to the temperate Eurasia-Mediterranean region
(i.e., in the genera Glycyrrhiza and Wisteria). In P51,
26% (and 28%) of species occur in the R- and G-biomes
repectively and c. 51% in the TN-biome. The
remaining, largely warm to cold temperate or montane
tropical, IRLC tribes (P52 - P56) comprise the Galegeae,
Hedysareae , Cicereae, Trifolieae and Fabeae. Asia-
Australia has c. 59% (in Galegeae), 18% (in
Hedysareae) and 8% (in the remaining tribes) of genera
endemic to the region. No genera are endemic to the
other intercontinental areas except for two (i.e., 9% of
genera) in the Galegeae in Africa. The Eurasian-
Mediterranean region has 77% (in Galegeae), 96% (in
Hedysareae) and 76% (in the remaining tribes) of
species endemic there, with c. 98% of species in each
of the groups endemic to the T-, mostly TN-biome.
Clades P52 - P56 are often centred in the Irano-
Turanian phytogeographical region of Eurasia
(Tahktajan, 1986) with montane tropical links to Africa.
Putative basally branching elements are often
associated with the S-biome west Asian, Arabian and
Horn of Africa regions, usually in montane refugia
from west Asia to the Sino-Himalayan region, and with
extensive diversifications in the Mediterranean, Europe
BIOGEOGRAPHY OF THE LEGUMINOSAE 53
and the cooler and continental temperate regions of
nLral and no1th-east As ia . The c rown of tJ1 Ga legca
J-1 dysarea P52 - 53) is ·. 33.4 ± 2.5 Mtl, whll chat o f
Hedysar a P53) is c. 29. 9 ± 2.7 Ma and Gal egeae
s 1~s. fc1t. (P52) . J 5.1 ± 2.2 Ma. The ~aleg '"fie ·em. !tit.
is mo~l wld ~s f')r ad wit:h major Jiversifl atlons in wann
temperaLe Ausrra.lnsia an l · uth Africa and all d ndes
(except P54. have t mpen.1le 01 I World - New Wor.ld
(North AmeriCI oftei linl ed LO montane South America
cli.~jun ctic.ms within s p cios ge n e ra, .g., Astraguius
0;,._111ropis Heci;ysa nmi Viciu, Lat/:Jyn1s and Trifoliwn.
Th ·rown ag o f the i e rea -Trffolfeae-F·1b ae ·lad
( in ·ludi ng Lhe I asally l ranching genera Pciro hetus
and Gale,o 1 is . 32.6 ± 2.'5 Mn, a nd th :ll o f th e
pa rapbyle tl · Trilo ll a (P55 including P56), c. 25.3 ± 2.2
Ma d1e Pab ne P56), " 17.7 ± 1.8 M::i and Cice reae
(P54), c. 15.4 ± 3 Ma.
Of nine inter-continental disjunctions noted in the
IJU.C, all •u·e inLragenc1·ic an I oc ·ur ~unong t mp ~ rate
rax;1 (,[ ahl ' . 'onr 42% of gen ra a re reslr.d d to
unly o n or ro ur maj r contin ntal regions (11th! ) the
lowest for <.my gro up of I 'gum s, indicating the more
highl y di s r ·rsed na tur · o f ma ny ternper·H ,
predorn.in;rntly hedya ·eou,<; g "llera . ·rhis is also shown by
the high p •rcenrages of g n ra o ·curring in (but no t
ndemic to the ·ontinenwl rcglo ns, i. e. 92%, . % and
31 % of IRLC genera are present in Asia, Europe-
Mediterranean and Africa respectively. While 38% of
gen.eJ':i a r ~ ndeml · to the t'ropirnl/subtropiL-a l A:ila n
3
r gion. C::l nd nm t the New World), , 78% of species
are e nde mic ro the tempcr.:\l Eurns fan- M •diterran •an
region and c. 15% to the New World (Table 4). The T-
biome accounts for 98% of IRLC species, with 91% in the
T - a nd 7% in Lhe TS~biornes.
In s umm;iry , WiLh the e x ce ption or th .. BaS<ll
1)~1pili on oid a a11d haphio l Is ha ving 76% a n I 63% of
Utxa in lbt: H-hiome res p tively I au othe r papil.iono id
·lades are p.re lominantly / G-biom d istributed, wirh
th e ge njstoids, mirbe l.l o ids ;i nd 1 l o loga l ·gina mrna
dive r11 In th e T-hi.om . All papllionoi.ds xcept the
B·1saJ Parilionoid ae have from . 70% 10 1.00% of th ir
lot · r· onti11 nwl tl.i.5junclio11s intragene r!c ( Tab! ~ 4 ,
54 LEGUMES OF THE WORLD
hence a relatively high proportion of genera are
distributed in more than one continental region. The
Basal Pap1Jionoidc;1 an I dalb rgio icls h av the g reatesL
divers ity o f taxa i.n tl1 New Wo rld (78% and 68%
gen ric , a nd 92% and 62% sp "cifi • nd em.i sm
Tribes and Genera of Leguminosae
resp t ive ly) while the genistoi Is and haphio ids ar
most Jjver e in the Africa-Madagasca r regi< n 32% 'Ind
57% generic, und 5 % a nd 93% sp cifi , end " miSLll
re p " tlv ly). The mirbe lioid s se n s. let/. . (wJtb an caesalpinioideae pp. 56-161
unexpe(;tedly o ld crown dive rsification) ar reslricted
almost ntirely to the TS-I i >mes o r Australia a nd So uth Mimosoideae pp. 162-213
.Mrica . The miJl e ttioicl s ·ens . lm . p ·trti ·ul arly th
millettioids sens. strict. and the phaseoloids) are more
~ve nly listributecl with maj r ·entr s f dlv rsiLy a r ss
Papilionoideae pp. 214-509
Lb e rropi ·s o f the three ma jo r tro p ica l o ntin ntal
regi ns (Tai le 4). ln l lologalegina the rohinio ids a r •
gen e ri ·al.ly mos t diver in the N"e w Wo rld 4 %
endemism) whil. .. tlP enlr o f g neri · di versity in th '
mL is th temp rate A"> i~11~ region 38% endemism }.
B th ha ve the highest le · Is of sp ci ~. ntle mis m in
L11 Euras ia n-Medite rra n an reg i n (50% a nd 78%
r sp <:tively . Th pcrce nrage. or g n ra r stri t d to
o nly on of fo ur major ontincn ral r gion de ·rea. es
from Lhc R-1 iome nasal Pa piliono id ae (95%) to Lhe T-
biome-wide IRLC (42%). This is due to the more
a pica lly bran bing d ad s pre lo mina ntly omprising
' hrubl y or he rba O LI S ta.xa f' cJri " f a nd coo le r
nvlr >nme nts, whi h c.lis p •rs m r · rea dily a ross
'(>n tine nrn l harriers. Within the contex1 · f a r la.Live!
W ,ll Sllppo 1t ·cl , . ( Ma da te for Lbe div rsilkmion of
lb famlly , To wn ages of ·omp H1 " 0L l:id ~s ,
particularly in the :\esalpinioideae a nd Mimosoidea
(Lavin et ul. in I r ~ss , a re Iii e ly ro be pus hed I a I
with more intensive sampling. It is interesting to note,
how v •r, tlrnt ma ny o f Lhe pre I< inin antly H-biom
cbd c.:s, .g. Dc tnri cae Di aliin:1e, ln g ·a a nd th
baphio icls hav e r latively r · em ·rown d acJe ag s Browneo mocrophyllo Photograph by G. P. Lewis Zopoteco corocosono Photograph by G. P. Lewis
(Tai l , 4), ·o m pared to tl1 e Largel y S/ -1 io m • cl::tdes,
e. " ~ae. alpiniea sens. lat., 1b gcn.i to ids, da lhe rg io ids
~111d. millettio.ids .wms. fat.; an ob, e rvntio n supporting th
findin gs of Schrire et al. (2005) that R-biome taxa are
d rived in the family (Figs. 4-14).
Chaetocalyx longilobo Photograph by G. P. Lewis Pithece/lobium exce/sum Photograph by G. P. Lewis
TRIBES AND GENERA 55
and the cooler and continental temperate regions of
central and north-east Asia. The crown of the Galegeae-
Hedysareae (PS2- S3) is c. 33.4 ± 2.5 Ma, while that of
Hedysareae (PS3) is c. 29.9 ± 2.7 Ma and Galegeae
sens. lat. (PS2) c. lS.1 ± 2.2 Ma. The Galegeae sens. lat.
is most widespread with major diversifications in warm
temperate Australasia and South Africa, and all clades
(except PS4) have temperate Old World-New World
(N01th America often linked to montane South America)
disjunctions within speciose genera, e.g., Astragalus,
Oxytropis, Hedysarum, Vicia, Latbyrus and Tri:folium.
The crown age of the Cicereae-Trifolieae-Fabeae clade
(including the basally branching genera Parocbetus
and Galega) is c. 32.6 ± 2.S Ma, and that of the
paraphyletic Trifolieae (PSS including PS6), c. 2S.3 ± 2.2
Ma, the Fabeae (PS6), c. 17. 7 ± 1.8 Ma and Cicereae
(PS4), c. 15.4 ± 3 Ma.
Of nine inter-continental disjunctions noted in the
IRLC, all are intrageneric and occur among temperate
taxa (Table S). Some 42% of genera are restricted to
only one of four major continental regions (Table 4), the
lowest for any group of legumes, indicating the more
highly dispersed nature of many temperate,
predominantly herbaceous genera. This is also shown by
the high percentages of genera occurring in (but not
endemic to) the continental regions, i.e. 92%, S4% and
31 % of IRLC genera are present in Asia, Europe-
Mediterranean and Africa respectively. While 38% of
genera are endemic to the tropical/subtropical Asian
region (and none to the New World), c. 78% of species
are endemic to the temperate Eurasian-Mediterranean
region and c. 15% to the New World (Table 4). The T-
biome accounts for 98% of IRLC species, with 91 % in the
TN- and 7% in the TS-biomes.
In summary, with the exception of the Basal
Papilionoideae and baphioids (having 76% and 63% of
taxa in the R-biome respectively), all other papilionoid
cbdes are predominantly S/G-biome distributed, with
the genistoids, mirbelioicJs and I-Iologalegina most
diverse in the T-biome. All papilionoids except the
Basal Papilionoideae have from c. 70% to 100% of their
inter-continental disjunctions intrageneric (Table 4),
54 LEGUMESOFTHEWORLD
h e nce a rel a tiv ~ 1 y h ig h proportion of gen ra are
di.strib uted in more rha n one continenta l region, ·n1
Basal PapiUonoid 'a, and dalbergioids have ct1 · gr •atesr
livei·sity or taX'I in rh N w World 78% and 68%
ge n eric, and 92% ~ nd 62% s pe ·ifi c ncl mis m
1ribes and Genera of Leguminosae
re::;p ·tlvely, whil the g n istoic.L- and baphicids are
most diverse in the Africa-Madagascar region (32% and
S7% generic, and 54% and 93% specific endemism
respectively). The mirbelioids sens. lat. (with an caesalpinioideae pp. 56-161
unexpectedly old crown diversification) are restricted
almost entirely to the TS-biomes of Australia and South Mimosoideae pp. 162-213
Africa. The millettioids sens. lat. (particularly the
millettioids sens. strict. and the phaseoloids) are more
Papilionoideae pp. 214-509
evenly distributed with major centres of diversity across
the tropics of the three major tropical continental
regions (Table 4). In Hologalegina, the robinioids are
generically most diverse in the New World (44%
endemism) while the centre of generic diversity in the
IRLC is the temperate Asian region (38% endemism).
Both have the highest levels of species endemism in
the Eurasian-Mediterranean region (50% and 78%
respectively). The percentages of genera restricted to
only one of four major continental regions decreases
from the R-biome Basal Papilionoideae (95%) to the T-
biome-wide IRLC (42%). This is clue to the more
apically branching clades predominantly comprising
shrubby or herbaceous taxa of drier and cooler
environments, which disperse more readily across
continental barriers. Within the context of a relatively
well supported, c. 60 Ma date for the diversification of
the family, crown ages of component clades,
particularly in the Caesalpinioicleae and Mimosoicleae
(Lavin et al., in press), are likely to be pushed hack
with more intensive sampling. It is interesting to note,
however, that many of the predominantly R-biome
clades, e.g. Detarieae, Dialiinae, Ingeae and the
baphioids have relatively recent crown clade ages Brownea macrophy//a Photograph by G. P. Lewis Zapoteca caracasana Photograph by G. P. Lewis
(Table 4), compared to the largely S/G-biome cbcles,
e.g. Caesalpinieae sens. lat., the genistoids, dalbergioids
and millettioids sens. lat.; an observation supporting the
findings of Schrire et al. (2005) that R-biome taxa are
derived in the family (Figs. 4-14).
Chaetoca/yx lo .
ng1/oba Photograph by G. P. Lewis Pithece//obium exce/sum Photograph by G. P. Lewis
TRIBES AND GENERA 55
TRIBE
Cercideae by G. P. Lewis and F. Forest
-===~==~;~
Tribe Cercicleae 131 nn 11L2
Tribe Bmtbiuiuae Benih. 0840)
Trine C r id a is basally I ranching iJ1 th
L gumin ae (Brun au el al., 2001; HerencJ en et al.,
2003:1), ::is pr di t -·d by Wund din et al. 1981), and
,'ercis Is th m st basally branchlng genus in th Lrib .
While much taxon mi· work l·l<1 , b n carri d out n
rhe Lril io the pa st thirty y ~ars .g. , L;.irsen et al.,
1 80, 198 ; Wunde rlin , 1976, 1979; Wuncl rlin et al.,
J981 , 1987; Zh~mg, 1 c 5; Vaz, 2 03· Vaz & Tozzi, 2 . ,
fe s pe i s hav ' I e n includ ed Jn phylogen Li
an:ilyses and inter- a nd inu·a~ge m:ri relationships a r
Lill largely unre olv •<l with Lb cprjon of C rci ·
(Hao,, al. , 2001· Davis el al. 20 2b .
Wund rlin J 97 and Wunc.I Jrl in el al. 1981)
di id I th tribe into two sul u·ibcs, Cer idinae and
Ba11hiniint1e, I as don se cl fl ml and rruit ·ha ra rs.
Walpers 1842) b~id already down-ranl e l Bau hiniea
!Jenth . ( I 0) Lo uhtrilx d sr:uus, rhus the ombinatioo
lhuhiniinae B nth. Wun le rlin 1 7 is superfluou -.
P lhill Cl 99'1 J ~pt I he ,er id ae un ·hanged with two
·ubt ri bes a nd five genera . Whil th CercidintJe
c nlalns three small distin t gen ra, 'erck , 'rij]on'ict
a nd Ade11 l bus, th Ba11h1:1iiintP hou ,·e the
m nospc ·1n Madagas :rn g1::nus Br >11i >r a an I th
large divers pantwpical g nu Htutbi11ia sens. !Ctt.
whkb has I e n segr g t cl into a· many as tw nty-six
gene ra by vari )US authors (Wund rlin 1976).
While ma ny of th IUwbini 1 scgr gat s are bas "cl
on minor morphologi ·ll c.liffe ren · s thers are
distinguish <l morphologi ally by a suit of chara te rs.
13rillo n and Ros l . 0), in th ir a· ·ounl or th ~
Cae 'tlpinia ·e:1 r r th orlh Ameri ·an Flora, di id ·d
Bt11tbi11ia into sev ral s gr ga te g n "ra in ·luding
clmel!a Raddi whi ·h h r is Lr :-uecl as a synonym c I'
Phm1era , hut might pr v ~ l<> I e di tin t as indicated
in ree<.:111 molecular amil.yse!'I by r r ·st (unpublished
data . l:MtL n and Killip 093 r ·ognis d 'cbriella as
clistin :t from Bauhinia in ol mbb . 1 Wit 195 ,
tr <i ting 'Malaysian Tiauhinieae', r ·ogn ised
Bra tl/ola nthus, Ly, tpbyllum, Gigasipbon, Piliostigma,
l.astohema and Pbtt1Pra a. separa te gene1 n nd this
Was larg •ly foll wed by subsequent flora writers in
Africa and New uinea (e.g., Brenan, 1967; Coetzer
& Ross in Ross, 1977; Verdcourt, 1979). Others have
retained a more inclusive Baubinia proposed by
Wunderlin et al. (1981 , 1987), e.g., Macbride (1943:
207 -220) for Peru; Larsen et al. (1980) for the Flora
LEFT Bauh/n/a madagascariensis Photograph by D. Du Puy
of Cambodia, Laos and Vietnam; Larsen et al. 0984)
for the Flora of Thailand; Chen (1988) for China, and
Larsen & Larsen in Hou et al. (1996) in Flora
Malesiana. Zhang 0995) published a morphological
cladistic ;.inalysis of the series of Bauhinia sens. lat.,
but few species of Bauhinia have been included in
molecular studies. It remains equivocal as to whether
Bcmhf.ni 1 sens. lat. is monophyletic, but preliminary
molecu lar results indicate that some elements should
be reinstated as distinct genera (Bruneau et al., in
prep.; Forest, unpubl.). This runs contrary to the
findings of Larsen & Larsen in Hou et al. (1996) who
concluded "that Bauhinia in the sense of Linnaeus,
Bentham, De Candolle, T;.iubert and Hutchinson is
an evolutionary unit and a ve1y natural genus''. Larsen
and Larsen also noted that Bauhinia sens. lat.
presents a reticulate p all rn or variation across its
pantropical range (this appar ntly ·onflicting
somewhat with its status as a "n:m1ral gen us ") . While
this is undoubtedly true if the genus is considered as
all-inclusive, recent studies of legume distributions in
genera l (Schrire et al., this volume and 2005) have
revealed repeated patterns of generic distribution
which appear to be duplicated by at least some of the
segregates of Bau.hinia . If these segregates are
recognised as distinct genera (as several are in this
treatment) then the reticulate pattern of variation of
Bauhinia is far less pronounced. More sampling at
the species level in molecular analyses and more
morphological studies are needed across the full
pantropical range of Baubinia sens. lat. before inter-
and intra-gen Jrlc relationships are clearly resolved. In
the current ac ount genera that have been recognised
as distinct from Bauhinia in at least one flora
treatment that post-dates De Wit (1956) have been
treated as separate genera, especially where these are
supported by the preliminary results from a
ch loroplast trnl Cintron and spacer) sequence analysis
(Forest, unpubl.). The reader's attention is also :ilerted
to the detailed infra-generic division of Baubinia by
Wunderlin et al. (1987) in their reorganisation of the
Cercideae which also forms a sound basis for
sampling in future studies.
Palynological studies of Bau.hinia (Larsen, 1975;
Schmitz, 1977; Ferguson & Pearce, 1986) have all
stressed the considerable variation in pollen
TRIBE CERCIDEAE 57
morphology within the genus sens. lat. and there are
clear correlations between pollen exine ornament-
ation, floral morphology and pollination. It remains to
be seen just how closely these correspond to
evolutiona1y relationships of species. Nevertheless,
Schmitz (1977) made several new combinations in
segregate genera of Bauhinia based on palynological
type. These included new names in Lasiobema,
Lysiphyllum, Pauletia, Perlebia and Phanera (Pauletia
and Perlebia here considered as synonyms of
Bauhinia). Zhang 0995), who analysed morpholog-
ically the series of Bauhinia proposed by Wunderlin et
al. 0987), concluded that while some supraspecific
segregates of the genus were supported, none of the·
subgenera appeared to be monophyletic. Several
realignments were proposed.
- Tylosema
I
I Barklya
:
? = position uncertain
FIG. 20 Diagram of relationships in tribe Cercldeae based on a recent preliminary molecular analysis (Forest, unpublished)
58 LEGUMES OF THE WORLD
The Cercideae as presented here (Fig. 20) includes
12 genera and (322)- 335-(348) species. This treatment
differs from Wunderlin et al. (1981, 1987) and Polhill
0994) in that Bark/ya, Gigasiphon, Lasiobema,
Lysiphyllum, Phanera, Piliostigma and Tylosema are
considered distinct from Bauhinia. While some of these
may well be reincluded in Bauhinia after further study,
yet other genera may be reinstated from within
Bauhinia. Bracteolanthus, treated as distinct by De Wit
0956), is here included in Lysiphyllum following
Wunderlin et al. (1987), while Bark/ya, considered
congeneric with Bauhinia by Wunderlin (1979) and
Wunderlin et al. (1981, 1987) is considered distinct
following George Cl998b) and Forest (unpublished
data). The reinstatement of Lasiobema appears least
well supported (Forest, unpubl.).
Cercis
Adenolobus
Griffonia Cerds sll/quastrum Photograph by R.8.G., Kew Adeno/obus garlpensls Photograph by P. Van Wyk
Cercis L 1753
Brenierea n
10 spp,; Eu1asia, China and N Ame1'ica (1 in E USA to NE Mexico, 1 in western N
Ame1ica 1 2 in the Mediterranean 1egion to C Asia , and 6 rest1icted to tempernte
Bauhinia IT1 China in E Asb)
;;o Derived from cerkis, the ancient G1eek name used by Theoph1astu.s fo1· C.
n
Gigasiphon - siliquastrum L.
Shrubs and small trees; warm temperate <ind Mecllterrane<in climate zones of N
0 America and Eurasia in mesophyric forest, meclite11anean bushland and thicket
OJ IT1
OJ )> (chapar1-al and maquis), lower montane forest ;incl mixed deciduous forest
c:
IT1 Refe1ence(s): Hopkins 0942); Li 0944); lsely (1975: 134 -150); Robertson 0976);
::J"'
Robertson & Lee 0976: 48-52); Hao el al. (2001); Davis eta/. (2002b)
::::i A membe1 of subtribe Cercidi11ae (Wunciedin et al., 1981; 1987). In the molecular
OJ analyses of Davis et at (2002b) 1 C. canadcmsis L from eastern N America is 11101e
Vl
I~ ~
closely rehued to C siliquastrum from western Eurc1si::1 than to C. occidenta/is
:J Toirey ex A.Gray from western N Ame1 irn The pseudopapilionoicl tlowers of
\fl, Cercis species sha1e many speci<11isations found in flowers of typic:.11 members of
subfamily Papilionoideae 1 but these are convergences and are not indicative of
Lysiphyllum 0
:--.. dose evolurionc.11y reh1tionship (Tucker, 2002b). Desides, Cercis flowers have
'ascending cochlear' co1ollc1 aestivation, nol 'descending cochlear' as in
I Phan era Papilionoicleae. Neve1theless, the flowers of Cercis differ ma1kedly in st1uctu1e
Lasiobema and development f1om those of Ba.uhinta (Ttu:ke1, 2002b). In the moleculai
analyses of Bruneau et al. (2001; unpubl.) and the combined molecuial'-
?Piliostigma morphological analyses of He>encleen et al, (2003a), Cercis is basally b1anching
both within lribe Cercicleae and the whole of the Legurninosae
Some species, e.g., C. st/iquastntm (Judas tree) OI C. canadensis L. (redbud) are
Detarieae (see page 69) widely cultivated as ornamentals; also used for medicine, human food (flowers,
young pods), bee forage (honey) and timber (the Indians of N America made
Cassieae (see page 111) bows from the wood of C cmwdensis)
Caesalpinieae (see page 127) Adenolobus (Harv ex l3enth & Hook.f) Tol're & Hille. 1955
Mimosoideae Ba11bi11ia sect. Ade110/ob11s Harv . ex l3enth. & Hook f. (1865)
2 spp . (1 wirh 2 subspp )j Ka100-Namib Regiomd Centre of Endemi~m in N:rn1ibia,
Papilionoideae Angola, Botswana and NW Cape Province of South Afri<..."i.1
From ade110- (Gk : gland) and /obos (Gk : Jobes or pocls), alluding to the often
(but not always) glanduhll' f1uits
Shl'tlbs (erect or p1ost1ate) 01 small t1ees; twpical lO subtiopical arid sh1ubland on
coarse gmvel, stony slopes and sandy 1ive1 beds
Refe1ence(s): To11e 0963); Brummitt & Ross (1976); Coetze1 & Ross in Ross (1977:
55 - 58)
A member of subtribe Cercidi11ae (Wunderlin et al. 1981, 1987), and a number of
m01phological characters suggest a 1el<1tionshlp with Cercis , Only recently inc\udecl
in molecuh1r analyses (Forest, unpubl.) where it i.s si.ster to ~1 polytomy containing
Griffoultl, Brenierea, and Bauhinia sens Int
Adeno/obus garlpensls Photograph by P. Van Wyk Both species g1azecl by livestock and game
TRIBE CERCIDEAE 59
 Brenierea lnslgnls Photograph by D. Du Puy Bauhlnla grandidierl Photograph by D. Du Puy Bauhlnia hildebrondtii Photograph by D. Du Puy

Griffonia Bai ll 1s65 Baubinia L. 1753

Ba11deiraea Welw ex Benth. Pt111lelit1 Cav, (1799); Amaria Mutis (1810); Casparia Kunth (1824); Perlebit1 Spix
4 spp.; WC Africa (Liberia to Congo (Kinshasa] and Angola) & Mart (1828), non DC (1829); 'f'e/esfl'it1 Raf. (1838); Alwsit1 Welw (1858), non
Named after M. Giiffon du Bellay, surgeon in the French Imperial Navy, who Welw. (1869); Caspareo}Jsis Britton & ltose (1930)
collected plants in Gabon (around 1850) c 150-160 spp. (plus seve1"l undesc1ibecl), panuorirn l, most abundant in the
Scandent shrubs and lian:is; tropic<.11 humid forest, swamp forest, and rhicke[ in Neotropics (c. 75 spp in S America, c. 35 in N & C America), 32 in Afri ca and
coa.'ital wooded g1asslancl Madagascar (of which 14 endt:!mic to Macfagasrn 1), c 15 in Asia. Section Bmrbinia
Reference(s): Aubreville 0968: 18-23; 1970: 19-23); Lock 0989: 43 -44) c 20 sp p. Mexico, Texas, no1thc1 n C Amc1ica, Gre,1ter Antilles and NE I31:.1zil; sect.
A member of subtribe Cercldinae (Wunderlin et at., 1981, 1987); the inflated fruit Pauletta c 80 in tropical AmericJ and 2 in S Asia, S Ch ina and Malesi;;1; sect.
of some species is a chaiacte1 unique in t1ibe CerclcleHe. Only l'ecently included in Amaria c. 15 f1om N South America to Mexico; secl. Alvesia c 6 in southern
molecular ana lyses (Forest, unpubl.), where it forms pa11 of a polytomy with Af1 ic.:<1, S Asia, S China and M'ilesi,1; se<.:t. Micrulvesia 9 in Africa, S Asia 1 S China
Brenierea and Baubinia sens lat in a clade si.c;te1 r.o Adeno/obus ;md Malesi<:1; sect. Te/est1ia 3 in S As ia; sect. PseudopJ:)(mcra, 2 in S Asia and
Stems used 11s cord age In hw constmction cind baskeuy; used for medid ne as ;1 Ma lesia; sect, Aji·obaubinia c. 20 in southern Aflica a nd Madagasca r)
purgative and a ph1 odis iac, fo1 delousing and as a Cllle fo1 m:iny other ailments; Named fo1 two Swiss botanists, the brothers ] e"n 0541-1613) and Caspar
cnished leaves of two species yield a black dye 0560-1624) Hauhin; the often bilobecl leaves suggestive or a brotherly
re latio nship
Reference(s): Britton & Rose (1930: 203-205, 208- 217, including Alwsit1,
Brenierea Humbert 1959 Casparit1 and Ct1spareopsis); De Wit (1956: 390-4 ·15); 13rcnan 0967: 207- 213);
1 sp.; endemic to SW and S Madagasc;JT Coetzcr & Ross in Ross 0977: 47 -55); Fo11unato (1986: 542 -555, under
Named arrer J. B1eniere, Director of 1he Agricultural Srntion in Betio ky, Bt111bi11ft1 section Paulelia) ; Lock (1989: 4 1- 43); Zhang (1995); Larsen & Larsen in
Madagascar in the 1950's, who collected nowe1ing material which allowed Hou el at 0996: 444 - 456, as Bm1hi11it1 subgenus BtJJ1bi11ia); Du Puy &
1-Iumbe11 10 complete his descriptions ltabevohit 1a in Du Puy el al (2002: 104-125, excluding B bum/1/olimw =
Shrub or uee; largely subtropie<il xe1ophyt!c scrubl<.1nd, o n metamorphic and non- Cigt1sipbo11); Va7. & Tozzi (2003, under Brlllbinia ser. Ca11se11ia)
ca\careous rocks T1ees and sh1ubs (sometimes semi-sc~1ndenr); seasom1lly c111' tropical bushland,
Refe1ence(s): Du Puy & Habevohitra in Du Puy eta/. (2002: 125 - 127) woodlnncl, wooded grassland (savanna and cerrndo), thrn n scrub (including
A member of subtribe Baubiniinae (Wunderlin et al., 1981; 1987); the Gl;Jtinga) :ind coastal forest, t\everal on sand or limestone; less than 20 spp in w et
cl<.1ciodinous habit is unique in tribe Ce1cidcae. The inclusion of Drcmierea in the habitms, infrequently in evergreen humid forest and rain foresl
molecu l:J r analyses of Forest (unpt1bl.) suggests it is cl osely related to BcJJ1b i11it1 Baubi11itJ in its broadest sense luis been divided by varioL1s aL1thors into 26
sens strict segregate genera (Wundedin, 1976). Wunderlin el lll (1981 , 1987) took a broad
view or the genus and placed in synonomy ;ill the seg1egate genera recognised by
De \Xlit (1956). Most recent Aflican Florns have ;1doptcd a less inclusive appio:1ch
which is followed here. Bt111bi11fa subgenus Baubinia was divided into 9
sections, 2 subsections and 19 st:ries by Wunderlin el al 0987) but section
Gigas11Jbon is he1e conside1ed ns a separate genus pending further study. Tht:
neotmpkal taxn included here might 1equire fu11he1 reinstatemern of distinct
genc1e1 hut we await a comprehen sive phylogenetic an<tlysis of the who le genus
to compl e ment the numerous 1egional 1norphologit'al studies
Sevt!wl species (ca mels-foot) used a.s orrn.11nenral s h111 bs ~ind trees of st 1ee1s, parks
and go1rdens throughout the tropi cs; severa l h~vl.:! local medicinal uses; seeds of B
pelerslana Dolle used as a coffee !:iubst iture; the w ood of B acumiHala L (known
a:-> mounrain ebony in Inclia); some .spec.:ies usecl !'01 house posts and fi1c.::woocl,
the hark u.o;ed as rope

Brenlerea lnslgnls Drawing by M. Tebbs Bauh/nia purpurea Photograph by s. A Mori

60 LEGUMESOFTHEWORLD TRIBE CERCIDEAE 61

 Glgaslphon macroslphon Photograph by A. McRobb
Gigasiphon Drake 1902
Ba11binia L. section Gigasipbo11 (Drake) Harms (1917)
4-5 spp.; Wand E (Swahelian Regional Centre of Endemism) Africa 1-2 spp., (1
in Angola, Gabon and Congo [Kinshasa], I in Kenya ancl Tanzania), 1 in
Madc1gascar, 2 in Malesia (1 in the Philippines, and 1 in Timer, Papua New Guinea
and Irian Jaya)
From giga- (Gk.: giant or large) and -siphon (Gk: tube or pipe), in reference to
the very long hypanthium of the flowers
Mostly trees, occasionally large shrubs, I sp. (G. gossweilerl (!3aker f.) Torre &
Hille.) a climber; tropical lowland 1ain forest, sometlmes in marshy a1eas or
flooclplains
Reference(s): De Wit (1956: 418-422); Ilrenan 0967: 204 -205); Aubreville 0968:
24-25); Verdcourt 0979: 110, 112; 1982b); Lock (1989: 43); Larsen & Larsen in
I-IOlr et al 0996: 447-448, as Bcmhinia); Du l'uy & Habevohiti>< Jn Du Puy el al
(2002: 1l6-1!7, as Bauhinia)
Gigasiphou was placed as a synonym of Bauhinia and ~1scribed to an informal
Ilauhinia group by Wunderlin et al 0980, then reinstated at sectional rank in
Bm1binia subgenus Baubinitt by Wunderlin et al, 0987). Du Puy & Rabevohiti:r
in Du Puy et al. (2002) follow Wundetlin et al (1987) in not recognising
Cigasiphm1 as a distinct genus, but commented that Bauhinia httmblolitllUI Baill
(= Gigasiphon humblotiam1m (Ilaill) Drake) b a very distinctive species amongst
Maclagasccin Buuhinias in having a very long hypanthium, 5 rree sepab, and a
small indistinct stigma , It is also l of only 2 Madagascan species growing in
evergr·een humicl forest. Brenan (1967) suggested that the generic position of G
gossweileri needs reconside1ation; Schmitz 0977) placed it in Tournaya as T
gossweileri (Ilak.f.) Schmitz, and Wunderlin el al, (1987) placed It in their series
Africa nae of section Lys1jJbyllum subsection 1'ournaya In the moleculai· analyses
of Forest (unrubl.), G. 111t1cros1pho11 (Harms) Ilrenan is sister to a clade
comprising Tylosema, Bark(yct, Lysiphyl/11m, Phanera and Lasiobema
Gigasiphon schlechteri Drawing by unknown artist
62 LEGUMESOFTHEWORLD
Tylosema fassoglensis Drawing by L. M. Ripley Tylosemo fassoglensis Photograph by A. E. Van Wyk
Tylosema (Schweinf.) Tone & Hilk. 1955
Ba11hi11ia L section Tylosema Schweinf (1868)
4 spp; Africa, mostly in the So1m1lia-Ma!mi Regional Centre of Endemism (2
restiicted to Kenya and Somalia), 1 in the Kalahari-Highveld Regional Transition
zone (Uotsw<ina, Namibia and South Africa), 1 widespread
From tylo- (Gk : knob or callus) and sema (Gk: sign or mark), the upper or
.o.;randard petal has 2 basal callosities
Trailing OJ climbing herbs or lianas from large underground tubers; seasonally dry
tropical wooded g1a.ssland (savanna), gr.assland 1 deciduous bushland and open
scJ'ub, often among rocks (where they are protected from <1nimals that would
mherwise dig up tubers)
Reference(s): Il1enan 0967: 213-216); Cuetzer & Ross in Ross 0977: 61-64);
Lock (1989: 44); Van Wyk & Gericke (2000: 26-27, including photographs)
!lr enan 0967) noted that heterostyly is a feature of the genus. Inclucled in the
info1 mal Phanera group of Baubinia by WLinderlin el tll, 0981), and as section
Tylose11w of Baubinia subgenus Plxmera by Wunderlin et. al. 0987), but
maintained as H distinct genus in most African floras . In the molecular anafy!;es
of Fore.st (unpubl ), Ty/osema is sister to a dadc containing Bark/ya, Lysipbyllum,
Pbanera ~ind Lt1siobema. Based on habitat and geography, however, a closer
link to Ba11hi11ia sensy sl1"ict. and Brenierea, than to Pbanera and allies might
be exrected [Author's postscript: Castro et al. (2005) revise Ty/osema, describing
one new species from Angola]
Seeds and tubers a1e used as human food; T esc11/ent11m (Burch.) A.Schreib.
(nwrnma bean, gemsbuck bean), is highly rared as a survival plant because the
tube1s sto1e water and the seeds form the staple diet of the Kalah<iri bushmen;
seed.s are rich in protein and oil, and when roasted they can be eaten as nuts or
ground to make coffee and porridge; potentic11ly a valuable crnp plant for semi-
arid lands
lYlosema escu/entum Photograph by A. E. Van Wyk & S. Malan
TRIBE CERCIDEAE 63
 Lysiphyllum binatum Drawing by P. Halliday Phanera williamsil Drawing by P. Halliday Phanera bidentata var. fraseri Photagraph by G. P. Lewis

Bark/ya F,Muell, 1859 Phanera Lou r. 1790

Ba11/Ji11ia L subgenus Bark/ya (P.Muell) Wunde11in, KLarsen & S,SJ,arsen (1987) Sc/me/la Raclcli (1820); Cmilolrel11s Rich. ex Schott (1827); /3a11hinia oubgenus
1 sp.; endemic to Queensland (possibly in NE New South Wales), Aust1<ilia Pbcmera (Lour..) Wunc\e11in, KJ,a rsen & S#S,La1sen (1987)
Nomecl afte1 Sir Herny lla1 kly (1815 -1898), Governo1· of Victoria, Australia c. 120-130 spp.; most abundant in S Asia, Malesia and S America (section
(1856-1863) Pbcmera c, 55 spp. in S Asia, Malesia & Chin::1; sect, Auslrocercis l in New
T1ee; uopical vine Frnest and thicket, on sandstone, basalt ~mcl g1•anocli0Jite Guinea; sect. Palmatifo/ia c. 18 in Malesiai sect. T1thicalyx 4 in Indo-Chine.se and
Rcference(s): George 0998c) Malay Peninsulas; sect, Semla l in India, Pakistan & Nepal; sec t. Scb11e/la 8 in
Indudecl in a broadly circumsc1ibecl Baubinia by \Vunderlin et al., as eithe1· the tropical Ame rica; sect. Ccmlolrelus c. 31 in tropical A111e1•ica; .<;eel. Pseudobattbinia
informal monogeneric I3a1klya gioup (1981) 01 as subgenus Barklya \Vuncle11in, l in S China, this hette1· placed in Baubinia based on molecular analyses [Frnest,
K.Larsen & S.S.Larsen (1987), bul consicle1ed w01thy of generic recognition by unpubl.])
George Cl998c) based on a suite of dislinguishing morphological characte1s , From pbcmero- (Gk: evident), p1esumahly alluding to the ve1y showy flowers of
The p reliminary molecular results of Fore:-;t (unpubl.) supprn t Bark(ya :JS a .some .species
distinct genus Lianas find scandent sh!'ub.s (1mely t1ees); p redominantly of trnpical lowland
Bark/ya syringijO/ia F.Muel l. occasionally cultivated as an ornamental evergreen humid or rain foiest, also peat swamp forest, mang1ove swamp,
seconcl::uy sc!Llb and along 1iverbanks, less fiequent in mixed deciduous forest,
heath fo1est, and in d1y forest on limestone, occasionally in liana woodland and
Lysiphyllum Cllenth J de Wit 1956 thorn woodland (rnatinga); the genus has a wide altitudinal i:Jngc; the habitat
Bm1/Ji11ia L. section Lysip/Jyllum llenth. (1865); Bracleola111b11s de Wit (1956)
prefe rence of seveial species is unknown
c, 8 spp.; 4 endemic to Australia, l throughout Malesia extending to N Aust1a\ia
Reference(,): I31'itton & Rose 0930: 205-208, as Sc!JJ1e!la); De Wit (1956,
:rnd Thailand (and prefening a seasonal climate), 1 ende1nic to Thailand, 2 435-530); Vc1clcou1t 0979: 121-123); Vaz 0979; 1993, as Battbi11ia); \Vuncle1lin
et al. (1987, as Baubinia); Larsen & Llrsen in Hou el al. 0996: 4/i2-535, as
endemic in N Borneo (Saraw;1k)
F1om (ysi- (Gk,: loosing) and pbyllo- (Gk.: leaf); the leaves have two free leatlets Baubinia)
Trees, sh1ubs (serni-scandent) and lianas; seasonally d ry t1opical frnest and
Pbcmera w;Js ph1ced as an informal g10up of Bcmbinia by \Vundcrlin et al. 0981)
and as Baubiu;a subgenus Pbcmera (\Vundc1 Jin el al., 1987), the latte! was ft11the1
woodland, vine thicket, floodplains, alluvial flats, tidal forest, scrub, occasionally
.subdivided into 11 sections, 10 sub.sections, and 11 seiie.s. Sections Lasivbenw,
on dunes and crnal islets
Reference(s): De Wit 0956: 431-435); Pedley 0977: 32-33); Ve rdcm11t (1979: Lysipbyllum and Tylosema ate he1e consicle1ecl ::is distinct gene1 '< L, removing
108-110); Larsen & Larsen in Hou e/ al 0996: 501-505, as Bat1bi11ia sect £1pp1ox. 30 spp. f10111 Pbanera, although molecular analyses (Forest, unpubl.)
LysijJbyl/11m); George (1998b, 163-165, as Bm1bi11ia)
suggest that Lasiobema might be best retained within Pbanera
A .second Thailand endemic, Bauhinia slrycbnifolia Cwib, was t1ansfe11ed to Severn] species used ~1.s omamentals, in local medicines (especially !he 1oots), also
LysijJ/Jyllttm by Schmitz 0977) ba.sed on pollen type, but it lacks the two separate
as tying materials in const ruction <.1ncl fo r fish fences
le<1flets otherwise Lypical of the genus and its position is thus equivocal
Lysipbyllum was placed in the infonnal Phane1a group of Baubinia by Wunde l'lin
et al. 0981) and as Ba11binia subgenus Pbcmera section LysijJbyl/11m by Wunderlin
el al. 0987), who fu1the1 divided the section into 2 subsections and 5 selies,
including Se1•ies Ajrlcmwe for the single specie.s B . gusswei/eri (here lentatively
retained in G'lgasipbon, although see note under that genus). De \Vit (1956) st:nccl
th~1t Lysipbyllum winifii (Craib) de \'V'it is clea rly a link species between
Lys1j;byll11m and Gigasipbon In the molecula1 analyses of Bn.meau el al, (2001)
and Forest (unpubl ), and the combined molecular-11101 phological analyses ol
Herendeen el al (2003a), th1ee species of Lysiphyllwn (as Baubiuia in the fiist two
analyses) from Austialia fo rm a well suppolted clacle, hut much wicle1• sampling
anmngst Ba11hi11ia 5e11s lat is requi1ed before relationships can be clearly resolved
Some species lrnve local medicinal uses Phanera sp · l'/8 auh'm1a
. flexuosa) Phatograph by G. P. Lewis
Lysiphyllum cunninghamii Photograph by A. 5. George

64 LEGUMES OF THE WORLD TRIBE CERCIDEAE 65

 Pillostlgma thonnlngli Illustration by 0. H. Coates Pa/grave

Lasiobema (Korth ) Miq. 1855

8a11bi11ta subge nus LasiolJema Kort h (1841)
c 15-20 spp ; Asia (including Bengal, Assa m, Sikkim, Jndo-China, China and
Japan; Malesia 15 spp I in Malay Peninsula, Sumatm, Jaw, Madura , Sumba; 1
restricted to India, but this perh aps misplaced generi ca lly)
From /asio- (Gk.: shaggy) and bema (Gk.: a step or mised rlace), perhaps alluding to
the ladder-like (ster1>e<J) robust lianas which often have ha iry stems when juvenile
Lia m1.s and scandenl shrubs; seasonally dry rropical forest, d1y thorn scrub,
evergreen forest, many on limestone
Refe 1ence(s): De Wit 0956: 422- 431); Verclcourt 0979: 121); Zha ng & Chen
0 994, as secti o n lastobema)
O ne o f the less well ~ upportccl segregates o f Ba11bfuit1; h<:::1e given gene1ic status
follow ing De Wit 0956) and Verdcourt (1979), btr t rerhaps better oeated as an
infmgeneric 1oxon of Pbcmera b;1sed on the molecula1 analyses of f'orest
(unpubl.) Wunderlin el al 0981) placed subgenus lnslobema in the informal
Ph::rnern group of Bt111bi11ia, and larer 0987) placed it as 8<111bf11ia subgenus
Phanera section Lasiobenw ;ind fu11he1 divided ir into three subsections totalling
15 spp. The mo nospecific subseccion Pul/ae \Vundeilin, K .Llusen & S.S.L·usen w:1:-;
rejected from Sectio n lasiobema by Zhang & Chen ( 1994)
Local medicinal uses: some species used to make 1opes

Piliostigma Hochst. 1846

Piliostlgma thannlngll Drawing by 5. Ross-Craig
Ba11bi11ia L secti on Piliosligma las 'Pileoslig11w'I (l·Iochst) Benth. (1865)
3 sp p.; Africa (1 wic.lesp re"d from Senegal and Sudan to South Africa, 1 from W
and C Afrka to Eth iopia <.md Sud<in), 1 widespread from Asia to Austra lia
P1om pileus (L: cap) in 1eferenc.:e to the large peltate stigma
T1 ees and shn1 h!ii se~1 so na ll y dry tropical woocl\and 1 wooded grassland (savan na),
bush land, dry monsoon forest
Reference(s): Milne-l!cc.lhead 0947); De Wit (1956, 530 - 535); ll renan 0967: 206):
Coetzer & l!o" in Ross (1977: 58 - 60); Thulin (1983: 32-33); Lock (1989: 44);
Laisen & Larsen in Hou et a l 0996: 456-457, <rs Battbluia); George 0998b: 162.
as Ba11hf11ia)
A dioecious genus; species from t1opical Amelica tentatively refo1red 10 che
info1mal Piliostigma g1oup of Baubinia by Wunderlin et al. (1981) were excluded
from 8a11bi11ia section Piliostigma by Wunderl in el al ( 1987) Most African fiorJs
retain Pifiostigma '1.!:i <I distinct genus although Wunde11in el al (1987) recog nise
section Pifiosligm(J in Bau/Jinia su bgen us Elayuna . Species of the genus have yet
to he indudc.:::d in molecula1 ana lyses
The bark and roots of P. th on11i11gii (Schumach.) Milne-Recl h, (monkey bread) arc
Jic.:h in tannin and have nume1ous meclicinal p1 ore 1ties, the boiled 1uots yield a
reel dye, th e rmds tJncJ seeds a blue 01 bl<1ck dye, the b;.11k is used fo1 c..:orciage, the
species i."i ;dso ust:!d as livestock fodder <md is alleged to have 1rn1gic.li powers; P
reliculalum (DC.) Mochsl. ;dso has numerous uses acros.'i its 1;,mgc Bouh/n/a weberbauerl Photograph by G. P. Lewis

66 LEGUMES OF THE WORLD TRIBE CERCIDEAE 67

TRIBE
D etarieae by B. Mackinder
Tribe 1>e micac en.~. fat . Polhill 1994
Tnlx: D ' tmieac DC. C1825)
Trlbe ynomctrt::ll! I! •nth. (1840), emend]. Leonard (1957)
Tribe Phyllocarpeae Britton & Rose (1930)
Tribe Amherstieae Benth. (1840), emend]. Leonard (1957)
subfamily Brachystegioideae Hutch. (1964)
Tribe Macrolobieae Breteler pro maj. parte (excl. the Amherstia group)
The Detarieae sens. lat. are pantropical in distribution,
with c. 58% of the genera confined to Africa (incl.
Madagascar), c. 20% to the Neotropics, and c. 12% to
tropical Asia. Only Copaifera, Crudia and Cynometra
are pantropical (and all possibly non monophyletic)
and Afzelia, Guibourtia, Hymenaea, Intsia and
Sindora are native to at least two of these regions. The
apparent high level of diversity in the African tropics
may in part be an artefact of the (relatively) greater
taxonomic effort that has been invested in the study of
the African taxa. Characteristic of African Detarieae
taxonomy has been the splitting off of disparate
elements as segregate genera, while this has often not
been the case in the Neotropics. Both regions,
howeve r, currently contain large paraphyletic
assemblages requiring detailed species-level analysis.
Eighty-two genera and from (729)- 747 - (765) species
are treated here in Detarieae sens. lat. (Fig. 21). Of the
132 (extant) species so far assessed for IUCN reel data
status, 97 have categories of threat. Of these 73 are
assessed as vulnerable, 13 are endangered and 11 are
critically endangered.
The remarkable range and complex patterns of
floral modifications found in the Detarieae sens. lat.
have proved a considerable challenge to the
establishmen t of widely accepted and clearly
circumscribed generic groupings. Based on the work of
Leonard 0957) and Cowan & Polhill 0981a & b), ten
Inf rm:il gr ups of g a ra wcr propo ~ d ; Lh ·
Cynom e Lra , H ym nost i~ , H ym na 'a, Cruck1,
Det~1rium an I nrown e~1 groups in trib Derari a", and
lhe B •rl ir1ia , M;icrolobium , Amh •rslia an I 13ra hystegia
groups in tribe Amherstieae. Genera with imbricate
bra l ·ol ' aestivation were assigned to Detarieae whilst
rh se With valvme bra -r •ole a stivmion w r placed in
Amh 'rstieal!. P U1iJl 199 retain cl th se gen ri ·
group ings with a f a ldlti ns t a ommodaL
recentl y cl •scril l gen ra, and merg d Lbe two tril :
lnio a sing! broadly cl Gned u·ibe, Detari ae en . lat .
Bret l r 199-) proposed the r · ogniLion of Lwo
trlhes Wilh in Detariea ei1 . lat. , s paraLecl according
l lhl' r •!alive size and po itioo of d1e paired I ractcolc
LEFT Ba b
rne ydendran riedelii Photograph by G. P. Lewis
before anthesis. Essentially, this resulted in the
reassignment of Cowan & Polhill's Amherstia group
genera (Amherstia, Tamarindus and Humboldtia) from
Amherstieae to Detarieae with the remaining genera
forming tribe Macrolobieae. In 1999, Breteler (pers.
comm.) proposed a modified Breteler 0995) tribal
system in which Macrolobieae was maintained, the
circumscription of Detarieae was greatly narrowed and
the genera newly excl uded from Detarieae were
together recognised as Cynometreae sens. strict.
The first comprehensive studies of phylogenetic
relationships in tribe Detarieae sens. lat., were the
analyses of Bruneau et al. (2000; 2001), based on
nucleotide sequence data from the chloroplast trnL
intron. They found that tribes Detarieae a nd
Macrolobieae formed a well supported monophyletic
group, which included all genera placed previously in
Detarieae sens. lat., excep t Umtiza. Bruneau et al.
(2000) examined 71 genera, with the African taxa most
widely sampled. The key results of the analysis were
that none of the generic groupings proposed by
Cowan and Polhill 0981a & b) and Polhill 0994) were
suppotted as strictly monophyletic, and the majority of
the members of tribe Macrolobieae (although not
Macrolobium) were placed as a monophyletic group
derived within Detarieae sens. lat. These analyses also
repeatedly recognised a second group of related
genera made up entirely of resin-producing taxa, with
the exception of some species of Guibourlia. The
resins can be seen as translucent gland dots in the
leaflets and (sometimes) other organs. Within the resin
producing taxa, the genus Prioria and several
members of Cowan & Polhill's Crudia group were
consistently placed together (see taxonomic notes
under individual genera in main text). Another
subclade within the resin-producing Detarieae
comprising six members of Polhill's Detarium group
was repeatedly recognised (Bruneau et al., 2000; 2001;
Fougere-Danezan et al., 2003). In addition, in the trnL
intron analysis, the five sampled members of Cowan
and Polhill's Brownea group were consistently placed
together with Macrolobium, although this grouping
TRIBE DETARIEAE 69
 Cercideae (see page 57)
was not upheld in a more recent molecular and base of the clade, and within some subclades, greater Neoapaloxylon?
combined molecular-morphological analysis resolution was provided but as several groups were Schotia
(Herendeen et al., 2003a). Within the exclusively not well s upported , it would be premature to Barnebydendron, Goniorrhachis
African Macrolobieae of Bruneau et ct!. (2001 ), a well emphasise the details of this greater resolution. Brandzeia
supported subclacle of six genera was recognised by Elsewhere the addition of morphological characters
Oxystigma, Kingiodendron, Gossweilerodendron, Prioria clade of Bruneau
Gervais & Bruneau (2002) and as the 'babjit' clacle produced weaker resolution and a less robust Prioria, Colophospermum, Hardwickia et al. (2001) ;;o
rn
sensu Wieringa & Gervais (2003). Bruneau et al. (2000; phylogeny clue either to conflicting phylogenetic signal Vl
Daniellia
2001) confirmed the view of Polhill 0994) that the two or increased hornoplasy in the morphological data or z
both. The order of taxa followed here (Fig. 21) Eurypetalum -0
tribes Detarieae and Macrolobieae (sensu Breteler, ;;o
1995) are best considered a single entity. Evidence represents a synthesis of the present understanding of Eperua 0
0
from ontogenetic studies by Tucker (2000, 2001, putative relationships within this (perhaps most Augouardia, Stemonocoleus c:
2002a) challenged the validity of bracteole aestivation morphologically diverse) tribe in the Leguminosac. Peltogyne, Hymenaea, Guibourtia
n
rn
;;o
as a criterion for subdividing Detarieae sens. lat. and Unsampled genera in the combined analysis arc Hylodendron, Gilletiodendron Vl
identified a set of character states associated with two inserted into this order where morphological evidence Baikiaea, Tessmannia, Sindora, Detarium clade of Fougere -
modes of floral development (Circular and Omega) appears to suggest close relationships. Sindoropsis, Copaifera, Detarium Danezan et al. (zoo 3)
Detarieae
whose distribution amongst cletarioid genera does not Whilst significant progress has been made since sens. lat.
support Polhill's groups. Polhill Cl 994), further studies (particul::trly including Endertia, Lysidice, Saraca, Leucostegane '
The analyses of Herendeen et al. (2003a) united a the non-African members of the larger and less wcl l
Talbotiella
morphological dataset with the chloropbst trnl intron understood genera) are needed before a new
comprehensive classification of the Detarieae sens. lat. , Scorodophloeus
dataset of Bruneau et al. (2001). The combined analysis
provided mixed results within Detarieae sens. lat. based on a synthesis of all available data, can he Crudia

compared with the molecular dataset alone. Near the established. Lebruniodendron 1
Plagiosiphon
Micklethwaitia :'
Maniltoa, Cynometra
Tamarindus
Amherstieae clade
lntsia, Afzelia, Brodriguesia 1
of Bruneau et al.
(2001) Loesenera
Neochevalierodendron
Normandiodendron
Zenkerella
Humboldtia
Hymenostegia
Leonardoxa
Amherstia

Ecuadendron, Paloue, Paloveopsis 1 , Brachycylix' , Brownea clade of

Heterostemon 7 , Elizabetha, Brownea, Browneopsis, Bruneau et al.
Macrolobium pro parte (2001)

Macrolobium pro parte

Paramacrolobium, Cryptosepalum
Dicymbe
s)>
Polystemonanthus n
;;o
Pseudomacrolobium? 0
Englerodendron '
0
co
Anthonotha rn
)>
Berlinia rn
0....,,
Librevillea co
Didelotia ro
......
ro
Pellegriniodendron, Gilbertiodendron ro
lsoberlinia
Oddoniodendron
Microberlinia
? ::=: genus not
sampled in the Julbernardia, Brachystegia, Tetraberlinia, 'babjit' clade of Wieringa
cited ana lyses Bikini a, Icuria, Aphanocalyx, Michelsonia ! & Gervais (2003)

Remaining legume tribes

~1: · 21Diagram of relationships in the Detarieae sens. lat. after Bruneau et al. (2000; 2001); Gervais & Bruneau (2002); Fougere-Danezan et al. (2003);
rendeen et al. (2003a); Wieringa & Gervais (2003)

70 LEGUMESOFTHEWORLD TRIBE DETARIEAE 71

 Neoopoloxy/on tuberosum Photograph by D. Du Puy Schotia brochypetolo Photograph by P. Van Wyk Bornebydendron riede/11 Photograph by G. P. Lewis

Neoapaloxylon nauscherr 1982 Schotia Jacq . 1787

Ajm loxy /011 D1:1ke (1903) c 4 spp.; E Z im babwe, S\V Mozamhiqul", S N:imibic1 , S A f1ica [excludi ng centt--;.ll
3 spp; N, \YI :Lncl S Mc1dagasca1· 1egions] and Swazi lancl; cll ylctncl disjuncti o ns o ccu1 between \'i/ & E Cape [>mvincc.-;
rrnm 11eo- <Gk.: new), bapa/o- (G k.: soft, WL'~tk) ;\ml :\J/011 (wood), rrn ils sofl- and NE Cape-Namibia (S q/ht (L) Thunh.) <tnd bL:tween E Cape and the Sekhu -
tcxtu red wood k lrn nelctml Centre of Enclemi.<;m in Nrnthc1 n Pmvin c.:e, S Afric1 (S latifrJ!ia .J acq )
Trees oi .'i h1uhs; .'ie:ascrnally d1•y lmpical wooclbnd lo xc1nphylit: scniblancl on N~1111 ed :1fte1· Richa1d van de1 Schol , a fwmc1 Chic! Ga1ckncr at SchlJnhiunn
limestone o r sanely so ils lmpe1i<li Ga1den, Vienna; plant co ll e<.:trn , hot;rnic;_il aitist cmd hiend of Nikolau.'i
RcfetenceC-.): Du Puy & Habevohitra in Du Puy el al (2002: 1'18 - 1)2) Jacquin ( 1726-1817)
The biogcographical paltein of this and the following four gene1a appears to T1ces 01· sh1ub.s; t()1est margins , wc>odlancl, sc1't1h frnc:·;L, bushland and tlHJJll

pan:dle l that of the (,"/edilsia clack~ of the Cacsalpinicae; Neoaj){f/o..\.y/u11 (not yet thicket, often on 1ive1• banks 01 on te1 mite mouncl.s
sampled in any molecula1 analysis) and 13randzeirr are from Madagascar, Schotit1 Rde1ence(s): Ross (1977: 25 -33): Estc1 huysc ( 199:1)
occur."i in south{.: Jn Af1il:a and Har11el~yde11dru11 ~111cl Go11 iorrlwcbis arc in the H yh1idisalion occur's between species in the E C 1pt: 1egion; thL: postions of
Neot1(1pics Sebo/la, !3c1rJ1ebyde1uho11 and c,'onirwrhacbis wit hin the 1csin-p1oducing Dc t;11icac
The fibrous hark is used frn rope and the tuhcrow; 10ot.s a1 e edihll" ;11c uncc..:1rn in ; SL'C t<1xo11omic notes undL'1 Neuajulloxyhm
Variou.o; .'ipecies (t1ec..: fuchsia , l)()er-lx::1 n , Arric..::111 \Valnut) ~11c LIS<.:cl as shade t1i.:cs
01 Ollli\lllentillS, browse for li vestock, tirnbe1 (fur nillllC, norn ing, wagons), cclihk
si.:ctls (a fte1 coo king), med icine, tann ins and dyL'S

Barnebydendron.111 Kirkb1 1999

Pbyltow17n1s !lieclel ex Tu ! (IH~j)
l sp; clisju nct 11eot1opiGd clist1iht1ti n n fro m C Am c iica ( t;uatcmab to P~111:1111a),
VL'nczuela, the Purl1s basin in SW Am<1zu nian B111zil and nea1l)y Pcrt1 and Bolivia ,
:incl o n the Atlantic slopes of B1azil f1om S Bahia 10 Hio cle_Ja11ic10
Named afte1 Hupctt C, lfa1ncby (1911 - 2000), at1thrnity in the systematics ul Nc\V
World Lcguminosac (New Yoik Botani c; d (~a1clen) and deJ1dro11 (Gk: tree)
Trees; seasonally d1y tiopical fo1L'.'il, woodland and Atlantic f'o1cst, often on slopes
Hd'cience(.s): Barnehy 0996); Ki1khiicle C1Y99h); \Varwick el fll (suhmillcd)
Bornel~pdendron is a nc\v gcnerk name /'01 · JJbylloc({ljms Hiedd ex Tul which is
a latc 1 homonym of Phyllucmjms Hiedel ex E11cll. Pl~1ccd in a moc.lc1~1tcly .'iupprntc.x.I
dac.le with (,'rmiorrhachis by I-IcrL:ndL'l!ll el al. (200)<1), si rnil:uly (hut without suppt>11J
IJy Fougt::1e-Da 11ez<1 n el al . (2003J; see taxonomic note.'i undc1 NeoajHt/O_\y/011
!Jar11ehyde11dro11 riedelii tTul.) j.HJ\:irkb1 (gu:uihd1 0, gu<tcamayo) is cultivated <1.'i
an u 1n;.1mental in .'i1>uthe1n f1lrnicb, C::1ril1l)c.::1 n, C Amct ica and in 'J'ha ilanc.1

Goniorrhachis Taul> rnn

I sp ; ll 1<1zil (Bahia)
F1om -go Hia (Glc: angle) and rbacb is lC;k ,: ;ixis), foi the 'zigz<ig' <.1ppca1;:11u.:c of
the innrn cs<.:cncc 1hachis
Tn::cs or."ihrubs; sc~1somilly dry tropical wood land, wooded g1a..,~land ~111d sciuh
Rcle1e11ce(s): Cowan (!'J8lh); Lewis ( 1987: 11)1)
Sec taxonomic notes under Neoapalo.\Tlon and Jkt1 ·11d~)!dt.!11dro11; two va1 ·ietiL:s,
one :l sh1uh and one a tic~. occw in Bahia , although lhL:.'iC inf1~1..,pecilics may not
Neoopoloxylon madogoscoriense (A-E); N. tuberosum (F-L); he \Vo11hy o f 1ecognition
N. mondrorense (M-P) Drawing by M. Tebbs Goniorrhochis morginoto Photograph by G. P. Lewis lbcc l l'rn ti111he1 (but known to C:lllSC a[IL:1gic ICSj)OllSCS)

72 LEGUMESOFTHEWORLD TRIBE DETARIEAE 73

 Brandzeia filicifolia Photograph by D. Du Puy Klngiodendran alternifolium Drawing by F. Owner Gossweilerodendran balsamlferum Drawing by M. Boutique

Brandzeia Baill 1869 Kingiodendron Ham1s 1897

Batbiaea Drnke (1902), under which it W<IS known until Du Puy & Habevohitl<:l in 6 spp; Asia (\Y/ Peninsular India, Malesia [Philippines and Moluccas], Papuasi::i
Du Puy et al (2002: 146) found the earlie1· name Bm11dzeia [New Guinea ancl Solomon Islands] and the Pacific [Fiji Islands))
1 sp.; N and W Madagascar Named fo r Sir Georgl' King (1840-1909), Superintendent of the Bot<:mical
Named frn P1ofessrn• D. Ilranclza 0846-1895), an authrnity on the the1·apeutic Ga1clens, Calcutta and P1ofes.so1 of Botany, :.rnd dendron (Gk: t1ec)
p1ope1ties of Gentianaceae and founde1 of the botanical gardens in Bucharest Trees; t1opical lowland 1•ain forest, often along rivers rn on flood plains
Tree; season<:lily d1y tl'Opical woodland, p1 incipally on limestone, sometimes in Reference(s): Knaap van Meeuwen (1970: 46-50); Vcrdcou1t 0979: 93-98); Hou
seasonally flooded a1eas in Hou, el Cil, (1996: 627-630); Il1eteler 0999) uncle1· PrioriCI
Reference(s); Du Puy & Rabevohit1a in Du Puy el al (2002: 144-148) See tc:1xonomic notes unde1· Oxysligma, from which it is sca1cely distinct (Cowan
Placed as siste1 to the 1est of the Deta1ieae se11s strict clade (without .suppo1t) by & Polhill, 1981<1) Initial tesults indicate that Kingiudendron is not monophylctic
Fougere-D£1nez:rn et al. (2003), but also ,o;;ee taxonomic notes uncleJ but fu1ther sampling is needed to resolve gene1 ic delimitation
NeoapaloAJ'lon Ki11giode11dron }Jinualum (Roxb,) Ha rms (pula, kulavu em1) is used fo1' limber
The wood is used in house construction (construction and fumitu1'e), rneclicine, and exploited for oleo-resins (balsam)
which are an ingredient of incen/'ie

Oxystigma Haims 1897

Pte1ygopodi11111 Haims 0913) Gossweflerodendron Ha rms 1925
5 spp; WC Africa (4 spp. in Nigel'ia, Cameioon, Equat01ial Guinea, Gabon, Congo 2 spp; WC Af1ica (Nige1ia, Came1oon> Equato1ial Guinea, Gabon, Congo
[Il razzaville], Congo [Kinshasa] and Angola) and E Africa (1 sp , Kenya and [Kinshasa] and Angola)
Tanzania) Named for .John Gossweiler (1873-1952), an autho1i ty on the flo1<:1 of Angola, and
From o.'\y- (Gk ,: shaip) and stigma, fo r the pointed stigma de11dro11 (Gk,: tree)
T1ees; tropic~d lowland rain fo1est 1 often along 1ive1s, mang1ove swamps and T rees; t1opic::il lowland everg1een rain forest
ground-water forest (in E Af1 ica) llcfcrence(s): Aub1eville 0970: 146-148); Bretele1• 0999) u11de1· PrioriCI
Reference(s): Leona1-d 0952b: 369-375); Aub1eville 0970: 127-131); Il1etele1 See taxonomic notes unde1· Oxysligma
0999) uncle1 Prioria Gossweilerode11dro11 balsamij'erum (Vennoesen) Hai ms (agba, emonga, tola
Scarcely distinct from Ki11giode11dro11 (Cowan & Polhill, 1981a); B1etele1• (1999) bianca) p1oduces <I gene1al pu rpose timber treated as a substitute fo1· 1rn1hogany;
unites O:.:ysUgma, Gossweilerodendron, Kingiodendron and Pte1ygopudi11 m also usecl as a source of oleo-resin.s (balsam)
(included in 0.~ystigma by Cow'rn & Polhill, 198la; Polhill, 1994) with Prioria
Molecular evidence (Bruneau el al., 2000; Fougere-Danezan el al, 2003; He1endeen
el al., 2003a) suggests that the 111embe1s of this g1oup are closely 1elatecl but fu1the1 Prioria G l'iscb . 1860
sampling is needed to assess generic delimitation. O~\.:ysligma and the next fou 1· 1 sp.; coastoll a1e~1s of C Ame1'ica (Nica1agua, Costa llica and Pomama, .south to
genera constitute the well suppoitecl P1 ioria c\acle of Biuneau el al, (2000), which Colombia); also cultivated in the Ca ri bbe~m
togethe r with the Detarieae sens. strict clade (from Daniellia to Delarium) comprise N~unecl fo1 H.icha1d Chandle1· Alexancle1 Pl'iol' (1809-1902), British botanist
the resin-p1oc\ucing members of Deta1ieae sens lat. Initial 1esults indicate that Trees often in rn.:atly pu1e stands; coastal swamp fewest, along livct cstu;.uies and
Oxysligma as currently ci1cumsclibec\ is not monophyletic in seasonally inundated a1eas
Used for timbe r, e.g ., 0 oxyphy/111111 (Harms)] Leonard (tchitola), in furniture, Reference(s): ll1eteler 0999)
cabinet work, joineiy, deco1<1tive venee1s, canoes, construction and household See taxonomic notes under 0."Cystigma
utensils; also used for medicine, shade trees, bee forage <incl as'' sowce of oil Used fo r timbe r, e g» P. copaij'era Griseb. (cativo, camibar•) fo r interio1· t1 i111,
(balsam) in the preparation of shellac fu1 niture, cabinet work, joine1 y, venee r and plywood; also used few medicine and
as a sou rce of oleo-resins (balsam)

Priorio c ,,
Oxystigma msaa Drawing by L. M. Ripley opa1,era Drawing by W. H. Fitch

74 LEGUMES OF THE WORLD TRIBE DETARIEAE 75

 Calophospermum mopane Photograph by P. Van Wyk Don/ell/a olsteenlono Drawing by M. Boutique Doniellio oliveri Drawing by L. M. Ripley

Co'lophospermum 1.Ki1k ex) Leonard 1949 Daniellia Benn. 1854

1 sp.; South T1opical and sou thern Africa (in lowe1 alt itude rive 1 va lleys from S c. 9 spp.; Africa (cent1ed in Gt1inea-Congolian WC [7 spp J and W (2 spp.I regions,
Angola, N Namibia, N Ootswana 1 Zimbabwe, S Zambia, S Malawi , N South Africa with 3 spp extending to the drier Sudanian and Zc1mbezian centres, south to
and Mozambiqt1e) Angola and east to Sudan and Uganda)
Alternative etymologies a1e: from colopho11i11111 (Gk,: resin) and -sperma (Gk: Named after Dr W.F. Daniell (1818 - 1865), British milita1y surgeon and collector
seed), eluding to the numerous scattered 1esin glands which cover rhe seed, rn of plants in Senegal and Sierra Leone
from colopho- (Gk : top) and -sperma (Gk.: seed), in 1eference to the above Trees; tropical rain frnest, often in swmnpy and inundated areas, and 2 spp in
g1ound gel'l11ination of seeds whilst they 01re still in the pods seasonally dry forest, wood land and wooded grassland
Trees or shrubs; seasonally chy tropical woodland, bushland and thicket Reference(s): Leona1d 0957: 110 - 115); Aubreville (1970: 162 - 170); Blllkill
Reference(s): Mapaure (1994); Ti mberlake 0995; 1999); Bretele r et al. (1997); (1995: 93 -100)
Leonard (l 999b) Placed as basally branching within the Detarieae sens strict clade by Ilruneau et al
Colopbospem111m mopc111e (J Kirk ex Benth.)J Ki1k ex j .Leonard (mopane) (2001); the fruit is a rnre example of a 'hemi-legume'; tile seed and endoc;up of
woodland is dominant over an aiea exceeding 50,000 km2; the species is one valve foll together as a uni£i genus currently under revjsion
g1egarious and thought to be wind-pollinated A proposal by Bretele r et al. 0997) Used as timber, e g., D oliveri (Rolfe) Hutch. & Dalziel (West African copal , gum
to unite Colopbosperm11m with Hardwick/a was contested by Leonard (1999b) copa l, ogea, fo 10) for plywood, joine1y, furnitu re components, boxes, crales
and Smith el al. 0998), but their unification is a va lid phylogenetic hypothesis as and decorative veneers; fibre (for paper), copal-resins and gums (used in boat
clemonst1ated by Fouge1e-Danezan el al (2003) and supported by studies of bL1 ilc\ing, preserving 1ailway sleepers, as medicines, glue, va rnishes, polishes,
pollen morphology (llanks & Kli tgaard, 2000) and chromosome number cosmetics and for ceremonial purposes), cha 1coal, reaffo restation and as
(Goldblatt, 1981). In the analysis of Fougere-Danez.111 el al. (2003) , shade tiees
Colopbospen1111111 and Hardwick/a together fo11n a strongly supponed clade
placed as .:;ister to a clade complising representatives of Gossweilerodendron,
Oxysllgma, Prioria and Kingiodendron Eurypetalum Ha11ns 1910
Used for timber (constniction, fencing posts, cra fts), fuelwoocl, livestock browse 3 spp; WC Africa (Ca meroon, Equatoria l Guinea and Gabon); may occur in
and fodde r, human Food (edible moth la!V'ac called mopa ne worms), medicine, Nigeria (Keay et al., 1964)
tann ins, 1esins and soil srabillsation; the ba1-k is a sour<.:e of fibl'e From e111y- (Gk .: wide) and petalo11 (Gk.: petal) for the well-developed abaxial
petal which is wider than long
Trees; tropical lowland min forest
Hardwickia Roxb. 1811 Reference(s): Aub1eville (1970: 108-110); Breteler & Obiang (in prep.)
1 sp,; in relatively isolated blocks in the drie1 parts of Peninsular Jndia (map in Placed in the Detarieae sens. strict. clade (Bruneau et al., 2000; Fouge1e-Daneza n
llreteler ef al (1997)) et al., 2003) and strongly suppo1ted as sister to Epema; ft11ther sampling,
Named for Thomas Hardwicke (1757-1835), Blitish bota nist however, is requiled to confirm rhis affinlty; genus currently under 1evision
Trees; grega1ious in seasom1lly chy tropica l forest, woodland and wooded Used for ti mber, e g, E batesii Baker f (anzilim) and as a source of a 1ecl dye
grassland
Refere nce(s): Knaap van Meeuwen 0970: 50-51); ll1eteler et al 0997); Leonard
0999b)
See notes unde1 Colopbospermum
Htu-dwickia binala noxb. (anja n) is used for timber (const ruction and ag1icultt11,.1l
implements), g1een manu1e and livestock fodde1 1 fuelwood, resins (as a wood
bark r1 oduces a st 1ong nbre used fo1· ropes
p 1eserver) and the

~~'Y~etolum unl/ugum (1 -4); E. tessmonnl/ (5); E. batesli (6-11)

Hardwick/a binata Illustration by unknown artist awing by/. Soussotte-Guerel

76 LEGUMES OF THE WORLD TRIBE DETARIEAE 77

 Eperua grandiflora Photograph by S. A. Mori Augouar<lia letestui Drawing by/. Sausotte-Guerel Stemonocoleus micranthus Drawing by/. Sausotte-Guerel

Eperua Aulil. ins Augouardia Pclil'g1 . 1•12.1

l 1 i ..;pp.; E J\m:1zo 11i ;111 S A1rn..:ri e:1 (l\1.1zil , Cui;111 :1s. E VL'llL'Zud:1 and N l ~ I sp .; \VC: Al1h.1 lC~:tlxrnJ
Oil1unlli:1J N:um.:cl al"lc1 Mon sdgnot J\ugoua1d, Ard1hi.-.hop of Congu t lk.1zz:1villL•J in 1l1c
Frt1111 l'/><'rU , :t lrn:al (;uian an ( lrnlian } 11:1111c !"or lhL' ,-;li:1p1..• nf' the !ruil of/:" fitlt :!lfl late IHOO's
Auhl, whic h rl.! ... t:1nh lcs :1 sah 1c '1'1t..:L'; !IOj>icli lmvl:t ncl 1<1ill Crnvst
'['1 c.:i.:s, c11"tc11 in 1111w1odo111i11:111l sl:tnd"; lJ1\1Ji t<1l lci wl;1m l 1:1in f'll1L'~I , c)f"icn :1lcmg l kl'~tL'ntl!lsJ: Auh1Cvillc (lCJ(iH: 182 - JH~)
1ivcrs and in inumlatl!cl an:a-;, :rnd in se.1,..,on:illy d1y l"on.: ...;t, woodl:1ml .'iciuh :llld l'l :H:1..:d in the f)cl<1liL·:ic ~t· 11s slrhl , eb<lc hy Bn1nc..::iu el al CWOO) anti sl1ongly·
\\CH>d i.:1..I g1:1...;sla11d supprntc c. I a-. sisll'J tu S1< muJ1oc.:oh.'11s hy 1:ougC:.· 1c- Danez:in et al (200.1)
1

JkJt..: 1cm·dsJ: Cu\v:tn ( l ~J7S I>); Cmv:1n .._\ lkrry in Bl'l'l'y t'/ rt!. ( J1J1)8: (vl-(17); l).11ilc-;
0:.: Hico ( l t)lJ<))
S1..·c: nut''·" undc:1 h'ffly/iet11/u111 Ste111011ocoleus 11 :11111s 1•Jos
Planted :is rnn;111K·nt:tls; .'ii:vc ul spi.:l' ics (wall:1h:1, ap:1, w.1m<11<1) a1 L' used 1"01 l .-.p.; \VC Af1·ic.1 (Jvrny Co;tsl , ( ;h;1n;1 1 Nig1.:1ia, C:1mc1oon, C:1bon :inc.I O ..·n!i;i]
ti1nlw1 in lu.::iv y to nst ru t't irm , 1oors (.-;hinglt:"). c1 1pc n11 y, j<iirn.:i y, fu1nilllrL: :iml Afr'ic..": m lh::pulilic)
~l!lClling; ro1 (i1 1.;worn l :ind d1;moal, :rntl rm n:"in ()I' w.il l;1b:1 oil (e.g.,"'" oh'!l;'l d l'rolll S(t!ll/UI/ ((;k ,: .'>!:imcn) <lllcl c;pfoo~ (Ck: .-.h1.:alliJ , Cor llie prolonged Sl:1111in:d
l>udu:) di sk to which lh12 fib11K 11l-; art; :ilt ;td1L'd
0

' [
0
/L'CS; !ropk:tl lowland r;till rrncsl
l kk:1L'11n.~(s): l,,l'.011<.1rd ( l tJ57: I 'iC1 - t 'i~); Auh 1t.'"vill..: l 1970: 17,f -l7(l)
l'l:lc.. ..:d in thi...: l1i..:t;11iL':1i...: se11x. strhl . l•hc.lc l1y B1urn..::1u l'I ol (2000) :111c1 sl!nngly
.-.upprntl'd a.-. si ...,lt..:1 to /wgulfrtrr/i(I h y Fm1gl'tl~ - n:111cz;m t'I a( (200.))
Sten w11rn. :11fv11s 111it:rt111tb11s 1 latms lahi:in:m:I) is llM.; d rm li111he1, JC.'iill :me.I
nh: dicint..:

Peltogytte Vogel 18:17

c..: . 2'i :-. pp.; cc nl1L·d in A111:tzoni:1n S J\1n t..: 1il,1 lllr :1zil [al.'in A!l~1nlic fort:sl :ind c .1
spp 10 th e c.l1il'1 Nl~J, (;uia11:1 <..;, Vent..:zuda, <:c,lcimhi:1 :ind nolivia), 2 spp. in C
A111c.: 1i<.: a, one extending lo Mex it(); 1 lllL' S J\nie1 ica n .'iJllX.'.il's Lo T1 inic.lad
[!1 o rn pelto (Ck: Slll:ill l'OllllCI ,-; liiekl) :ind ,'{Jlllti· (Ck,: l'L'in:tle), rrn·
the .... h<.lflL' o!
lilt..: .'iliglll<l
'l'rt..:cs olkn in monrnloinin:inl .... 1:111ds; typically in 11,.1pil;ll l!1wla11d 1:1in frnL' ....,\,
ollcll :llnng tiV<.'1.'i; ll''iS t·oi111ncmly in SL':t-;on:dly c.fty l"rnL'Sl, wootll:tnd OT" lhnm
.. t:l'llh
l<l'ktt:m.d ...;): Dw ki: ( 1919: 99 -102 ); Si lv:1 ( 1CJ7h); Cowan & lkny in B1:11y C'/ ({/
( [<Jy8, '!.i-'J7)
11 1<11....:d in thL' DdatiL':tc ·'"'If"
sllitl ~ t:l: 1dc hy Biunt'ill1 l'I al (2UOO) ~me.I fo1111s a
wdl suppo1tL'd suhcl:1cll' (with (/u i/Jo11 rtia ~1ml /~ )lme1wer1) in lll.'.1c11dt..:cn ('(al
llOllb), a 1c lalionsl1ip :!111..::ic.ly imli<.:alctl hy pollen 111rnplmlog)1 (Banks 0:.:
l.\.lit ~ aa1d, 2lJ00i
SC\'L'l:il spetic..; :tie Ulilisnl ror thei1 high quality ti111IJc1 (:1111: 11 ':.llHli, a111;11alllL',
pu1plchc:ir!, violclwood, p:tu 1oxo ), L' 1g, in l'u1niLu1 i..:, c :1hinet wo1·k, lloo1i11g,
to11s 11u ct io11, lt11'1le1y, clec01.1tive v~llCL'l.'i, 1rn.iH1u eliy, 11n1...;iul inslt'umenls
:ind L:irving
Pe/togy111• 11 "fl
Eperua jenmanii subsp. sandwithii Photograph by P. /. M. Maas · auc1 ora Photograph by G. P. Lewis

78 LEGUMES OF THE WORLD TRIBE DETARIEAE 79

 Gulbauttia coleospermo Photograph by P. Van Wyk Gu/bourtlo co/eospermo Photograph by P. Van Wyk
Hymenaea courbarll Photograph bys. A. Mori

Hymenaea L 1753
Guibourtia Benn. 1857
Pse11docopaiva Boitton & Wilson (1929)
'{iy1 by/v//111111Ilny nl!I 1827)
c. 14 spp.; Africa (Guinea-Congolean WC Africa 17 spp I: dryland S tropica l
14 •PP·' 1101l 11 S1\111,•1ir.n wlt l1 tl'.:nlf!S Jn A111no11in and coasml A1inn1 c forest (c
Zambezian (c. 3 spp.) and N tropical Sudanian (1 sp.) regions, and c. 2 spp. in
s pp.) nnd Lirylan I 1l 1n ' J! !31 .11.il (5 ~pp.; nc of whk ll c>.tcnds IO l':im11u11 y
"'P'"""
1111 tl Argl!lllln:t); I ~l'· wh.I In 1h • N1!1~ropic (c.<1e11Llini: to • Amcrk.1. the
Zanzibar-lnhambane E Africa), with a single neotropical species disjunct betwe en
Cuba and seasonally dry E llrazil, Paraguay and Bolivia
C:irlhhe:m um.I M,c.<1wl; I 'I" e ndemic ltl ub:i , '1 111L·d 10 n •in11lc spc<it:'S 111
Named fo1 Nicolas Jean BG . Guibou1t (1790-1867), Professor of Phaimacology
coaslal E Africa, Madagascar and 1he Mascarene Islands
Derived from Hymen, the Greek god of marriage, referring here to the twinned in Paris
Trees or sh111bs: tlopical lowland (some1imes swampy 01• seasonally inundated)
pair of leafle1s comprising each leaf
rain forest, seasonally chy forest, woodland, bushland and thicket, often along
Trees or shrubs: iropical riverine and inundated forest to seasonally dty forest,
rivers and on sandy soils
woodland, thorn forest, bushland and thicket, often on slopes
Reference(s): Leona1d 0950: 1957: 137-156): Boenan (1967: 136-138); Aubreville
llc fc1"'0Lc(.'I): Laogcnli ·im & Ll>c (\97·0 : I.cc l.•ni: •nh •11 11 Ci• 751
Modcra1cly "ell upporll!d as sl.,lc r 10 ,uf1J011rtla h l.lc remk-c n <'I <II. lll03a) (1970: 120-127): Lock (1989: 53-54); Barneby (1996: 182-183)
Moderately we ll supported as sister to Hymenaea by He1endeen et al. (2003a):
olthouj,111 :mn plini: lncu111pl<-'lc: Poillllr llll'J I I tlcscribc.."Cl n f0>:<il spc es t ll
see taxonomic notes under Hymenaea Barneby (1996) united a number of S
j/m/~rrt J•o lnllr), fl (llll th ' l10111111il.lll itc pul Jj wh ich h ' C( llsitfco'Cd ·loscJy
American species occupying discontinuous, seasonally dry distributions under che
resembled the African H. vern1cosa Gaertn
single species G bymenaeifo//a (Morie.)) .Leonard

0
Resin (referred to as S American or Zanzibar copal, depending on the species) is
used for incense, glue, varnish, shellac and as traditional medicines; other uses
Various species used for limber (bubinga, akume, ovangkol, hyedua), e.g. in high
quality furniture, cabinet work, joine ry, panelling, veneers, heavy caopentry,
include edible fruits, timber (algarrobo, jatoba, courbaoil, guapinoD for
consmoction, high quality furniture, cabinetry, veneers, joinery, panelling, turnery. implement handles, boat masts, for firewood and charcoal; gum-copal (for
incense, coating pills, varnishes, illuminants and mosquito repeJlents), medicine,
111u<lc11I instruments and boat building, the bark is used for canoes, and some
fish poisons and (in at least one species) edible seeds
species are cultivated as 0 1namentals

Guiboutt/a schlelbenll Drawing by E. M. Stones

Hymenaea verrucasa Drawing by L. M. Ripley

TRIBE DETARIEAE 81
80 LEGUMES OFTHE WORLD
 Gllletlodendron p/e"eanum (1-4); G. mlldbraedll (5-6); G. klsantuense (7,8)
Drawing by M. ·F. Adam
Hylodendron Taub. 1894
I sp.; WC Af1 ica (Cameroon, Gabon, Congo (Kinshasa] and Nigeria)
From bylo- (Gk.: forest) and dendron (Gk.: tree), 1efeiring to the habit of the
species
Trees; t1opical lowland rain forest
Reference(s): Leonard 0957: 75 - 76); Aubreville (1970: 140 - 142)
Position unresolved within the Deta1·ieae sens strict. clade (Bruneau el al., 2001)
Used for timbe1 (carpentry, implements and construction), fuelwood and as a
traditional medicine
Gilletiodendron Vermoesen 1923
5 spp ; centred in WC Africa ( 4 spp. Ivory Coast, Cameroon, Centra l African
Republic, Equato1 ia I Guinea, Gabon, Congo (Kinshasa] and Angola); 1 sp. in Mali
Named for Abel Gillet (1857 - 1927), French botanist, and de11dro11 (Gk : tree)
Trees; t.ropical lowland Guineo-Congolian rain forest and Sudanian seasonally
dry forest
Refe1ence(s): Leonard (1952b: 283 - 287; 1957: 61 - 66); Hutchinson & Dalziel
(1958: 455); Aubreville 0970: 71 - 75)
Position unresolved wilh in the Detarieae sens s/l"/cl. clade (Bruneau et al., 2001)
Used for reafforestat ion, timbe1 (house construc1ion) 1 1esin, edible seeds and
bee plants
Baikiaea Benth 1865
c. 6 spp ; centred in WC Africa (4 spp., Gabon, Equatorial Guinea, Congo
[Kinshasa] to Angola; B inslg11is Benth widespread in the Guineo-Congolian to
Lake Victoria regions); 2 spp. in drier habitats, one in lowland E Tanzania , the
other widespread in southern tiopical Africa
Named for Dr William Balfour Baikie (1827 - 1873), Royal Navy Surgeon, explorer
and missionary in West Africa
Trees; tropical lowland rain forest, seasonally dry forest and woodland on well
drained soilsi B plurijuga Harms forms near-monoclo1ninanl scands on the
Kalahari sands of the Zambezi Basin
Reference(s): Brummitt (1986: 61 - 73)
Placed in the Detarium clade sensu Fougere-Danezan et al (2003) within the
Dernrieae sens. strict clade
Used comme1 cially for timber, e g., B plurij11ga (Rhodesian teak, Za mbesi
redwood, utngllsi), maJnly for Roaring, furniture, c~binerry, veneer, turnery,
ca1ving and construction; also cultivated as an ornamencal (.Incl introduced in Asia
(B. lnslgnfs, or nkobakoba)
Sindoron I I
Balklaea plurljuga Photograph by G. R. Nichols
82 LEGUMES OF THE WORLD
Tessmannla macrontha, ined. Photograph by J. J. Wieringa
,.s s etestu/ Drawing by M. -F. Adam
Slndora cochlnchlnensls Photograph by L. Averyanov
Tessmannia Hanns 1910
c 12 spp ; centred in WC Af1ica (c 7 spp. (Cameroon, Central African Republic,
Equatorial Guinea, Congo (Biazzaville), Gabon, Congo (Kinshasa) and Angola]
ancl 1 sp, in W Africa [Sierra Leone, Liberia, Ivory Coast]); 2 spp. in Zambezian
south tropical Africa (Tanzania, Zambia and Angola) and 2 spp in Zanzibar-
Inhambane E Tanzania
Named for G. Tessmann (jl. 1904-1926), German missionary, ethnographer and
botanical collector in Equatorial Guinea and Gabon
Trees; t1opical lowland min forest, seasonally dry forest and woodland
Reference(s): Leonard 0957: 66-72); Brenan 0967 : 106-108); Aubreville (1970:
155-161)
Placed in the Detarium clade sensu Fougere-Danezan et al. (2003) within the
Detarieae sens. strict clade; the south tropical and E African taxa are poorly
known; genus currently under revision
Used for copal (resin) and timber (carpentry, railway sleepers, tool handles, posts)
Sindora Miq. 1861
c. 18 - 20 spp.; SE Asia (most diverse in Malesia with c. 15 spp., 2 spp. e ndemic
to lndo-China); WC Africa (a single species in Gabon)
From slndor, the Malay name for sever•! species of the genus
Trees; tropical lowland rain forest, seasonally dry forest and woodland, along
1ivers, in swamps, occasionally on hillsides, often on sandy soils
Reference(s): De Wit 0949); Hou in Hou el al. 0996: 691 - 709)
Placed in the Detariurn clade sensu Fougere-Danezan el al, (2003) within the
Detarieae sens, strict. clade
Various species are used for timber (supa, sepe tir, pctir) fo1 construction,
furniture, joinery, veneer, plywood, flooring, ship building •nd utensils; also for
medicine and oleo-resins (used in perfumes, for illuminacion, varnishes, paints,
transparent paper and for the adulteration of other oils)
Sindoropsis ) .Leonard 1957
1 sp; WC Africa (Gabon)
From opsis- (Gk.: appearance) and Sindora, for its resemblance to this closely
related genus (although it differs in petal indumentllm, androecium structure and
fruit morphology)
Trees; tropical lowland rain forest
Reference(s): Leonard (1957: 81-84); Aubreville (1968: 171-174)
Placed in the Detarium clade by Fougere-Danezan et al. (2003) within the
Detarieae sens. strict. clade; first described in Data1twn and then Copaifera before
being rJ.ised to generic rank
Timber of S letes/111 (Pellegr.)] Leonard (gheombi) is used for furniture, flooring,
joinery, veneers, boxes and crates
TRIBE DETARIEAE 83
 Endertia spectabilis Drawing by unknown artist lysidice rhodostegia Drawing by P. Halliday
Copoifera sp. (Carvalho 2174) Photograph by G. P. Lewis Copalfera palustrls Drawing by P. Halliday

Endertia Steenis & de \Xlir 1947

Copaifera r. 1762 I sp; Malesia (13orneo IKalimantanJ)
Pseudosindom Symington 0944) Named foi [)1 F1edelik Hendrik Enclen (1891-1953), Dutch (Java born) botanist
c 35 spp; must diver:-;e in S America (cenl1•ed in B1'c12il) with c. 25 spp . (c 10 spp and authrnity on Malaysian trees
in Amazonian Guianas, B1azil, Venezuel<1, Colombia :irnd Bolivia; c 15 .<;pp in T1ees; tlopical lowland forest, somelimes occu1ring on sandy, loam soil 01 on
dric1 h;ibitats in B1<1zil to Bolivia and Pa1aguay); 3 spp . in the C.11 ibbcan (one limestone
disjt1nct to E B1azil, one e xtending from Panama to Venezuela); 2 spp in C Reterence(s), Hou in I-Jou et al (1996, 616-619)
Americ,1 (Panama and Costc1 Hic.i) 4 spp. in Af1ica (3 spp. in \Y/ and WC n:gions Hnderlia and the following two genera are placed in a strongly supported clade in
~ind 1 sp in Z:rn1bezian south tropic.ii Aflio1 [Angola, Zrnnbi<1 Hncl Congo the analysis of 13runeau el al (2001)
(Kinshasa)}); a single ..;;pecies o<.:cu1s in Malesia (Urn neo) Hnderlia spectahi/(,· Steeni.s & de \'\fit (sua'alat) is used frn timbe1
F10111 copailJa (Anrnzonian Indian name frn this rree) andj(!r. (L,: rn11ying),
1efc11ing to the resin of Cojx1ifera t1ecs
Tree.., 01· shrubs; tropical lowland 1ain fo1~st (sometimes inunclalecl), seasonally Lysidice 1-l;mce 1867
d1y forest, thorn forest, woodland and sh1l1hl:md
2 spp; Inclo-China (Vietnam) and S China
Refe1ence(s), Dwye1 0951); Aub1eville (1970, 131-135); Enrech et al 0983); !loll
Ftrnn /ysi- (G k.: loosing) and di- (Gk : two), few the two f111it va lvt>s that curl back
(1994, 320-321) at clehiscence to 1elease rhe seed.s
Placed in the Dctarium clacle sensu Pougen:-D<rnez,m ef al. (2003) within Lhe
T1ees or shrnh.s; t1opical rive1ine frnest and open rocky slope.s
Det;:1rie<1c sens stricf clade. Includes Cupaifera paluslris (Symington) clc \Vit, the
Reference(s} Wei (1983)
sole Male:.;i;m species, mainwincd ;1s a sepa1atc t.:enw; Pseudosfudora by U:on<1al
See taxonomic notes l1!1cler EnderJia
(1957, 86), but consicle1ecl doubtfully distinct by Cowan & Polhill 098la, l;l2); Cultivated ;.1s ornamentals (L rbodusleMia Hance) <Incl intn.xluo.~d in tropical Af1ica,
Brazilian species n111ently undc1 1evisi<>n
C America, Caribbean and rlrn ida
Commerci::1lly p1oducecl high quality resins (copal, copaiha, copaivc.1, Jesuit's
b.;lisam) are used for medicines (scented gums are c1lso used as unguents),
varni~hes, lacque1s, paints and fuel (Lile oleo-1esins <He stdd to he a substi1ure 1'01
diesel); oth~1 uses a1·e timber, c .g. for construction, blidgcs, shipbuilding,
Saraca i. 1767
c , l l spp; mostly in Malesia to Sulawesi (8 spp ); 3 spp endemic co lndo~Chi1w
fu1 nitu1c, joine1y, panelling, turnery and hJockboa1·d
(induding Myanmat (llll!ma)); introduced in the Philippines and P<lpua New
Guinea
Detarium;uss. 1789 De1ived from the Indi<1n ve1nacular name asvl.!a, (sara in S<lnsk1it), me;;1ning
3 spp .; \V/ ;rnd \VC Af1ic<1 (.south to Congo !Kinshasa]), 1 sp widespread in the coloured or spotted; 1~fers to the 'So1Tmvle.ss' tree uncler which Gautama 13uddh;1
suh-S::iharan Suclani<m 1egion was thought to be horn
F1om detar, a loc1l \Volof n;.nne (Senegal) fo1 D seuc~c:r1le11se j F.Gmel.; the edible Trees 01 sh1ubs; tiopical lowland (sometimes swamp) r~1in forest, pa1ricu\arly
sweet fiuits rnc Gilled dalacb ~dong 1 ivers
and on hill slopes
Ttees or shrubs; lropical rain forest, sc:.1:-;onally dry frncst, woodhmd and wuodcd Hefe1ence(s), Zuijde1houdt (1968); I-lou in 1-lou el al. (1996, 660 - 673)
g1 ;1s.'ih1 nd See taxonomic notes under Enderlitt
Hefe1ence(s), Hutchinson & Dalziel (1958, 457); Aui>1cville (1970, 136-1/iO) Cultivclted :1s ornamenlals and shade rree:.; (seventl :-;pedes), ;tnd sornctitr1c.s
Placed as si.ste1 to Copa{/era in an unsuppo1 lccl clade within the Del<irieae sens phmted ::i.s sac1ed trees near shrines; used as a tr~1ditional medicine (named
s/licl. daclc (I-Je1endeen el al,, 2003a); these gene1ic 1clation.ships cilso agree wirl1 ashoka, asok;H ista in the t1ade) and timber (babai) for light constn1ction and
Cowan&. Polhill (198la: 132-133); the position within the D~tailu111 cladc of packing G1ses
fougl:1 c-D;1neza11 el al (2003) is unresolved
Used fm timbc1 (hoiie, rnllow t1cc, dirah) in const1uction, frnnirure, ca1pcntry
cmd r~nces; meclidne; human food (t:dible leav~s. fruil:.; ancl seeds ollthough
some frnms of D se11egale11se a1c toxic (the .seeds ~ue ckph<1n1-dispersedJ);
1·esin.s (for prnm1de.s, .soaps, fumigants and glue), live.'itock forage and as ;i Soraca sp. Photograph by G. P. Lewis
Detar/um senegalense Drawing by unknown artist mosquito 1epelle11l
-----~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
TRIBE DETARIEAE 85
84 LEGUMESOFTHEWORLD
 Talbotie//o batesil (1- 3); T. eketensis (4-8) Drawing by M.·F. Adam Crudia gabonensis (1-2); C. kloinel (3-8) Drawing by J. Saussotte-Guerel Crudia choussyano Photograph by c. E. Hughes
Leucostegone grandls Drawing by P. Halliday

Leucostegane Prain 190 1 Crudia Sch1eb, 1789

2 spp.; Malesia (M<da y Peninsula an d Borneo [Sarnw;tkD c. 50-55 spp.; p:rntior ical; most dive rse in Asia, particulady Malesia (c 30 spp ..
F1om /euco- (G k , w hite) and slegano (Gk., shelter), fo1 the w hite calyx lobes tlrnt extending to Papu<1sh1) and c. 3 spp endemic to Inda-China, Andaman-Nicobar•
enclose and protect the two small petals in bud Islands and Sri Lmka; c , 11 spp. in S Ameiica (c. 9 spp. in Amazonia and 1 sp
T1ees; t10pica\ lowland (.sometimes swamp) and hill forest, sometimes on each in the Cari bbean and C America); c. 7-9 spp . in Wand WC Africa (~enegal,
limestone Gu ine~ 1, Sierra Leone, Liberia, Ivory Coasl, Ghoma , Nigeria, C:.imeroon, Equatorial
Reference(.•), Hou in Hou el al_ (1996, 635-638) Guine<I, G:1bon, Congo [KinHhasa] and Angola)
Not yet sampled for molecular analysis but consicle1·ed closely allied to Saraca Named fo1 01) W, C1ucly 0753 - 1810?), collector o f plants in the Bahamas
and LysidiCC!; may be confused with Saraca when sterile Trees or shrubs (1arely scrambling); tropi<:cd lowland rain and swamp fo1est,
thicket and sci ub forest, often along 1ivers
Reference(s} Corner (1952, 248-253): Aub1eville (1970, 152-155); Ve1dcou1t
Talbotiella I3ake1 r. 1914 (1979, 98-103); Vieira 0990): Hou in Hot1 et al (1996, 573-597)
Genus pooily uncle1stood ancl in need of 1evi:;ion; several specie:.; are known only
3 spJ1.i \Y/C Aflic.:<.1 (S Nigeria and Carneroon extending to Glwna)
Named fo 1 PA- Talbot (1877-1945), a botanical collectrn and o ffi cer of the from their type collection.sand ;J m1mbe1 <He l111clescdbecl The position of Cruc/ia
is unresolved within the Amherstieae claclc (scmsu 131 unt:au el al ., 2001); African
Nigerian Administ1ative Se1vke and his wife Dorothy (1871-1916), bornnical
s pecie:-; currently llllder revision
collt=ctor
T1ees or shr ubsi C<n1s tal ;ind freshwater swa mp foH~sl and sec1sonally dry frncsl, on The timber is extremely hard :ind heavy, with limited known uses
sa ndy soil, so1netilnes fo11ning stilt roots, and on 1ocky hills
Reference(s), Hutchinson & Dalziel 0958, 467); Aubreville (1970, J48-151l
Generic affiniti es are unknown; B1uneau el al (200 1) placed Talboliella in an Lebruniodendron J.Leona•d 1951
unsupported clacle with lecmardoxa and Hymenustegia bl1t this was not l sp.; WC: Af1 ica (Cameroon, Gabon and Congo (K inshasa))
st1ppo1ted by the analysis of Herendeen el al, (2003") Named J'or OrJ-P.A Lebrun (1906- 1985), a uthority on the Afdc;in llrn'1, and
Used for timbe1 (construction) ancl charcm1l; '/'. gentii Hutch. & Greenway d e11dro11 (Gk., tree)
(charcoal tree) is on rhe IUCN Red Data List as critically endangered; the Tie~; tropical lowland 1·a in foresL

rem~ining species are also rare 01 endange1ed Hefei·e nce(s} Leornucl (1957, 96 - 97): At1b1~ville (1970, 86 - 88)
Not yet sampled fo1 n10lecula1• analysis but con~ide1ecl to be closely allied to
Scorudopbloeus and Scholia (Leonard, 1957)
Scorodophloeus Haims 1901
2 spp.; \VC Af1ica (1 ~p. in C1mei-oon, Gabon, Congo [Kinshasa] and Angola; l sp.
in Kenya ~incl Tttnzani~1)
From scorodo· (G k., g;ulic) :incl pbloeo (Gk' ba1 kl, for the ba1k which emits"
garlic-like odou r
T1 eesi tiopical lowland rain forest and seasonally dry foiest, scr ub forest and
wooded grassland, often along 1iver bank:.;
Refe1 ence(s): Brenan 0967, 122-124); Aubreville (1970: 83 - 86)
Considered a llied 10 Schol/a and Hymenoslegia by Leonard (1957, 102-104).
[Auth01,'s postscript: Lock (.submitted) describes <1 new species of Scorodupb/ol!llS
f1 o m coastal fo1est ;:111d thicket in C to N Mozambique]
'rhu lxtrk and le r\. CS f'mm wh d i ~~sc u tial u ll' •ne :1lso exti ;H.lt'tl) :.uc ll \l.1..1 a.s ' 1
su l1sdt111c fo r ).llJ rhc te.;1., S. re11korl I Jorms ih umh" IJ: timber <II' .j'ls<:liurl (J'auliJ
LMR
J.1.Corrarcl (m):todonlil, mm1nd ··) is u:st:e l for rm1).lh f.=onstru ctlnn 1 l'intw·o1x l ;111d tool
handles; the tr~e is used fo r shade and bee forage
Lebrun/0 d
endron leptonthum Drawing by G. Chypre
Scorodoph/oeus fischerl Drawing by L. M. Ripley

86 LEGUMES OF THE WORLD TRIBE DETARIEAE 87

 Micklethwoitio corvo/hoi Drawing by E. Catherine Cynometro obrahomii Photograph by D. Du Puy Cynometro crossifollo Photograph by B. B. Klitgaord

Plagiosiphon !-l;mns 1897 Cynometra L 1753

1i'ipetalm11l111s A Chev (19~6)

c H0-90 spp.; c. 26-28 spp in Africa (Guineo-Congolian W ,111<1 WC regions [c
5 spp; centred in \VC Aflio1 (1 spp Cnneroon and Gabon, l sp. extend ing to 16 s pp.), Zanzibar-lnhambane E Afric• [c 9 spp.) and 1 sp. each in the Sudanian
Ivory Coast, Liberia and Sierrc·I Leone)
::incl Lake Victrnia 1cgions); 10 spp endemic to Madagasca1· ~ind 2 spp , to the
f 10 111 fJICl~io- (Gk,: oblique) ancl sij>bon (Gk: tube), for the obliquely cylimlriul Comoros; c.:. 26 spp. in SE Asi;t (c. 22 spp , centred in Malesi<1 <tnd Papu asi:i to
Inclo-China <llld Aus11.11ia ~md the Pacific, c 4 spp endemic to SW ~tnd E Indi<1); c ,
calyx
T1ees oi• sh1ubs; gregm•ious in tropic;tl lowland forest, often ;tlon~ rivers , on rolky 23 spp. in the Neotiopics (c. 7 spp in Mexico, Cai ibhean <.ind C Ame1 ica; c. 16
banks or hillsides spp in S Ame1irn centred in Am~1zoni::1)
Reference(s): Aubreville (1970: 90-97 & 336-337); Lock (1989: 58) From cyno- (Gk.: dog) and metro· (Gk: womb), apparently fo1 the fll'shy pods u f
Gcne1ic position unresolved within the Amherstie.ie dade (se11s1t l31une;n1 el llf, some spedes, which resembl e a clog's womb
2001) hut placecl (withm1t supprnt) by Hc1enclee11 el al (2003a) in their T1ees rn sh1uhs; t1opical luwland 1ain and swa mp forest, often along l'ivc1s ;:111d
subliuornl; St'<:1sonall y dry foie~t . woodhind, hushl~ind 0 1 thicket, often on white..:
Cynometra dade
sands; some spcdes g1ow g1egariously fo1111ing dominant st::inds
Jlefe1ence(s): Dwye1· 0958); B1en:1n (1967: 'J l 1-J 19); Knaap va n Mccuwcn 0970:
Micklethwaitia G I' Lewis & Schrire 12-31); Ye1dcou1t (1979: 77-87); Koste1mans (1982); L; trnen el al (1984: 45 - 19);
Tavares & Silva (1992); Hou in Hou et al ( 1996: 597 - 608); Du Puy & Rabevohit1a
1.Jre11t111iudemlron .J .Lc!onard (1999)
in Du Puy el al (2002: 129-141)
1 sr; Zanzihal'-inhambane region of SE Afric.:a (Moz~imbique)
Closdy rel~ted to A1m1iltoa (see l<1x:onom it: notes thc1ew1de1); frn genelic
amed for John Pat1ic.:k Middethwait 13renan (1917-1985), an authority in
delimitation sec Kna:.ip van Meeuwen 0970); genus in need of revision wirh
Leguminos:1c system atics and former Dire<.:tor uf Lhe Hoyal Botanic Gardens, Kew
seve1al species undesc1il>ed (Ve1d<.:ou 1r , '1 979)
Trees; seasonally dry t1opical crn1stal mosaic forest
Vaiious species used fo1 timbe1 (kek<1tong, grn1pinol ncgio, nganga, hawk<i,
lleference(s): Leona 1cl 0999a); Lewis & Schdre (2004)
muhimbe) in c.:onst1ucrion, ships keels, r.1ilw<1y sleepers, tl orning and cupcntly;
Not y ~ , :unpled for molec..:ular am1lysis but considered closely alli •r;I w
~1lso used for fi1ewood, as shade !lees :Jml bee foiage
Scorodt•/Jbftx•lfs and /.uhrHniodendrcm (Leonard, 1999a); previously tn:.ate<l under
C'.yitomelra (see Ltwis, 1996b) hut 1aised to generic 1ank by Leonard Cl c..:.)
Bre1wniodendrrm .f r.eomud required <1 nmne change because of Brcmmulendroll
I-I.Rob, a genus in th e Compo.'ii tac Lhat was published 8 clays before l.~ona1 d's
!Wille

Maniltoa schdf 1876

20-25 spp; SE Asia (1 sp. in India, Inclo-China to Malcsia; all other species
occ..:rnring in Papuasia [P;.tpua New Guinea or l1ian J1yal extending ro Malesia,
Austialia and the Pacific Islands)
Fi om mt1uiltou, local apau Nr.;w Guinea 1w111e for M grmuliflora Scheff, the type
species of the genus
Trees; tmpicod lowh1nd rain frnest, often in swall'"tp me.as, along 1i vers and
tKGlsic>na lly in mang1oves
Refercnce(s): Knaap van Meeuwen (1970: 31-46); Vcrclcoult ('1979: 57-77); I-l oll
in Hou el al (1996: 638-650)
Genus in need ol revision; closely 1elated to G:vnometr" a.'i in<licalecl by l3ninc:1u
el al. (200 I) ancl l-lerencleen el al. (2003.,), liut the 111onophyly llf both genern "
1
unconfirmed (foi discus..o;ion of generic ddimit;Jtion see Knaap van Meeuwen (19701
Used ::1s rnn;,1mcntals (several species), liml>c1 and in agiofrncsuy Cynometra
- several Madagascan species. Drawing by M. Tebbs
Mani/too megolocephola Drawing by unknown artist

TRIBE DETARIEAE 89
88 LEGUMES OFTHE WORLD
 ~·
rr- ,·

o-
.j
r-- '

Tamarindus indica Drawing by l. M. Ripley Tamarindus lndica Photograph by G. P. Lewis lntsia bijuga Drawing by l. M. Ripley Afzelia quanzenzis Photograph by G. R. Nichols

Tamarindus L. 1753 lntsia (text on previous page)

1 sp.; the natural range is unce1tain because of widespread cultivation th1oughout
the tropics (especially India); apparently native to tropical Africa and Madagascar
but introduced to Asia in ancient times Afzelia sm. 1798
From ta mar (Arabic: d ried elate) and hindi (Indian), refe rring to the acidic pulp of Pahudia Miq. 0855)
the f1uit c 11 spp.; 7 spp. in Af1ica (cent1ecl in Guineo-Congolian \Y/ and WC Afiica with 1
T1ee; tropical seasonally dry frnest, woodland, wooded g1asslancl and bushland, sp. each to Suclanian and Zambezian to Somali::i-Masai 1egions); 4 spp. in SE Asia
often along rivers (2 spp in Indo-China to S China; 2 spp. in Malesia)
Refe1ence(s): Hutchinson & Dalziel 0958: 467); B1enan 0967: 151-153); Bu1kill Named fo1 Adam Afzelius (1750-1837), last pupil of Linnaeus and late1' Professor
0995: 169-176); Du Puy & Rabevohit1a in Du Puy el al. (2002: 152-153) of Medicine :H the University of Uppsala, who collected plants in Sie1ia Leone
The position of this genus is unresolved within the Amherstieae clacle (sens11 Trees; tl'Opical lowland rain forest and seasonally dry frnest, woodland, bushland
IJ1uneau eta/., 2001) and thicket
Tamarindus htdica L, (tamarind, clakkar) is widely cultivated for its edible fiuits Reference(s): Brenan 0967: 124-128); Aubreville 0970: 110-120); Illll'kill 0995:
(with pleas~mt tasting acidic pulp around the seeds) that ate eaten 1:nv or used in 50-55); Hou in Hou el al. 0996: 438-442)
confectionery, as an ingredient of curries, pickles and in fermented cll'inks; the See taxonomic note under fntsia
seeds are edible 1aw rn cooked, and produce a useful oil. Dyes are produced Various species used as imprntant commercial timbers (afzelia, cloussie, pod
from the bark and leaves, and an ink is made from the burned ba1k; the timbe r is mahogany, mahogany bean, chanfuta, bolcngu, apa, tindalo) for heavy
used in joinety, construction, fu1nitu1e, utensils and posts, <ind the wood fo1 construction, joinery, f1ames, stai1cases, flooring, furniture, implements, boat
firewood ancJ charcoal; plants a1e also used as medicine, shade t1·ees, building and turne1y; also used for dyes, cha1coal, medicine, cattle fodcle1, edible
orn~unentals, bee fornge and frn tannins seed a rils (although the seed of some species are poisonous); the seed.i.; are also
used in necklaces, curios and as charms; a number of species <ire vulne1able or
enclange1ecl
Intsia Thmws 1806
c. 3 spp. ; centred in SE Asia (Malesia and Papuasia); /. hij11ga (Coleb1.) Kuntzc
from coastal E Africa, Madagasca1, Indian Ocean Islands, E India, Inda-China,
Malesia, Papuasia, N Aust1alia cmd the Pacific
P1obably delived from Hintsi, one of seve1al similar local Mah1gasy names foi' !.
bijuga
Trees; t1opical humid coastal foiest on sand, rain forest, nrnng1ove f1inge.-; and
tidal river mouths, occasionally inland on hills
Reference(s): Ilrenan (1967: 128-130); Ve1dcou1t 0979: 90-93); Hou (1994:
322-328); Du Puy & Rabevohitra in Du Puy eta/. (2002: 141-142)
Closely related to Afze/ia (Fouge1e-Danezan el al., 2003) with which it is strongly
suppo1ted as sister in the analysis of Herendeen et al. (2003a)
It1tsic1 bijuga is a highly desi1able and over-exploited timber (ipil, rnc1bau, nwbcca
teak, miiabow, kwilau, vesi) used for, e _g_, construction, furniture, cabinetry,
joine1y, panelling, musical instruments, flooring and turnety; also used for dyes
;:md as traditional medicines; I. bijuga is sea-dispersed and the seeds are comr11only
washed up on beaches; it is listed in the lUCN Red Data List :Js vulne1able

(ext1a illustration on following page)

Afzella quan .
lntsia bijuga Photograph by D. Du Puy zens1s Photograph by J. Dransfield

90 LEGUMESOFTHEWORLD TRIBE DETARIEAE 91

 Brodriguesia santosii Photograph by G. P. Lewis Neochevalierodendron stephanii Drawing by M. -F. Adam Normandiodendron romil Drawing by P. Halliday

Brodriguesia it s .Cowan 1981 Neochevalierodendron; .Leom11c11951

1 sp.; l31azil (Bahia State) 1 sp.; WC Africa (Gabon)
Named forJ , l3a1bosa Rocl1igues (1842-1909), 13rnzilian botanist Named fo r Prof, Auguste Chevalie1 (1873-1956) collector of the type of the
Trees; t1opical coastal Atlantic rain forest genus, and dendron (Gk: tree); the genus name Cbevalierodendro11 had alteacly
Reference(s): Cowan Cl981b) been used in the Moraceae, hence the prefix neo (Gk.: new)
Brodriguesia santosii RS.Cowan was not sampled in the molecula r analysis of Trees; tropical lowhmcl rain forest
Herendeen et al (2003a), but is considered by Cowan 0981b) to he related to Reference(s): Leona1d 0957: 119 -120); Aubreville 0968: 94-96)
!11/sia and Afzelia Considered allied to Loese11era and 1-fymenostegia (Leonard, 1957) cincl weakly
supported as sister to Loeseuera by He1encleen el ol (2003a); the position of the
genus within the Amherstieae clade (senstt Brunem1 et al, 2001) is unresolved
Loesenera mums 1897
4 spp; centred in WC Aflica (3 spp; Nige ria, Cameroon and Gabon) with 1 sp. in
W Africa (Liberia) Normandiodendron J Leonard 1993
Named for Ludwig Eduard Theodo r Loesene r (1865-1941), German botanist 2 spp.; WC Africa (Congo (Kinshasa], Gabon and Cabincla)
Trees or shiub5; t1opical lowland min fo l'est, commonly along 1iver banks and in Named fot• Didier Normand (1908-2002), authority in wood anatomy of t1opical,
swampy valleys especially caesalpinioicl trees, ;.incl dendron (Gk: t1ee)
Refeience(s): Leonard (1957: 120-122); Aubreville (1968: 90-93) Trees or shrubs; t10pical lowland rain fo rest and seasonally cl1y forest, o!'ten along
Consiclerecl 0:1llied to Hymenostegla by Leona1d (1957) but its gene1ic po.<.;ition 1ivers
within the Amherstieae clac\e (se11su Bruneau el al, 2001) is uniesolved Refel'ence(s): Leonaid 0993: 446- 451)
Loesenera kalantba Haims is used fo1 timhe1, medicine and in witchcraft Normandiudendron was erected to accommodate two species frn·medy placed in
Scholia; the genus is suppoited a.s monophyletic (Fougere-Danezan el al., 2003;
Herendeen et al, 2003a) but its position is unresolved within the Amherstieae
clacle (se11su Bruneau el ol., 2001); Brown el al (2001) place Nornumdiode11dron
in a group with J~yme11ostegia, loesellera and Leouardoxa, while Herendeen et al.
(2003a) have it as sL'lte1 to the remaining genera in thei1 Hymenostegia clade
Used as 01 namentals, medicine ancl the leaves a1e eaten as a vegetable

Zenkerella Taub 1894

c 5 spp.; Africa (1 .sp. in \'V'C Africa [Nigeria, Cameroon, Equatorial Guinea and
Gabon]; 4 spp . in Af1omontane to Zanzibc1r-Inhamhane E Africa [Tanzania])
Named fo1 .Jonathon Kad Zenker 0855-1922), Ge1 man botanist who collected
extensively in Cameroon
T1ees or sh1ubs; tropical lowland rain frll'est, Iiveline and .swamp forest (2 spp. in
E Africa in mont;me iain frnest)
Reference(s): 131'enan (1967: ll9-122); AulJl'eville (1970: 75-76)
T he single species sampled by Herendeen el al . (2003a) wa.s placed a.s sister to
1-lumholdlia in thei1 Hymeno.'itegia clade (although with no bootslr<lp support) so
gene1 ic position within the Amherstieae clade (sens/I Bruneau el al., 2001)
1emains unclea r
Used fo1 wood (fiiewood, ch<t1coal, utensils, tool handles and poles); also a.s
ornamentals and shade trees
Zenkerelt · ·
Laesenera walkeri (1-5); L. gabonensis (6-10) Drawing byM.·F. Adam a c1trma Photograph by M. Cheek

92 LEGUMES OF THE WORLD TRIBE DETARIEAE 93

 Hymenostegla a{zelli Photograph by D. Kirkup Leonordoxa afr/cana Drawing by M. -F. Adam Amherst/a nobllls Photograph by P. Cribb

Humboldtia Yahl. 1794 Leonardoxa Au brev. 1968

6 sp p ; SW Ind ia (W Ghats) wi th 1 sp. extending so uthwa rds to Sri La nka 1 sp.; WC Af1ica ( Nige ria, Camernon, Gabon <ind Equ ato rial Guinea)
Named for Friedrich Alexan der von Hu mbo ldt 0 769-1859), scientist and ex plor er Na med frn Prnf.] Leonard, Belgian botanist, Legumi nosae specialist and au th o rity
of C and trop ical S Ame ri ca on the no ra of Congo (Kins hasa), and d oxa (G k.: glory)
Tl'ees or sh ru bs; tropical lowla nd and upla nd rai n forest, commonly along rive1s T1ces 01 sh1ubs; tropical lowla nd ra in fo rest in the unders tory
Reference(s): Sa njappa 0986) Refere nce(s): Au breville 0 970: 88-91); Leon ard 0993); McKey (2000)
Sanjap pa 0 986) tr a nsfe n ecl the sole African species (H. aji'icm w llaill.) to Scholia Laonardo.'\·a was erected to accom modate o ne species fo rmerly placed in Scbolia ;
Jacq . (subseq uentl y placed in Leo11ardoxa (Leonard, 1993)); ge ne ri c position it comprises four (mostly) allopat1 ic subspecies o f whi ch all but one a1e
within th e Amhersti eae clacle (sensu Bruneau et al., 2001) remains unclear but rn yrmecophytic (Brouat el al, 2001), Ge ne ric affinities are unclear although
placed by Herendee n el al (2003a) in their Hymenostegia clacle ll1ot1at el al. (Le.) groups the L. cifriccma (Baill.) Aubrev complex with
Used for human food (pods), timber, as fuel, medicine and o rnamentals Hy 111enostegia and Loese11era; placement within the Amherstieae clade (senstl
I3 run e;.1u et ar, 2001), howeve1, is unresolved
Used fo r timber
Hymenostegia (Be nth.) Harms 1897
c. 17 spp.; centred in WC Af1ica (c 14 s pp , mainly Cameroo n and Gabo n bu t
sotlth to Angoh1 and no 1th to Nigeria); 3 .spp .1 in W Africa (Sie1ra Leone to Amherstia w a11. 1829
Nige1ia) l ... p.; S As ia (e nde mic to the Tanin tlrn yi [Te nasserim] region of S Myanma1 [I3u1 ma]
From by me110- (Gk,: membr.mo us) and siege (G k.: shelter), for the fragile Named for L;1dy Sarah Am herst 0 762- 1838), artist, pa tro n o f bota ny and
bracteoles th at enclose the nowe1 in b ud collector, w ho was the wife of \'(lill iam Pitt, Loi d Amherst, Gove rnor-Ge neral o f
Trees o r sh ru bs; tropica l lowland mi n fo rest and seasonally d ry fores t, bushk111 cl Ind ia w hose 1em1re spa nned the first Anglo-llt1nnese wa r (18 24- 1826)
and th icket, often al o ng ri vers; some species ci re g1egarious T1ees; tro pical lowland fo rest along rive rs
Refe rence(s): Ht1 tchinson & Dalziel 0958: 464); Auh rev ille 0970: 99- 108); Lock Re fe re nce(s): Corner 0952: 377 - 378); Lyte (2003)
(1989: 55-56) Ge neric affinities a1e unclear; its place me nt w ithin the Amherstieae clade (se11su,
The mo no phyly o f the ge nus was questioned by Brunea u el al. (2001) but not by Brunea u et al 1 2001) is umesolvecl
Herendeen el at. (2003a) who placed it in their Hyme nos tegia clacle ; generic Amherstia nobilis Wall. (amhe1stia 1 pride o f Burma) is widely cultivated as a
position within the Amherstieae clade (senstt Bruneau el al , 2001) Is unresolved .specta cular ornamental and shade tree through o ut the Asian tropics but is
J-Iymen ostegia afzelU (Oli v.) Harms is used for timber (co nstru ctio n and p robably extinct in the wild
im pleme n1s), fuelwoocl , medicine and as an orna me ntal

Ecuadendron DA.Ne ill 1998

1 sp.; S Ame ri ca (\YI Ecuador, lower Andean foo thills)
Named for the count ry of o!'igin, Ecu<1cl o r (sig ni fy ing 'equator' in Spanish), and
de11dron (G k.: tree)
T1 ees; tropiccil lowland wet to moist fo rest (with considernble fog-associated
preci p itat io n)
Reference(s): Neill 0998)
Placed w ithin the I3rownea group ( Biunea u el al , 200 1) w ith
1 closest relatives
th o ug ht to include Brachycylix, Heleroslem o n and Elizabetha (Neill, 1998)

Ecuadend
Hymenostegla f/orlbundo (1-3); H. normandii (4- 6) Drawing byG. Chypre ron acosta-solisianum Drawing by J. Myers

94 LEGUMESOFTHEWORLD TRIBE DETARIEAE 95

 Paloue gulanensis Photograph by 5. A Mori Brachycylix vageleri Drawing by N. M. Goar Heterostemon mlmosoldes Photograph by A. Gentry

Paloue Aubl. 1775 Brachycylix (Harms) R.S.Cowa n 1975

Heleroslemou Oesf. subgen Bmcbycylix Hanns (1925)
Palovea juss. (] 789)
4 spp; S Amerirn (Amazonian B1azil and the Guh1rn1s) 1 sp : S Ame1ica (NC Colombia)
Derived from the local Galibis name in Gtli~ina frn P. guimwnsis Aubl F1om bmcby- (Gk , shrnt) and ~y/ico (cup), for the shoit cupula1 hy panthium
T1ees and s h1ub.s; tropica l lowland rain Forest (te1ra flnne a ncl iminclated), ofte n Trees; ti opical lowland 1ain forest, often 1iverine
along rive rs
Re ference(s): Cowan (1975a)
Previo usly treated <IS c1 subgenus of /-leleroste mon cmd not sampled in the analysis
Reference(s): llodligues /!.: Linrn (1990)
Placed in the Brownea g1oup by Bruneau el al (2001) of He 1endeen el al (2003a)
Used for o rnamenta ls and timber

Paf.oveopsis Rs Cowan 1957

1 sp; S Ame1ic;:1 (Amazonian Guya na and Dr:nil)
Heterostemon o esf. 1818
F1om -opsis (Gk.: appearance), fo r its resemblance to the genus Pafoue (synonym 7 spp.; S Amelica (most dive1se in the Upper Am<Jzon in N\V D1azil, S\Y/
of Palo11ea) Venezuela and SE Columbia with 2 spp in Guyana)
Trees; t1opical Juwbncl r;-1in fo rest ( tern.1 firm e), o ften ;.dong 1ive1s
From belero- (Gk: differe nt) an d sle111u11 (G k.: stamen), fiom the p 1esence o f both
llefe1ence(s): llodligues /!.:Lima (1990) fe 1tile and ste1ile st~1mens
Sma ll trees or sh1ul>si u opical lowland (sometimes inundated) r..ii n forest,
Not sampled in the analysis o f He1 endeen ct t1I (2003.i) but conside red closely
understo1ey and fo1est margins, o ften live1 ine ::ind on s;indy soils
allied to Pttlotte (Hodligues & Lima, 1990)
Hefe1e nce(s): Cowa n (1976a); Silva/!.: Ci111 ei1a (1995): Cowan/!.: Be11y in !le11 y el
al (1998: 67 - 69)
Not sa mpled in the analysis o f Herendeen el al (2003a) bu t conside 1ed closely
allied to Bracbycylix and Elizabelba (Cowan, 1975a; Neill, '1998)
Used <IS 01 namentals and fo 1 timber

Elizabetha Sd1omb . ex IJen th . 1840

c 11 spp ; S Ame1ic-J ('H ylc1ea' phytogeog1aphic p1ovince fro m SE Colombi<t, S\'V'
Venezuela, Am azonian Brazil to the inte1io1 of Guya na and Su linam)
Con fl icting etymologies are given; Cowan (i976b) notes that Bentham adopted
the 1m111uscript name o f Richard Schomburgk who wanted to honour 'H .R H [he
Plin ccss Roya l of Pn1ssia ', but \'Vittstein (1856) stcttes tlwt it was named fo1
Elizctbeth Fitto n, who togethe1 w ith Sa1ah Mary Fiuon were authors of lhe
Co11versaliot1s on Bolnny, published in 1840
Trees; t1opic:a\ lowhind rJin fore.st (terttl fi 1me and inundated) often along livers
Re fere nce(s), Cowan (1976b): Cow•m & Jleny in lleny el at. (1998: 62-64)
Th e si ngle species sampled was placed in the Brownea group by Bruneau et al,
(2001); conside1ecl closely allied to Ec1 wde11dron and Bmcbycylix by Neill (1998)
The bark of E princeps Schomb ex 13enth is burned to produce an ash mixed
with snuffs oi hallucinogens (known as ebena), prepmed from the resin of the
genus Virola Aubl. (My1 isticaceae)

Elizabeth 0
Paloveops/s emarglnata Drawing by P. Halliday cocclnea va r. oxyphylla Photograph by G. P. Lewis

TRIBE DETARIEAE 97
96 LEGUMES OF THE WORLD

Brownea macrophylla Photograph by G. P. Lewis
BrowneaJ"cq 1160
12 spp .; westl.!Hl S A111e1ica (Ve nezul'ia, Columbia, Pelll :me.I Ecuador with 2 spp
more widcsp1e;1c.I to I3razil, the Guian;;1s ~incl extending to C America [P<tnama ~ind
Cost<1 Rical and the C:a1 ibbea n)
Named for Ptll1ick 131owne 0 720-1790), Irish doctui, 18th CL'l lllll1' pionee1 and
naturalist working in J:imak:a and latc1 Cu1atrn o f Oxford Bot<inic Gardens
T1ecs rn· sh 1ubs; unde1stor·~y in uopical lowland 1ain fo1est
Heference(sJ: Klitga:utl (1991: 437-443); Qui11ones (1995; 1997); Cowan & lk11y
in Ile1ryl'/a/ (1998: 14-17)
Se<: ta;-:onomic notL'S u ncle1 Brow11eojJsis
Scvc1·al spet:ies (pa lo d e cruz, bois 1ose, palo rosa, monta dd rosa) arc used f01
timber, mc...:clicinl:', handic1;1fts (frnm dried pods and se1.xl.11) and c.: uhivated :JS
0111<llllelltii 1S
Browneopsis Huber 1906
6 .spp.; western S Ainc1ie<.1 (Pe lll , Ecu~uJ or and Columbia), extending nrnth
Panama
Fmm -opsls (Gk .: appr.:a 1ance) :ind Bruwn ea, for it:-; si111i l:11i ty 10 this closely
rclall'd genus
'J'rees; principal ly tropical lowl1:1nd 1·;.1in forest (tc11·;:1 fi1 me and inundated), but :1lso
in sub-mont;rne frnest on thl' foothills of the Andes
Rde1en cds): Klitgaa1tl (1991: 443-447)
Not st1ppo1tc.:d as monophylctic by Bruneau el al (2001); indications liom the
Herendeen el al. (2003c.1) analysis are that Hru1(11UJ(l may be paraphylctic if
/Jrowneopsis is excluded but more comprchcn~i\tc .sampling is required to conf11lll
its :-;tatus
U.'ied frn timber
Browneopsis disepala Photograph by 8. 8. Klitgaard
98 LEGUMES OF THE WORLD
Macro/obium unijugum Photograph by T. D. Pennington Macrolobium latifo/ium Photograph bys. A. Mori
Macrolobiutn sch 1eh , 17H9
c 70-80 spp .; cc n11r.:d in S Americ a (mnst divel'sily in Amazon ia with 2 spp to
lh..: J\.th.tntic.: co;1st in SE B razil, also west f1om Vcnr.:zuehi, Columbia , Pelll to
Enwdrn in foothills on either• side o f the And c.'i); few spp in C Americ1 (Co:-;ta
Hica, Hondu 1as ;i nd P<:inam;1)
P1om 11/flU U- (G k,: lmge) ancl /o/Jo (Gk.: lobe..:), fo1 the..: si ngle \Vl' ll-dcve loped
aclaxi:d petal
T1ecs rn• shrnbs; t1opical lmvbnd 1;:1in frncsl, c>flc.:n along 1ive1s <incl in seasonally
inundated pl~tces, or in seasonally cl1y woodland ;:ind wooded g1<1sslancl
Refe1cncc(s); Ll:ona1d (1952<1); Cowan 0953) ; Cowan & Be11y in Be11y el al
( 1998: 74-HH)
Macrulohitfl/1 W<lS fi1'!'it desctibecl with a single ncot1opic1I species and in 1806,
Palisot c.k Beauvais clesc1ibecl A11lb01wlba with a single Alrican species; frn m.:aily
1')0 years the two genc1a were lhc.:n tn.:cttecl ;1 s cu ngene1ic ( umle1· Mm.:rolubi11111)
until Louis (1949) suggested that the Afrk:;1n and neot1o pical species ~1gain be
t1c...:atccl ;1s sepa 1ate gene1::1 , The.: Af1ican species of Mt1c:10/ohi11m were t1ansli:1rccl
m fou1 exclusively Anican gcne1;1 by Leon0:ucl who rd11st1.1ted J111lbo11 01Jw
to (Ll:onc1rcl, 1955: 201-202) ~ind clcsc.:J'ibccl th1ec new gc.:nc.:1<1, Gi/herliode11dro11,
Parm1w<.-rn/vb i11m and PelleR1'i11iodenrlro11 . Cowa n ( 19'53) explit:itly cxduckd all
rh c Af1il:an t;1xa f10111 his 1evision of 48 species of n ~o tiupic::ll /v/ac:mlu/Ji1tm Th t.!
gem1s is much in need of revision; Macroluhiwu as p1es..:ntly cirnunsuibecl is
ve1y dive1sc: 111rnphologically and is dm1btful ly monophyle tic, as inclicatccl by
Brune:tu el al (200l) and I-k1endL~n el al. (20U3a), although fu1the1 ' sampling is
needed
Usc.:d for timbc1 ~ Ji.sh poi.sons ~111d a.'i orn:uncnrab
Macro/obiu .. .
m acac1ifo/1um Photograph by G. P. Lewis
TRIBE DETARIEAE 99
 Cryptosepalum sp. Photograph by/. Anton-Smith Dlcymbe a/stonii Photograph by L. Y. Th. Westra Dlcymbe hymenaea Drawing by B. Angell
Paramacrolobium coeruleum Drawing by l. M. Ripley

Para1nacrolobium J Ll'onaiCI 195-i Dicytube Sp1uce ex Be111 h , JH65

00 1 sp: WC Afri ca (Sic1r.1 ~one and Guinea to Congo, C:1me1oon , Ccnt~d Afiic111
llepuhlic.:, Congo lKinsha.sa] and Angola) and disjum:t lo E Af1i t::.1 (lmvh1ml E
Kenya and E Tanzanic1)
From para- (Gk.: beside) ;111d lv!acru/obi11111, •1flc1 the genus f10111 which ii w.L,
scg1 eg:ited
Dicy111ho/Jsis Ducke tl950)
16-20 spp; S Ame1ica ( most
!..'.
Brazi l .inti Guyan;.i)
divcrn~ ill Amazoni;111 VenezueJ.1 to Colombic1,

F1 om di- (Gk : lwo) and i..:ymhi- (Ck: .'i1rndl cup) ,

enclosing the llowe1 in hud
rm
the two COllGl\le h1;1C[eolt'S

T 1ecs; uopirnl lowland 1<.1i n forest uswil ly ~d ong rivc 1." and in inundated pl~1ccs to
seasonally d1y wcHJcllancl and woocl ed g1assl<1nd in 1ipa1ian vegetation
Refe1cncc\sl: ll1e11<111 ( 1%7: l 'i5-1 57)
Scg1egated f1o m M({crulvbirtm by Ll'onaicl ( 1954); 1cccn t cvidc 1Ke sugges ts~'
close affi nity with CJ:)'jJ/osepalum (B runeau el ell., 2001; Jlc 1cnclcc n el a l , 2003:1)
Tiecs; often nmnmlumin;rnl in uopic;ll lowlancl r;1in fore.st, g<il le ry forest , in
valleys and (1;_11ely) un 1ocky slopes
Rere1ence(s): IJucke ( 19'i0); Cowan ( 1957; 1961); llarncby 0990); Cow"n & lk 11 y
in llc 1ry et al (1998: 'i6-5H)
G~nc 1 'ic affinities ;ue unclear, but p o.s.s ih ly assuci<tl ccl w ith Pol)!s/emo11f/Jllb11s

Parm1wcrolu/Ji11111 coc:r11/e11m Cfi1u b) J L0011a1d (sa -gbcmbe) is tiscd for ti mbc 1 (Bru neau el al, 200 1) Ma ny specie.'i a1e 1:1re, known o nl y f10 ;11 the typ e co llection
(1a il way sleepe1s, ru1 nitl11c, e<upen t ry and gongs) and shade t1ccs; the seeds <llL' or· <1 hanclful of g~1t h e1 ings, an d JJicym/Jo/JSL\· may he disti n ct
Usecl fot timhei
used ;.is cm11lte1s in games

Cryptosepalum llenih . 1865 Poly stemonanthus 1-Li1111s IH97

l .' ip.; W Altica (lvoiy Coast aml Libcli;l)
Dewi11dlia De Wi ld t 1902); 1'.y11aertiode11dm11 lk \Viki, (19l'iJ
c 11 spp; Afrirn (c. 7 spp ill Guillca-Congoli<ln \'\IC 15 ~pp] and \YI 12 spp.J Fro111 po~y- \Gk : many) ~mcl S/('mo11 (Gk : .'it<1111cnJ, fo1 the many C:m oi mo1e)
slamcn.'i pe1 Jluwer
1egi o 11.'i: ..i sp p in Z:1 mhezic1n SE AfriG1)
Til...'l:S; 11opic..:;d lowland r~ 1 in forest, ll.'iually in th~ unclcr.-;toiey
F1om c1y/1/u- (G k.: hiclclcn) ;.11ld sejHi/11111 (L: :-;ep;.d), frn lhc minute calyx hidden
by the wdl -developcd 1>1 al:lcolcs
llefe1enc:ds): Hutchinson & Dalziel l 19';8: 467 -·168)
T1ce.s . .shn1bs and .'itill 1utices; t1opica\ lowland (:mmdimcs inundatcd) fo1cs t lo '!'he Sl <lllll'll nullll)CJ g1 c atly exceeds lilac or C>lhe1 llle1nhers of t1i b e D et<1 1ie:1e;
gene1ic affinities a1e u11 ce1t<1in hut see taxono m ic notes under Dicym /Je
SC<l.'it111a] Jy d 1y, t11~-p 1 onc WOOdJa ncl and SC I Lib !'01L'Sl
l(cfe1e nce\s): Lfo na1d (1 957: 270-276); I-Iutchi nso n IX Da lziel (1958: '\HO);
D uv igncaucl & B1 enan (1966); B rcn~1n 0967: 198 - 203); Au lnL'vil k ( 1970:
2 19-224)
Sec l'H)tcs undc1 Para111acro/o/Ji11111; the sh1ubby Z.imhezian reg ion g1m1ps
ccnt1ed arm1ml C. c'."!f/'fJfittl11111 De W ilcl and C maravie11se O l iv., ~Ill: consiclc1Ld
t<1xono1nically cc )mpll'x with species cliffe1entiatio11 diffk:u lt (D uvigneaud &
llrenan, 1966)
Usc cl fu1 timbt:r (L:onsl1uct ion), fih1e (f1om b;11k fo 1 cloth , bindings and 1i~1skcLs)
bee frn age, and med icine

S.R-C.

Cryptosepalum pseudotaxus Drawing bys. Ross·Craig

100 LEGUMES OF THE WOR LD

--
Po/ystemon th . .
an us dmklage1 Drawing by W. E. Trevithick

TR IBE DETARIEA E 101

Pseudomacroloblum mengei Drawing by M. Boutique Englerodendron usambarensis Drawing by L. M. Ripley
Pseudomacrolobium Hauman 1952
1 sp ; WC Africa (Co ngo [Kinshasa])
f rom pse11do- (G k.: fa lse or spuriow;) and Macro/obium, for its appare nt si milarity
ro J\1rtcrolobi1tm
Trees; t1opica l lowland rain fo1est
Refe1e11ce(s): Hauman (1952); Leonard 0957: 179-180)
Not sampled in the analysis of He1encleen et al, (2003a) and position unce11ain
within th e Mac1olobieae senstt Bretele1 0995)
Pseudomacro/obi11m. mengei (De \Viki) Hmurnm is used for timbe1 and as lish
pol.sons
Englerodendron Ha rms 1907
1 sp; E Africa CE Tanzania)
Named fo 1 H,G A Engler (1844-1930), German botanist, and de11dro11 (Gk_: tiee)
T1ees 01 slu ubs; [mpic1l Zanzibar-In hambane lowland 1c1in forest
Refer ence(s): Drenan 0967: 149-15 1)
Englerodendrmi usmnbarensis Harms is p:.ut of th e exdt1.o;;ively African
Macrolobieae c\acle (sensu 13runeau et al. , 2001)
Anthonotha P.[Je.ruv . rno6
Macrolobium au ct. non Schreb; fsomacrolobium Aubrev. & Pelleg1 . 0957);
"Ji"ipliso111eris Aubrcv & Pellegr. 0957); Leo11arde11dro11 Aubrev (1968)
c.:. 30 spp.; Afric.1 (centred in Guinea-Congolian WC 1egions [24 spp., one
extending to \V/ Tanzania]; c. 6 spp. in W Afric<i, one extending into Sudanian dry
forest)
F10111 antbu- (G k: flower) and 11otho- (Gk: fol.se) , fo1 the ~1pp~1rent re/'iembi<1nce
of the Rowers to those of ot her genera
Trees or shnibs; tropical lowland , riverine and upland 1<.1in forest, season<illy cl1y
forest and sci ub fo 1est, woodhmd and wooded g1assland
Reference(s): Leo nmd 0957: 215-229); Hutchi11so11 & Dalziel (1958: 471 - 474!;
Drenan (1967: 153-155); Aubreville (1970: 188-199); Lock (1989: 1-4)
Uni ted with, and t1eated under A1acro/obirtm Schreb fr om 1865 Llntil LComud
0955: 201-202) re-established A11thv11otba Forming part of the exdtrsively
African Mac1olobieae clade (sensu Brune:-1u el al., 2001), the genus as currently
circumscribed is ve1-y diverse mo1phologically <md may not be monophyletic
(Dnrneatr et al, 200 1; Wieringa & Gervais, 2003)
U/'ied fo r timhe r, e.g., A 110/deae (Rossherg) Exel\ & Hille. (1111 mh •la), for
constrnction, poles ;mcl tool handles; as part of agroforest1y :s.ybt<.•10.s; for fi1ewoocl,
cha1coal, withies, as dyes, medicine and human food (M:'eds)
Didelotia u ., .
Anthonotha conchyllophora Drawing by M. -F. Adam
102 LEGUMESOFTHEWORLD
Berlin/a bruneelii Photograph by D. J. Harris
ni,o110/ata Drawing by P. Halliday
Librevlllea klalnei Drawing by J. Sausotte-Guerel
Berlinia Sol ex Hook.f. 1849
Weslia Vahl (1810)
c 17 spp .; cent1ed in \VC to \Y/ Afrirn ClS spp , one extending to SL1da nia n and
o ne into Zambezian wood land); 1 sp in Zambezian S Co ngo (Kinshasa) <1nd 1 .o;p ,
in Sw:1helian E T<1nzania ancl E Mozambique
N;1111ed for the Swedish bot;rnist A.I-I [Jerlin (1746- 1773), pupil of Linnaeus who
togethe r with Henry Srneathman Cf/ 1750- 1787) , rn llectecl one of the three
specimens from which the genu.'i W<IS desc ribed
TJ'ees or shn.1bs; tropical lowland (sometimes inundated) rain forest, occasionally
in mang1ove; seasonally dry fo1est, sclllb forest, woodland and thicket, often
along rive1 banks
Refe rence(s): Hutchinson & Dalziel 0958: 469-471); Brenan 0967: 143-145);
Aubreville (1970: 242-254); Lock (1989: 4-6); Mackinder & Harris (submitted)
The cu1rent gene1ic ci1cumsc1iption follows Lt:one1 1d (1957: 180-184); pa1t of the
exclusively African Macrolobie;ie clade (se11su Oruneau el al., 2001)
V:1rio us species L1sed locally for timbe1 Cbedinia , ebiam, ekpogoi, essabem, abem)
for heavy construction, Aooring, fuinitu1e, ca binetry, joi nery, veneers and tl11n e1y;
also used for medicine
Librevillea Hoyle 1956
1 sp .; WC Afl'ica (Cameroo11, Gabon and Ango la)
Named frn Lib1eville , Gabon, loc,1\ity of th e type collection
Tree.s; uopical lowland rain fo1est
Refer e nce(.,): Aubreville (1968: 283 - 285)
Pait of the exclusively Af1ican Macrolobie;1e c\:ade (sensu Bruneau et al, 2001), its
generic affinities a1e unclear. Considered s imilar in some aspects to Bracbystegia
cy11ometroides Harms by Chikuni (1998) and placed as siste r to Oddo11iode1ulron
in n weakly supported clade by Wiel'inga & Ge rvais (2003)
Didelotia Baill. 1865
Tcmbav11te Aubrev. & Pellegr 0958)
l' 11 spp; cenr1ecl in Guinea-Congolian \Y/C Africa (c. 8 spp) south to Al1gol<1,
with c 3 S[Jp . in \VI Atiica
Named fo r Adm ir al (Baron) Didelot, Commandant-Superiol' of Gahan, 1862 -1864
T1ee.s; tropica l lowland 1ain forest, often along ri vers; so me species form
m o n odo min~1nt sta nds
Reference(s): Leonard 0957: 265-270); Oldcman (1964); Aubreville 0968:
247-260); Lock 0989: J0-11)
Some species limits are contentious; D , minutijlora (A.Chev.) ].Leonard is poorly
known <Ind may be a liana. Pa11 of the exc\w;ively Af1ican Macrnlohieae clade
(se11su. Bi uneau et al., 2001)
Val'ious species used for timbe1 (gombe, sapo, angok, bondu, tirnba) for
const1uction, carpentry, furniture , joine1y, pa ne lling, venee1s, musical instruments,
plywood ancl blockboarcl; also used for charcoal, sot'cery and medicine
TRIBE DETARIEAE 103
 Gilbertiodendron bilineatum Drawing by f. M. Stones
Pellegriniodendron.r.Leo11a1cl 1955
l sp,; \YJ ~ind \V/C Af1ica (lvory Coast, Ghana, Nigeii;,1 , Crnne1oon and G<llmn)
Named fo1 F. Pclleg1in (1881-1965), french botanist and au thrnity on Afiic111
Leguminusae, and de11dro11 (Gk: t1ee)
T1ees; tropical lowland rain forest
l!cfe1ence(s): Leonard (1957: 245-248); Aub1eville ('1970: 185-187)
Seg1 eg:.1tecl f1om Macrolohium by Lc2on:ircl (1955); part of the exclusively Al 1ic1n
M<.1crolobieae dade (sensu I31uneau et ol , 2001) ::ind possibly not distincL hrnn
Gil/Jerllode11drun (Bnineau el al, 2000)
Used fo1· timbt.!r (joinery and dorn fram es)
Gilbertiodendron .r Leonm11 952
c 26 spp ; cenu cd in Guinea-Congolian \V/C Afric:<1 k. 19 spp.) south to Angola ,
with c 7 .spp. in \Y/ Af1 ka
N:rn1ccl fo1 Prof. G Gilbe1't, Belgian botanist and contributrn to Flora du Cungo
/3e/JJ.e et d11 1?wauda-Ur1t11di, and dendrcm (Gk: t1ee)
Trees; t1upical lowland win frnesl (sometimes im1nd;.1ted) ancl seasoncdly d1y
frnest, often along livers; some species form monoclominarn stands
Hefcrence(s): J.eonaicl (1957: 232-245); Aubieville 0970: 199-219); Lock C 1989:
11-14)
Ci/bertiodendron was erected lo accomm odate a numbe r o! species previou<>ly
treated under Macrofohiu.111 (see t:axonomic notes uncle1 th<1l genus) The genus is
patt of the exdusivd y Africa n Mac1olobie<1e dade (seusu ll1uneau <'f al , 200I> hul
is not suppo1tecl as monophyletic in th e analysis of I-Ie1 en deen et al. (2003a); sec
note under Pel/egriniodemlron
Va1 iou.'i species are used frn timber (limbali) ro1 conslnictirn\ floo1 ing, staiis,
c.1rpent1y, joine1y, vc..:nCt..'rS ancl boat.s; ;.1L"io used as medicine ;mcl ln111wn food
(seeds) in limes of short.ige
Isoberlinia Crail) s, Stapf 1911
c, 5 spp ,; AfriG:t (4 .'ipp. wicksprcacl in Suclani;rn to Z;unhcz ian woocllancl; 1 sp in
Z<1nzilx1r-lnha1nbane E T~1n Z<Jnia)
Fmm iso- (Gk : equal) ancl JJerlinifl ; the gen us is allied 10 1Jerli11ia hut cliffe1s in
(usually) having petal.s of equa l length
T1ees or sh1ubs; seasonally clry t1opi Gil frnesl :rnd wooclland (the E Tanz<ini:in sp.
occu1s in lowland to upland min forest); speci es often oc:cul' in monoclo1nin: 1nt
stands
l!cferencds): Leonard 0952b: 376-383); B1enan 0963); l.l1enan (1967: 13H-llt3);
Luck (1989: 14)
Placccl within the exclusive ly Af1ican Maciolobieae cl;1cle (sens11 B1uneau el({/, ,
--
2001) but genetic affiniti es remain um::Je;11
Va1 ious sp~dcs used frn timhet (mbarik~1 , mtonclo, ;_1bogo, dolrn) , e,g, fo 1 hc:tVY
/ulbernardia b / ·
conslruction ancl 11001 ing; also used for medicine <mcl human food (leaves)
Jsoberlinio angolensis var. losioco/yx Drawing by L. M. Ripley
104 LEGUMESOFTHEWORLD
Oddoniodendron micronthum Drawing by/. Sausotte-Guerel
r ey1 Photograph by/. J. Wieringa
Mlcroberlinla blsulcato (1, 2); M. brazzavlllensis (3-11) Drawing by G. Chypre
Oddoniodendron ne w;1c1 I925
c 3 spp; W'C Af1i<.:<1 (C:.1meroon, Galion, Congo [Kinshasa! ~md Angola)
N(1111ed frn the colleclOI' (Oddon) of the holotype o f 0 micr<wlbum Oicll'lns)
Baker f, and de11dro11 (Gk : tree)
TrcL:s; t1opicaJ lowland r;Jin forl.?St
Rde1ence(sl: Aub1cville 0968: 260- 263); Lock (1989: 18)
Placed wilhin the exclt1.sivdy AJ'J kan Macrolohic;1e cladc (se11s11 Ht uneau el al,
2001) hut genetic :1ftlnitics remain unde<u ; Oddouiode11dron romerui Mende:-; is
pooily known. {Aurhrn 1 .s pos1sctipt: Ngok 13annk & Bretcle1 (2004) t\!cogni se 6
spp, in ; 1 1evision o f the genus, dcsc1 ibing 2 new spp f1om Gc1bonl
Oddoniodendron micrmztbum is used frn timber
Microberlinia A.Chev _ 1916
2 spp; WC Ar1i c.1 (C11ne1oon .ind Gabon)
From micro- (Gk: snull) and 1Jerli11ia, frn· its 1cscmhlancc to the genus /Jerli11ia
but with smaller leaves
·1·1ees; t1opic:1I lowland 11.1in !'orcst; fo1ming mon<Kl<1111inc1nl st<1nds
l!eference(s): Aui>rcvillc (1968: 286-288; 1970: 266 - 268)
lts position is unresolved within the exclusively African Mac1olobieae d ~1d e C1w11srr
Bn1ne1u et a/. 1 2001)
1Hicroherli11ia 1Jrazzaui/le11sis A Chev (Aflican zcb1awoocl, zeh 1a110, zi11gam1) is
l\.'ied commcn.: ia\ly for ti1nbc1 (co11sl1uction, c1uality fu1•nitu1e <lml joinery)
Julbernardia Pelleg1 1943
1l1y/m.:a11th1ts Tul (J844); Paralierlinia Pellegr. 0943); Pse11do1Jerli11ia PA l)uvign .
(1950)
c 11 spp.; c. 5 spp in Guim.•a-Congolian \'(IC: Afric.1 (Nige1fa, Carnctoon , Congo,
Gabun, Congo (Kinsh<1sa] cind Angoh1); c . 5 spp. in Z1.11nbczi<111 woodl;.md (1 sp. in
\V/ Tanzania) <11)(1 l sp in Z:tnziba1-lnhamb:1nc E Keny<t and E Ta nzania
Na med frn Julc.'i Bcrna1d (/1. 1924-1931), former Governor of Gabon
'J'rces orshruh.s; t1npical lowhtncl 1ain frncst <.tncl 1·ive1i11e forest, OI se;1sonally cl1y
fo 1est, woodlancl , bushland ;111<1 thicket, often fo1·ming 111011oclominanl slamls
Hcle1e!lcc(s): ll1enan (1967: 145-149); Aui>1cville (1968: 289-29'>); Lock ( 1989:
15-16)
This <tnd the follow ing five geneia fo1m a suongly supporlL'd dacle (Gervais &
Bn111e~1u, 2002; also called the 'babjit' clmle .w:11s1t \V'iering~1 & Gervais, 2003), within
the exclusively Af1ic111 Ma c:rol ohi~ae dade (se11s11 11runcau el ol, 2001) The position
otj. jx;//egri11imw Troupin (= Pamb<'rlinia Pclicgr.) w ith i11J11/lx•rnardia is uncetrain
(Bruneau el al, 2001 ; \Vielinga & Ge1vais, 2003» as s;11npling remains porn;
7l~JJ/t1c:w1tlms Tul., though! to he a 111011otypic genus f1om Brazil, is basl'd on a single
co llection o!J. pr111iculatc1 (Benlh) T1oupin J'1om Mozambique (Bretder,j;er:a, comm.)
Va1 i(>LIS species used for timl x:T (zeb1ali, awour:.1, heli, rnuw;.1) foi construclil)n, quality
fumitun::, c;1q1cnlly, juine1y, p<tndling and tlcx)Jing; also us~cl ti.>1 fibre (1oofi11g, lm~es
and ba11ds), fin:wo<xl, cha1 coal, dyes, ;1s OJ namentals, bee plants and medicine
TRIBE DETARIEAE 105
 dPA10A1cl.klr
O<J~(':)
'1~Q~ ai
Brachystegia torrei Drawing by J. Chandler Tetroberlinia longlrocemoso Photograph by J. J. Wieringa Blklnlo evrordii Drawing by W. Wessel Blklnio pe/legrinii Photograph by J. J. Wieringa

Bracbystegia Ben1h. 1865 Bikinia Wie1 inga 1999

I
26 spp; Africa (cento ed in Zambezian wood land [18 spp.]; Guinea-Congolian 1l1011opetalantbus ~met. non 1-l;ums, pro pa rte
forest 16 spp. in \VIC and 1 sp i n W Afr ica] a1KI 1 sp in Zanzibar-l nhambanc E 'JO spp .; WC Af1ica ( possibly noilh to SE N ige1 ia, 1h1 o ugh Cameroon, Congo
Mozambique) {llrnzzavi lle), G;ibo n, Congo (Ki nshasa ] sou1h to N Angola)
From bmcby- (Gk.: sho1t ) and stego (Gk : shel ter), for 1he brncteoles which From bikini, the two-p iece ba thing su it itself named afk1 the P;-1citk isl<ind, Bikin i;
enclose ~ind p1otcct the fl owe r buds before anthesis for the rused ;1ch1xial sepcds wh ich consist of two pi eces
Trees and sh1ubs (nu ely Sllffllltices); lowhrn cl t1opical rain fore.st and se~1.sona lly T1ee:;i trop ical lowland (~omet im e.s peri odically inu ndated) rain fo 1est :and 1iveiine
dry fore.st, woodland, wooded grassland and bush!<Jnd, often along 1ivers, nw rgins forest; often occu11 ing in monodominant s t ~ind~
of wetlands tind on upland plateaus; a nu mber o f species frn 111 munodomiru nt Jkference(s): W ie1inga (1999, 187-240)
sta nds The ge n us Bikinia was erected to <1ccornmodate six t~1 x.i previously placed in the
Re fe1ence(s): Aub1 evi lle Cl 970: 258 -265); Chik uni (1998) now disb;rn decl Monopetalanthus, toget he1• w ith 4 new species cl escl'ibed by
Twenty three ptuarive hybiids have been published among the Zamb..:zian W ieiinga 0999); p1obably monuphyletic and sister to TetralJerlinia (Wie1inga &
domain taxa, illustrJt ing the p1oble1ns uad itionally encounte1ed with cl elim il ing Ge1vais, 2003) but fu1the1 sampling is needed 10 confirm the monophyly of
species in Bracbystegia (Hoyle in !31enan, 1967: 157 - 196) Howeve1, BikinJa; see taxonom ic notes unde1ju/lJenwrdia
m oi phomel 1ic analyses by Ch ikuni (1998) 1evealed 1hat there was Jinle evidence U.sed fo 1 ti111be1 (ando u ng, ekop 1ouge, mayo) frn co nsu uction, poles, e<11 pentry,
o f the phenorypic intennediacy distingu ishing hyb1 ids; rnther the d ifficulty with joine1y, r>Lllp, plywood <ind venee1; also used fo1 cha1coal and medicine
species li mits was due to irn1deq uately defined states of highly va1iable chawcters;
.see 1axonomic notes underjulben wrdia
Used for ti mber (msasa, mtundu, okwen, bom angH, arielht, naga, meblo), e.g .. I curia Wielinga 1999
fur construction, flooring, furni ture, jo inery and plywood; <dso used for fudwood I sp.; SE Ah ico1 (Za nzibar-Jnhambane E Moz:1mb i4 ue)
(fi rewood and d i arcoal) , fib 1e (containers, clot h, mars and bark rope), human F1om icuni, one of several simila1 locc.11 Mozambique nmn es frn I du nensis
fo od (seeds), medicine, dyes, livestock fodder, bee pl ants, 0 1namentals and \X'ieringa
shade plan ts T1ces; t1opica l Jowhtnd coasta l fo1est and littoral th icket on older sand du nes,
fo11ning (almost) munodo1ni nan t stands
l tefere nce(s): Wie1inga 0999: 24 1-248)
Tetraberlinia (Harms) Hauman 1952 See ta xono mic notes unde r.fulben w rdia; positio n wirhin 'babjit' clack: sensu
7 spp.; WC Af1ica (6 ' PP· from Came1oon, Equa 1orial Guinea, Congo [Jl1azzavil le], \X'ielinga & Gervais (2003) is unce1 ta in
Gabon, Congo {Kinshasa] to Angola); I sp. in W Afrirn ( Liberia) Used fo r timbe1 (construction) , fitewoocl Ctllc.I bark (canoes)
From tetra- (Gk.: four) and Berlin ia, for the fou r sep als (adaxial pai1 are fused)
compa 1ed to five sepals found in Berlinia
T1 ees; tropical lowland m in forest and ri veri ne forest; o ften occl 111ing in
mo nodo1 tl irntnt stands
Uderc m:<:(s): \Y/l.•nngn 1 1' '.J'J: l55-l92)
~ II supported as ~I'll<!< to JJ1k1mr1 in 1hc unulysis of I lc1 ~nclC'en ,, , al. ! lfl 15al and
th wmbin I :umlysl• of W1on11g'1 ti< rv~~' (2003); see 1~x< 111 nuc no1c, under
j 11//Jr.•nu11·rlft1: W I rltig .1 1 19' 9) d csuil •cl 1wo new sp x·ics and tn1nsfernd a 1hird
f1 o m the disbanded genus Mon opetalantb us
Used for timber (ekaba , ekop) in cons trucl io n , ca1 pen11y, joinery, panelling,
venee rs <tnd blockboa1cl

lcur/o dun •
Tetrober/lnio po/yphy/lo Drawing by J. Sausotte·Guerel ensis Drawing by W. Wessel

106 LEGUMES OF THE WORLD TRIBE DETARIEAE 107

Aphanocalyx obscurus Drawing by w. Wessel Aphonocolyx richords/oe Drawing by L. M. Ripley

Aphanocalyx otivc1 IH7o

Mrmo;1t•fl'llt111lbmi 1lnrrns 118971 f"'C> fJll11<'
H 'llP•; c ·111retl In w e Af11l.1 111 .<pp. lw111 111 ·t1.1, C:nmcroon, E<1u111111l.1I
l :uln .1 , C1 1111u ln1.1u .1vlllcl, GnlJo n. ConAO (Kl nsh.1.:d 10 G Angoln I <p. hm<l •
suhsp o/';.t WC l<IX()l1) ill \V A/'1k ,1 (SiL'l'll.I L<..:< >nL', LilK:1i;.1 a11d TV()I~' c:(usl); 2 sp11
in /.~1111hczi<m SC Af1it.:a
Fwm a- ((;k.: wilhout) 1 pbouero- (tJk ,: cvid<.:nl) and ca~)ix, 1'01 iht: oh:-.olctc rn
rudi111cnl:1ry c dyx
Trees or sl1ruhs; t1tlpi t.:~1l lowla11d (so ml'li111cs pc1iudic1lly int111Lbli..xl) l()
StlhlllOlll:lllC l:til1 furc.'il and li Vcril1C ro1CSl , ulh.'ll C>ll 1Hll1ie11\ porn ,'O:tlldy soil.1, ; Id
...:L': t:-.onally d1y finest, woodl:l nd ;111d wooded g1<1:-.sbnd , often along li vc1s; usu :d l)'
f( >1mi11g 111C>nl>lhlminanr stands
Hcf'c1cncc(s): \Viet inga ( l 99'.J: 1 Vi - l8(J)
l7cnci•ic ;1lfo1itics ~lie wirl1 'Ji.:tm/Jer/i11ia ;iml HiN11ia (\Vieringa & CL' l'V<iis, 200->l:
see t<1xomnnic notes under /11//Je1•1wrclio \V1e1 ing: t ( 1999) l1;1nsfc11cd nine sp~( it:'>
from Mu110/JL'fala11th11s ro !lphw10Cll~JJ.\' and de.- ;nilK:d two ncw s1h..: '-: ic.o;
Vario us spc<:ics UM;d for limhcr (anclo ung) f'or co nslru ctio n, urpcntry, impk11 \l 1ll."
;incl vencc1s; al.-;o fo1 b 111k fboxes) , human foml (M_:cds) :mcl 111txlici nc

Michelsonia 11>1L11n"" 19'i2

I s p ,; Ciuinl·a-Congolian WC Al1i c; 1 (E Congo Jl\:inslu.•.:d on the c:1.'-'l c 111 rim u! tlw
/'.;iirc basin)
N~1111cd ror· MA MichL'l.'-'Oll ( 1807 - [~JO;-)), ~l lk!lgian ;1g1icultu1:d c11gi11ccr wlH>
lll:tde .'iOlllC llOU lOllcc1ions (111oslly lll'CS), the g1eal 111<1io1ily r1rn11 Co ngo
(l<i11.'\has:1), including the l)' IR~ spc<.. ies ol the genus
Trce.'i; tropical ~l1hrnoillane r<tin fotc:-.l, o lh.:n on hilltops <llld slopcs, in cx1c11si\1.:
~ind ;tlln<l~l 11Hinodo111i11anl staml'i
Hcrt.:1c11cds): \Vic1ing;1 (1999: 2·19 - :l'd)
Not sampled in the an:lly:-. i.s of I k:n..: nd~en el al (2t10j;1) hut lx1scd on
mrnpholngy, affinities arc possibly with gc·ner.1 of the 'lx1 hjit' cbdc (\X'icring:t N
Ccrv ~1is, :lU03)
Used 1"01 timbe1 (ungL' n..:dtK:cd l>y L'Xtc11siv c l()gging in the mid 201h ccnlu1y)

Browneo grondiceps Photograph by G. P. Lewis

Miche/sonio microphyl/o Drawing by P. Halliday

TRIBE DETARIEAE 109

108 LEGUMES OFTHE WORLD
TRIBE
cassieae byG. P. Lewis
Tribe cassieae Bronn 1822, emend. Irwin & Barneby (1981, and formally in 1982)
Tribe Poeppigieae Britton & Rose (1930)
Trib Cassie.ie, long ·onsid red to b an artill iaJ
group, w:is divided l Y__Irwln & B:i rn. I . 1 9~1 ..into
fiv sul tribes ( ratonunae [m nogenen J, Dt:1. lu rn1e
(1 3 gen •ra l Du parqueri inae [monogen rid, Cassii nae
[3 gen ra I, and 1.abi ·he inae [2 rcneral). They sta t cl
that tlvy wou ld have u ore! cl criba l rank to the five
subtribes if they had not been dissuaded by others.
The 20 genera of tribe Cassieae sensu Irwin &
Barneby (1981) remained unchanged in Polhill 0994).
Since then, Uittienia has been reinstated as a
monospecific genus in subtribe Dialiinae (Rojo, 1982)
and this was followed by Hou in Flora Malesiana
(Hou et al., 1996).
In the rbcl phylogeny of Doyle et al. 0997) the
Cassieae (represented by 6 genera) is not monophyletic.
In addition, Cassia did not group with Senna or
Chamaecrista, and Ceratonia grouped with
Gymnocladus (traditionally placed in tribe
Caesalpinieae). Kajita et al. (2001) similarly found the
Cassieae not to be monophyletic, with 4 genera of
Cassieae interspersed amongst genera of tribe
Caesalpinieae, and a well-supported clade composed
of Zenia, Apuleia (subtribe Dialiinae) and Petalostylis
(subtribe Labicheinae) basally branching to the rest of
the legumes except for tribe Cercideae. Chamaecrista
grouped with Senna (except for Senna alexandrina
Mill., represented in the Kajita et al. study by its
synonym Cassia senna L.), and Ceratonia grouped
with Acrocarpus of tribe Caesalpinieae (a relationship
also supported by the analysis of Doyle et al., 2000).
The molecular phylogeny of Bruneau et al. (2001)
largely supports the monophyly of the subtribes of
Irwin & Barneby (1981) but not of tribe Cassieae .
1-J ev r, the Dialiinae are monopltyletic only if the
bigencric Australian subtrib 1.abich •inac is included;
Labichea and Petalostylis grouping with the
Australasian Storckiella. The monospecific neotropical
genus Poeppigia, sole member of its generic group in
tribe Caesalpinieae, occurs as sister to the Dialiinae
~ens. lat. clade (the two share cym< se inflorescences)
m the Bruneau et al. (2001) analysis. The same analysis
sugg Sled thaL Ceratonia has a close relationship with
A.er wpus . upport:ing th fin ling. f Doy! el al.,
2?00 :'Ind I jita •/ al., 2001) anc.I tJ1e Maclagas ·an gen us
7etrciptcrocarpon tra liti na !J y pla eel in the
Dim rpllandrn gro u p >f t ribe Cae al plni a
LEFTS
enna leandrll Photograph by D. Du Puy
Ceratonia is clearly distinct from all other Cassieae.
The analysis of Bruneau et al. (2001) supported earlier
studies (Doyle et al. , 1997, Kajita et al. , 2001) in finding
that subtribe Cassiinae was not monophyletic, although
the exact relationship between the three genera,
Cassia, Senna, and Chamaecrista differed. Herendeen
et al. (2003a), in a combined molecular-morphological
analysis which expanded on the study of Bruneau et
al. (2001), found strong support for a Dialiinae sens.
lat. clade with Labicheinae nested within it, and
including Poeppigia from the Caesalpinieae which
grouped with Madagascan Baudouinia. The single
species of Dicorynia sampled, D. guianensis Amshoff,
nested within a clade of six Dialium species. Ceratonia
grouped with Acrocarpus, with Tetrapterocarpon sister
to these two, confirming the isolated position of
Ceratonia in the Cassieae. Cassia did not group with
Senna or Chamaecrista; neotropical, monospecific
Pterogyne, traditionally placed in its own generic group
in tribe Caesalpinieae, was sister to the Chamaecrista-
Senna clade, a relationship worthy of further study. In
the study by Herendeen et al. (2003b), Cassia, Senna
and Chamaecrista do form a clade, with Chamaecrista
sister to the other two genera. These authors also found
a moderately well supported Umtiza clade which is
basally branching in the Caesalpinieae sens. lat. (see
page 129), comprising the genera Umtiza, Gleditsia,
Gymnocladus, Ceratonia, Acrocarpus, Arcoa and
Tetrapterocarpon.
Five genera of Cassieae have not yet been included
in any published molecular studies: Androcalymma,
Eligmocarpus, Kalappia , Mendoravia, and Uittienia.
Koeppen & Iltis (1962) remarked on the absence of
silica in the woods of Martiodendron and
Androcalymma, a character that differentiated them
from the highly siliceous woods of Apuleia, many
(non-Asiatic) Dialium species, and Dicorynia
(Koeppen, 1980). They also commented that sclerotic
parenchyma cells, rare in the Leguminosae, are found
in Martiodendron and Androcalymma, further
evidence linking these two genera .
The Cassieae as presented here (Fig. 22), containing
21 genera and (729)-732-(735) species, is retained
largely in the traditional sense of Irwin & Barneby
(1981) and Polhill 0994), a lthough the tribe is
undoubtedly an assemblage of unrelated taxa which in
TRIBE CASSIEAE 111
time will be segregated into more natural groupings.
For example, Ca ssia, Senna and Chamaecrista
comprising (655) - 658-(661) species are more closely
allied to Caesalpinie ae sens. lat. (Fig. 22), and will
form the basis of a more narrowly circumscribed tribe
Cassieae . This will render the remaining genera of
Cassie ae sens . lat. in need of high e r !eve[
nomenclatural restructuring. Nevertheless, Ceratonia is
here removed from the Cassieae to the Caesalpinieae
with which it finds closer affinity, and Poeppigia i~
moved into the Cassieae (as a new member of subtribe
Dialiinae) from Caesalpinieae sens. lat.

Cercideae (see page 57)

Detarieae (see page 69)

* Duparquetia Cassieae: Duparquetiinae

Poeppigia
Baudouinia
?Eligmocarpus
?Mendoravia
(")
)>
Distemonanthus (./')
* Apuleia (./')

rn
)>
Storckiella
rn
Labichea Cassieae:
Petalostylis Labicheinae 0
)>
r
Koompassia
Duparquetia orchidacea Drawing by N. Halle Duparquetla orchldacea Ph otograph by D. Kirkup
Martiodendron
?Androcalymma Duparquetia na;11. rno5
Ol igvsle111 011 13cnt h (1 865), no n Tu rcz (1858)
1 .'ip ,; en d em ic to \VC Afric;1 (Liberi a to the lower Co ngo D;.1sin)
?Kalappia Named fo 1 the Revere nd PVA Duparqu et 0830 - 1888) w ho collected pla nts in
P1e nch Equato l'ial Af1 ica (as it wa s then kn o wn ) bt.::tween 1863 and 1887
Scandent 01 sa 1mentose sh1uhs, robust lianas (' hush 1opcs ') 01 (less fi equently)
Zenia small t1ees; evergree n fou~st especi all y ne;,11 1ivcrs, seco nda ry fo 1est, .sc1L1bl<1 ncl,
w:1ste g1 ou ncl and coastal wooded grn:-;sland
llcfc 1c 11cc(s): Aubrcville 0968: 47 - 49; 1970: 47 - 49)
Pla ced in th e 111o noge11e1 ic su bt1ibc Dupa1 q ueriinae by I1 w in & 13arneby (198 1)
?Uittienia
although they had 01 iginally inlenclecl lo est;,1b lish ~1 sepa1ate ll ibe to
Dialium <1cco111 m odate the ge nus, q uoti ng Bcnrlrn m - w ho ( in 1865) clescii becl it (under
Dicorynia Oligosfemun ) - 11s "pe1fectly i:-;ola ted, combini ng cl1e1racte1s of very cli ff<:! 1Cnt
11ihes" D11parq11elic1 orcbidt1cea Dc1i ll hc1s no parclllcl in the C~1 esalp in io i de:ic:;
esp ecially notewo1thy are its zygo mo1phic oichid-like ~l owe 1 s, the syn and li t1m of
4 -5 anther.s, th e cbstically clehiscent pocl ancl the cliporate pollen. In the
co1nbinecl 111olecul<1r-morpho logic;1l analys is of He 1en decn et al (2003;:1),
Papitionoideae DHpt1rquelia i.11 pl:tt:cd in an isolated b;;1s;ill y br:inch ing posit io n siste1 to the 1est
o f the Legt1111ino.sclt.:! (above tribes Ce rcidcac and Deta ricac sens lat ), suggesti ng,
at lea.st, th at scp<ll <JlC ti ibal St<ltus might be justi fied fo1 D11jJarq11elia
It h::is potential :as an rn•namental due to i[s sh o wy white and pinkish 1ed nowers;
Caesalpinieae pro parte (see page 127) the sinuous stems a1e used :.is ties 0 1 m ace1c1tecl and p u t into palm-wine to
at:t:de1ate fe1men t<-1tio n; the f1uits mi xed wit h tobacco serve <IS a snu ff; also used
as ;1 tr1.1clit imuil medicine

Chamaecrista
Senna Cassieae: Cassiinae
Poeppigia c P1esl 1830
1 ."ip,; N eot1of)it:s (Mexi co, C Ame1ica [H o nclu1as, G uatemala, El Salvaclo1J, Ct1 ba ,
Cassia Venezuele1, Colo m bia, Peru, l \olivia, ci1t: um-Amazo ni<1n Dr:.1zil (inclt1 clin g Acre,
llonc16nia, M ato G 1osso, M inas Ge1;:1is, Bahia, Pernam buco •rn d Piau f])
Named frn Eclt1ard F1ied1ich Poeppig C1798-JH68), Geiman botanist, zool ogist and
cx plo rc1 who t1 avellecl \Viclely in Latin Americ;.1; P1ofessor :a t the University of Leipzig
Caesalpinieae pro parte (see page 129) T1ees; seasonally cl1y Lm pirn l foiest ( includi ng caatin g:.1)
Mimosoideae Refc1ence(s): McVm1gh (1987: 73-75)
In Pol hill & Vidal 0981 ), PoejJjJigia was p laced as sol e 111embe1 of its gene1 ic
g1oup in t1ibe C 1esalpinieae, ;1 position 1ett1ined in Polhill (1994) . In the moleculai
;.111al ysis of 131unc au el al. (200 1) the genus is siste1 • to th <:: Dialiinae sen s lat clacle ,
? =placement uncertain due to lack of molecu lar data an d in th e comb ined molecul<u -mrnpho logicc1l an alys is of H e1endeen el al
* =position weakly supported (2003.i) it is nested wit hin the .s;.1111e clade (m~ m be rs sha1c cy mose innrnescences)
;Js si.s ter to Madagasca n 1Jmulo11 i11ia Removal f1 o m tri be Caesalp iniec1e to tlibe
Cassicc.1c (."iuht1ihe Dialiinac) is a logical co nclusion of the 131uneau el al ;1ncl
H erendeen el al ~ tuclies

-------
Poeppigia procera CJ)rc sl (tasa~I) is locall y used for const1l1cti o n timbe1, focwoucl
FIG. 22 Diagram of relationships between genera of Cassieae largely based on Herendeen et al. (2003a) Poepp; ·
gia procera Photograph by G. P. Lewis cmcl medici ne

112 LEGUMES OF THE WORLD TRIBE CASSIEAE 113

 Baudouin/a fluggeiformis Photograph by D. Du Puy
Baudouitiia I3aill. 1866
6 spp.; cnclcmk to M::1dagascat', 1 sp., B fluggeij(>rm;s fh1ilt, i!> widcspre.1d .ind
highl y va oiablc
Named for Ctptain Baudouin, :rn explore1 and planl collectrn in New C:dcdonia
Sh1ubs 01 trccs; ttopie<d humid everg1ecn coastal fo1t~.st (2 spp) to :-;eason:ll ly d1y
xc1ophyric woodland (4 ~pp)
Rde1~11cds), Du Puy & Jt;ibevohit1 a in Du Puy el al. (2002' 61i - 71)
A 111cmbe1· of subtribc Dialiinae; the species can be divided into 2 g1oups h<1~cd
on mode of fruit fragmentation , In the combined molecular-11101 phological
analysis of l-le1t..:ndecn el al. (2003a), 13a11clo11i11it1 g1oups with the neottopical
genul'i Puepplgio (<1 relationship wrn thy of further stw.ly, not Je,1.st because
PuepjJigio has tradirionally been placed in tribe Caesa lpi nicae), and the two
genera arc basall y branching in the Dialiinae sens. lat. dadt::. Du Puy &
Hab1.:vohitn1 in Du Pt1y l'I al. (2002) comment that n1J'ts of hairs on the anthc1s of
some spcck:s, togethc1• with cymose inflorescem.:e."i, sugge:;t a ielationship with
Eli;:mocaljms, also a Mad:agasGrn endemic; PuejJjJigia cd."io has cymo.-;e
intltn esccnces
The ha1d wood of some species is used in Gupentry, <md for poses an<l pole:-. for
houses and cattle endoswes; the deeply folded trunks 1.ind very hard wood ol It
rouxeviffui l·t.P1.:11ier a1e p1ized for making 01naments, lamp s1ands and walking
stit:ks and a1e sought :liter for their mystical propc11ies
Eligmocarpus capu1on 1968
1 sp.; n1.1rrowly r12striclccl endemic in SE Madag11sc 11'
From be/i14mo- (Gk.: winding or twisting) and ,:rutJO'f (Gk~ : fruit), 1cfe1 ling to the
folded zig-z<1g clrn1actc1· of the ovary and pod
Ti ees; seasonally chy tiopical deciduous or paltially evt.:1g1i...:cn woodland.
U- c. liOO 111
Jtefe1ence(s), Du Pt1y & lbbevohitia in Du l'uy el of (2002' 72 - 75)
Plac.:cd in Sl1ht1 ihc Dialiinae, ;_ii though uniqm: in irs co111bi 11<1tion of opposiri. .:
lea fl ets, a highly modified anclrnecium (5 st~nnens wilh fused filaments <\lld
tufts of long hai1s <II the an the r apices, 5 stamens free with glal>rou.s ::inthcr~).
<ind renrn1 kable tightly fo lded pods. As yet not induc.lcd in any published
mo\ccuh11· ana lyses
Eligmocarpus cynametroides Drawing by M. Tebbs
114 LEGUMES OF THE WORLD
Mendoravio dumazlano Drawing by M. Tebbs Olstemonanthus benthamianus Drawing by unknown artist
Mendoravia captnnn 1968
1 sp.; n:.11 1owly 1est1ictcd endemic in SE Madag.asc;.11·
De1'ived from the com nmn mime 'mendoravy' (f)u Puy &. H<ihevohitia in Du Puy
et ol., 2002)
T1ecs; lowland eve1g1een frnest, c.:. 50-JOOm
Jkfe1ence(s), Du l'uy &. Habevohilla in Du Puy el al, (2002, 71-73)
A monospecific genus in suh11ibc Dialiin:ie; of unknown generic affinity although
Irwin & B<11neby (1981) suggest .:1 relationship with nauduui11ia and pe1haps
Sfl,,.c:kiella
The lrn1•d du1;,1ble wood is used in c.:onsn ·u ction and c:11pent1y
Distemonatithus Bench . 1865
1 sp.; endem ic in \VC Af1ica (Sie1r::1 Leone to Gabon)
F1om di- (Gk.: two) and stemo11 (Gk.: stamen) and m1thos (Gk.: flowe1); thL:
tloWel has l WO COllS[>iC.:llOUS fe1 ti]e .st<lllli..:llS
T1ees; tiopic;,11 1ain frne.st
ltcli.:rcncds)o Aub1cville ( 1968, 14 - lo6); Koeppen (l 978)
Phu:ed in suht1ibe Dialiim1e, <I position supp01ted hy the analysi.'i of J-Ic1endeen
et al (2003a), Koeppen 0978) noted sliiking anatomical similari ties between the
woods of Distemona11tb11s ~tnd Apuleict f10111 S Ame1ka; hoth contain amo1phous
silica bodies deposited in the same hasir..: pattern , The.; smell of a fresh hark slash
of Distemo11a11tbus has been compaied with 1h ~1[ of boiled chicken
Distemo11a11thus /Je11thamia1111s Baill. (movingui, ayan, ayanran, Nigeii:rn rn
yellow satinwood) is ;:1 valw:1hle timhi:-1 well-suited to unde1wate1· and
unclergl'Ouncl const!'uction; also used for fumiture, cabinet wo1i<, joinc1y,
panelling, floo1i11g, 1oof-."ihinglcs, plankin~. house-post'i, diums and canoes;
scvc1<.il local medicinal uses arc known, a yellow dye from the roots has been
U."iecl fo1 body decrnation and it also h;ts allcgecl magical pmvers
Apuleia Man. IH37
2 (- 3) :;pp. (some aurhors consider the genus to be monuspcci tk) ; widesp1c~1d
~1c1oss S Ame1ic<1 in A1m1zonian ~md exua-Amazonian B1azil, Venezuela, Peru,
Paraguay, Bolivia ;ind A 1gentina, l sp endemic to Cea1·<'1 in NE Brnzil
Named frn Lucius Aruleus (124-c. 170 AD), Pl:tlonic philosopher and autho1·
1ernembe1ecl for '111e Golden Ass, a p1osc n<111ativc
Ttces; in a w ide range of tropical and subttopical h:1bitat.s, ft om rain frnest, 1ive1inc
and g:tllery forest to seasonally d1y fnreM ;111d thom-sc1uh woocll:ind (caatinga)
Jtde1ence(s} Koeppen 0978); Fernandes (1~)9t,, 284-287)
A memlx:1 of suht1ibe Dialiinac; in the 111olcc.:ula1 analysis of Kajita el al, (2001),
Apuleia is sister to a Pelaloslyfis-Zeuia dad<:: ~ind Diali11m is sister to these th1cc ,
\'\food ::111atomy suggests a rclarionsllip wi1h /Jislemonmtlb us (Koeppen, 1978)
The timhe1 o/ II /eiucmJHt (Vogel) .J ~f.Macbr (gi·:ipia, gat'a pa, yvy1a f)l'Je), is usL!cl
Apu/eia leiocarpa Photograph by G. P. Lewis <.:olllllH:.: 1'cially for~ e g,, high quality furniture, hctivy const1uction and flooring
----~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
TRIBE CASSIEAE 115
Storck/el/a austroliensis Photograph by P. H. Weston Storckiella pancheri Photograph by M. F. Gardner Loblchea lanceolata Photograph by B. Fuhrer Peto/ostylis labicheoides Photographer unknown

Storckiella Seem. 1H<>1 Labichea Gaudid1. ex DC . 1825

4 spp ., I in Fiji, 2 spp. endemic to New Call:donia, 1 sp endemic in Aust1<1li:I (N
Quccnsl:tnd)
Named for Jacob Stoick. the assistant of Bcithold C. Seenwn (aurhrn of rhe genus)
during his explo1;t1ions in fiji (1860-61)
Trees or Clmgc) shrubs; t1opkal 1ain fo1csl (S austrahe11sis J.H.Ro.-.s &: B Hybnd in
;m ;_uca whe1~ the rainfall is sc1icl to exceed 3.8 ma y1.:;u)
Refe1en ce(s), Hoss & Hylancl (1983); ltuss Cl998b)
Sturckie!ICI is a 1nem[)(:1 of subtrihe Dialiinae; a difh..:1c1KC in sr<.1me11 numl)t:I 111
the th1ec meas ol di.stJibution of th~ genus is inte1esting: thL: f'iji Sf). has lO (- [2}
stamens, the Austulian sp . 5 (-6), and the Nl'W Cak:doni;1n spp, ~ (- 5) In !he
comhinecl mokc:t11<1r-11101phological analysi.s of He1endcen el o/ (2003:;1), S
ou51/ro/ieusis is sister to a Lobicbea-Pe/oloslylis cl;1dc (;.di th tee demcnts thus hci:1g
Aust1alian)
Used for wood; of g1c;1t pmential ~l.'i or narncnlals due to the massed showy yd low
nowe1s
14 spp , endemic to Awar;.1lia, disjurn..:r in \V/ Auscralia, N Tcnitrny ;.111d Queensland
N<imed fc)I' J J Lahiche (1784 -1819), <1 French naval oftke1• who accompanied
Frcydnel on his wrn Id voyage ;:md died en ro11h• to the Molun:as
Sh1ubs; seasonally dry tropi n il roc k y sandstone 1avines, t:oasral sand dum~s, desert
and dry snub woodland, mainly on granite or sand.srone
Heferc ncc(s): Huss (1985; 1998d
Tt)gethe1 with Pela/ostylis frnming suht1ibe Lahid1 ei11ae l1win & Ba1·nehy (1981);
in the combined 11101l'cular-111rnpl10logica l ;inalysis of Hercncleen el ol, (2003a),
the suht1 ibe is nested within subt1 ibe Dialiinae, with Lahicbea and Pela/oslylis
forming a well-suppotlecl chide sister to S/rm;kiella (l/tslro/ie11s1\·
Some spedes culrivated as rn namcntals; used ;is pioneer plants in n.~vt'getation
Petalostylis it B1 . 1849
2 spr; endemic ro Au.strcdia (Queensland ;.111d New South Wales)
From jJt!lalo- (Gk: fl :.ll) and s~vlo (Gk: column or pillar), alluding to the
exrwrndina1y pet;iloid styles
Shrubs; tropi<.:al alid <1re~1s, sand plains, stony ridges and rocky outcrops, creek
heels and sc:iee s lopes
lkfe11encc(s): Huss (I 99Hcl)
Tc)geche1 with /.ahicbea form.<; sul)t1iht: l.abic.:hein ae Irwin & Bamel)y (J981) In
lhe c.:otnhi ncd molecular-rnrnphological analysis of H eiendecn el ol. (2UU.)a), the
subtrihe is nested within suht1 ihe Di<diinae, with Pe/a/oslylis and La!Jicbeo frn·min,g
a well -supported chick: sistet' to Store/de/la 011slra/ie11sr~
Alk:iloicls in leaves and stems might have medit:in:d potential; usl.'cl as pioncc1
plants in revcgctation

Koompassia Maingay ex Lkntil 1873

3 spp; M::tlcsb (Sum:ttra , M:day Peninsula, Boi-nL'o, Philippines ;md Ncw <~l1inea) ,
2 spp extending to S Thaihmd
Derived f1um the ve1nacular name 'kumpas'
T1 ees (of huge st;t1t11e>; tmpical (lowh1nd) 1ain frnesl, coastal plain foothills, liclgc

~ ~o ~
...;lopes and 1ivcr valleys, l sp occasionally in peal <.tncl f1esb watc1 sw~1mps
Heference(s): Larsen el al. (1984, 83-85); Hou in !lou el al. (1996: 631-63'i)
Phicecl in suhtribc Dialiin;1e; grouping with Morliode1fd1'f.m (f10111 S Ame1ica) in
the <.:ombined molccular-mo1phologic:al an;dysis o[ J-le1enclcen el al. (2UU3a), and

,,{;
.,
t/6 C1 lmlh :11e sister to a JJia/i11111-Dicrn:y11ia clad~ . /\ £'.\-celso (Lk'cc.) Tauh cm :lltain a
height of 75 (- 88) 111
Va1 ious specil~s (kL'rnpas, Lu:tlang, mengris, ta pang) usecl ror timhe1· with a wide
;11 ray of uses, e g, cthinet making, jc>inery, venee1s, boat-builcling, fl(H>I ing,

pa11elling 1 railway slecper.s 1 plywoocl , ch:1rco;d :111d fence post.'i; also as bee forage
Koompa .
Storckiel/a pancheri Drawing by C. A. Sobel ss1a ma/accensis Drawing by w. H. Fitch (for honey)

116 LEGUMES OF THE WORLD TRIBE CASSIEAE 117

 Martiodendron parviflorum Photograph by 5. A. Mori
Martiodendron Gleason 1935
Mm1i11sia Denrh (1840); Mm1ia Denth (1840), non Spreng. (1818)
4 spp; S Americ.:a in the Glli<m~s. Amazonian llrnzil (Rio Brnncu, P<1ra, Amazunas,
A c re) , S Venezuela, and NE l31azil (Bahia, Pi~1L1f, M<11anh:io)
amed fo r C.P. P. Martit1s Cl 794 -1868) renowned Ge1 man botanist and editor of
floia J31;1siliensis, ~ind dendron (Gk: tree)
Trees; tropical 1ain forest (often pe1 ioclically im1ndated) to seasonally dry forest
and wooded grassland (savanna), below 600 m
Reference(s} Koeppen & lltis 0962)
Placed in subtribe Dialiinae; the presence of scle1otic parem.: hyma cells (ra1e in
the Leguminosae) and lack of silica in the wood indicates a close 1elationship
with the Amazonian ilndrocalymma (Koeppen & !ltis, 1962; Koeppen, 1963) ln
the combined molecular-morphological analysis of He1encleen et al. (2003a)
klarlioclelldrml groups with M~ilesian Koompassia, the rwu sisrer to a chide
conta in ing Zenia, DiC1lium and Dic01ynia
Androcalymma Dwye r 1957
1 s p. ~ endemic to th e Lip per Am~1zon lla.<>in in l3m zil
F1om a11dro- (G k: m;in ) and ca~wnnw (Gk : a cowl oi coveli ng), alludi ng to the
anthers, which are turned inward over the style; when deflexed they 1esemble a
pulled-down hood
T1ees; tropic.ii Amazonian terra firme rain forest
Refe1ence(s): Dwye1 C1957b); Koeppen 0963)
A ra1e\y collected genus in subtribe Dialiinae, <IS yet unknown in f1uit and noL
included in :my molecula1 ::maly.ses Sclerotic pa1end1ym:-1, uncommon in
leguminous wood, is found in thi.s genus and in Martiodc:11dro11, also in the
Dialiin;ie (Koeppen & lltis, 1962) Dwyer Cl957b) states that seve1a l flor.J!
chmacters suggest a relations hip with Dicvrynia and Marliudemlron (;is
Marliusia ); Koeppen (1963) also comments that several featurc:.s of the flowers
s uggest :1 possible lillk with Apuleia Dt'.cu1y11ia ~incl Ajm/eia, however, have
highly siliceo us woods while A 11drocalymma and Mw1iode11d1011 do not
(Koepren & lltis, 1962; Koeppen, 1963)
Ulttien/a modesta
Androcalymma glabrlfolia Drawing by P. Halliday
118 LEGUMES OF THE WORLD
Kalappia celebica Drawing by R. A. Soelaiman Zenia inslgnis Drawing by P. Halliday
Kalappia Kostenn. 1952
l .<iip ; endemic around Malili in Sulawesi (previously Celebes)
Derived from the local name 'kalappi' for K ce/ebica Kostenn ~me! it.s timber
T1ee.'i; lowland t1opical 1<1in forest
Reference(.,): Koste11nans 0952); Hm1 in Hou et al 0996: 625-627)
Poss ib ly now an endangered species and the gem1s has LlOt been included in ~my
molecular studies Tcnt<itively placed here b::1secl on Irwin & lfarneby (1981) and
Polhill 0994)
The timber is highly esteemed fc)I cabinet wo1 k and fu1 niture making because of
a bem1tiful grain, also once used fo1 ship building, bt iclge con.st1uction, and for
1
boa1ds ancl stiles
Zenia Chun 1946
1 .o;p. ; endemic to S China and N Vietnam
Named fo1 l-1 C Zen, executive se<.:1et<1ry of rhe C him1 Foundation f01 the
promotion of education and culture
Trees; uopical forest, 1ocky hi lls in forested ravines, 200- 750 m
l!efe1ence(s): Chun (1946); ulfsen el al (1980: 11 2- 113)
A poorly known and little-colle<.:ted red-flowe1ecl .o.; pecies . In the molectli<1r
:tlla lysis of KajiL<1 el al. (2001) there is 100% suppott for a 1elationship between
Zenia and Peta/uslylr\· f10111 Australia, with Apuleia f1om S Ame1ica weakly
supported as siste1 to these two. In the combined morphological and moleclllar
amtlysis of Herendeen et al, (2003a), Zenia is s istc1 to a Dialium-Dicotynia clade
of s ubt1ibe Dialiinae sens. /al, which in their study includes Pelalostylis
Zenic1 insignis Chun yields a high quality l'imbe 1 used for ca1pent1y, but the
supply is too small to be of any com111e1cial value; of omamental potential ;ind
c ultivated in Hawaii
Uittienia Steenis 1948
Dia/i11111 subgemis Uillie11/a (Stce nis) Steyae1t 0953)
I .'ip; W Ma lesia: Slltnat1a and Borneo (Sahah ;;incl I<a linrnntan)
N<1 mecl fo 1 the Dutch botanist Hendiik Uittien (1898-1944) who actively engaged
in rhe undergrmind movement ag~1inst rhe Ge 1mall occupation during the Second
\'Vrn Id War; he was executed in 1944
T1ees; t1opiGd iain fo1est
ReJ'e1ence(s): Hm1 in Hou el al. 0996: 714-716)
Rojo ( 1982) reinstated Uillie11ia as a monos pecific genus in subtribe Dialiinac,
rejecting it from Dic1/i1tm largely because uf a distinctive fruit type (this i.s not
supprnted by wood anatomy [Koep pen, 1980] as Uilllent'.a and Asian species of
Dia/ium both lack silica accumul:.1tioll); a.s yet not included in molecular analysis
T imber used in construction
Os
Drawing by}. H. van
TRIBE CASSIEAE 119
 •

(•)
Dialium orientate Drawing by L. M. Ripley Dia/ium guianense Photograph by 5. A Mori D/carynia guianensis Photograph by 5. A Mori Dicorynia guianensis Photograph by 5. A. Mori

Dialium 1.. 1767 Dicorynia 1icnth. 1840

/Jan.w1<1 Stccnis (19'18); Sdt1j1/1:t1 lt.1uschcrt (1982) 2 spp: l .sp e ndemic lo the Guianas , rhe othc1 in Amazonian n1oizil (cspecblly
c 28 !->pp ( b<1s\.."d larµc l y on Steyacit ( 19'> I) and Rojo ( 19821 although pct haps :111
common along the Hio Ncg10) and just cxtencling to che Colrnnhian-Vcnczuclan
underestimate; p1cvious mtmber:> 1angc Imm liU - 70); panlropical (f10111 19°N in S ho1dc 1
Mexico and Belize to 2J 0 S in Math1ga.<;car) with 1 sp cxua -tmpictl in Moz:unhique F1om di- (Gk ,: two) and cwy11it1 (Gk: dubs), app·a 1ently alluding to the two cluh-
and S Africa; not extending E of \'V'allacc's Linc ( lol :11ly absent fwm Au~tr<ilia and sha pi.:d stam~ns
Trt..~t!s; esse ntially <1 riparian genus, typically nf inund:lled 1ain fores! 01 swc1mp
New Guinea); 111;1in <.'entll:'S of cliveisity in \VC: Afr ic;1 ( 14 spp; 3 spp also in E
uu.st~tl Africa , 3 spp. cnclcmic to Madagasca1) and \V Malcsia (I{ spp. in S !'ol't..::.St (v<llZL'a), hut also on ten~1 fi1111t.: and in 'L'vergrccn seasonal forest' on wcll -
Tlwil;111d, Laos, Ca111hodi~1 and S Victna111, 2 spp in 1301 n~o. St11rnH1:1 and MaLiy clt ;.1i ncd sites; below 600 111
Peninsula, l (- 2) spp . in S India 1.111cl Sl'i Lank;:1), and l sp. wiclesp1eacl in the Heferencds): K(>eppcn ( l967)
Ne<H1opics fr11m S Mexico th1(a1gh C A1ne1ica t<> the A111azlm and Atlantic fu1est in In s uht1ihe Dialiinae; the Am::1zonia11 sp~des Dicw:p11ia pr1rae11sis Benth . is
Bl'azil, externling into Colombi;1, Pe1u, En1ad01 ancl l\0Jivh1 divided inl<l () V<11icties; in the Ulllll>ined llHJJe cu ]:\J - !ll(>!phc1logical :lnaly:->is ol"
The origin 1JI' the gt.:'11eric n:1rne is va1 iciusly intc rr 1·cted; tile most likdy l)eing I 1\ i1n Heienclt.:!en l'I al. (2003;_1), 1Jicwy11ie1 ~11iw1u11sis Amshoff nests within ~I <.:lade ur
dia~po, c/({i~vd11 (Cjk: thlClugh, a11d to loo.sen 01 sd fiee), 1eft:lling tn the Hxlu(. 1•tl Diflli11m sp t::<..:ies, sistc:1 to !Jiolilfm di111.!lt(~ei I-I:11111s fm111 \'\/ Ahica; K.rn:ppen
numlx~ 1· of pct;tls tc. <>U'!:1 of species me apetalous and c 20% h:1vc on ly l p<.:1:1D (1967) COllllllellt.'i that lhL' floWL'JS or !Ji<.:01y11ia ~ind the only A1nc1ican spt.:cics o!
The l.:1tin fo1m of the (;reek pl.int name 'dialion', used hy Linnaeus frn £he gcnn<;, JJialium , /) g11it111c11sc (Aubl.) Sc111dw., ;,uL' 11101phologie<dly simila1 in that hoth
1n:1y :1llc1rn1tivdy explain lhe de1ivali(Hl h ave only lwo stamens on ct hro;1d , tl<it receptadc. Borh also ha ve wood
Tte'-·s; ttopi<.•al 1;1in frn est ( mainly lowbnd eve1g1eenl 1 O(.t:a .o,;i o 11~ 11ly in pe;ll, i..:onwining <1m01 phou.s si lic 1 bodies; the /Jia/i11111 spec ies diffc1 s, however. in
swamp 01 heath t<>a:st, 01 coa~tal ; a few species in m1mso1 1n f(H\~Sl :ind wooded being apet~tlous
g1;tssl;1nd (S;tV;tlll1:t), hut in d1•ier v eget~ltion types the spe<.:ie... f1equently OC<. t11 lltl The c:xccllcnt, dui;.tbk timhe1 of /Jiccnynia ~uiu1tu11sis (angdique, hasg1\ocus) has
1ivl'I margins (>J' dl!<..!p S(>ils been widely used t(J1 f'urnittne and ( abim:t nm king, :mcl in he:1vy con,'it1uction ,
Refctenu:(s): Stcy;ic1t (I 951 ); Rojo ( 1982, ~tml in I Jou el al ( l 99fr (108- ()1(1); Du genc1al c11pcnl1~ 1 , 1;1ilto~1d tiL's and coopc:1;1gL', and espL'cLdly in in~11inc
Puy &. Hahl.'voh iLta in Du Puy l'I al (2002: 60 - (1')) nms11uctio11 due lo its high silic1 contc:nt \vhich 1t.:sults in a high 11~1tlll'al l'l'.',islanu.:
Irwin & Bamehy ( 1981) descril)(xl a new subt1•iln~ Di:diinae lo accommodate U to wood boie1s
genera (indudinµ f)ia/i11111) of the Cassieae and 1ecognised 'i suhgcne1c1 in
1Jiolift111; Rojo ( 1982) only n..::c•ognised .~ suhgenl.'1:1, 1educi11g one: Lo a section ol
~tnolhcr, and 1ei nsrating Uillie11ifl as a sepa1.11e genus, rn lite lllolecula1 ;rnalysis o!
K:1jit;1 I!/ ol. (2001), /Jiali11111 is si.o,;le1 Lo a clade \vhicl1 cont:1ins Ap11luit1, JJett1los1ylis
and Xl'!liU; in lhc: cnmhincd <lllctly.sis or l-k1ernl een t:! fll ('2003;1), Xt'lliel is sisfL'I [O
~1 cl:1de which includes seve1~t1 spc.x:ies nf JJirllilfm llL'Sted :1111011gst which is om:
species of JHc:01y11ia
' Lhe edible f1uils :11 e used mainl y in c hutneys, 1hc timber of va liou:-; species
( k1.:1 ;111ji, jutahy) i.s used for bo:lt building, house c•onst1t1ction, llorning, tnob ,
fitc\vood :tnd c hm <..:o al (:-.nme :-.pccics have a high silic t content and 1csi'"ir hrnl'I
altaek) ·1 he lx1rk and li::1ve.s :uc wid ely u.o,;ed in folk 1ncdicine in \V Africa k .g ,
!'or li...:vcrs and lot>1h:1chc); the l>atk o1 j) <.ochi11(.hi11e1rsu l'it.:1rc is~' suh..-tilutc fn1
hetcl nut in lnclochina; the h:11k of JJ jJach.JjJhyllum Il:t1ms lrns a poisonous 1ctl
gunHe.Sin llSt:cl flll ' <Ii'\ OW -poison in C:ongo (Kinshas;t)

Dia/ium madagascariense (A-J); D. unifo/iolatum (K-M) Drawing by M. Tebbs Dicorynia guianensis Photograph by 5. A. Mori

TRIBE CASSIEAE 121

120 LEGUMES OF THE WORLD
 Chamaecrista desvauxli Photograph by G. P. Lewis
Chamaecrista Moench 1794
Grlmaldia Schrank (1805); Cassia subgen. Lasiorbegma Vogel ex 13enth. 0870):
Gc1ssia subgcn. Absus (DC. ex Collad .) Symon ( 1966)
c. 330 spp, mainly tropkal. thinly extending into the subt1opics, a fow in cool
tem pe1ate N and S Amelica and E Asia. 266 species in the New We)[ kl with
g1e;,1test diversity in S Ame1ica (concentrated in 1he C B1a zilian planallO (with c 7 5
spp. centred in Goic1s and Mato G1osso], and in E Brazil [c, 72 ~pp . endemk in
Minas Ge rais, 26 ende mic in Bo1hia, a furthe1 12 spp. restricted to E and NE B1azil.
15 spp. in SE BrdzilJ, c 7 spp. Amazonian, 5 in !he Guianas, 1 sp. encle111ic in
Venezuela, 1 in Bolivia, 13 spp. 1estricted to th e Antilles and SE USA [of rhose lJ
e ncl~mic in Cuba], 6 spp . limired to Mexico and S USA, 5 wiclesp1 ead in S Amc1 ica
and 13 widespread on borh sides of the Pan:1nrn isthmus); a seconda1y ce ntre in
Africa wirh 36 spp (nminly widesp1ec1d in Z;t111bezian, Such.1nian , S<nnHlia-M<1sai.
Afromontane and Zanzibar-lnhambane regions), 3 - 4 extending to Mad<1g<1sc 11 ; <1
spp ~nclcmic in Mad<1g;1sc;11 and 1 in Ald ab111; 12 native in AusrrL1Jia, 5 spp
endemic in Indi:.1, 3(-•j) spp. in continental SE Asi<t,Java :ind New Guinea , l sp
endemic in the Philippines, the genus extending into Korea and Japan; :-;cveral
·"PP· widely introduced as pastu1e he1bs, 01 inadvenently ::1s weeds
Frain cbmnae (Gk.: low-g1 owing, dwa1 f) and c:rista (L: a c1est), a combination ()f
g1owth fo1m ~md the 1'escmblance of the stamens ~or pc1'11ars the finely pinnate
leaves) to a c1 est
He1bs (perennial or monocu•pic), sluub.s 01 (occasionally) trees: man y are weLds
of o pen siles and colonise1s o f disturbed areas, 1o ac1sic.les, Jivet nrn1gins, cu ltivated
area.sand g1assland; many on sand, (including coastal dunes), some in semi-a1id
~11ec.1s, some in seasonally watel'loggecl areas, in S A111eric:1 common in wooded
grnssl:md (cerraclo, sav;rnna) ;.rnd on stony or sa ndy grassland (campn), vt!1y few
in tllle rain forest
Hefetence(s): Irwin & llogeis (1967); Irwin & 13atneby (1977; 1978; 1982:
636 - 892); Lock 0988. 1989: 30 -33); 13at reto & Ya kovlev 0990); Singh 0992);
La!Sen & Ho u in Hou et al (1996, 565-570); Pedley 0998); Baineby (1999);
Concci~:10 et al. (2001); Du Puy in Du Puy et"'· (2002: 94-103)
One of three gene1a in subr1 ibe Cassiim1e; divided into 6 section.s in th e New
Wo rld (!twin & Bat neby, 1981; 1982); see und e t• Cassia Fot comments o n
phylogenetic placement: Co1by 0974) and Sprent (2001) noted that while species
of Cassia and Senna do not noc.lulate, those of l'lxmwecrista do; 24 spp.
described ::1s new to the Neotropics .since 1982
Sonie species widely used ;ts traditional m edicines in Africa, cspedally as purg<-Hivc.-.
and for trcaring wmmds and sores; in Senl'gal a paste of Cb nbsus lL) Irwin &
Darneby, mixecl with butler, is used as a supposito1y fo1' piles, also used frn syphilh;
th e seeds conwin :.1 toxalbumin, ~1bsin, simila1 to ;i()l'in f1orn Abrusprec.:olori11s L
(t1 ibe Abreae) and this is tboughl to be the active .suhst;.mce used fo1' eye cnnclitions
such as cata1act; G1J mimosoides (LJ G1eene is llsed <1s a tea suhslitute in China <ind
ja p~1 n, <inc.I is u.'icd against snake bite and .-;crnpion sting in Tam:ani ~ 1
Chamaecrlsta nigricans Photograph bys. Collenette
122 LEGUMESOFTHEWORLD
Senna austrolls Photograph by G. P. Lewis Senna form taxon 'flllfalla' Photograph by R.B.G., Kew
Senna Mill. 1754
Cassia subgc n Se1111a (Mill ) 13enth (1871); for a long list of generic syno nyms
sec !twin & !larneby (1982)
c. 295-300 spp., circumtmpical, mo.st nume1ous in rhe Amclie<1s, well
rcp1escnted in Aftic:1 and Australia, pornly .so in Asic1 and Oceania, some
extending into w~irrn and (few) dese1t or cold tcmpe1;,lle N ;.ind S Ameiic,1; 206
spp.. native to the New Woild (c , 45 spp. restricted to C and N Ame1ica, 10 spp. in
the Greater Antilles [including 6 endemic in Cuba], c , 30 res tricted to N\V South
Americ;.1 (Venezuela, Colombia, Ecuador, Pem], 3 re.'it1 ictecl to the Guianas Hnd
Venezuela, c. 28 in southem S A111e1ica [Argentina, Urugut.1y, Pa1::1guay, induding 8
spp endemic in Chile), 5 spp . endemic in Bolivia, c. 20 spp. 1estlicted to SE B1azil
and 13 to NE 13rnzil, c 25 srr. widespread across s America, and C, 25 - 30 on
eith er side of the Panama isth1m.1 s [eit h er~ disjunctly or widesp1eadl ), c 18 - 20
spp. n~Hive to Af1ica (many in Somalia-M;.1sai E ;md NE Ahica (including 2
extending to Socolla; I sp. endemic on Socoti·a and 3 exlending to the Middle
East and A.sit.1), othe1s mrne wiclesp1e;;1d in dryl<Jnd phytod1oria) , 9 spp. endemic
in Madagascar, 3 spp . indigenolls in Malesia, 3 spp . 1e:-;t1icted to lndia, l sp in
Myammu [Burma] and Thailand (and possibly al.so narive in Cambodi~1, Laos and
Vietnam); 33 spp. endemic in Austi alia plus 16 endemic .so-c1lled 'form tax<i '; c
3-5 spp . widely ctil tivated and of unknown ce1t;:1i11 oiigin
Delived fro m the Arabic smw 01 samw, for s p ecies w hich h~1 vc leaves ;111d pods
with cathartic and ht xative p1ope1ties
Trees, sh1ubs 0t he1bs (some monocatpic); rnid scrubhrnd, 1ocky hillsides •tnd
deep dese11 sands, deciduous woodland, thrn n woodland (in cluding ca;:1tinga),
wooclecl grassland (.savanna and ceriado), few in Anrn zo nian frnest; some
endemics on limestone (especia lly in Madagascar), coas tal t1opical and .subtro[1ical
foresl ; many in disturbed sites
Refetence(s): !twin & 13arncby (1982: 64 -635); Zocl\ne1 & San Mattin (1986);
Lock ('1988, 1989: 36-40); Singh (1992): Larsen & Hou in Hou el al, (1996,
673-691); Randell & Ilatlow Cl998b); Du Puy in Du Puy et al. (2002, 79-94)
One of three genern in subtdbc Cnssiinae; see unde1 (,'assia fo r comments on
phylogenetic placement. In iilid a1eas in Australia taxonomic distinctions ;:ue
blunecl by polyploidy, hyb1iclisa tion and apomixis (Randell & Barlow, 1998b)
Senna a/C!xcmdrina Mill. (= Cassia senna L,), 01 trn e s<:!n na is well-known as a
laxative; 1rnmy species a1e multi-ptupose sh1ub.'i with numerous mec.licirnd uses,
especially ~Ls purgative!i; S (I/ala ( L) lloxb. is univ~rsally used against p;u asitic
skin disc~lscs; rimher of some species is used in consuuctio n and frn charcoal;
occ1sionally seeds a1e ioasled nnd g1ound ;u; a coffee substitute, less often l 1sed
as fish poi.sons and for tanning 1e i11h c 1~ many arc 0t rnunentals and shade trees
widely int1 oduced throughout the tropics; for a detc1iled account of uses in \V
Arrica sec Burkill 0995)
Senna plstaclifa/la var. picto Photograph by G. P. Lewis
TRIBE CASSIEAE 123

Cossio /ovonlco Photograph by H.
124 LEGUMES OF THE WORLD
Cossio moschoto Photograph by G. P. Lewis
Cassia L. 1753
c 30 spp; circumtropical; 12 - 13 spp . nat ive in the Amelicas (mostly of the Am:1 zon
Basin bur l sp. endemic to Mexico, 2 in SE I31 azil , I in NE Bmzil and 2 w idesp1cad
o n eilher side of the P:rnanrn isth mus, indt1ding in the Ant illes), 10 spp. native in
Af1irn S of the Sa hara (2 spp. in Gu inea-Congolian, 2 spp. in Suda nian, 2 spp in
Zanzibar-Inhambane and 4 spp. more widespread in Z<rn1 bezian, Somalia-Masai and
Z:rnzibar-lnhamb•rne reg ions, one extending into Madagascar); 1 sp endemic in
Mad<.1gasc;11•; 1 sr. endemic to Myanmar [I3u1 ma) and Tha iland; 3 spp. native to
Indi a, S1i L:mka and SE Asia a1e widely cultivated th1ougho ut the tropics and
subtiopics; 2 spp. endemic in Australia (Queensland and New South Wales)
Derived f1om the ancient Greek n<1me casia fo1 an aromatic and fi.1grant plant; the
type species, C.fis/11/a L (purging cassia , Indi;.m lc1buinu111 OI golden showei), has
~1 long his101y of he1bal use, the pith of its lol"'lg, woody pods yielding a rurgativt:
medici ne; not to be co nfused with the true 'ca.s.sb' of co mme1ce wh ich is
Cimwmomum aromalic11m Nees (L:rn 1aceal!), although Linn~1eus used 'cassi a' in
the gene1 ic sense to inclt1cle various plants wi th similar medicinal uses
Trees and arbort:scent shrnhsi tropical in 1.1 w ide range of habitats including 1~1in
fo 1e.')I (Ol"'I re1ra fi1m e), 1ive1ine and g;.iJJe1y frne.st (includ ing nooded tive1i>anks
'ind v;.lr7.e:1), se~1 so nally d ry forest, woodland, wooded g1asslancl (s<1v;.rnm1 and
cem1clo), d1y sc1t1b, th ickets ~in d co;..1st::1 I eve1green bushlcrnd
Heferencel,): !1wi11 & llaine by 0982: 4-63); Lock (1988, 1989: 27-29); La1sen &
Hou in Hou el al 0996: 556 -565); Rande ll & Barlow 0998<1); Du Puy in Du l'uy
et al, (2002: 74 - 78)
Prio1 to the seminal w01ks of Ilwin & Barneby (1981; 1982) most Floras ti eared
Cassia in its bwadest .sense to include Se1111a and Cbamaecrisla ; these three
gene1:-1 a1c now l:.irgely accepted and togethe1 make up subtribe Cassiinae (Irwin
& l3arnehy 1981; 1982) Cassia differs from the othe1 two genera pl'incipally in
stamen organ isation 131uneau et al, (2001 ) found t h~lt th e th1 ce gene1;.1 did not
fo 1m a c.:lade, Senna ,wouping w ith Cassia, while CJwmaecrista wa.s isolated In
the combined molectliar-11101phological an alysis o f Herendeen et al (2003;;1),
Cbamaec ,.isla 1e.'iolvecl as sistet to a claclc of Senna species wh ile Cassia gra11dis
L. w:is b<Asally b 1anching to a l<irge Cae.')<ilp in ieae-Mimosoideae clacle in a posiLion
isolated f10111 the Semw -Cbamaecrisla cl ade (i e., ag1edng w ith the findings nl
Doyle et al., 1997 and Kajita el al. , 2001). More 1ccently in He1enc\een el al_
(2003b) rh e three gene1;.1 frn1 11 a clacl e, w ith Cbamaecrista si.')ter to the orhe1 LWO
(mrne in line with m01pho logy ~in cl noclulat ion data)
Pou r sp ecies o f Cassia are widely cultive1ted as 0111<.1men1als throughout th e tio p icS
and sl1bt1 opic'i: C j/slula, C. ~rmulis, C.jaum1.ica L and C. 1·u:dmrgbii DC.; nwny
species have numel'Ous medicinal u.ses (e g., C. sleberimw DC, m; laxatives and
purgatives) ; C', arereh Dclilc is used as a fish stupifi e r; in Malaya C fistula was
once u.sed as a fraudul en t substitute for opium, C r oxbw]Sbii and C. sie/Jeria11a
ha ve w oods used w idely fo1 turne1y, ca1pent1y , cahinet1y, tool handles, pcstk.'i
and mrnt<ll S 80 udouini fl ugge1.,.1orm1s.
o. c. de Wit o Photograph by D. Du Puy
TRI BE CASSI EAE 125
TRIBE
caesalpinieae byG.P.Lewis
Tribe caesalpinic::1c Hchb. 1832
Tribe Ceratonicae Rchb. (1832)
Tribe Dimorphandreae Benth. (1840)
Tribe Sclerolobieae Benth. (1865)
Tribe Moreae Britton & Rose (1930)
Polhill & Vidal 0981) divided the Caesalpinieae into 8
informal generic groups: the Gleditsia group (2
genera), the Acrocarpus group (munog neric), the
Sclerolobium group (3 genera), the P Jr ph rum group
(13 genera), the Caesalpinia group 06 genera), the
Poeppigia and Pterogyne groups (both monogeneric)
and the Dimorphandra group (10 genera) . They
commented that the tribe is "a remarkable mixture of
relics and complexes of relatively recent speciation,
providing many pitfalls for formal systematics and
biogeographical interpretations" . Polhill 0994) added
a ninth informal group, the monogeneric
Orphanodendron group, and placed Cordeauxia as a
synonym of Stuhlmannia (both genera recognised in
the present treatment) so that the total of 47 genera in
the tribe remained unchanged . Within the tribe ,
Parkinsonia (including Cercidium) , Conzattia and
Lemuropisum were moved from the Caesalpinia group
to the Peltophorum group (Polhill, 1994) in agreement
with the subsequently published works of Lewis &
Schrire 0995) and Du Puy et al. Cl995b).
Since 1994 several studies have cast new light on
intergeneric relationships within the Caesalpinieae,
necessitating the restructuring of some of the nine
informal generic groups presented by Polhill 0994). As
pointed out in the introduction to tribe Cassieae, the
genus Ceratonia has been removed from that tribe to
the Caesalpinieae, and Poeppigia has been removed
from the Caesalpinieae to the Cassieae (for further
detail see discussion under each genus). In the rbcL
phylogeny of Doyle et al. (1997) the Caesalpinieae, as
traditionally circumscribed, was shown to be
paraphyletic with members scattered throughout a
clade which also included genera of the Cassieae and
one mimosoid genus. The molecular analysis of Kajita
et al. (2001) also found the Caesalpinieae to be non-
monophyletic. In the molecular analysis of Bruneau
et al. 2001 om or tb in ~ rmal g ~n ric groups of
Polh11l CJC 4) w r suppo rted as 111 mo phyletic I ul the
tr ib~ as a who le wa clearl y d t:! mon. tral d to be
Par:1 phy lcti . With regard to lnterg n ri relation.ship!>,
PteroJ{YIU! rcsolv ~d as ·istcr tu a aesa lpi.ni·1 group
clad '; /Jatesia and 011.acctpoua fe ll outside a core-
LEFT Caesa IPInta
. madagascarlensls Photogroph by G. P. Lewis
Peltophorum group, and the Dimorphandra group was
clearly shown to be a diverse assemblage of genera,
many of which share certain characteristics with the
Mimosoideae, specifically with members of tribe
Mimoseae (Bruneau et al., 2001). Erythrophleum was
sister to a clade that comprised the majority of the
Mimosoideae sampled, and Pachyelasma was sister to
the two mimosoid genera Pentaclethra and Catpocalyx.
Herendeen et al. (2003a) in a combined molecular-
morphological analysis which expanded on the study
of Bruneau et al. (2001), presented an 'Umtiza clade'
containing Gymnocladus and Gleditsia (the two
members of Polhill and Vidal's Gleditsia Group),
Umtiza (traditionally included in tribe Detarieae),
Tetrapterocarpon (from the Dimorphandra Group),
Acrocarpus (the sole genus of the Acrocarpus Group),
and Ceratonia (from subtribe Ceratoniinae in tribe
Cassi~ae) . This new generic grouping raises some
fascinating phytogeographical questions (see Schrire
et al., pages 21-54, this volume). Pterogyne resolved as
sister to a Chamaecrista-Senna clade (of tribe Cassieae)
a relationship worthy of further study; Batesia and
Vouacapoua again fell outside the core-Peltophorum
group; Dimorphandra grouped with Mora as sister to
all Mimosoideae, and Pachyelasma grouped with
Erythrophleum as sister to the Dimorphandra-
Mimosoideae clade. In the phylogenetic investigation
of Haston et al. (2003), the Peltophorum group of
Polhill (1994) was non-monophyletic but there was
support for a core-Peltophorum group comprising
Peltophorum, Parkinsonia, Schizolobium, Conzattia,
Delonix, Lemuropisum, Colvillea and Bussea. Pterogyne
resolved as sister to a clade containing Haematoxylum
and Cordeauxia (both of the Caesalpinia group), thus
supporting the earlier findings of Bruneau et al. (2001)
rather than those of Herendeen et al. (2003a). Haston
et al. (submitted) have further refined the relationships
of the non core-Peltophorum group genera. They place
Arapatiella and jacqueshuberia with Tachigali in a
newly defined Tachigali group and find strong
molecular support for associating Batesia with
Recordoxylon and Melanoxylon in a new Batesia group.
Moldenhawera is placed in its own monogeneric
TRIBE CAESALPINIEAE 127
group sister to a Tachigali group-core-Peltophorum
group-Dimorphandra group-Mimosoideae clade. In an
analysis testing the monophyly of the Umtiza clade
(Herendeen et al., 2003b), Arcoa (traditionally of the
Dirnorphanclra group) from the Dominican Republic
The Caesalpinieae as presented here contains 56 Cassieae: Diali inae (see page 111)
genera and 2 -436- 448) s pecie Fig. 2 . Thirty
two of the g nera contain 3 or fewe r species each
.
---
with 23 mo no p e cific (a ond ·p c ies of Papilionoideae
Orphanodendron has apparently been discovered in

was added to the group of genera in that clade.
Without do ubt , the genus w ith the greatest
taxonomic and nomenclatu ral complexity within the
Caesalpinieae is the type genus Caesalpiizia , which in
its broadest se nse comprises c. 140 spp. and contains
25 generic names in synonymy. Of these 140 species,
12 -15 predominantly Asian taxa have still to be
inc lu ded in molecular studies and cannot yet be
assigned to any generic segregate recognised in this
treatment (see notes under Caesalpinia L.). Studies by
Lewis & Schrire 0995), Simpson & Miao (1997), Lewis
(1998), Simpson (1998, 1999), Simpson & Lewis (2003)
and Simpson et al. (2003), have clearly demonstrated
that Caesalpinia, as traditionally circumscribed, is
polyphyletic . In this treatment Hof(mannseggia is
recognised as distinct following Simpson & Miao
(1997) , Simpson 0999) and Uli barri (1979, 1996);
Pomaria is also segregated from Caesalpinia sens. lat.
---
Colombia, but is, as yet, und scribed [Cogollo Pach co Gymnocladus
pers. comm., 2002]) . A n w g nus, tentatively n ru ci Gleditsia
as Heterqfl.orwn by Sousa & Delgado 0993), but not Umtiza
yet formally published, is a monospecific Mexican Tetrapterocarpon Umtiza clade
endemic closely related to Conzattia. It is not dealt Arco a
with here . The informal ge neric groups of tribe =-- Acrocarpus
Ceratonia
Caesalpinieae presented by Polhill & Vidal 0981) and
Polhill (1994) are retained in part in Fig. 23 which
accompanies this treatment, but there are some Cassi eae: Cassiinae
(seep age 112)
noteworthy exceptions. The Gleditsia and Acrocarpus
groups are both subsumed into the 'Umtiza clacle';
Dip~ychandra is rejected from the Sclerolobium group Pterogyne Pterogyne group
which now becomes the Tachiga li group and includes
Arapatiella andjacqueshuberia; Poeppigia is moved to
Haematoxylum; Cordea uxia; Stuhlmannia;
the Cassieae ; several genera are removed from the Mezoneuron; Pterolobi um; Tara; Coulteria;
Peltophorum group leaving a core of nine related """" Caesalpinia; Pomaria; Erythrostemon;
gene ra (if Heterqfl.ontm is included); Batesia, together
with Recordoxylon and Melanoxylon constitutes a new L Poincianella; Cenostigm a; Guilandina

following Simpson (1998) and Simpson & Lewis (2003).
The genera Cou.lteria, Etythrostemon, Guilandina,
Lihidihia, Mezoneuron , Poincianella and Tara are also
reinstated following the findings of Lewis & Schrire
0995), Lewis 0998), Simpson et al. (2003), Lewis &
Bruneau (unpublished) , Lewis & Lavin (unpublished)
and Sotoyo (unpublished). Caesalpinia sens. strict. is,
in consequence, reclucecl to a genus of 25 species.
l~
Caesalpinia group
Batesia group based on the work of Haston el al.
Libidibia; Stahlia; Hoffmannseggia;
(submitted). Moldenbawera is placed in its own group
Stenodrepanum; Zucca gnia;
as its generic relationships are currently unclear Lophocarpinia; Balsam ocarpon; Moullava
(Haston et al., submitted) . The Caesalpinia group
increases in size from 12 to 21 genera. Five genera :ire
currently too poorly known for them to be placed wi th Batesia
confidence: Campsiandra, Chidlowia , Dipzychandra, Recordoxylon Batesia group
Orphanodendron and Vouacapoua. Melanoxylon

Moldenhawera Moldenhawera group

Tachigali
Arapatiella Tachigali group
Jacqueshuberia

Sc hi zolobium; Bussea;
Pelt ophorum;
Parki nsonia; Conzattia; core-Peltophorum group
Delo nix; Colvillea;
Lem uropisum

UNPLACED TAXA
- Pach yelasma
Eryth rophleum
Dim orphandra
Mora Dimorphandra group
Campsiandra Bur kea
Chidlowia Sta chyothyrsus;
Diptychandra 'Sym petalandra
Orphanodendron
Vouacapoua

7_ Mimosoideae
- doubtful placement in generic group

~~: :3 Diag~am of relationships between genera and Informal groups of Caesalpinieae based on the analyses of Polhill & Vidal (1981); Polhill (1994);
1
et at (& Schnre (1995); Kajlta et al. (2001); Bruneau et al. (2001); Herendeen et al. (2003a & b); Simpson & Lewis (2003); Simpson et al. (2003); Haston
• 2 003 & submitted)

TRIBE CAESALPINIEAE 129

128 LEGUMESOFTHEWORLD
Gymnocladus dialca Drawing by K. S. Velmure
Gledltsla caspica Photograph by R.B.G., Kew
130 LEGUMES OF THE WORLD
Gleditsla japonica Photograph by G. P. Lewis
Gymnocladus Lim 17H5
6 srp; I in E orth Ame1ica, Sin E and SE Asia (l in Ind ia [A.c;sa m], 1 in My.1nma1
[Bl1tm<il, 2 in Chin a, 1 in N Vi~rnam)
From R.JJ11"/Cl- (Gk : naked) and c/ados (Gk : b1;ilnch), the b 1:1nches lea fless !'0 1
most o f the year
T 1ees; tempe1<1t~ 0 1 mo ntane t1opical low woodhrnd ;rnc.1 wooded hillsides, some
by 1 ivcrs
Refe1ence(s), Lee (1976); Robertson & Lee (l976, 21-26); Huang & Yao (1<)80);
birscn el al ( 1980' 73 - 74)
An essentially dioccious genus; placed in the Gleditsia g1oup together with the genus
Gledllsia hy Polhill & Vidal (19Hl) and Polhill 0994), a 1elationship confi1 med hy the
molecu la1' analyses of Kajita el al, (2001), B1une<.1u el al (2001) and the com bined
molecular-mm phologi cal analysis of I-Lca.:nclcen el al. (2003a); c. lO extinct spp of
Gym11udad11s (most ly fossil leaves ;rncl fruit ) have been de.sc1ilX::d f1om C and \V
U.SA., Mexico, Eu1opc and Chi na. qymmx:ladus cl ifft:: r."i fi om G'ledilsia in its . . :11J(Hlin
cheinisl1y cmd mncle of frllit dd1iscence (Gymnuclrulus o pens along the placcnr;d
slm1re), tht:: l;u..:k of spines on the ln.1nk and lm.mches, and by i[S e1ec:t mccmuse
intlore.i.;ccnccs with larger !lowers Gynmodadus dioica (I.) K.Koch (Kentucky uiffee
free [as masted seeds we1e once used as a coffee substi tute]) pl;rnted as an
orm1inc11tnl in pa1ks and gardens; also used frn wood (employed kx:ally in a v.11iL:ly
of mino r uses), in soi l reclamation, for medicine, inseclicicks, oils (t::. g , from llll'
~eels of (J. cbim:11sis Bail\., used as a lc1tex in making p<.1int), soap (G. cbi11e11sis and
G. as.samicus Kanjilal ex P.C.K:rnjilal, used in China and Incli<1 respt..'lth•cly,
. arc iich in
saponins); \cavt::.<> ;tncl pod pulp contain the alkaloid cytisin t:: fatal 10 ,-.t me livesttH..:k,
otherwise ."iec:cls cincl pod pulp edible after cooking
Gleditsia L. 1753
c 13-16 spp .; 2-3 in E Nrnth Ame1ka (of which 1 also rt::c.:rnded from
Tamaulipas in Mexico), 1 in S South America (N Argcnlina and adj<1cenl Urugu:i y,
Paniguay and Brazi l), l (or subsp.) ;1rou11d the Caspia n Se:1, l fro m E Ind ia, 1 in
Malcsia (Philippines and Sul<iwesi [C:dcbes]), 7-9 in Asia Cfirnn E, C ancl S\XI
China to Ko1('<1 1 .Japan, Vie ln:rn1 , !.~10s and Thailand)
Named frn jolrnn n Goulieb Gled itsch (17 l 4-17H6), fticncl of Li nnaeus, Gc1m:1n
bornni st and Oil t::CtOI of lhc Bol<ln ische1 Ga1len, nerlin
Trees (anti 1 sp ii shnib); tempe1ate ~rncl subll o pica l d1y woodland and thickd on
sandy and rocky ."ilopes, lowland wet forest and sw;;nnp forest
lkfercncc(s), Gordon (1966); Robertson & Lee (1976, 26-32); bii"sen el al (l'JHO'
68-72); lli nggui (1982); 7.h i-Cheng 0987); Llli ones (19HH); L1rsen (19H'J); I-Jou in
Hou el al. ( 1996, 6J 9 - li22)
A dioeLious gt:: m1 s; placed in the Glcditsia g1oup toge th er with the genus
(~y11111odml11s hy Po lhill & Vidal 0981) and Po lhill 0994), a 1clationship
--
Umt/za r
(cont inued o n next p;.1gc)
Umtiza listerana Drawing by P. Halliday
tsterana Photograph by P. Van Wyk
Umtlza llsterana Photograph by A. E. Van Wyk
Gleditsia (conti nued)
confirmed by the molecul<11 amilyse.s of K<1jila el al, (2001) , 1311111cau el al (2001)
and the combi ned molect1\a r- mo1ph o logica l an~ilys is of Herendeen el al. (2003a),
(see notes unde1' Gynmocladus); c 40 extinct G'leditsia 01 Gledilsia-like spp have
been 1eprntecl i11 the fossil li te1atu1e ex tend ing the histrn ic~d 1<1nge of the genus
in to Eurnpe e1nd \Y/ Nonh Amerirn , The molccu la1-biogeog1<1 phical studies of
Schnabel & Wendel 0998) and Schnabel et al (2003) concl uded that there is on ly
a singlt! Asian-N Amc1'ican disjuncti on within the genus, no Asian-Ame1ican
.species pairs, and ;;in unusual A.sian-S Ame1ican d isjt1nc.:tion
Gledilsia triacm1tbos L. (honey locust) is wide ly cu ltiv<1tecl as an ornamental, also
used in agrofrnestry (excellent sou1ce o f animal fodder), frn soil 1edamation,
timbe1 (fence posrs, const1uction and rnilw~ 1 y tics, the Che1okee Indicins of
Te nnessee alleged ly used the wood lo make bows), ed ible fru it pulp (used in folk
bcc1s), rhorns (:.1s needles and fc>I ca1d ing wool , ;incl tied to sticks frn hunting
hl1 llf1ogs); soa p (fruits of some species 1kh in saponins anc.1 used as soap
substitutes in China, .J ~1 pan and Indochina), bee f<>l'<igc (honey) and medicine
Umtiza Sim 1907
1 .~p .; endem ic to a nar1ow cuea ::11 o und lhe Duff;llo River region (Albany Cen tre)
of the E Cape Province of S Af1ka
Derived from umthiza, th e African vernaClilat name frn the type species
U. listerrma Sim
T1ee rn shrub; subtropical chy forest, bush land and thicket (including valley
b ushvekl)
l!efe1encds), Palmc1 & Pitman (1973, 857- 859); J{oss (1977)
Ross (1977) 1emarked that the exact position of Umliza in the Cac:salpinioicleae
had not been este1blished , bu t a cu1 ious fe::iture o f the genus is that the calyx lobes
al'e open even when the flower.sate in young bud The flowe rs of Gleditsia ancl
G)mmoc/adus share th is condition , Cowan & Polhill (1981~1 ) place d Umliza in the:
Cynometrc1 group of tl'ibe Detadeae bur co mmen ted that its finely 1etict1\ate pollen
is mrne characte1isric of t1ibe Caesalpine:it: than o f Cy11omelra Morph ology
(Leona1~1, 1957, 279-280), pa lynology (Fci guson & flank s, 1995) and molecula 1
evidence (D 1lmcal1 el al., 2001) shed fu1th e1 <loubt on the pl::1cement of Umtizn in
the Deta1 ieae. In th e comb ined mo1phological and molecula 1 analysis of
I·Ie1endeen et al (2003 a), Umtiza was nested in a daclc containing Acrocwfms,
Cerato11ia, '/'elrapterocarpon, Gledilsia ancl G)im11.oc/ad11s, the i1 so-ca lled 'Umtiza
dac.le'; in a paper fu1 ther testing the monophyly of the Umtiza clacle (He1endeen
el al., 2003b), the genus gtoup.s with C7ym11oc/ad11s :.ind G'ledilsia in a clade si.ste1
to a Tet1<.1ptcrocc11pon-A1-coa-Ac1uc;,11pus-Ce1atoni:.1 chicle Removal of the gen us
from the Dctaiieae sens lat is thus strongly suppo1ted
Used locally frn medicine ;Jncl has alleged magk::il p1ope1ties, witchdoctors have
used sticks of the tree as heali ng wands; the wood w:as once usecl to house
p1opdlor sha fts in small boats because il s o ili ness p 1oviclcd constant lub1irntion
TR IBE CAESALPINIEAE 131
Tetropterocorpon - various taxa Drawing by M. Tebbs
Arcoo gonovensls Drawing by
132 LEGUMES OF THE WORLD
Tetropterocorpon geoy/ Photograph by D. Du Puy
Tetrapterocarpon Humbert 1939
2 spp.; endemic to S, W, SC & N Madagascar
From tetra· (Gk: four), ptero- (Gk.: winged) and carpos (Gk : fruit), the Fruits
have 4 dry, papery wings in unequal pairs
Trees o r· shrubs; seasonally dry tropical to xerophytic Forest and thicket, o n
limestone, basalt or sand
ReFerence(s): Du Puy & Rabevohitra in Du Puy el al, (2002: 56- 59)
A dioecious genus placed in the Dirnrnphandra group of Caesalpinieae by Polhill
& Vidal (1981); sister to Ceralonia and Acroca1pus in the molecular analyses o[
Bruneau el a l. (2001) and Herendeen et al. (2003a); grouping with Arcoa in a
detailed study of the 'Urntiza clade' (Herendeen et al, 2003b), the two together
sister roan Ac1ocarpu s-Ceratonia clade
Tetrapterocarpongeayi Humbert is used locally for carpentJy, cart const1uction
and to make charcoa l
Arcoa urb 1923
1 sp .; endemic to Hispa niola (Sa nto Domingo, Haili and the Island of Gonirve)
Named for Count George von Arco (jl. c. 1903), who was highly esteemed in the
fi eld of wireless teleg raphy in Germany
Shru b or small tree; arid tropical vegetation on limestone hills and cliffs
Reference(s): Urban (1929: 34-37, t. 1 & 2)
Placecl in the Dimorphandra group oF tl'ibe Caesalpinieae by Polhill & Vidal
(1981 ) ; Ur ban (1929) considered Arcoa to have a close botanical affinity with
Gledilsia; included in the 'Umtiza clade' (Herendeen et al., 2003b), which includes
Gledilsia, as siste1• to Tetrapterocarpon from Madagascar
Ceraton/a sl//quo, 9 flowers Photograph by G. P. Lewis
c. Speight
Acrocorpus froxln/fo/lus Drawing by P. Halliday Cerotonlo siliquo, d' flowers Photograph by G. P. Lewis
Acrocarpus Wight & Arn 1838
1 sp .; India, Bangladesh, Nepal, Bhutan, Myanma r [13unna), Thailand, Lws, Java
and Su1natra
From aero- (Gk.: summ it or top) and ca1pos (Gk.: fntit), most probably alluding to
lhe long-stipitate ova1 ies a nd fruits
Trees; tropic~! and subtropical broetd-leaved rnin forest and evergreen gallery rrnest
Reference(s): Hou in Hou et al, 0996: 435-438)
Placed in its own Acrocal'pus gl'oup of tri be Caesalpinieae by Polhill & Vidal
(1981); sister to Ceratonia in 1ecent molecu lm st udies (Kajita et al ,, 2001; I3nme.1u
et al. 1 2001) and in combined molecular-mo1vhological studies (Herendeen el al,
2003a & b); a large range in flower size thrDLrghout its distl'ibution formerly !eel to
the recognition of two species
Timber of A ji'axi11ifoli11s Wight ex Arn. (pink cecbir tree) is used to make tea-
boxes, ful'nitu1e and plywood; widely grown as an ornamental (for its beautiful
red flowers), also used for foddel', gums and bee forage ( honey)
Ceratonia L 1753
2 spp.; 1 nmive in E Medite1ranean (Turkey, Cyprns, ls1ael, W Syria, Lebanon, S
Jordan), Arabia and N Africa (Tunisia and Libya), widely cultivated and naturalised
in the Mediterraneorn and elsewhe1e, l in Ambia (Oman) and Somali<I
From 'ce1atonia', 'ce1onia' 01· 'ceratea', Greek names for C. siliqua L., or possibly
from ceras (Gk.: horn), referring to the long curved pods of C. siliqua
Trees or shrubs; mecliterranean scrnblancl (city hillsides in gaiigue and coastal and
submaritime maquis); rocky limestone slopes and gullies (C. oreothauma Hille,
G.P.Lewis & Verde.), 0-2000 m
Refere nce(s): Meikle 0977: 589-591); Hillcoat et al. 0980); Thulin 0993: 356- 357)
Irwin & Bomeby 0981) placed Cerato11ia in its own monogeneric subtribe
Ceratoniinae. Polhill (pe1s. comm. in Hillcom el al., 1980) "'ggestecl a possible
relationship with Gledilsia and Gym11oc/ad11s of tribe Caesa lpinieae. In the
molecular analysis of Doyle el al (1997), Ceratonia did group with Gy111Hoclad11s
and in Kajita et al. (2001), it grouped with Acrocaipus, also of tribe Caesalpinieae.
In the combined morphological and molecula1 a nalysis of Herendeen et al,
(2003a), Ceratonla also grouped with Acrocc11p11s in a clacle with 7'etmpterocwpo11
from Madagascar and Umliza from southern Africa These four ge ne~ 1 were sister
co a Gleditsia-Gymnodadus cleide, bearing out Polbill's earlier prediction
Ceratonia siliqua (ca rob tree, locust bean or St. John's bre<.1d) is widely cultivated
in the Mediterranean foothills for forest-forage, fodde r and its nutritious pods;
carob pods unearthed in the store ho uses of Pompeii show that the Romans were
ha1vesting the species as early as 79AD, the seeds are said to be the original c:1rnt
used <JS a standard weight by jewellers; also used fo1 wood, as chocolate and
coffee substitutes, in Sicily turned into alcoho l, carob seed gum is used in foods,
cosmetics) medicines, photogiaphic film emu lsio ns, adhesives, p<iints, inks,
polishes and has numerous othe1 uses
TRIBE CAESALPINIEAE 133
Pterogyne nltens Photograph by G. P. Lewis Pterogyne nltens Photograph by G. P. Lewis Haematoxylum dinteri Drawing by P. Halliday Haemotoxylum brosiletto Photograph by G. P. Lewis

Pterogyne Tul. 1843 Haematoxylum L 1153

I sp. ; E Brazil, ParJguay, Bolivia, N Arge ntina Haematoxy /011 L. (1764), onhogra phic variant; Cymbosepa/11111 Baker 0895)
From ptero- (Gk.: winged) and gyno- (Gk.: woman, female), the fe1tilised 3 spp. ; 2 in C Ameiica (Sa lvador to Costa Rica), Mexico, S America (Colombia and
gynoecium develops into a s~mara Venezuela) and Caribbean (perha ps introduced), 1 in Af1 ica (Na mibia)
Tree; seasonally chy tropical to subt1opical woodland and thorn scrub (including From baemato- (Gk.: bloody) and xy/on (Gk ,: wood), alluding to the blood-reel
caatinga), semi-deciduous sub tro pical forest and liana forest hea1 twood or H. campecbicmum L, which produces a brilliant red dye
Reference(s): Lewis 0987: 42), Centurion (1993: 411) Trees and shrubs; seasonaHy d1y tropical semi-deciduous scrub and thorn sc1t1b,
Polhill & Vidal (198 1) and Polhill (1994) placed the gen11s in its own Pterogyne sandy liver beds and dry rocky hillsides, 1 sp. in swamps
group in tribe Caesalpinieae; in the molec ula1 analysis of Bruneau et Cll (2001) it Hefe1ence(s): Standley & Steyennark (1946); Hoss (1977: 112-114)
occuired in the same c\ade as Caesa/pinia and relatives (inclucliog Placed in the Caesalpinia group of the Caesalpinieae by Polhill & Vidal (1981) and
J-Jaemato:i..ylttm), a position confirmed by the molec\ar ana lysis of HHston el al. Polhill 0994); sister to Pterolobi11m in the mo lecula1 analysis of Bruneau et al
(2003) in which it grouped with Haemaloxylum and Cordeauxia; in the combined (2001), and in a clade with P1ero/obi11111 and Cc1esalpi11ia sens lat. On pa1t) in the
molecular-morphological analysis of Herendeen et al (2003a), however, combined molecular-morphological arn.1lysis of He1enclee n el al. (2003a); in a
l'lt11Ttgj'' JU clu 1 r.i. w ith (.'/,utmaecrl'</11 and . ~·w10
. in u C1ssiln:t • . _, Jade clade with Cordec111xia in the molecular study of Haston el at, (2003)
/'lemi:.111111 11111111•· Tul. (vlrnn , amenduh n, tlp.1 or 1.l p:i mlur~1lol yields an excellent The heartwood of H. campechianttm (campeche, logwood), is the source of a
thnlJc r for cnhi11c1work, fu r1111111·e, 1n1 ~rior OnL~hlug, t11 rncry. nooring, railway colourless chemical haem:Hoxylin which on oxidation turns to haematein, a
eras.sues and cooperage commerci<ll dark violet dye used for wool, silk, cotton, fur, leather, bone and
synthetic fibre dying, ::tncl with i1on chromium mordanrs co obt<tin red and black;
also Lisee\ as a stain in microscopical preparations (pa1ticuh1rly to show up cell
nuclei), an ink fo1 writing and painting and rhe lich red colotw has been used to
adulterate wine The species lrns minor medicinal uses; H. brasiletto H.Karst
(brazilette), p1ocluces brnsilin used as a bright reel dye; both New Wo rld species
used as rn namentals ;md living hedges (H. campecbituu11n cultivated in Af1ica and
Maclagc1scar); minor uses for furniture and carrenll)'i bee flowers yield a high
quality honey

Cordeauxia Hems! 1907

1 sp.; NE Africa (Somalia and Ethiopia)
Named for Captain H E.S. Co1deaux (1870-1943), one time H.M. Commissioner in
Somalia
Shrub or t1ee; seasonally d1y tropical (semi-dese1t) bushhind and thicket o n sand
l!eference(s): Hemsley 0907); Thulin 0983: 20 - 21; 1993: 348)
Placed in the Cl1esalp inia group of the Caesalpinieae by Polhill & Vidal (1981);
1ecluced to a synonym of St11/J/111a11nia in Polhill (1994) and Lewis & Schrire
0995), but reinstated as a separate genus in Lewis 0996a); weakly suppo1ted as
sister to Haematoxytum in rh e molecular analysis of Haston et al. (2003) 1 who did
not include Stublmcmuia (which has 1eddish, secretory, multi<.:ellular glands in its
leaves, very similar to those in Cordeauxia)
The seeds of C. edu/is Hems! Cyeheb nut) are t1.sed as huni:in food and have great
potential as an arid-land food; also used for livestock foclcler, a red dye (goats that
eat Cordeauxiu leaves have the dye depo.sired in their bones which become
CordeauXia· e dulls stained blight or.1nge), medicine, wood, an insecticide and soap substitute
Pterogyne nitens Drawing from Mart. Fl. Brosi/ Photograph by M. Thulin

134 LEGUMES OF THE WORLD TRIBE CAESALPINIEAE 135

Stuhlmannia maavi Drawing by E. M. Stones Mezoneuran hi/debrandtii Photograph by D. Du Puy Ptera/obium stellatum Drawing by L. M. Ripley Ptero/obium stel/atum Photograph by P. Van Wyk

Stuhlmantiia Taub. 1895 PterQ[Obium R J3 r ex Wight & A111 1 8:l~

J sp.; E Ar1il"il (co;1sti1l Keny<1 aml Tanzania) ;md N Madagasc1r Ca 11t11[(a J.FG rncl (1791); lleicba rdio Roth (1821)
N:-im ecl fo1 the Ge1 1rn111 ncttt11alist Franz Ludwig Sluhlmann (1863-1928) 11 spp .; 1 in S tropical Africa, E Af1ica and A111bi:1 , 10 in SE Asia (1 cncll'mic ro
T 1ee; se a.'iorlillly clry tropical for~st. woodland on lime."itone ancl riverine fo11.._'')l l ncli;1, 2 in Chin~1, ·i in lndo-China [l emk:mic to Thaila nd, 2 extending to MalesiaJ,
L~cferencds): B1 en:in (1967: 45 - 47); Lewis 0996a); Dll PL1 y & lbbevohit1c1 in Du 3 rcst 1i<.:led to the Malayan Peninsula an d A 1chipel:igo (l endem ic to the
Puy el fll (2002: '18, 50, Lmde1 Coesalpinia l11sulitt1) l'hilippinesl)
Pla ced in the C:aesalpinia g1oup of the Caesalpinieae by Polhill&. Vidal (1981) :tncl F10111 plero- (Gk: wing) ~mcl lobion (Gk.: pod, f1uit), in 1efe1ence to the .'i;.lllWJOic.1
Polhill (1994); the closely related genus Cordecll/.'\'iu induclccl in synonymy in f1uit
Polhill (1994) and Lewis & Sch1i1e (199')) hul not in Lew is Cl996a); Du Puy (,'\: Sh 1ubs {usually sc1c11nhling rn cl imbing) and lio:mas: seasonally d1y t1opical upland
Rtb~vohilT<l in l)u Puy el ol. (2002) recognised S moaui Taub, in Madagascar l'ver~ree n forest, rivel'ine and hurnicl fo1c.s1, \Voocllancl and woodl'd g1assbncl,
uncle1 one:: of its .synonyms: Caes(/ljJinia i11so!itC1 (Harms) Ikenan & ] n GillcLt 0 - 2500 m
The wood has pot~nt i al cat pcntry uses Hcfe1cnce(s): B1en<1n 0967: 40-42); Vi<l<1l & Hui Timi ( 1971, 1976a); L-lul Tho! &
Hideux ( l 977 !; Hou in Hou e/ al ( l 996: 61~ - (i(10)
Phicl'd in the Caesalpinia g1oup of the C::ie.salpinieae hy Polhill & Vidal ( 1981)
Mezoneuroti Dcsf. 1s1s ancl Po lhill ( 199·1); sister to llaemaloJ.:ylum in the mo lecula r study of Bruncctu !!/
Ml•zo11e11ro11 Desf. ;rncl Mezu11e11r11m DC. ( 1825) rn1hogr•1r hic va 1i:1nt.'i al (2001 ), and in a clade with H c1emcflo.\:Ylt1111 ;iml Caesc1 (/ ,iHia ( 'ieJts fol) as
CaesnljJillifl .'iubgen us Me.zo11e11rn11 (De.'i f,) Vida l ex H t!t~nc.I & Z: uu cchi U9'>tJ) sisre1 to Caesafj,i11ia subgcrn1s l14ezo1fem'V11 in the comb i n~d rn olccul a1·-
c. 26 spp.; mainly in Asi;1, extending to Aust1~.t1 ia , Poly nesia, and thin ly to moi phological ana lys is of IJeiencl ~t:n el al. (2003:1); in rhe molecular 1111al ysis o/
Madagascar and Afric;1; 2 in At'rica (1 wicle."irreacl in \VJ Africa, the o the1 from \Y/ w Sim pson el al (2003), P. slellotum l Fu 1t;sk) B1t::nan (as P. lacerans (Hoxb) \'qight
E and SE); l endemic in Madagascu, 4 endemic in New Calccloni;;1, 1 endemic in & A1 ll ) is si."i le1 to Caesalpi11ia decapetcdo (l~oth) Alston. an Old \Voild spec.:ies o l
L-Iaw:1ii, possibly 2 restricted to Chin~t, l to Vietnam, •i endemic to Australia unknown alfinity
(Qm~ensland and NS\'(I) , l endemic in the Philippin es, l in Austr:alia and Papu.1 The h.:avcs of JJ. slella/1(1JI are used in part.<> or Af1 iGt as ~1 lemhcr• dye , frn ink and
New G uinea , 9 mrn\.: wiclesp1·ead th1oughout Asia :-ts medicine; plants are giown as living !'cnccs
Piom meso- (Gk,: middle) or meizon (Gk.: greater) and neuron (Gk.: nerve). the
uppe1 sutu1e of the fruit is brncle1ed by a usually b10ad lengthwise wing so th:it
the suture appe:us as a subcentral p1-ominent ne1vc or vein
P1 ickly climber...;, scanclent sh1 ubs and small trees; tropirnl <ind subtropictl 1iverine
forest, lowland 1;1in fo1est 1 swamp fo1est, seasonally dry forest, thicket, vine fo1L'ST
and woodecl g1assl:111d, especi::tlly along foa~st <tn d live1 margins
Hefercncc{s): Brenan (1\)()7: 38-40); Hanink 0971); Vidal & Hui Timi (!9761>):
Verdcmni (1'179· IR - 20); l.cx:k 0989: 25); I ll!Nfl(ieen & Zarucch i (1990); Pedley
(1997); 1.c ~Ill" (199Ha: 59--07); Du l'uy & l!al"'vohitra in Du l'uy el of. (2002: ci8 - ·19)
Most dosely related m PlerulCJln'um :rncl Caesalpinio sens slril"I . :ind pe1haps hL'St
lrL'ated 11s a st1bgenus of Caesalpi11 ia (Vidal&. Hui Th o!, 1976h; Herendeen &
Zaiun:hi, 1990; Hcrendet:n & Dilcher, 199D. Other wrnke1.'i have rec.:ognisecl
Mezone111wf as a distinct genus (Brenan, 1967; Verdc.:m1n, 1979; Lock, 1Y89).
Afrzml<'lll'Oll is absent from the American tropics and subtropics today, but fo.-:"iil
rect11ds show lhat the taxon was widesp1e~1d aci1>ss Nrnth Ameiica clu1ing the
Te1'tia1y (l·le1't!mlee n & Dilcher, 1991) and C{U'.wlp1'.11ia sens strict has sever·.:tl
extant species in the Caribbean . Mezo11e1tron rm1.:ole11se \'\lclw. ex Oliv (as
.~I j a ljilnia anRole11sis (Oi l .) Herend. & Zarucchi) i.s si.'iter to Pterolobilllll
s.ttil/uwm (Porssk.) Brenan fr'\ the combined an:ilysi."i oJ' H erendeen el al. (200j:i)
Used I'm medicine :ind the woocl of /v/ l•mwie11se ( M;11111) Hilleb1 (uhiuhi) f1oin
Mezaneuron scortechinii Drawing by E. Catherine Hawaii 'vas o nce used loully frn spears and in house co nst1 uction Ptero/obium stel/atum Photograph by 5. Collenette

---- ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~-
136 LEGUMES OF THE WORLD TRIBE CAESALPINIEAE 137

Tara spinoso Photograph by G. P. Lewis
Tara Molina 1789
C'u11/te1ia Kunlh C1824), in lmgc pair (exduding C 1110//is Kunth): NicaraJ.:V Hutton
& Hose (1930); R11s~el/ode11d1tm B1itton & Rose ( 19~0)
3 spp; 1 1est1ktec.I to Mcxit:o, l in Mexit:o (Yucatan), Nicuagua, Cuba, I la iti
jamaic;1 ancl 1he Bahanws, l in S America (Venezuela, Colombia, fa.: uadrn, Pc1u ,
Bolivia ~ind Chile) <Ind culliv;Hed both within ;-rnd oulsick: (including M~ilt~t . lhe
Canal)' Islands, India, E and NE Afi ica) its native 1ange
Derived from the vcrrn1cular name 't;.11a' in Peru, Bolivia and Chik
T1ce.s and .shrubs; seasonally cl1y t1opical fo1est to se111i-t11'ic.I thorn scruh
Refel'encds): IlriltDJ1 & Rose 0930: 320); Sp1aguc 0931)
Two of tile 3 .c;pcx.: ies in the genus cu11·ently lac k combinations in Tara . Niu1rap,u.
Tara ~111c1 }(11ssellucle11dro1t wete all 1ccognisl'd as dcments of the infrn m;tl
Russellodc:ndron ~1m1p of Cl1esa/jJi11ic1 senJ lat. hy L<..:wis ( 1998). See taxonc )111ic
notl'S t111ck:1 Co11lteria which is the sistl'r' gl'ml.s to J'ara
Planted as living fcncdincs, the ba!'k of two !-ip(.x:il's willl 1rn..: dic.:in;il p1opcrlil .,,
thal or r SjJillOStl (Molina) B t ilton & Llose O<ll<l, spiny holdback) 1ich in l:mnms
and used in Pen1 frn· t:urning lecHher, dyeing and ink p1oduc1ion (vmious
unnme1cial p1oduc.:ls sold), also foi foc\voud and gum.-.
Taro (Caesalplnia) caca/aco (unpublished combination)
Photographby G. P. Lewis
138 LEGUMES OFTHEWORLD
Coulterio (Brasilettia) ve/utina (unpublished combination) Coulter/a (Brasilettia) velutino (unpublished combination)
Photograph by G. P. Lewis Drawing by E. Catherine
Coulteria Kunth 1824
13rasilellia se11su B1i1ton & Hose (1950); (,'uaymo.'>ia Brillon & Hose (JlJ30)
9 - 10 spp; Mexico and C Aml.'1ica, 1 exlcnding 10 Cu ha , Jamaic<-1 ;ind Cu ta\·:10, l
to \'en<..:zucla (including lsla Marg;uita) ;.ind Colombi;t
NanH::d for the hish bota nist Thomas Coulte1'(179.3-18·1<1) who c.:ollccted in C
Mexico (1821 - 1834) and was cu1at01 of the hc1halium at Tiinity College, Dul)lin
T1c1..•s and sh1uhs; scasrn1<.li1y d1y l1opica l forest, deciduous woodh1nd ::ind d1y
1ho1 n scrub, some species on limestone
Rct'c1 e 11cds): Kunth (!82/i); ll1itro11 & H1Jse (1930: 320 - 322)
The gcnc1a Caesa//Ji11ia L and Brasllellia (])C) Kuntzc a1c based on the same
type, Cae.wlpi11ia 1Jrasi/ie11sis L, so tllat Brasilc:llitl sens , strict passes inlo
synonymy under Cae.wljJi11ia \'V'hcn Ku nth Iii st dcsn ihed Cu11//eria he included
th1cl' sp<..:ci<..:s, hut did not typiCy the genus Two of Kunt h's species <ire synonyms
of (.'aesa//Ji11ia sjJi11osa (Molin::1) Kuntzc (here placed in the genus Tara) Thl'
rl'llli.lining species, Cvulleria mollis T<Linth, is c...:ongcne1ic with /Jrasi/ellia .'WllS/f
l\Jinon & Hose (19,30) and C'oulteria is thus the oldest available name r01 ;1 s1rnil l
group of srecies the1t were 1cfe1rcd to by Lewis 0998) as the Hrt1silellia group of
<.'aesa/j)illitL I n the molecular analysi:-> o l Simpson el al. (2003) rhc ·i spcc...:ics of
(,(,11/lel'ill sampled ( as mcmhc1s of the /Jrasi/ellfa spct:ies g1oup of C"aesalpinia)
tinm c1 wdl-supprntcd monophylt:tic g1oup sistc1 to 'Fan1 (as the J<11ss!!llode11dru11
species g1oup of <.'aesalpi11ia) Such a 1el;11 ionship supprnts rhe cailie1
phytochcmkal findings of Ki le&: Lcwi.o.; ( 199'i) whid1 suggested that rhcsc two
dcmcnls of Caesalpinia se11s, lat might he.: l'losl'ly 1cbtcd , Some (if not all)
Crmlteria spccil's arc clioecious, and the Jlor;il moi phology wilh a fimb1 iatl! lowei-
ca lyx: lobe is .suggestive or ~I pseud o-copulatory insect pollination syndrome . The
genus is c.:ha1 'aclc1isccl by a uniqt1e cnmh i11 ;1Lion of flowe r, fiuil and seed Lypc
comhined with a distinctive phyloc.:hl'mislly: it is unck:1 revision by J.L. Cont1c1<1s_J,
Cll ll(:ntly in Puebla, Mexico; combination ... in Omlteria <lll' not yet av;:1ilablc frn <Iii
the species 1ccognised in the gl'nll.'i
P1de11ccl loc.1lly for fiic wood; .'iome specie~ used <IS. 0 1namcnt;.ds in living
fl:nl'.elines
Cou/teria sp. Photograph by G. P. Lewis
TRIBE CAESALPINIEAE 139
 Caesalpinla cassia/des Photograph by G. P. Lewis
Caesalpinia L 1753
Pui11cit11w L. (1753); 1Jmsilellit1 mc.l Kuntzc 0891)
c 25 spp ; Ca lihhc;.m ( 10 in I·Jispanio la . Cuha and rhe Bahamas). Ml.'xico Cl-i
spp., l extend ing to C Ame1ica), l in Colombia, E<.'uadrn and Pe 1u , 1 in P~11 ;1 guay
and Argentina, 1 endemic to MacbgasGll', 1U in Ah ic1 (1 cndcmil: in Congo
lKinsho1scd, 5 r.;stJii.:recl to .sou thern Afrirn, 4 1estJ icted to the E and NE, with 1
extending inro Oma n, S Yemen and Arabia )
N~1111t:!d f(ll' Anclie;1 Ces::ilpino (1519-1603), ft;.ilian n<HuJ;dist, lmt~1nkal collcdor,
systcmalist <ind philosopher, physician to Popr.: C\c111cnl Vlll, profcssol' of
medicine ~111d botany in Pisa ~ind Rome
Trees and shrnbs; seasonally dry tropical wooclbncl, wooded g1;1ssland. co;1.-;L:tl
thi ck et. hush land and thrn n scr·uh, d ry plains '1!ld r ip'11 i ~in woodland, many
.speci .... .s on lime.stone or .sandstone
llefel'encc(.<): ll1 ilton IX !lose (1930: 32.'\-32(1); Ro." (1977: 122 -DO); Thulin
(1983: !6-18; 1993: 3•14-347); Uliha11i ( 1996)
few spet:ics of Cae.wlpi11ia SC!llS, slricl have be<..'n indudcd in 111olecul;.ir an:dyscs
to datc, hue Simpson l'I al (2003) found .'>ll ong suppo11 frn a g1ouping of C rosl'i
U1h fi o m th<: Dominican Jk:puhlic, (,' sessilf/lura S \Xlatson f'iom Mcxko and <:
11uufaJiasca1icmsis(R,Vig) Sene:-;se f10111 M;,1dagasc.11', as we ll a.s st1cmg ~uppo 11 fo1
C, r:assioides \Ville! from N\XI Soulh Amcrk;1 being sister to C, p11lc:herrimt1 (l )
Sw fiom Mexio) and C. Ame1ic.1 , An additional 12-15, predominantly Asian ,
species whk·h do not belong to CaesaljJi11ia sens slrlU nii..: provision~rlly placed
he1 e (set: intJoductoiy text to tribe), Thei1 <lffinity is Jalhc..:r to a clustc1 of
associated lu xa which rcqui1c f11rthc1· study before rhc..:y can be placed into gcnc1a.
The."ie rmw include Caesa/pinia sect Sappm1ia Benth, sL:ct N11µ.aria (DC) Hcnth.,
sect, Cinclidn1.:mp11s (Zoll.) Bcnth ., and the p1cvio11sly 1ccog niscd gene1a
/3i{/11 C.:ctl'({ Tncl, C'r.1mjJecia Adans , :me.I 7i'cc111to Ad.ins
C{/esa/jJinia }Jlfl<:herrinw (p1 idc of Barbados la I though nor native in lhc
C::rdhheanl, Jed l)itd of paradise..:) is widely LllltiV<lted as ~I wuden On1<llllLll!:ll ill
rh e t1upk: ..., and subt 1opics, ;rnd has varimrs medicinal p1ope1tit:s
E'Yfhrostemon g 1·ii·1es11.. Photograph by G. P. Lewis
Coesolpinia bohomensis Photograph by G. P. Lewis
140 LEGUMES OF THE WORLD
Pomaria stipu/oris Drawing by E. Catherine Erythrostemon (Caesalpinia) colycino (unpublished combination)
Photograph by G. P. Lewis
Pomaria cav . 1799
Me/a11oslicla DC. (1825): Clado1ricbi11111 Vogel 08;17)
16 spp.; 9 in N America (SE USA, C and N Mexiro) , 4 in S Amel'iG1 (SE Brazil.
Pa1agl1<1y and Argentina), 3 in southern Af1ica (Namibia, Uot~wctna and S Af1ka)
Named frn Dominic Pomar, bot~rnist from Valencia, and doctor to Philip HI
(1598-1621), King of Spain
Sh1ubs or pcrcrrnial herbs; mainly in subt1opkal cl1y areas of grassland and
wooded g1;1ssland and in degraded sites, many on limestone
Hefcrcncc(sJ: 13l'urnmitt & Ross (1974, as l-Jc!f/i11mi11seg~ict): Simpson 0998);
Si111pson & Lewis (2003)
Nnt 1ecognisecl by l'olhill IX Viclal (l981) m Polhill (1994); Simpson IX Miao 0997)
and Simpson el al, (2003) have clearly clemonstr;tted that Ponwrir1 (including
(.'/culolricbium from S Amedca and Ml!la11oslicla from southern Africa [the latter
dealt with u11c\e1 J!q/Jiummse~ia by Bnimrnitl & Hoss, 1974D is monophyletic
and more closely rebted to the Poinci<1nclh1 - E1yth1oscemnn g1oup of C(u•.wlpi11h1
sens /t1J., th::tn to lfoj}i11mu1se.l!.Mill with w hk h it has previously been confused
Erythrostemon Klotz.<d1 184/o
Sc/Jrammia B1itton & Rose 0930)
12 -1 3 spp.; 1 in S USA ancl Mexico, 11 - 12 in S Ame1ka (I in NE Biazil, 2
enclemir tn Chile, 7-8 in Argentina (3 of which extend Lo Bolivia, l tu Bolivia ancl
Peru, 1 l<> U1ugu<ty ancl l to Pa1<Lguay], l 1cst1ictecl to Peru and Ecuador)
F1om e1ylbro- (Gk:: 1ed) <incl ste1110n (Gk : staml'n), the type species L', gilliesii
(W;.dl ex I look) Klotzsch has long 1ed exse1•ted swnH:::ns
Shnrbs and woocly-hased pe1e11nial hc1hs; season: tlly chy tropicil ancl subtiopical
semi-;1 1id Lhor n sci uh (including caating~r), spiny <.:actus sci uh, \V(HKllaml and
g1assl:1nd
HelC1cnce(s): Brillon IX l(ose (1930: 326); Lewi« 0998)
B1irton & Hose 09:m> recognised the genus as di:-;linct from Ca£·sa/pinio; Lewis
(1998) 1·eviscd the combined Poinci;u1elb-Eryth10stemon group ol CaesaljJl11ia
hut did nor rei nstate eithe1• element <ll generic rank . Most /:,'Jytbroslem on species
;ue cuncntl y to be found in the lite 1alllle and in he1ba1ia under <.C1esaljJiHitL In
the molt:<..:ula1 analysis of Simpson el al (2003), 4 species of E1ytb1 ·osle11um fu11ned
;1 cl;1de within a wdl-."iupprnted Poi11ci~111ella-Eryrh10stemon clade , Species
1dacion.ships within the genus a1e .still IO he resolved
/~'lylbroslumun gilliesli (yellow, or· dl'Sert bi1cl of p<11:rdise) is widl'ly cultivatL'd as ;1
g:11c.IC:'n rn namcnt;d <rnd is used in r•evc::::getation
.
TR IBE CAESALPINIEAE 141
 Poincianel/a exostemma Photograph by G. P. Lewis
Poincianella B1itto11 & Hme 1030
c. 55 spp.; Neot1 opic.:s, 2(1 in N Amctica ancl lhe Ca riblJern (lit -1 5 1esL1 ic lt;d Lo
Mexico [of which 2 endemic Lo Baja C:tlifol'llial, 2 c:-;Lencling f'i 'o111 Mexico \(l S
USA, 'fin Cuba [of which 2 enclcrnic, l exrcncling to IIis1x111iol:1 and l lo i\11 \i(ol,
I enclernic in 1-lispaniola, 1 cndcmic lo Nic:11·~1gua . .1 wiclcspre~1d ac10,-.,s 1\k;·, i( o
and C A1nc1iG1), 7 in E and NF B1~1zil ( l e:-;tending to C l~1~1zil and I to\'\: l\uzil.
Bolivi;i, P:11<1guay <1ncl N Argcnli11:1), J umlcsc1il>cd sp , cmll'mic to Pe1u
Dc1ivccl from Lile di1nint1livc o r H1111 t ia11i1, (] gl:'llLIS o r legume now in syll(lllVlll)'
unde1· (.,'t1eso//Ji11io ;incl confused with 1Jt:lo11ix by V<tr·ious :tulhors; /)ui11c i11w1
j)(l ...,sil)ly n;1mcd r()! Loui.'i de Poinci ([)<>incy,jl c 1(1)0), l'IL'llCh (~(JVC!l(JI ()I llll' \'o/
Indies :.111cl 1x1trnn o[ botany
Tice"" and shrubs; ,..,c:.tsonally d1y 11opicil :1ml suhlropi<.•;.il woocllancl, arid tl1()111
and t:tclus sc1uh (in c lucling t:taringa), hc;ich 1hic ke1 (including aihrnc;il 1t ~ .,1111g;i)
<tml coa.~ tal dutll'S, -"::Indy w~1slK'.S :1ml pl:tins, ,scvc1al splxies cm .s~111dy S(Jil..., <•l
liml'stonl', less frequent in wooclL"d g1'< tsslancl ( <..' el 1';1do), gal let y forest ~rncl 111( lLsl
L·oast:il fon..·st, l sp, in mixed pine-oak fot'L'sl, m:tny on highly dq.,\1<1C\ecl silt•"
Hl'fl'1c11cl'(.s): l~1 illon & Hose (I ')30: .127 - .136); J,ewis ( l 99H)
B1 itron & l{ose (I t):)O) 1ecognised lhc genus as dislincl holll (.'(fesdlj>i11it1: I ( \Vi"
(ll)98) tevisecl lhe co1nlJinl'd Poinl'ianella-E1·y1h1 '( isternon g1oup ol ('rt('Strl/1f11fd
hut cl id not 1einst;1le cithe1' l'lcmcnl al gene ril' l'ank , Sevl'1-,tl Foi11 c id11d!d ."pt•t il">,
especially llm-"e l"rom s Allll'li l':l, :ire lll!IL'lltly lo he round in the lill'l~l[Ull: u1d in
hcd)<u·ia uncle!· (.'(1t'sa//'i11ia ln the molecul:tr :1nalysis ol Simpson el t1! (2tHl ')) • ll
species o f' H1i11 c ia11el/o and H1ythrosh•111011 togethe1 fc)l'mcd <1 well-supp< )rft•cl
dad!.=', IJLtl 2 wiclcly clist1·ihured spp. of J1oi11cfrme!/d (Cl1t•salpi11id e1•fos/(lt/1J''
lknlh . and C pl11oiusu DC:.) clu ...,lc1cd logl'the1 in ~tn UlllL\'iolvecl positirni rn11-..idc
this <. l:.1de. Spel ie.s 1elationships \Vithin the genus arc still lo he rcsolvccl
0
A few species u.-,cd :ts g~11lk'n 01·m1mcnl:tl.", dccrnalivc sl1l'l'l l1ecs :incl livi11:.•,
femelinl's; Caesa//Jillid cd Jilldld I.am. (pau h rasil) is tcnl<tlivcly placnl ill
H!i11cim1cl!d :ind is lhc 11el' tl1:il gavc ils name to l~1ar.il; omc Vl'IY i111prn1:1111 ;ts ' 1
M>urcc of a reel clyc it 1cmains 1he only sou rce of~ wood usecl f"o1 high qu<ili1:
violin hows; :-;0111e spe cies have kll:il 111eclicinal uses
Cenostigma Tul 18'13
:S spp.; C:, NE ~tnd 1x11l o r Amazonian B1;tzil
[•'mm Ct'//0- ((;[(: e111ply) and sligllla, j)ll'Slllll:tlJly ;tlJuding lo the cf1a1lllll'll'tl
-"tigrna Cf'ound in many specie" of Lhe Cae.<.;;tlpini;1 group)
Trcl's ancl ,<.;hruhs; SL;asonally d ry tropical hushl:tncl and tliickl'l (c1:.11ing<1, ,-,l·1 11 i: 111 d
lh(lnl scrulJ), \Vl)(Jclcc] g r:t..,<.;J:tncl (Cl'lraclo) :tncl tl'll"..l f'i11lll' rt11est
lkfc1cm~ds): Lewis ( PJ87: 34-j'i); Frd re ( 199/i)
Placed in lhc C:tcsalpini:t group ol the Ctesalpinie:tl' by Polhill &. Vicl~tl ( [ ll,'.; I)
;ind Polhill (l9t) O; nesting williin the Poinci:mell:t g1oup o f' C't1estr//Ji11it1 ill \Ill""
1rnirvh(1logical :1nalysis <ii Lewis&. SLh1•i1•c ( ll)C)'i)
Cenostigma gardnerianum Photograph by G. P. Lewis
142 LEGUMES OF THE WORLD
Guilandi11n bonduc Photograph by 5. A. Mori Guilandina bonduc Photograph by G. P. Lewis
Guilandina 1. 17'5
/Jo11d11c: Mil l. 07'::i•D; <.'t1esa//Ji11it1 suhgl'nus <,'uilmulina (L) CJillis &. fJioctrn·
(197/i)
7 to :1s high as lH spp . (the lowe1 figu1c 11101c 1c<1listk); pant l'Opictl, f"i om :is 1":11· N
:is Jap;in south lD S A l"1ict, .~ in the Cal'ihhean, I in China, India, Myan111:11•
[Bu1 lll~1J. lnclo China, I Iong Kong and Taiwan, l cnclemil• lo M:1d:igasc11, l in
Aust1alia, ~mcl 2 widcsp1l':tcl ae1os:-; the Olcl ~md New Wrnkl l1opics
Named ro1· Mekhi(l1 \XlieL1nd (1')1'1-1')8t)), Pru.':isian n<1lu1;.1list, ll:tvclll' r ;tncl sclH1la1•
f'mm Ki)nigsherg, who ,-,eulecl in lt<tly :.111cl itali:u1ised his name to ·c~uilandini' :ind
btini.scd ir as C~uil;1nclint1.s; he w~1-; sent Lo Lhe Levant, Asia ~incl Af r ica (l'i'il)-J')()(J),
w;is th,1plurecl IJy pi1Jh.::.;; :incl linally 1<111,-;omccl l)y Cahriele F:tlloppio
Sc1~ 11nhling p r ickly vines o r sctnclcnl .-;hrubs; coast.ii s,mds and thicket, ,-;econcla1 y
foil'sl, lowl;tnd 1 ~<1ill f'rnesl, some on lirnl'slone, 0-1770 Ill
l~cfc1encc..:o(s): B1itton & Hose (lt)j0: :-1d6-Yil); Heald 09l)·0; J)u Puy &
l~:1hevohil1a in nu Puy el d/ (2002: ,fh-,18)
Gillis&. l)1oc ru1 (1')7'i) 1ccogni.scd <r'11i/(//u/i1w <1s :.1 suhgcm1s ol C(1t'salpi11i(f ,
Ve1'lluH11'l ( 1~)7l)) .suppo1tcd the 1cinst;1tcmcnl of (,'11i/(//1Cfi110 at gcnc1ic 1ank .
Polhill&. Vid~d ( 1{)81) placed it :1s <1 .synonym of' <.t1esdljJi11id hut 1'Cm<11kecl upon
tl1l' unique nillll)in;ttion <if uni."il'XU:tl ~lt1wc rs, ha I'd gl<1IJulal'" seeds \Vilh st( w,1gc
cotylL"clons ~incl a clislinclive accumulation of non-ptnlein a111ino adds which
togethe r ch<tLlclcrisc the genus and scpa1~lle il l1'om other elernl'nls of
Caesi1/jJi11ir1 se11s /t1/ , llealcl ( ltYH) 1educecl lhc genus Lo six spcl'ies and fou r
va1ielics, l)Ul hc1· Wt)lk is unpulJlishecl and 11ccc"s'11y new eo111hinalit111s in
<i'11illl11dilltt ail' oulsl:tncling. Du Puy &. lbhl'vohitta in Du Puy el al. (2002)
clcsu ihecl (.'dl'so//Ji11io delpbi11e11sis ])u Puy & H.Rahev , ~is ~1 nl'w clo.':ie 1"elarivc lo
(/. hmu/11c. Only c;_/Ju11r/11c L. has been included in 1ccenl molecula1 studies ~111d
:t specic.<;-leve] mollXLlla1 su1vey o !" :tll (,'11f/(//tc/i11c1 hinomi~ds <1m] rcl<tll'd
Ct1es11/jJi1ti{f spl'cics would seem app1op1·iale
The .seed:-; o! all <i'11ilaiuli110 spl'.' cics (nicka1' beans, ni e k~11 • nuts 01 nickds), aie
comnHm d1·ill ,-,eccl:-; eap<1hle ()I llo~1ting :tcl'oss oceans The seeds and ll':1vcs of <,'
/Jom/11c have many medicinal uses; the seeds we1e once used in Eu n )pe ro make
IJul!<)JlS, :.tncl 11uny ltictl crnn1nunities Ll-"l' llK•tn HJ 111:1ke ncckL.1l'l'." ancl lw,ttclds;
roas1ecl .-;eccls h~1vl' hecn u.sed :1:-> ;i co!fee suhstitute Tile seed-; o f' some spl'cics
a1e used as etluntc1.s in h<>a1d g:ime.s o r as lhe equivalent to 111;1dJll''> in :1
chilcl ren'.s g;imc in lite C:11 ihhe:tn
1
Gui/andit la Cl·1·tata Drawing by P. Halliday
TRI BE CAESALPI NI EAE 143
 Libidibla sc/erocarpa Photograph by c. E. Hughes
Libldlbla (Caesa/plnia) glabrata (unpublished combination)
Photograph by G. P. Lewis
Libidibia (DC ) Schltdl. 1830
Caesalpiu/a .<ectio n lihldi/Jit1 DC. 0825)
6-8 spp.; Neotwpics (l in Mexico, 1 widesprc<id in Brazil (w ilh p~rhaps a second
sp. more restricted ), 1 in Colombia, Venczuel;.1 Hnd the Antilles, 1 in Colombia,
Ecuado1· and Peru, 1 in Pa1aguay, Dolivia, Argentinct and SW Bwzil and L curiaria
(Jacq) SchltdL wiclesp1ead throughout Mexico, C An-1e1irn, the Caiibbean ;md NW
South Ame1ka, 1 possible add itional sp. restricted to Colombia)
Derived from the ve1 nacuh.11 name 'libi-clibi' OI' 'cl ivi-clivi' used fo1 ,o;ome speci es
Trees or shrubs; seasonally dry t1opical forest ::ind scrub, tho1 n fo1esl (incl tiding
caatinga) and .sav,mna woodland
Reference(,): Britton & Ro.<e (1930: 318-319); Fo rd 0995)
Lewis & Schrire (1995) recognised grm1ps A and Il within the Libidibia grour ol
G'aesalpi11fa sens lat G i oup A contains o nl y the generic type species, l coriarit1,
while group B contains ;tlJ the othe1 speci~s. l. corfmi't1 differs in h~tv ing wh ite
(nol yellow) flowers; exsertcd stamens with red :tmhers; rough , fissllled ba1k
whkh Oakes ;1way in vc rti rn l strips (not smomh, patchwo1k bark exfolhlling in
woody plate-<), and tlowers hocking the te ntade-like p;opillae of the o ther specie'
All Lihidl/Jia species have indehiscenl woody, bl;ock 01 d;11 k brown pods, but
those of l coriaria ~irt:: distinct in being cu rled und twisted
Used as ornamental park and srrect crecs; pods rich in rannins and used
coir1mercia lly in the tanning industry, pods of sorne species also used for anilrn1I
fodder, ink ::i ncl local medicines (du e to astringen t prope11 ies of the t<.mnins);
wood prized in turne1y, ancl for parts of gu itars an cl violins, tha t of CaesaljJinia
glahra/11 Ku nth a nd C parag1wrie11sis CD.Parodi) Lllll kart (both species of
LibidilJia [pa11ritlge w oocl] but without combinati ons in the genus) u.sed in
clecorative inh1y and cabinet work; some spedes used in heavy co nstruction
(1"ilway sleepers, beall\O, bridge supports), for to o l lrnndles and as firewood
Hoffmonnseggia g/ouca Photograph by 8. 8. Simpson
Libidlbla cor/arla Drawing by E. Catherine
144 LEGUMES OF THE WORLD
Stahlio monospermo Drawing by P. Holliday Stohl/a monosperma Photograph by M. F. Gardner
Stablia Bello 188 1
1 sp.; Cai ibbcan ( Pue110 Hico and Dominican Repuhlic)
Named in honot11 of Augus1in Stahl 0842- l917), Puetlo Rkan ph ysidan <.tnd
nallm1lis1 of Ge1man-Outch p<1ren1 ~•Ae
Tice; uopkal m~mg1ove swamps (margins) and nu1rshy deltas (tolL"1atcs sale),
fore."il remnants
Refe1 en~e(.s): Britton 0927); Il1 iuon & Rose 0 930: 221)
Placed in the Caesalpinia group of the Cacsalpinicae by Polh ill & Vidal (1981 ) and
Polhill (1Y94); siste1 to a claclc of 3 species o f the Libidibi:1 group of genus
Caesalpiuia in the molecular analysis of Simpson et al. (2003), a position
suppoi'ted by similar imlchisccm woody fl'uirs, Jlmv c1· mrn phology ancl foliage
glands
S1ablit1 mu11ospe1•nw (Tul.) Urb w(<.:6banc1 m:g1<1) is va lued fu1• its bl~1ck, he~1vy,
durabk wood
Hoffma11nseggia cav. 1798
f,<m WI O rtega (1797), non Larrea Cav (1800); Moparia Il1itton & Rose (1930)
Z·i spp.; 10 restricted to N Ame 1ic._1 (SUSA and Me xico), 13 1estlictcd to S Amc1ic;1
(Pt:lll and Bolivia lo soulh-o:nti;tl Argcntin~• ;_ind Chile, many Andean), J sp. ( /-1.
glauca (Ortega) Eife1t) th1oughout 1hc 1angc of 1he genus
N~1me<l frn the Geiman bol<rnist, entomologist and u mithol<>gist, Johann Ct!ntu1ius
Graf von 1-Ioffma nnsegg (1761>-1849)
Shrubs or perennial hL"1bs; subt 1opic:d dc:;eit and scmi -dcscit g1"<1ssbncl, oft~n in
OfH.:n a1eas ancl on distu1b~d sites, on sanely, iocky rn c;.1 Jca1eous .soils
lk fe1ence(s): lJliln111·i (1979, 1996); Simpson (1999); Simpson el al. (2004 ; 2005)
Placed in the Caesalpinia grour of the Caes;ilpinicae by Polhill & Vidal (1981) and
Polhill 0994); Simrson & Lewi' (2003) place Af1irnn species of f-lo[/t11c11111sel!l!ill in
Pomaria (1ecently reinswtcd as a distincr ge nus); thc gen us is !-iister to Z11C:cap,111<1
Ho.ffman11sep,git1 glauca (Indian rush pea, hog pota10) p1oduccs tube1s once eaten
hy indigenous g1oups in N Amc1•ica (hu t .sometimes beco mes :a noxious weed);
the ioots of II i11lricalt1 Jlr;.mdcgee piodut:e a reddish b1own d ye
TRIBE CAESALPINIEAE 145
 ~
!. .. _/

Stenodrepanum berg/I Drawing by G. A. Larsen and E. A. Ulibarri Zuccagnla punctata Drawing by c. A. Sobel Ba/samocarpon brevifo/lum Photograph by M. F. Gardner Mou/lava spicata Photograph by M. Sanjappa

Stenodrepanum Haims 1921 Balsamocarpon clos 1846

1 sp , endemic to C and W Argentina 1 sp.; S South America (Chile)
From steno- (Gk.: narrow) and drepano- (Gk: sickle), in allusion to the m111ow From ba/samo- (Gk.: balsam) and cmpos (Gk ,: fruit), the pods yield a sticky resin
sickle-shaped fruit t1aditionally used for tanning
Shrub o r perennial herb; subtropical wooded grassland and SCl'ub, especially on Shrubs; c\ese1t scrub, rocky hillsides
salt pans Reference(s): Burka1 t 0940: 162)
Reference(s): Uliball"i 0978; in Kiesling et al., 1994: 285) Placed in the Caesalpinia group of the Caesalpinieae by Polhill & Vidal 0981) and
Placed in the Caesalpinia group of the Caesalpinieae by Polhill & Vidal (1981) and Polhill 0994); siste1 to a Zuccagnia-I-Ioffmannseggia clade in the molecula1
Polhill 0994); as yet not included in molecula r analyses, but mrnphologically analysis of Simpson et al. (2003); sometimes placed within Caesalpinia sens lat .,
close to Caesalpinia (Poincianel\a group) and Hoflincumseggia but cle<irly distinct from Caesa/pinia sens, strict.
The fruit resin is used for tanning

Zuccagnia cav. 1799

1 sp.; S South Amelica (Chile, NW and central-W A1gentina) Moullava Adans. 1763
Named for the Italian physician, trave\le1 and plant collector, Attilio Zuccagni Wagatea Dalzell (1851)
0754-1807) I sp.; enclen1ic to India, int1oclucecl as an rnnamental in Hawaii and Z<Jmbia
Shrubs; cllJ' temperate upland and montane brushland, thicket and sanely plains De1ived fiom the ve1•nacuh1r name 'mullu' (Malayalam: spiny), a spiny climber
Referencd,): llurlrn1'l t I 52; IR~- 18~). Ullb11rri (hi Ki ling el al., 199': 281!) Liarn:1 or scrambling shrub; seasonally cl1y twpical semi-everg1een frnest margins
Placed 111 Ill<! tnc•a lpi nf:i !tl'llliJl or the Oll:!>il lp luleac J1y Polhill & Vl<lul ( 198 ll and Reference(s): Nicolson 0980); Sanjappa 0992: 33)
Polhill C I ~!)<\) ; sb tcl le> rll'ij})11111111.<egs f(I l11 1hc Ill b ;1J lar analysis of SimpsJJn el al. Placed (as Wagatea) in the Caesalpinia group of the Caesalpinieae by Polhill &
(2003) Vidal (1981) and (as Mo11/lava with Wagatea in synonymy) in the same g1oup by
Mino1 loc<il medicinal uses; the leaves yield a yellow dye Polhill (1994); few molecula1 analyses have included the genus, but the molecular
study of Simpson el al (2003) found weak suppo1t fo1 placement of Mou/lava as
siste1 to a neotropical Balsamocarpon-Zuccagnia-Hoffmannseggia clacle
Lophocarpinia Bu1brt 1957 Mou/lava spicata (Dalzell) Nicolson (false thorn, 1at bean) is used fo r medicine
1 sp., S South America (Argentina and Paragu<Jy)
From lopbo- (Gk.: combed or crested) and cmpos (Gk: fruit), the f1uil has 4
crested wings, the ending -inia signifies a close 1elationship with Caesalpinitl
Shrub; seasonally d1y t1opical to subtropical woodland (chaco)
R •forc m:d.~): llurk uit 0957)
Pi:t c<:Jcl in 1h e <1c;" Jpinia grou 11or1 l1c Caesalpinieae by Polhill & Vidal 0981) and
Pc!ll111l (1994): •istc r to a Lemu mpl6u m-Colvillea-Delonix-Cenostigma clade in the
molecula1 analysis of Simpson et al. (2003), but for fu rther comment on the
relationships of Cenosligma see notes undei that genus

Lophocarplnla acu/eatifolla Drawing by c. A. Sobel Mou/lava splcata Drawing by E. Catherine

146 LEGUMES OF THE WORLD TRIBE CAESALPINIEAE 147

Batesia f/oribunda Dra wing by w. H. Fitch
Melanoxy/on brauna Drawing from Mart. Fl. Brasil
148 LEGUMES OF THE WORLD
Recordoxylon speciosum Drawing by P. Halliday Moldenhawera b/anchetiana var. blanchetlana Photograph bys. A. Mori Maldenhawera nutans Photograp h by G. P. Lewis
Batesia Sprt1cc ex Bcn th & I look.f. 1H65 Moldenhawera schmcl 1821
l ·"P; Am;1zon ia n Brnzil , C:olombi<t a nd Pc1t1, and Fn~ n ch Gui an a LJCJ!icbo11em ill ees (1821)
1amcd foi Hl!n1y \'Valte 1 Bates ( 1825 -1 H92), l ~nglish e n1 011mlc,g i:·a, 11 :ilu 1 ~1 l i:-.1, 9 sp p ; endemic Lo E B i::tzil (') spp in Bahia)
tic1ve ller rrnd ph1 n t c;olle<.:tOJ, com panion or
Alfn.:d lhl.'iSC!l \Val\ace Named fo r Jo ha nn J aco b PaLil Molclenhawcr (1766 - 1827), Gc1man p rofessor or
T ree; trn p ic:.11 no n-inu nchtted 1ain fo iest, mainl y o n sand hot:in y al K id a nd o ne of the fou n ders o f p lant ;inatomy
Hdcrence(s): llentha m 0870: 56, t, 16); Duckc (1949: 129 - 130) T rees and sh1'l1hs; di verse r~t n gc o f hah it:tts: tropica l moist coast<:ll forest,
Placed by Polhill & V i d~ il 098 1) in the Pd top l10n 1111 g 1oup o f C:~lc!'!<llp i n i e ::1c In n wood land a nd scrub on p ocl.sol ised so il.\ just ah ov~ the beac h line (resting;-1),
no t incl uded in th e 'unc-Pcltopho1·um g1oup' in lhc molecular an<il ys is ol l-h 'i lon coastal d unes, low mountain sc ru b 0 11 sa ndsto ne, wooded g1;1ssland ( cena do),
el ri/, (2003); in th e co mbine.x i morphological <ind 111okcul:i1• ::111alysis of I rc1ernk:cn and rock y grassland (campo l\lpes tre)
el e1I ( 2003 a) si.'it(..' 1 lo Vo1wcc1jJ01w, but this is nol suppoited by th e analyses of Refetence(s): Queimz et al (1999)
L-Ja sto n C!/ llf ( suhmiltc d) who found I.I wcJJ .suppo1tt.::d 1d::itio nship between Q uc iroz et at. (1999) found (in a p1 eli111in a1ry mrnphological analysis) that th e
Hofesi ll, l?el:urdu.\:)llu 11 and 1He/c1110.'.ylon gene1a Melmw.'\ylun <Incl N.ecordo.'\:y/on a r~ prob abl e s iste 1 tax;J to Molde11baweru ,
The ti111bc 1 o f 13 jlo r i lmuda Sp1uce ex Bcn th is Ll s~d lm::lll y !'01 furn itu1 c a11cl Placed b y Polhill & Yicl~il (1 981) in th e Pd to ph rn 'l11n g10up of Caesalpini eae b u t
small c_11·p cnl 1y items not inclu ded in the 'core-Peltop hrn um g1oup' in the molecu lar anal ys is of l-1 :1sto11
el lll (2003), and p laced in its ow n J.;IOUp by Haston el <1/ (su b mi u ed) <L'i sister to
a Tac; higal i g 1o up - co1e-PeltophrnL1m g1m1p- D i11101phaml 1a g 1oup - M i mosoide;_u~
Recordoxylon D ucke i93;, clacle
3 spp; Alll<17.onian Bmzil, Guyana, French Gu iana and Ve nezuela
a med for Sann1cl Ja1m:s Recrn d l 1881 ~ l lJlil), Ame1 iG111 h ota n i.-;t, dend1ologisl
and wood ;.1natom isl, whose o b sc1 v at ions of t he wood st 1uclu rL· led Duckc to
H.:visc hi.'i rnigi n11 \ classificH ion, and .\v/0 11 (Gk: wood)
Trees; non- tl oo ded 1ain frn est o n tC ird film e, 1110 11\a ne rrne.s l , sea.son ally tl oo cl cd
1iv c1 inc rrn1.:st (v;cl 1zea)
l(cfer encc(s): Du cl< c 093/ih) ; Mar iaux & Normand (1 96.'IJ; ll;11 neliy ( 1993);
Ayrrn11d i11 l k r•ry el o/, (1998: 98-99)
p[,1ced by Po lhill & Vidal (1981) in the Peltopl101um g1 u up o f Caesalpinicae, !Jul
not inc lu ded in th e 'c01 e-Peltopho1l1m g1oup' in the mol ecuh11 an cll ys i.'i ol Has!< >1l
el(//, (2003) in whic h it is siste1 to Melmw.\y /011
Melanoxylon schott 1827
l sp., E Bro1z il lplincip:a ll y Hio de Ja nei ro, Min~ts Ger.tis ;1ml Bah ia)
From 111e/0 11rJ- ((; k.: !'lack ) ;1 nd x yluu (Gk.: woc)d); the.:: heart wood i.'i alnui.-;I l)btk
T 1ee; seasonally d1 y l 1t1pical .'iem i-dc.::cicluous forest, li:1na frnc.::.'i l (mata de.:: c ipc.l),
d1y woodland, tho1 11 .'ic1u b (c;1 ali nga) <ind thicket
Rcferencds): Lewis ( 19H7: 30-32)
Pbced hy Polhill & V ida l U 9H lJ in th e Pd topho1L1 m g1 u u p o f C;u.~s::tl p in i c;i e hul
not incl u de d in the 'con.~- Pcl tu pho1 um g1oup' in the m o lccu \<1 1 :ina lysis o f l h<>!Oll
el al, CW03) in w h k: h it i.s sister to Recordo.y y/0 11; co n t1a1y to so me lite1atu1e
(wh ere sp cdcs o ! Necordo.\JJ/rJJI wc1e inclu~lccl in Melm1 0.'\y/o11 ) th e genus is nol
in the A1rn1 zc111
Th e timbc1 o f M h rmuw Scholl (ht alma, ga1 :1l1na ) i.s h<t1 d ;ind du1able :md u.st.:d
in co nstru clio n ~in cl frn fin e fu1nitu1c; the ha1k is a so u r<.:c o l tanni n; :,tl so u.scd for
1ncc.lkinc Mo/denhawera papillanthera Drawin g by H. Green op
TRIBE CAESALPINIEAE 149
 9

Arapatiel/a psllophylla Drawing by C. A. Sobel }acqueshuberla brevipes Drawing by P. Halliday

Tachiga/i versicolor Photograph by R. Foster

Tachigali Aubl 1775

Arapatiella Rizzini 8: A Mattos 1972
2 .spp ,; endemic lo SE l31<1zil ( L3ahia)
'f'acbigaJia, 01'thog1t.1phic varfa nt first used by Jus.sicu in 1789; Sc/ero/ohi11m Vogel
Dcdvcd f1on1 th~ loGtl name ·ai:tpati' frn A j>silojJbylla U-l;.ums) Il.S Cowan
(1837)
'!'ices; t1opica\ moist coastal frnest (mat:t higa">fib)
c. 60-70 spp; Neotrnpics, l\1l)Stly in S Amt.:1ica with 1rn1i11 centte ofdive1sity in
ltdei-encl'(s): Rizzini & Mattos filho 0972); Cow:in 0973: 447-450); Lewis (1987: 32)
the Amazon ll<lsin, hut extending S to P<1wguay ancl Argen tina ;rnd N into C
Phiced by Polllill & Yiclal ( L981) in Lhc Pelwphorum g1oup of Caes;1Jpinieae but
Amc1 ica (I ende mic in Costa Hica, 1 mon..: wide.spread); l ') in Yenezm~ lan
not included in the 'co1e-Pellophrnurn group' in the 111olecub1 analysis of llaston
Guayan•1. of whid1 3 endernit.:, 1·1 in Peru, of whid1 5 endemic, sevend spr
et ol. (2003); f-b1."it<m c:I al (submitted) I ind strong molecular support grouping
widcsp1'\:';H.I in S America)
Aropaliel/a wil'hjau111esbHIJeria and 'Jltcbt;C!,ali Mo1phologically the gt::m1s has
De1·ived f1om the Turi-Guarani Indian ve1rn1cular name 't;tchi' or 'tad' fc>1 slinging
been associated with T'{l(.:hip,ali (and Scle1t)lohium)
ant; many .-.pecies have a c.:ln."ie 1·elationship with ants and are 1efe1·1ed lo loctlly as
t~1<..:higali (ant-loving t1ees); Sch:rolohium (t ile gene1ic synonym) is f1om si.:lero-
(Gk: h;11d, d1y) and lo/;io11 (Gk: small llllil) Jacqueshuberia Durke 1922
T1ees, sometime.'> monc1Gt1pic; mainly of !l(>pic~ll 1•;1in frnest, lowland 1rn >ntanc 7 spp; northern S Ame1ica C2 in Amazonian B1<1zil, l in Guyana, 2 in Venezuelan
forest, SCiJ."iO!l<Jlly nomled ttml non-floock:d l!Verg1CL'll lowland foieSl <tile\ (Jua y;;1ria, 1 in Colombia ;tnt.I I in Pe1't1)
woodland, galle1y aml 1ip<1rian foa,.st, sometimes on white sands, also in N~111K·d for.Jakob (Jacques) E Huhe1 (18(17-1914), Swiss botanist who ~xplorcd
seasonally d1y woodl<1nd (including ce11<1do) and rocky g11.1sslancl sci uh (including the Amazon fron1 1894 to 190·1, .'iettled in Belem. P;:11~·1 , lhazil in 190') and
n1rn p< l 1upcst1e) developed tlle Museu Gocldi (Museu P:1men:-;e)
Refercnc<:::(s): Dwye1• ( 19')4 , J9')7a); Lewis ( 1987: 28 - 30)i 7.~1run:hi in lk.1k<> & T1·ees; tmpic;tl seasomdl y inundated 1iveiine fo1est on white s.and , mon1ane frn'Cst
Za1'ucchi ( 1993: 521-522; in Bel'I")' et a/_ (1998: 114 -I 20); Zarucchi & l-lc1cndccn on sandstone, sav<rnna, upen s~mdy a1-ca.'i by hbck-water streams ol th e Amazon
in Jl1·ako & Z;11ucd1i (]993: 1251); Pipoly (1995); Lla1n eby (1996: IH2) (caating;-1 and campinas)
Polhill ~I\: Vid:d (1981) and Polhill 099ii ) iecognised '/'acbigali and Sclerolohlll/J/ <l.'i lk!'e1ence(s): .Silva & G1aham (1980); Cow<in (198')); Ba1nehy 0990); Stergios &
separate gcne1~ 1 and pl<u..: ed lhcm, togethe1 wilh 1Ji/Jlydimulra, in lheil' llc11y 0996)
Sdcmlohium gtoup of t1 ibe Ctesalpinieac At least 35 spcdes we1e origin.illy A wide wnge ol"llowe1 colou1, including 1cd, pu1ple and ye llow exists in this
de.-.c1·ihed in Scle1·ulolJi/Wf Orhc1", rnoie 1eccnt works (e g, Zaiucchi in Br<tko 0. sm all gt.'nus . Pl:1ced by Polhill 1..'X Yield (19H1) in the Peltophrnt1m group of
Zarucchi, 1')93, & in Llcrry et al., 1998; l'ipoly, 1995; B:11ncby, 1996) ltavc Caesalpini e ae hut not included in the 'co1e-Peltopl10111m group' in the molecula1
conside1L:cl Sclero/o/Ji11m to he congene1ic wilh 'J'acln'J.:ctli, ln Lhc combined analysis c£ I Iastern el al (200~); I bston el rd (s ubmitted) plan~ Jacq11esbuheria
molen1l<11-111orphological analysis of Herendeen el al t2003a) 'f'ach1~~ali is ."iislc1 10 with Arflj)(tfiel/o and 'J(tc/J(~a/i in 0:1 Tach igali g1oup
a Pte1ogy ne-Chamaec1ista-Senn<1 cl;ide. 'fi1cbigali and <.1;cwwecrisla ~u e unusted
in lhe Ctt::salpinioidcac in both h~tving species that noclubte, IIaston el al Schizolobium vogcl IH.~7
(sul)miucd) rc>und a st1011gly suppo1'tecl 1clationship hdween 'J'achip,alt, I sp .; Allanti c SE B1azil (1 va1 'ie ty), Amazonian B1·azil, Bolivia, Pa1aguay, Peru,
!lro/Htlidla aml jaC(jlf£'Sht1huria E<.:u;iclor, Colomhit.1 and Vcnl.:'zuela, th rough C: Ameii<.:a lo SE Mexico (the nthe1~
Various species used for limber (djedoc , y;1wa reclan , suicitk: tree lhet:ausc some
mrnc w ide~p 1 1.:a d v;1riety)
~pecies 111onrn:a1pid) fo1 consl ruclion , e1110<:s and cha1coal; the bark of '/'. From scbizo- (l;I<.: split nr divided) and lvfJion (Gk: :1 s111:1\I pmD, th e inner :tnd
tim.·torio (Benth) Z:11ucchi & Herem! used in tanning and as ~1 dye m1[e1 laye1s ol the pod ,..,eparalc at 111:.Jlurily
Trees; t1opit::1I 1ai11 fo1est, 111ix.(.!d and seco11cla1y fore.'it
Hcfe1enceM: Ll:il'llehy ( 1991>)
A fost-giowing, sh<)\v y t1ee, tl<1wc1ing when lea tkss; P<1lhill &. Yi<..h1l ( L981) placed
the genus in th e Pehophonun g1oup orC:1esalpinie11e; included in the 'crne-
Pelloplmrum g1oup' in th e moh:cula1 analysis of I laslon el al (2003)
Scbizo/obi1f111 pc11c1byfJa (Vt:JI ,) SF Blake var 011wzo11iclfm (Ducke) lbrn ehy
(qua rnwood, guapu1uvu, 1e;1ch-frn-the-sky, 1e~1the1-clu:;te1 tiee) is widdy planted
:.is <Ill 0111ament:d Cinduding outside tlll! New Wol'id) ;ind <is shade lr~es <e.g.,
Schizotob"1um parahyba var. amazomcum
. Photograph by C. f. Hughes f(u 01ffcc)
Tachiga/I pan/culata Photograph bys. A Mori

TRIBE CAESALPINIEAE 151

150 LEGUMES OFTHE WORLD
 Busseo mossalensls Drawing by L. M. Ripley
Bussea Hanns 1902
7 spp.; 5 from Africa (1 endemic to Mozambique. 1 in W Afric.:a [lvoiy Crn1st,
Ghana, Libetia, Sie1ra Leonel, 2 in Tanzani<.1 (l extending to Zambial and 1 in
Angola and Congo (Kinshasa]), 2 endemic to W and N Madagascar
Na med for W<1lte1 Cad Otto Busse (1865-1933), German bownist and ecologist,
who collected specimens of the type species in E Africa in 1901
Trees or shrubs; seasonally dty tropical forest and thicket, moist semi.dcLiduous
forest, rain forest
Referencc(s): Brenan 0967: 25-28); Du Puy & Habevohitra in Du Puy el al
(2002: 21-24)
Polhill & Vidal 0981) and Polhill 0994) included the gem" in the l'eltophrn um
g1oup of tlibe Caesalpinieae, a placement confirmed by the molectila1 analy.sis of
Haston el al (2003) in which Bussea is sisteo to Pel1opho111m
The hatd wood of the 2 Machtg.ascan species is used in constlllction; in Af1 k a
some spedes used for building poles and firewood ; the seeds or B occide111alis
Hutch. me 1oas1ed and eaten in \Y/ Aftica , although unroasted tht::y are toxit: ;ind
have been used as H fish poison; c1lso used for medicine and for..1ge
Peltophorum (Vogel) Benth 1840
Caesulpi11/t1 sect Pellophomm Vogel 0837)
5-7 spp.; p<1nl1opica l (into the subnopics) with 2 native to che Neot1opics 0 in E
Bra zil, N Uruguc.1y, NE Argentina, P<iraguay, Bolivia Hncl the Caribbe;in, rhe u1he1
restricted to Vi:nc7.uela, although this p1obably conspec ific with the first), 1 rnHive
to southern Afn a1 1 in M;.ilesi;.i (Sabah, Sarawak, Kalimantan), and 2 widespread
in A.sh1 (Th~li lan c\, C.unbodia, Laos, Vietn;.:1111, and Peninsula M:daysia, 1 exLending
to Sumatra and )<tva, the othe1 to Sri Linka. the Philippines, ctnd N AL1strnlia);
nwny species widely cult ivated
From pelle- (Gk : a 'hidd) and p/Joros (Gk : healing) allllcling to the la rge pclt:ote
(mrl!Ct-,haped or ' hielcl-like) stigma, centrally attached to the style
'1 '1"~..,. seasonally
d1y tropic.:al <llld evergreen (mainly lowland) forest, beaches and
m;111grove forcsr, coasl<tl monsoon vine thicket, nooc.I plains and tkl.11 flats,
bushveld and woodlt1nd
lteference(s): Isley 0975: 176-178); Rrn;s 0977: '130- ·133); 1..<1rscn el al. (1980:
59-64); Cardena' <'I al. (191!6); Hou in Hou l'I al. 0996: 650-654); Barneby (J<J<J6)
Po lhill & Vidal (1981) and Polhill 0994) indlldc I the genus in the l'dtophorum
group of tribe Caesalpinieae, a placement n ml nnecl by the molecu lar analysi~ of
Haston et al. (2003) in which Peltopborum is sister to Bussea . Peltopbor11m
v<mez1re/ense L.Clrclenas, Roclr.-Rodr. & Varela prob:.1bly best treated as conspccifiC
w ith P. dublum (Spreng.) Taub, thus limiting the Neocropks to <I single
indigt:nous species
\Videly cultiv~llccl for ornament and shade; some species used for c.11'pentty,
planking, furniture nnd fuelwocxl; leaves Li.sect fc)I live.stock fod<le1; some
Parkins
Peltophorum dub/um Photograph bys. A. Mori medicina l uses and dyes
152 LEGUMES OF THE WORLD
Parklnsonio aculeoto Photograph by D. Du Puy Porklnsonlo oculeoto Photograph by c. E. Hughes
Parkinsonia L. 1753
Cercidium Tul. (1844); Peltopboropsis Chiov_ (1915); Cercldiopsis B1itton & ltose
0930)
11-12 spp. and 2 named hyboicls; 5 in SUSA :incl Mexico (2 extending to Baja
California :-tnd 1 widely cultivc1ted tlu oughout the tiopics), 1 in N Argentina and S
Bolivia , 1 widesp1ead from Mexico th1ough Venezuela, Ecuado1 <Jncl Pel'll to N
Argentina; 4 in Africa (1 endemic to Kenya, 1 to Somalia, 1 more widesp1ead in
the E and NE and I in Sand SW Africa); 2 natmally occu ning hybrids (both in
M~xi co ancl Baja Califomia, 1 also reco1cled F1om Hondu1as ancl Ecuador)
Named for the British apothecary (to King James 1) and herbalist John Pal'kinson
0 567-1650), author 0629) of Panidisf 111 Sole Pamdisus Terreslr1~' ('Pal'k in sun''
emthly r<11adise') and 7bealmm Bo1a11icum (1640)
Shrnbs and trees; season<Jlly d1y tropical bllsh lancl, semideseit scrub, coastal
dunes and flood plains
Refeoence(s): llrenan 0967: 43-45; 1980); Caller (1974); Burk;11t & Caotel'(1976);
Robertson & Lee 0976: 32-34); Hoss (1977: 109-l 12); Hawkins el al. 0999)
Polhill & Vidal 0981) placed the genus in the Caesalpinia gooup of C:oesalpinieae,
hut Polhill 0994) t1ansferoed it to the Pcltophorum gooup in agoeement with the
subsequently publi,hecl wook of Lewis & Schoi1e (1995); a position confoomecl by
the molecular analyses of llruneall el al. (2001) and Hc1encleen el al. (2003a)
whe1e it is siste1 to a Delonix-Lcmu1opisum clade; also nested in the 'core-
Peltophorum g1oup' in the m olcculm ana\y."is of Haston et al (2003) as siscer to a
Conwttia-Delonix-Colvillea-Lemu1opisum cla<le; 2 species and 1 hybrid in
Cercidium cur1ently hick comb inations In Parkinsonia
Parki11so11fa ac11/eata I.. , (Jea1salem thorn, palove1de) i.s widely cultiv:Jtecl
th1 oughout the tropics JS an 0 1 namenral and tho1 n hedge but can escape to
become a weed; also L1sed fo1 medicine and hunwn food (flowers eaten in
Senegal, the seeds of P. aji'icmw Sond used as a coffee sL1bstitute)
· .
onio scioono Drawing by L. M. Ripley
TRIBE CAESALPINIEAE 153
 Delanlx reg/a Photograph by o. Ou Puy Co/villea racemosa Photograph by o. Ou Puy

Conzattia Rose 1909 Colvillea Bojer ex Hook , 1834

1sr ; e ndemic to Me xico
N:.u ned for the Italian-born M exican botani st C'-!SS ia no Conzatti (1862- 195 1), planl
collecto1• in Mexico and Chile
Trec.:!.s ;;r n d s hrnbs; seasorn1lly d1y tropical to su btropical wood la nd, fort:st and
thorn scrub
Refo1t!nce(s): Sta ndley 0922: 427 - 128)
Polhill & Vi dal (1 981) placed the genus in l he Caesalpinia g1oup of Caesal p in ieae,
but Polhill 0994) t1ansfe1red it to the Peltophorum group in 1:1g1eemen t w irh the
st1bsequently publi shed wrn k of Lewis & Schli1t:! (1995); also nes ted in the 'crnl'-
Peltophorum g1oup' in the mo lecular an alysis o f Has ton et al (2003) as si.stc1 to a
lcirgely M<.iclagascan Delonix-Colvillea-Lem lll opisum d ad e; I-l<lston (pers co mm ,
2003) conside1s the g:enus to be mo nospeci fic A new genus, tentatively muned as
1-/eleroflomm by Sousa & Delgado 0 993) bul no1 yet frnm;dly published, is"
1 sp.; e ndemic to Madagasc::u
Namecl fo 1 Si1 Charles Colv ille ( 1770- 1843), distingu ished Scottish office r unde 1
\Vell ington in the Napoleonic Wa is, <incl Goven o 1 of Mm11 itius (1828-1 834)
T 1ee; ~e<1 somd l y d1y tropica l frnest a nd woodl~111d
Refe1encc(s): Ou Puy & Rabevohi1 ra in Ou Puy el al. (2002: 42-44)
Polhill & Vida l (198 1) and Polhill (199'1) i ncl uded 1he genus in 1he Peltophol'l11n
grou p of tribe Caesalpinieae, a p lacemen t co nfirmed by the molecu lar ana lysis of
Has ton et al (2003) in whi ch the genus grou rs wit h Delonix and Lemuropisum in
the 'corc-Peltopho!'um g1·m1p'
Co/vi/lect racem osa Boje1 ex Hook (Colvillc's glo1y t1ee, whip tree) is ct1\tiva ted ~s
a garden 01 namenw l in .'ieve1a l tru piGtl countries and is used locall y a::; a
decrn~llive shade t1ee; its wood is used frn conslnu.:tion pos ts, fences and iough
and its tru nks arc ho llowed ou( ro m;1kc ca noes
c.11 pe nt ry

monospecific Mexican endem ic closely 1elated to Conz attia

Co11za ll ia multijlora (13 L Rob,) Stanc.11 is used for me dic ine, co nstruction timhc1
and fue l; often g1 o wing ne;1r ancient temples indica ting early ex plo it ation hy local
com munities

Delonix Haf. 1837

Poi11cia11a se11s11 lla ill (1883), sensu H Vig, (1949) & se11s11 auct., non L.; AjJrctHlia
Baill (1884)
11 ·"PP ·; 9 endemic to Me:u l<1gascai, 1 wid esp read in E Africa, A 1abia and India , and
1 endemic to N K en ya , Somalia and Ethi op i ~1
From de/o- (Gk.: evident, visible or conspi ct1 m 1s) and onyx ( G k.: a claw 01 nail),
the pct::1 ls have conspicuously lo ng cl aws
Trees; seasonally d1y t1o pical forest, xerophyric woodlan d, sc1ubl<1nd and
bushland, especially dive1se o n lirnesto ne
Hcference(s): llren;i n (1967: 23 - 25); Thulin (1983: 15-16); Du Puy el al. (J99'>b);
Du Puy & Rabcvohi tia in Ou Puy el al. (2002: 27 -'12)
Polhill & Vidal 0 981) ancl Polhill 0994) included the genus in the l'eltophorurn
group o f tribe Caesalpinieuc, a placement co nfim1ed by the molecular ;malysis of
Hasto n et al. (2003) in w hi ch the genus g1m1ps with Co/vi/lea an d lem 111 opis11m
in the 't:rne-Pel to ph oru m group'
Cu lti va ted for shade a nd a:-; liv ing fence lines; D regia ( I3ojer ex Hook.) Ib f
cna mboyanr, toyal po inciana, name t1 ee) w idely pl amed th1 o ug ho ut the tropics
:incl subt1opics as 1.1 showy o rn amental of !Meets, pa1 ks and ga rdens; othe1 spl!de5
w ith high ornamenta l potentia l; some species pioclucc resin used ns g lue, rn
st1t:k ecl as sweets; the trn nks of a few species ;ire hollowed out to 1m1ke Grnocs;
seeds a1e c1ushecl an d eaten as a snack

Delonlx f/oribunda Photograph by o. Ou Puy Co/vii/ea racemosa Drawing by M. Tebbs

TRI BE CAESALPI NI EAE 155

154 LEGUMES OF THE WORLD
 Lemuroplsum edule Photograph by D. Du Puy
Lemuropisum H.Pe1rier 1939
I sp.; e nde mic to SW Madagascar
Derived from le mur and Pisum. (genus containing the edible pea), the seeds
allegedly relished by lemu1s
Shrub; seasonally d1y tropical xerophytic scrubland on exposed limestone 01
white sand over limestone
Reference(s): Du Puy & Rabevohitra in Du Puy et al. (2002: 25-27)
Polhill (1994) followed the proposals (published one year later) of Du Puy el lll
0995b) and Lewis & Schrire 0995) and moved Le11111roplsrm1 from the
Caesalpinia group to the Peltophorum g1oup of ttibe Caesalpinieae; Du Puy &
Rabevohit1 a in Du Puy et at. (2002) commented on the morphological similarity
between Lemt11'opis11m and Delon ix, a 1elationship confirmed by the combined
molecular-m o1•phological analysis of Herendeen et al. (2003a); the genus grouped
with Delon ix and Co/vii/ea in the molecular arntlysis of Haston et al (2003)
Young (us ually raw) seeds of L. ed11/e H.Perrier provide human food; great
potential as a food cl'Op for arid lands although mature seeds contain toxins
Pachyelasma Harms 1913
1 sp ; W Africa (Cameroon, Gabon, Equatrnial Guinea, Nigeria and Congo
[KinshnsaJ)
From pacby- (Gk : thick) and e/asma, e/asmos (Gk.: plate) , alluding to the stmll
woody r od
T1 ee; rropic:.i.J everg1 een 1ain forest
Rererence(s): Aubreville 0970: 321-323)
Placed in the Dimorphandra group of the Caesalpinieae by Polhill & Vidal (1981l
and Polhill 0994); in the molecular analysis of B1uneau et al. (2001) Pllchye/llsma
is siste1 to two mimosoicl genera: Pentaclethra and Calpocalyx, suggesting a close
relationship between the Dimorphandr.t group and tribe Mimoseae (as cunently
ci1cumscribed); in the combined molecular-morphological analysis of Herendeen
et lll. (2003a) Pacbyelasma is sister to Erythrophle11111, the two together basal to a
clade containing all Mimosoideae plus Dimo1pba11dra and Mora confirming the
link suggested by Bruneau et al (2001)
Crushed pods of P. tessmamrii (Harms) Ha11ns a1e used as a fish poison,
medicinally and in magic; in W Camernon cn1shed pod extwct administered by
injection is used as an abo1-tifacient and women who wish to miscarry sometimes
sit in fish~p oiso ned wa1e1
Pachye/asma tessmannll Drawing by unknown artist
156 LEGUMESOFTHEWORLD
Erythrophleum couminga Drawing by M. Tebbs
Dlmorphandra gardneriana Photograph by G. P. Lewis
----~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
Erythrophleum laslanthum Photograph by G. R. Nichols
Eryt1'rophl.eum Afzel. ex R.lk 1826
E1ylhrophloe11m is an rnthog1aphic variam
JO spp.; Paleotropics (5 in Africa [I restricted 10 Angola and Gabon, 1 to Mozam-
bique, Swaziland and S Africa, l widespread in \YI Africa, ;rnd 2 throughout much of
Africa, 1 extending to Asia]; 1 endemic to W Madagascar; 1 endemic to N Aust ra lia; 2
in Thailand and Cambodia [I extending into Laos], I in Vietnam and China)
From erythro- (Gk.: red) andph/oio (Gk.: bark of trees), the lnne1 ba1k is reel
Trees; tropical lowland 1ain forest to seasonally dry (and riverine) forest,
woodland and wooded grassland (savanna)
Reference(s): Brenan 0967: 18-21); Larse n et al (1980: 14-19, 1984: 51-3); Lock
0989: 23-24); Ross 0977: 10-13, 15; 1998a : 70-72), Du Puy & Rabevohitra in
Dtt Puy el at. (2002: 53-55)
Placed in the Dimorphanclrd group of the Caesalpinieae by Polhill & Vidal 0981)
and Polhill 0994); sister to the Mimosoideae in the molecula1 analyses of Bruneat1
el al (2001); in the analysis of He1endeen et al (2003a) sister to Pllchye/asma , the
two togethe1 sister to a clade indl1<ling Dimmpbcnulra., Mora and all
Mimosoideae; in the molecular study of Luckow el al. (2003) E1ythroph/e111n is
sister to Dinizia suggesting that the latter is not mimosoid
All species toxic (some very) due to presence of the alkaloid erythrophleine
which causes heart failure; timbe1 of some species used for railway sleepers, boat
bt1ilding :and canoes, heavy construction and joinety, firewood and charcoal; bark
gum used as a tanning agent; crushed bark as a fo;h poison; crushed seeds as a
component of arTow poison; also used medicinally
Di,morpbandra Schott 1827
26 spp.; S Amel'ica 04 in Amazonian Brazil [6 of which extend to C Biazil, the
Gt1ianas, Venezuela, Colombia or Peru], 1 endem lc to Peru, 1 to Venezuela, 5
cent1ed in the Guianas {extending inro N Brazil, Venezuela and Colombia], 1 in E
and NE Brazil, 2 in SE Brnzil, and 2 widespre'1d th1oughout Brazil and extending
Into 130\ivia and Paraguay)
From di- (G k.: two), mo1pho- (Gk.: slu1pe) and tmdro- (Gk.: man or stamen), in
refe1ence to the two types of stamens (5 fe1tile and S staminodial)
Trees; tropieil Amazonian min fore~t on terra firme, periodic<1lly inundated forest,
permanently humid forest , wooded grassland (cerrado), thorn scru b and
woodland (ca1Hinga) and o pen sandy sites in forest
Rererence(s): Silva 0986)
Placed in the Dim01phanclrn group of the Caesalpinieae by Polhill & Vidal 0981)
and Polhill 0994); sister to Mora in the analyses of Bnmeau et al (2001), Luckow
et al (2003) ancl Herendeen et al (2003a), in 1he latter the two are basally
b1;10ching to a mimo.soid dacle including Pentacle/bra and Calpocalyx
The wood of some species (e.g., D exaltata Schott) used in cabinetwork and fo1
tool handles; the fruits of some specie~ highly toxic to cattle (e.g., D . mollis Denth.
and D. gardneriana Tul.); the bark of D . ·mo/Jis used in tanning; seve1al species
used ;is medidne (e.g., for bioflavonoids) and 01 namentals
TRIBE CAESALPINIEAE 157
Moro exce/sa Drawing by P. Halliday Burkea africana Photograph by A. E. Van Wyk & s. Malan
Mora Schomb. ex Ilenth . 1839
6 spp; C America, N South America and the G1eate1· Antilles (2 fiom \V./ Surinam
th1ough G uyana to the 01 inoco delta in Venezueh1 [l extending to Tlinidt1dl, 1
from the Pacific coast of Costa Hica to Colombia fllld N coastal Ecl1<1clor. l f1oin
the de lta of the Amazon in Br.azil, 2 endemit: ro the Dominican Hepublk)
De1ived f1om the widely used Araw;1k vcrmicular name 'Mora'
Ttee!>; t1opic~1l dverine forest, pe1iudically inunda1ed or nut, or swampy a 1cas ,
b1oad-leaved humid forest, 1 sp just behind thl: mang1ove zone, hill slopes
Rcfe1ence(s): Steege 0990: 18-22)
Plci cc.::d in the Dimorphandra group of th e Caesa lpinieae by Polhill & Vidal ( L9Hl)
and Polhill 0994); sister to DimorjJbmulra in the molecut1r analyse:; of Biuncau
el t1/. (2001l, and Luckow el t1/ (2003), ;1nd the combine d molccu1"r-
morphological ;imdysis of l-k1endeen el al. (2003:.1), in the latte1 the two h:.1s:d to a
mimosoid dade including J>enJac/etbra and Calpoca(yx
Mor<1 exce/sa Bcnth and M gonggr(ipii (Kleinhoonte) S<mdwith {mrna,
mcm1bukea [refening only to M gonggr(ipi:il, n:ito, pr<1cuuba) a1e major timlK'I
tree!-i in the Guianas, especia lly used in heavy e<>nst1uction, indwmial florning :incl
frn cha1coal; the .seeds of M megislosperma (Pittier) Britton & Ho.')e are thought to
be the hugesl (18 x 12 cm) dicotyledonous s~ccls known, they a1e a local souice
of a red dye; the seedti or t\1 f!.Yce/sa are also Jmge (up to 12 X 7 <.:Ill)
Burkea Benth. 1843
1 sp,; widespread in Aft ica (except fo1 the foi ~st region.s) extending into Namibia,
Bot-;w~ma ancl S Afri ca (T1ansvaal)
Named for the Briti.')h botanist Joseph Burke (1812-1873), who collec..:tcd pl:1nt'>
:cincl aninrnls (especially in South Africa) frn the Earl o f Derby; he emig1atccl to the
US A. in 18..fH ~ 111 d died in l-la1risonvillc
T1ccs; seasonally d1y ttopictl woodl and and wooded g1e1ssland (buslwclcl)
Reference(s): llrcnan (1967: 21-22); Ross 0977: 11-16)
Placed in the Dimorphanclra g 1oup of the Caesalpinieae by Polhill & Vidal ( 19Hl)
and Polhill 0994), and grouping with Mora in th e molecular :inalytiis of Simp"'on
el al, (2003)
Burkea t!/i·icmw Hool<.. (wild or red syring<1 1 mukarati) yields a strong
c.:onstn1c.:tion timber, the wood also used frn tool h:.u1dles, 1ough c11penl1y ;md Jrn
t:hmc.:oal; twigs used as chew-sticks; bark used in l~tnning :.md as <1 fish slllpdk:i;
foli<1ge toxic tu :tll stock; many medicinal uses including as ;111 antidote co airow
p(>lson
Burkea afrlcana Drawing by L. M. Ripley
158 LEGUMES OF THE WORLD
Stachyothyrsus staudtii Drawing by/. Saussotte-Guerel
Camps/andra sp. Photograph by T. D. Pennington
Sympetalandra borneensis Drawing by M. Smith
Stachyothyrsus 1-1arms 1897
Kaoue Pelleg1. (1933)
2 spp.: W Afril:a (1 in Ivrny Co<tsr, Libe1ia and Sierl'a Leone, l in Came1oon 1
Gabon, Equ;.1torial Guinea and Congo [Kinshasa])
Prom stachys (Gk: spike) :md tbyrsos (Gk.: wand, p<inicle), in reference tu the
spicate inOrnescences agg1<eg<1ted inlo showy panides
T1L--es; seasonally dry tropical forest and wooded g:ras.c;fand (savanna)
Reference(,): Leonard & Yoo1hoeve 0964); Aub1cville (1970: 312 -314 )
Placed in th~ Dimorphandr~1 g1m1p of the Cae.salpinieae by Polhill & Vidal 0981)
and Polhill (1994); as yet not im:luded in any molecu lar analyses
Stacbyotbyr~·•11s stapjlrma (A Chev) J .L6onarc\ & Voorh. used for timber
Sympetalandra Stapf l901
5 spp.: resrrkled to Malesia {Surnatta, Malay Peninsula, Bomco, Philippines, and
Lesser Sunda Islands [Flrnes])
P1om -~vm- (Gk,: rogethe1), petaln- (Gk.: petal) and mulru- (Gki: man rn .'>t<1mcn),
the 10 st~nnen.s joined at their bases to the 5 petals
Trees; lowh1 ncl l1'opical forest
Reference(.<): Steenis 0975); I-Iou in I-Iou el al (1996: 709-714)
Ph1c.:ed in lhc Oimrnpha nd i:.1 g1our of tht.:": C~1esalp i nieac by Polhill&. Vida l (1981)
cmd Polhill 0994); ~1 s yel not induded in :.my molecul::11· analyses
The hark of S scbmulzii Steenis is used as a fish poison
Campsiandra Benth. 1840
19 spp.; S Ame1 ic:.1, mainly in the Amazon and Orinoco basins 03 endemi c to
Ven ezuela, 5 in Venezuela and l3razil [of which 2 extend to Colombi<1, 1 to Bolivia
and l to Colombia, Bolivia ancl Peru]. l 1esriicted to the Guhinas)
frorn campso, camjJsis (Gk .: bending, a bend) and andro- (Gk ,: man, anthel')
refeiring to the long wavy stamens
Trees: tropical rive1ine and swamp foiest (both blac.:k and white wate1• 1ivers) on
:.dluvia l plains, white sand beaches and embankments
Llefcren ce(s): Stergios 0996, ancl in !le11y el t1/., 1998: 18 - 30); Stergios&: Stergios
(1997)
Polhill &: Vidal 0981) and Polhill 0994) included the gemis in the Peltophrnum group
of tribe Caesalpinie<ie .:md 1ecorded only 3 species, a number greatly inc.:n.:·;;1sed by
Stergios 0996) and Stergios & Stergios (1997) who desc1ibecl 15 new species, most
endemic to Venezuela mid some known from ve1y few collec.:tions; the genus w1.1s not
inducled in the molecular analysis of Haston r!l al (2003) nor in the molecular-
mrnphologit:al :.111alysis of Hcrencleen et al (2003a); in [he molec..:ular analysis of
Simpson el al (2003) the single species of CamjJsiandra Included is urnesolvecl with
respect ro siste1 taxa OJ' group position and currently generic..: .iffinities <11e lmknmvn
The set.xis of <I few species are Utie<I loca lly to 111ake a n ucle flou1; also used
rneclicin;il\y
TR IBE CAESALPI NI EAE 159
 Diptychandra aurantlaca Photograph by G. P. Lewis Orphanodendran bernalii Drawing by B. Angell Vouacapaua americana Photograph by o. Daly

Chidlowia Hoyle 1932 Orphanodendron Bameby & J W,Grimes 1990

I sp.; W Africa (Ivory Coast, Ghana, Sierra Leone, Liberia)
Named for the British botanist and silviculturalist Chidlow Vigne (b. 1900), of the
rhen Gold Coast Forest Se1vice, the first to recognise the plant as a new genus
Tree; tropical, often upl:ind evergreen forest
Reference(s): Hoyle (1932)
Placed by Hoyle in tribe Amherstieae (this now part of the Detarieae sens lat)
but moved to the Dimorphandia group of the Caesalpinieae by Polhill & Vida l
(1981), a position retained in Polhill 0994); as yet not included in any molecular
analyses; the wine-red flowers in long pencllilous panicles and large woody
explosively dehiscent pods ate sttiking features of C. sanguinea Hoy le
Di.ptychandra Tu!. 1843
I sp.; C, E and NE Bmzil, Paraguay and Bolivia
F1om diptycho- (Gk: doub led, twice folcled) and andro (Gk,: man, stamen). the
sramen filaments are twice folded in bud
Tree or shrub; seasonally dry tropical forest, gallery fo1est, wooclecl grnsslaml
(savanna, ce1 rnclo) 1 thorn scrub or woodland (c;rntinga), 10cky grassland and
cha co
lte ference(s): Lima el al. (1990)
Polhill & Vidal 0981) and Polhill (1994) incluclecl the ge nus in the Sclerolobium
g1oup of tribe Caesalpinieae, together with Sclero/obi11111 and Tachigali; in the
molecular analysis of Haston et al. (2003), the generic affinities of Dfp1ychm1dm
are unresolved and the genus did not grm1p with Tachigali 01· Sclerolobiutn; D
cwrantiaco Tu!, contains 2 geographically separated subspecies; the pollen is in
tetrads (an unusual feature in the Caesalpinioideae)
1-2 spp.; 1 sp. endemic to NW Colombia, 1 possibly new sp f1om Colombia as
yet und escl'ibed
From 01plx1110- (Gk.: otphan) and de11dro11 (Gk.: t1ee) refening to the ctment
unknown affin ity o f the genus
Trees; tropical lowland forest
Refe1ence(s): llameby & G1imes 0990)
Polhill 0994) p laces the genus in its own Orphanodendron g1oup of tribe
Caesalpinieae; it has not yet been included in any molecular analyses
Vouacapoua Aubl. 1775
3 spp.; Guianas and Amazonian Brnzil
Derived from the local name for V. a me11ca11a Aubl.: wakapu or vouaca pou (used
by the Galibis in French Guiarnt), elsewhere: wacap or arn.pl1
Tree.s; tropicc1 I term Finne 1ain forest
llefe1ence(sJ: Ducke (1949: 130-131)
Placed by Polhill & Viclal 0981) in the Peltophrnum g1oup of Caesalpinieae but
not included in the 'core-Peltopholllrn g1oup' in the molecular <1n<1lysis of Haston
el al. (2003); its generic affinitie• <ire unclear; nrnny b inom i;lis ot iginally published
in Vouacapoua are now in Andira (the ftuits can look supe1fidally si milcu') but
the1e is no close 1elationship between the two genera
Vo11acapoue1 americana (pa1tlidgewood in the N American timber trade) has rot-
1esistenr wood, is used frn house beams and is a valued commercial wood in
Par{t, B1uzil; the wood of all species is used for cabinetwrnk, p<1rquet flool'ing an d
furniture

Vouacapoua americana Drawing by J. A. Lowe

Diptychandra aurantiaca Photograph by G. P. Lewis

160 LEGUMES OF THE WORLD TRI BE CAESALPI NI EAE 161

 oseae byM.Luckow
... ~se;:lle Bronn 1822
"''"'m k.·•..,, .. (\; i"ht & Arn.) En II. (18 0)
1i il?C PSI' 1,,P "
n d h111h crei1c (13enth . 13enth. He> k.f. (1865)
Tribe A en.. -
friht.' Piptad •niea B -nL11. (187 )
Ir ii Je Ml' mose'
ne (sensu Bentham, 1875) is
.
ioed here simply as a matte r of convernence. All
h logenetic analyses mdteate that Ingeae and
nt p yare derived from wit· l110
cie.'I · M'unoseae Cl1app1·11
Maslin, 1995; Kass & Wink, 1996; Luckow et al.
OOO; Bruneau et al., 2001 · Luckow et al., 2003;
erendeen et al., 2003a) , making it a paraphylet1c
up at best. The mosr recent_ studies indicate that
may not even be mo nophylet1c with respect to the
esalpinioicleae (Luckow et al., 2000; Bruneau et al. ,
I; Luckow et al., 2003 . Although the outline of a
new tribal cla si fication o f the mimosoids is
emerging, we await better-supported phylogenies
(based on more extensive data) before formalising
new stable and useful groups . Some parts of the
• assification proposed here are better supported than
~11.ers. Notably, the basal branches in Fig. 24 are
P.©orly supported in most analyses and the
elationships among the groups are likely to change
a we acquire more data. As presently indicated
~ uckow et al., 2003), the type genus Mimosa falls
· thin the derived Piptadenia group which is in turn
isten, and basally branching, to elements of Acacia
·Nill Ingeae (Fig. 24). A more narrowly circumscribed
imoseae sens . strict. will thus leave the bulk of
imoseae sens . lat. (i.e., as treated here) in need of
new tribal allocation.
TI1.t:: most c nspic:uo u liffere n b tween the
lflcatkm pr sent d he re and that of L wis & Elia
1981) is th in lu s ion of tribe Parki a wiLhin
. former wa • circums rib d based on
ncate a . stivation of the ca lyx and was considered
•basal trlb within the Mimosoideae (Elias, 198la).
ck~L phylogen tic analyses ( happill & Nh~slin, 1995;
. He~'''·· 2000; 13run ·'all et cit., 2001; Luckow et at.,
3
'ra . 1 ndee n el al., 2003a), i.ndicat that the two
. in lh Parkieae, Parkia and Pentac/etbra ar
st, l r ~xa (Fig. 24 · P.•n.tceclelhrtt is nested ith.in
<'lSeae m Luckow et cit. 2000 but i eitl1e1· 1·ste1·
ca al · · · ' ·
Cl'encle puu id taxa in Bruneau el al. (2001) a nd
11
h Mi el al. C2003a 1 r part of a I asal polytomy
mosea ·111d
• "esa Ipm1
ca . . id wxa (Luck w el al.,
llrt Mimosa ..
deb1/1s Var V t'
· es ita Photograph by G. P. Lewis
2003). Both Parkia and Pentaclethra are included in
the tribe Mimoseae pending additional data and tribal
rec ircumscription. Recent work (Luckow et al.,
submitted a) also indicates that the monospecific tribe
Mimozygantheae should be subsumed in the Mimoseae
near Piptadeniopsis and Prosopidastrum, currently in
the Prosopis group.
Otherwise, the informal groups within the
Mimoseae recognised by Lewis & Elias (1981) are
rel atively well-supported by current phylogenies and
only a few departures have been made from their
system. Where relationships are either poorly
supported or unresolved, the classification of Lewis &
Elias (1981) is retained. The Xylia group is dismantled
and the Adenanthera group recircumscribecl to include
Calpocalyx and Xylia. Desmanthus has been removed
from the Dichrostachys group, as has Neptunia , in
agreement with recent molecular and morphological
phylogenetic studies (Harris et al., 1994; Hughes, 1998;
Lu ckow, 1995, 1997). A new group is erected to
accommodate Piptadeniastru m which is well
separated from Newtonia in the most recent phylogeny
(Luckow et al., 2000; 2003), and another to
accommodate Cylicodiscus, which is more closely
related to the clacle containing the Prosopis, Leucaena,
Dichrostachys, and Piptaclenia groups than it is to the
Newtonia group. Neptunia is well supported as sister
to Prosopidastrum in recent analyses (Luckow et al.,
2003) and is included in the Prosopis group here.
Relationships of genera in the Prosopis group are not
resolved, but the group is reta ined here as there is no
evidence that it is not monophyletic. Genera newly
described since 1981 include Alantsilodendron,
Calliandropsis, Kanaloa, and Lemurodendron.
Alantsilodendron and Calliandropsis are placed in
the Dichrostachys group , and Kanaloa in the Leucaena
group based on phylogenetic analyses (Hughes, 1998;
Luckow, 1997; Luckow et al., 2000). Lemurodendron
is tentatively included in the Newtonia group as
suggested by Villiers & Guinet (1989). As treated here
the Mimoseae comprises 40 genera and from (859)-
869 - (879) species.
TRIBE MIMOSEAE 163
TABLE 6 Suprageneric groups currently recognised in Mimoseae
Ptptadenia group
Dinizia group SAmerica Dinizio
Anadenanthera .
Pentaclethra group Africa, tropical Sand C Ameri ca Pentaclethra P§eudoplptaden1a
Plptadenia
Aubrevillea group Africa Aubrevillea Paraplptadenla
Mtcrotoblus
Adenanthera group Africa, Asia, Madagascar Adenanthera Stryphnodendron
Aca cie ae (see page 187)
Tetrapleura Acfenopodla
lngeae (see page 193)
Amblygonocarpus Mimosa
Pseudoprosopis
Calpocalyx
Xylia
Prosopis
Piptadeniastrum group Africa Piptadeniastrum Dichrostachys Leucaena group
group group
Entada group Pantropical, including Madagascar En tada Prosopis
Elephantorrhiza Calllandropsis Leucaena Xerocladia
Gagneblna Schleinitzia Prosopidastrum
Plathymenia group S America Plathymenia Oichrostachys Desmanthus Piptadeniopsis
Alant sllodendron Kanaloa Neptunia Newtonia
Newtonia group Africa, Asia, Madagascar lndopiptadenia
group
Lemurodendron Cylicodiscus Fillaeopsis
Newtonia group group lndopiptadenia
Lemurodendron
Fillaeopsis group Africa Fi//aeopsis Cylicodiscus Fillaeopsis Newtonia

Cylicodiscus group Africa Cylicodiscus

Prosopis group Worldwide, mostly New World Prosopis

Xerocladia
Prosopidastrum Entada
Piptadeniopsis Plathymenia group
Neptunia group
Entada
Plathymenia Elephantorrhiza
Leucaena group Pacific Islands, the Americas Leucaena
Schleinitzia
Desmanthus
Kanaloa
Adenanthera
Dichrostachys group Africa, Madagascar, Mexico Calliandropsis group
Gagnebina Piptadeniastrum
Adenanthera
Dichrostachys group
Tetra pleura
Alantsilodendron Amblygonocarpus Piptadeniastrum
Pseudoprosopis
Piptadenia group Sand C America, Mexico, few Old World, Parkia
Calpocalyx
including Madagascar Anadenanthera Xylia
Pseudopiptadenia
Piptadenia
Parapiptadenia
Microlobius
Stryphnodendron
Adenopodia
Mimosa Dinizia Pentaclethra Aubrevillea
group group group
Dinizia Pentaclethra Aubrevillea
!
l

"6.~4 Dia
gram ofretar h. •
ions 1ps in tribe Mimoseae after Luckow et al. (2000; 2003)

164 LEGUMES OF THE WORLD TR IBE MIMOSEAE 165

Dinizio excelsa Drawing by M. Warwick Pentaclethra macroloba Photograph bys. A. Mori Tetrapleura tetraptera Photograph by G. P. Lewis

Dinizia Ducke 1922 Adenanthera L. 1753

1 sp .; Amazon ian S America (Bra zil, Guycma) 13 spp.; Asia (Sri Lanka [1 s p.1, Indo-China & S Ch ina [2 sppJ, Males ia [6 e ndem ic
Named for Jose P Diniz, a friend of the autho r s p p], Papuasia to SW Pacific [l sp ]), Australia (N Queensla nd, 1 sp ), Madagascar
Trees; t1opica l non-inu nd ated (often riverine) ra in forest (1 sp.) and 1 sp . throughou t tropica l Asia to Mela nesia and Australia, a nd widely
Refe rence(s} Ducke (1949, 64 - 65); Alle n & Allen (1981' 243) i ntroduced in Africa and the Neotrop ics
Appa 1ently closer to the Dimo rpha ndra gro up of Caesalpi nioicJeae than la lbr From adeno- (Gk., gla nd) and antbera (L ant her), in re fe re nce to th e gla nds on
Mimosoideoe (Luckow el al, 2000; 2003) th e anthers
Din.izia excelsa Ducke, (angelim ve rme lho, red angelim , angeli m pedra, pa Trees; tropica l primary and secondary ra in forest (i ncl ud ing swamp forest to
is used for timbe1 (heavy constnicti on, fl ooring , stairs, ship building, posts .:n&I coasta l th icket), and seasonally dry fo rest to o pen woodl and and scrub
railway slee pe rs) and in reforestation Refere nce(s} Nielsen (1992, 165-174); Nie lse n & Guin et (1992); Villiers 0995)
Mos t closely allied to Amhlygonoca1pus and Telrapleuta in the Adenanthera
g ro up (Lucko w et al., 2000; 2003)
Pentaclethra Benth . 1840 Adenanthera pavon ina L. (red sand alwood, red bead tree, saga, carolina) is wide ly
3 spp.; WC Africa (2 spp ), S and C America (1 sp., Amazo nas north to C A cultiv<i ted and natura lised througho ut the tro pics; used as shade t1ees , good q uality
and Caribbean) timbe r (fo r co nst1uction and furn iture), ornamentals (especially for the decorative
Fro m pe11tn- (Gk,, five) and -cleitbro (Gk , bolt), in reference to the five scarl et, glossy seeds used as beads in necklaces), fuelwood, medicine, human food
calyx lobes a nd the five pe tals joi ned at the base (cooked seeds), oil (ind ustri al lu b ri cant), dyes and soap substitu tes
Trees; tropic<ll low land ra in forest (o ften riparian) and secondary VCb.'(!t.'\l
seasonally dry woodland and wooded grassla nd Tetrapleura Benth. 1841
References' Ducke (1949, 70 - 71); Vill iers (1989' 18 - 26); Burk ill (1995' ZSl 2 spp.; W to E Africa (Senegal to Kenya and Tanzan ia, and S to Angola in the
!3arneby in Berry el al. (200 1, 665) I r. Guinea-Congolian and Lake Victoria regions)
Unresolved relative to other African taxa at the base of the 111i mosoid P ')' From tetra- (G k,, four) and pleu ra (G k., 1ibs), re ferri ng to the four- ribbed fru its
(Lu kow N 11/., 200)) .. 1 II<....,.. Trees; tropica l lowland rain forest, second ary thi cket and fri nging forest
10
Collsidcrctl" 11111lli-puq1osc 1re<: \illiltll)lc for :rgroforc.>11 ·; 1~ mfl<i n/W) d Refe rence(s} Bre nan (1959, 31- 32); Vi lliers (1989' 64-67); !3urkill (1995, 263-265)
oll lx:an It~. ugh:r , ow;rla) I usctl for 1inrhcr (l'Onw1r 1Ju11J, flrew~ (( Strongly s upported in a clade with Adenantbera and A mblygonoca1pus, in th e
rncclicinc. O!)re, dyes, oils, as so;1p suh.~tltut ', rtsh p<J!SOn.•, 111 "'"'° ~ fQCK" Olll Acle nanthe rn group (Luckow el al., 2003)
seeds) and orn:oment:ol:>: the mrongly elasti ll'J<il~ ;or · '"'~I ''·' sol~ or Tetrapleu ra tetraptera (Schumach. & Thon n ) Ta ub (a ridan, enzieze, akpa) is t1sed
lirnlJCr of/~ lllf/Cl'Olo/x1 (Wllld.) Kum1.c (1tm•llfo , pam'1·l'°Xil b U6CU as~ fo r tim be r (construction and genera l ca rpe ntry), numerous medicinal purposes,
fo r mahoga ny fish, mo ll usc and p m tozoan poiso n.s, human food (seeds and pulp of the two soft
wings of the fruit [ot her two w ings ha rd]), condi ments, firewood and ce remon ies
(due to its magica l-medici nal properties)
Aubrevillea Pellegr. 1933
2 spp.; we and \YI Africa ""'' Amblygonocarpus Harms 1897
Named fo r Prof. A. Aub reville (1897-1982), a noted French foreste r, c
I sp.; Africa (Gha na and Nigeria to Sudan and south to Angola and Mowmbique
taxonomist . . udan ian \\'00' in the Sudanian and Zambezian regions)
Trees; tropical rain forest and seasonally dry forest ou tliers ln S
Fro m a111bl~ (GL bl unt), gon ia (G k., ang le/co rner) and ca1pos (GL fru it), in
"nd wooded 8""' lond
re fe re nce to the b lunt-edged fru its
Hcference(;;), Villiers (1989: 3• - 38): !3urki ll (19'JS: 219 -UOl sidered ~
NOt yet sample<! for mok'<."1:11.rr :iru1lysis, b ut l.cwis & Ella> ( IY81) co n_ basal
Trees; seasona lly d ry tropical forest, wood land and wooded grassland
111<1S1 gcm': r.1 llscd nnd r.:r lhtr Isolated in th e tllhc: it is pl~c-'<I here
05 Reference(s} Brenan (1959, 32- 34); Ross (1975, 132- 133)
a
See notes under Adenanthera for re lati o nships
secti on of Mimoseae
A mblygo nocaipus andongensis (Oliv .) Exell & Torre is used for timber (joinery
Used as a shade tree , for tirnber and me dicine
and furnitu re), charcoal <md firewood, med icine, fami ne food (cooked seeds) 1 as a
Aubrevillea kerstingii Drawing by P. Halliday fish poison and in sorce1y

166 LEGUMES OF THE WORLD TRIBE MIMOSEAE 167

Pseudoprosopis fischeri Drawing by 0. Milne-Redhead Calpocalyx dinklagei Drawing by C. Speight
Pseudoprosopis Harms 1902
7 spp.; WC and W Africa (3 spp. Gabon, Congo [Brazzaville] and Congo
[Kinshasa]; 2 spp. Guinea, Sierra Leone and Liberia), 2 spp. SE Africa (Tanzania,
Zambia and Mozambique, one each in the Zambezian and Zanzib<ir-Jnhambane
regions)
F1ompseudo- (Gk.: false) and Prosopis, a rebted genus of mimosoid legumes
Shmbs, small trees or lianas; tropical 1ain forest, gallery forest, seasomdly dry
forest and dense thicket
Reference(s), Villiers 0983); Brenan 0984)
Strongly supported in a clade with Xylia and Calpocalyx (Luckow el al., 2003), in
the Adenanthera group
Used for fibre and fish poisons
Calpocaryx Harms 1897
11 spp.; Africa (9 spp., in WC regions, mostly in Cameroon and Gabon and 2 spp.
westwards to Siel'I a Leone)
From calpo- (Gk., um), for the um-shaped calyx
Trees; tropical rain forest (often riverine and littor<il)
Refercnce(s): Villiers (1984)
Related to Xylia in the Adenanthera group
Used for timber (construction, canoes, implements and firewood), edible seeds
(after cooking), vegetable salt (from bu int ashes) and medicine
Xylia Benth. 1842
9 spp.; Africa (1 sp. each in Wand WC regions, i.e. Congo [Kinshasa]; SE Africa,
3 spp. in Zanzibar-Inhambafle and 1 sp. in Zambezi<in regions); N Madagilscar CZ
spp.); S Asia (1 sp.) but introduced in E Africa, lndo-China and Malesia
From .\')'lo- (Ck" wood). in reference lO i1s use :is a 1Jmlx:r 1r'"'
Trees; uopiCnl mln frn'Os and riverine forest , scasormlly dry foresl, wcJQ<llnnd m!ll
bushland
Reference(s): 13renan 0959: 29-30; 1970: 33-35); Villiers in Du Puy eta! (200 2:
194-197)
Placed in a strongly supported clade with Pseudoprosopis and Calpocalyx
(Luckow et al., 2003), in the Adenanthera group
Xylia xylocmpa (Roxb.) Taub. ( ~ X. du /(lbrtfi>rmis 13enth.), commonly known as
pyinkado, pynkado, irul, sokram, ea rn xe or ll urmese ironwood, was one ot the
most important timber trees in Myanmar [Bu1"ma] (pmticula1 ly for railw<iy
sl<..~p<......S) ; other timber uses incl\ld e c:onstniction, floo ring, boat building,
fo rnim ~ . posts and wheels; this and other species are also used for fuel wood,
medicine, soap, human food (ro<isted seeds), tannins and oil (from seeds)
Entad0
Xylia fraterna Phatograph by G. P. Lewis
168 LEGUMES OF THE WORLD
Plptodeniastrum ofricanum Drawing by E. M. Stanes Entada polystachya Photagraph by G. P. Lewis
Piptadeniastrum Brenan 1955
1 sp.; WC AfriGJ (Senegal and Sudan to Uganda, south to Angola)
From -astrum (L.: incomplete J'e sernblance to) and Piptadenia, <inother mirnosoicl
genus which it pattly resembles
Trees; tropical lowland rain forest and seasonally dry forest (often riparian)
Reference(s): 13renan (1959: 21-22); Villiers 0989: 57-61); Bu1kill 0995:
255-257)
Clusters with the Entada group in the analysis of Luckow et al. (2003) although
this relationship is not strongly supported
Pipladeniastrum africanum (Hook .f.) Brenan (Af1 ican greenhe<itt, t<11ahuilca,
clabema, dahoma, ekhimi) is used for commercial timber (heavy construction,
floofiflg, decking, vehicle bodies, carpent1y and mine props), charcoal, medicine
(although also toxic), fish and rodent poisons and it has spi1·itual uses
Entada Adans, 1763
Entadopsis Britton 0928)
c. 28 spp.; p<rnt m piG1l: Africa with 16 spp., predominantly in Sudanian and
Zambezian regions, two extending to Madagascar; 3 spp . endemic to Madagascar;
5 spp. endemic to Indo-China and Malesia; 1 sp. in Neotropics; 3 spp. widespread
and dispersed by ocean currents (E. rheedii Spreng_ throughout the Palaeot1 opics;
E. phuseoloides (L.) Me rr. in Asia, Australasia and Pacific and E. gigas CL.) Fawcett
& Rendle in Africa to Neotropics)
Confusion exists as to whether Enlada originated f1om the indigenous name fo r
the plant in Mal::1ba1', India or from dentado (Po1tuguese: toothed), referring to the
prickles on the stems and leaves of some species
Trees, shnibs and lianas; tropical lowland and riverine rain forest, seasonally dry
forest, woodland <lnd wooded grassland, bushland, thicket and d1y scn_1b
References: 13renan 0959: 9-21; 1966); Lock 0989: 91-92); Nielsen 0992:
176-181); Lungu 0993); Barneby 0996); Villiers in Du Puy eta/_ (2002:
163-169)
Strongly supported as grouping with Elephantorrhiza in tl1e Entada group
(Luckow et al., 2003); the genus is much in need of revision <lS it is highly diverse
in habit, frnit size, intlorescence architectu re, presence or absence of tendrils <ind
armature; anther glands occu1 in all species except those in M;;1clagascar
Various species (S t. Thomas bean, matchbox bean, drinking vine) used as
livestock Fockler, ground cover, green manure, fibre (in 1ope and storage bins),
medicine, fish poisons, soap substitutes, firewood, charcoal and in tradition<il
ce remonies; the large round drift seeds of the species listed above emerge f10111
some of the biggest pods in the Leguminosae (over one metre in length) ancl <ll'e
often used to make jewelle1y ('sea hearts')
po/ystachya Photagraph by G. P. Lewis
TRIBE MIMOSEAE 169
 Lemurodendron copuronii Drawing by J. -F. Villiers Newtonia buchononii Drawing by E. M. Stones
P/othymenla reticuloto Photograph by G. P. Lewis

Elephantorrhiza Benth. 1841 Lemurodendron villieos 19s9

1 sp ; NE Madagascar (narrow endemic SSW of Vohemar)
9 sp p.; Afr ica S of the equator (Zambezian and Kalah•ri-l llghveld Regional
From 'lem ur' and -dendron (Gk : tree)
Cencres of Endemism; ce ntre of diversity in southern Afrlcn}
Trees; lropical mi xed evergreen and deciduous forest on igneous rock
From elepbas (Gk.: e lephant) and -rbiza (Gk.: root), re fe rring to the large storage
Refe rence(s): Villiers & Guinet (1989); Vill iers in Du Puy et al (2002: 160-161)
roots present in some species Placed in the Newto nia group, po.>sibly allied to Fi/laeopsis (Villiers & Guinet, 1989)
Trees, shrubs a nd suffrutices; tropical and s ubtropical wood land, bushland and
thi cket, shrubland a nd grassland; often in open rocky areas
Reference(s): Hoss (1974; 1975: 138-150)
Well suppo rted in a clacle with Enlada (Luckow et al., 2003), in the Entada group
Newtonia Bail!. 1888
The roots of various species (elands bean, intolwane) a re a major source of 15 spp.; Africa (11 spp. in WC and W regio ns, 2 spp. extending to Zambezian
regions; 3 spp. in the Sornalia~Masai, Zanzibar-Jnhambane and Zambezian regions
tannin s for tanning leather, and dyes; also used for medicine
of E and SE Africa)
N~med in honour of Sir Isaac Newton (1642 -1727), English mathematician,
philosopher, and scientist
Plathymenia Benth . 1840 Trees and lianas; tropical rain forest, both hig h and low elev<itions, riparian and
I ~p.: S Amurtcn (Or.1211, l'm••guoy, E Bolivia, :incl 1\Jl!Cntlrin) swamp fo rest; littoral oi sublittoral deciduous fo rest, woodland and bushland
P10111 /ifn1.1~ (Gk .; br(Y.1dl :11ltl by111<111 (Ck.: memhmm:). in reference 10 1h
Reference(s): Villiers 0990); Mackinder & Cheek (2003)
111 !111bmnou~ cndt><'lrp seed p:tckuges within the lcoibery c xoc:trp vo h·es
Lewis & Elias 098 1: 161) place this in the Newto ni a group with Cylicodiscus and
Trc : :«.-:t:;On:illy dry ll'O(} <'lll roic.i (~ ml S(;(.'()ncbry v~~Cnttion derived from moist
Indopiptadenia
fore-'1; W(toclland (c•=dn), rocky gr.osslnnd (L'll111po n 1pes1rc), bushlund oncl
Various species used fo r timber (e .g., N. bucbananii (Baker) G.C.C.Gilbert &
scru b) Boutique, lokundu 1 o rnutoyo) in construclio n, cabinetry , frames, joine1y, railway
Reference(s} Heringer 0956); Waowick & Lewis (2003) sleepers and canoe making; the bark of at least one species has reputed
In the Plathymenia group and somewhat isolated f1om orher mimosoid genera
aphrodisiac properties, also used for medicine and as shade plants
(Luckow er al., 2003); Warwick & Lewis (2003) conclude the genus is
monospecific, containing the ~ingle species P. reticu/atCl Benth.
Used commercially for [imber (vinhatico) in tl ooring, construction, posts and
furniture; also fo1 reforestation, shade trees, firewood and medicine
Fillaeopsis Harms 1899
1 sp .; WC Africa (Nigeria, Cuneroon, Gabo n, Congo [Kinshasa), Angola)
From apsis- (Gk : appearance) and Fitlaea, genus of legumes (now a synonym of
Etytbrophleum in Caesalpinioideae), so named for its resemblance to Fillaea
Indopiptadenia Brenan 1955 Trees; tropical lowland rain forest
I sp.; S Asia (India and Nepal)
Re fe rence(s): Villie1s (1989: 38-41)
From Indo- (denoting India) and Pipradenia (another genus in Mimoseae),
Placed by Lewis & Elias (1981) and Luckow et al. (2003) in the monogeneric
referri ng co the superficial affinities of the genus
Fillaeopsis group, sister to the Newtonia group
Trees; subtoopical forest and secondary vegetatio n in hills; 300-600 m
Fillaeop sis discopbora Harms (arbre a seme lle) is used for timber in constmction,
Refe rence(s): Biswas & Chandia 0997) flooring, f1 ames and for underwater construction; also used as a poison and for
Placed in the Newtonia group fo llowing Lewis & Elias (1981: 161); poorly
medicine
n:prcscntcd In 1110>1 h<:rb:orla
/11dop1~//tul1111f(I 0111/be11sl.< (Brandis) Brenan is cnd;1ngcred due to over-
c.,1llol1:orion for tlntb"r. II i> also used as livestock f0<klcr and medicine

Fi//oeo Psis
. d"tscophoro Drawing by P. Halliday
/ndopiptodenlo oudhensis Drawing by P. Halliday

TRIBE MIMOSEAE 171

170 LEGUMESOFTHEWORLD
 Xeroclodia viridiromis Drawing by P. Halliday Prosopidostrum globosum Drawing by P. Halliday
Prosopis polmeri Photograph by G. P. Lewis

Cylicodiscus Harms 1897 Xerocladia Haiv. 1s62

Prosopis ju/i{loro Photograph by D. Du Puy
1 sp~; southern Af1ica (Nama-Ka100 region in S Af1ico and Namibia)
1 sp.; W ancl \'(fC Africa (Siena Leone to Ivo1y Co::is t, Ghana, Nigeria, C<11ncroon
l'rorn xero- (Gk ,: dry) and c/adfon (Gk .: branch), referrin g to the habit of
and Ga bon)
photosymhetic bi•anches and tiny lea flets
Frnrn cylico- (Gk.: cup-s llaped) and discus (L.: clisk), in re feren ce to the Cllp -
Shrt1bs; semi-arid g1assy dwarf .shrublancl <incl thorn scrub , in sandy ii ver beds, on
s hapcd d isk in the flower to which the stame ns me attached
river banks, allu vium and saline flats
Trees; trop ical rain fo1 est and se~1 sonal l y dry foresl
Refer ence(s): Ross (1975: 130-1 31)
Referencc(s} Villiers (1989: 41- 45); Burkill (1995: 223 - 224)
Xeroclatlia vtridiramis (Bu rch ~) T~ub. is an isolated ende mic taxon in S\V Africa,
T1 e~1ted here as belon.g ing to its own group based on the anal y~ is of Luckow el
placed by Lew is & Elias (1981 : 164) in the Pmsopis gro up ancl sister to Pwsopis
al. (2003) which places it in ~in isolated position, more closely related to rhe
(Luckow, unpublished data)
LcuG1ena, Pl'Osopis, Di chrosta chys ;rnd Piptad cnia g1oups
Timber of C gabunensis J-b1 ms (okan, azobi, cl enya, adoum) is very dense and
used in heavy constru ction, Ooo1ing, rJilway sleepers and implements; Hlso used
J S charcoa l, medicine and a soap subscitute
Prosopidastrum Burka rt 1964
c. 5 spp.; N and 5 America , disjunct between Ivlexico (1 sp . Baja California) and
Prosopis L 1767 Argentina ( 4 spp.l
S1ro111hocaipa ( Benth.) Engelm & A.Gray (1845); Sopropis Britton & Rose (1928) From -as/rum ( L.: incomplete resembhmce to) and Prosopis, ano ther mimosoid
c 44 spp .; S Arner ica (31 spp. ce ntred in Argen 1ina, Chile, Paraguay Lo E Bolivi<J , genus whi ch ir resembles
Urugu ;:1y ancl S I31azil with a minor ccnt1e in the t1 o pical Andean regio n from Shrubs; subt 1opica l xerophytic bl1 sbland,_thicket, gras.s lancl ancl se111i-dcse1t
Pe1u, Ecuaclrn• :md Colombia; 2 spp- wiclc sp1'e;:1d in Neotropics; 7 spp. in Referencc(s): Burkart (1964); Pala cios & Hoc (2001)
Mex ico -SW USA); SW to C Asia (3 spp.), one exte nding to N Afri ca; Africa (I Placed in a clade with Neptwzia (Luckow el al., 2003) ~md as siste1 to
sp in Sudanian regi o n) ; cu ltivate d and n <Hura lised woilclwide Piptadeniopsis and Mimozygantbus (Luckow et al., submitted a); it is thu s
Possibl y from prosopon (Gk.: face), <m ancient rnune for the burdock, Arcti11111 lapjJa provisio nally plGi ced in the Prosopis group (as it w as in Lew is & Elias, 1981)
L , btll the meaning is obscttte. Pasiccznik et al.. (2001), howeve1 , give the 0 1igin
c~s pros- (Gk.: tow~1rds) and Opis (wife of Saturn , the G reek goddess of abund;ince
and ag1iculrn1e), hence 'towards abund;ince' for its wide ;:incl ancienl usage Piptadeniopsis Burkait 1944
T1 ees and sh11-1bs; tropica l to wa1 m tempcrnte season<1lly cl1y forest, wo ocll:rncl, 1 sp.; S America (N Paraguay)
woodecl grassland, serni-xerophytic woodh1ncl ancl sh1 ubland, tho1 n scrub ancl From -opsis (Gk.: appearance) ;rnd Piptadenia, another genus of mimosoicl
cleserr, o n .s;rndy plains or hills, ravines ;ind alo ng clry stre1rn beds legumes
Heference(s): Burkart (1976); Simpson 0977); Pedersen & Grainger (1981); Shn.1bs or small t1ees; sl1btropicll seasonally dry tl101n fo rest, thicket and scrub
Poynro n 0990); Pasiccznik el al. (2001) (cha co)
\Vhil e generic limits and the div ision into 5 sections are geneially accepted , Referencc(s): Burkart 094 4)
cl ebme co ntinues ove1 the relative rnnk of species; Bu1ka1t 0976) rnised man y P1ptade11 iopsis lomentifera 13urka1t is siste1 to Prosopidaslrum and
earli ei• infr::1specific taxa Lo species level and so me species complexes ::incl Mimozygcmtbus (Luckow et al., submitted a) ancl it1 thu s provisionally pla ce d in
hybridisation still <.:<rn.se taxonomic problem s. Pl ~i ce d here in the Prosopi s group the Prosopis g1oup (as it w<is in Lewis & Elias, 1981)
and pooi ly suppoitecl <:lt the base of the Dichrostachys <ind Piptad enia grours in
th e analysis of Luckow et al. (2003). Altbot1gh thi s nnd other genera in th e
Prosopis g1oup are not 1esolved c1s rnonophyletic by molecular darn (Luckow et
al., 2003), the Prosopis group is retained here pending frn1her sn1dy
Used since ancient times as human food and drink ( f10111 the sweet ~leshy pocls),
timber (e.g., ror constructi on, fence posts and furniture), livestock foclc.ler,
cha1 c0<d, foewoocl, .shade ti ees, ornamental s, gums, dyes for t<1nnin g, med icine.
co ffee subst itute, bee ph111ts <i ncl desert refor es tation; major speci es ru e P.j11lijlora
(Sw.) DC. ancl P pa/Iida (Humb & Bon pl . ex Wi llci .) Ku nth, known as mesquite.
:Jlga 110b::i and screw bean; some species are invasive and pernicious weeds Piptadeniopsis /omentifero Drawing by P. Halliday

TRIBE MIMOSEAE 173

172 LEGUMES OF THE WORLD
 Leucoena trichodes Photograph by G. P. Lewis
Neptunia Lou r. 1790
12 spp.; Aust1alia (4 endemic spp.), Papuasia th1'ough J\falesia lo I11do-Clil11a (2
spp.); India and Sri Lanka (l sp .), l sp. pantropiGd; New \'V'odd from S USA (1
spJ , Mexico, C Arnei iG1, Cu ibhein south to Paraguay and N Atgentina (3 spp.,
one also introclucecl in S Asia)
Named fo1· the Rorn~rn wate1• deity Neptune, referring to the aquatic habir uf
several species
Herbs; tropical to '\V~lrm temperate oren woodland, wooclecl grassland ~ind
gi-assL:mcl, floodplains, swa111ps and othcJ" wet a1eas
Rcfcrcncc(s} Windler (1966); Wiwller' & Windler• 0974); Nielsen (1992• 186-190);
Cowon 0998• 21-26)
Placed in a cl;1cle with Prosopidastru111 (Luclw'lv el al., 2003) and so inducled in
the Prosopis group
NeplHJlicJ uler{fceo Lour. is often ;1 troublesome weed (with sensitive leaves): usecl
for medicine, human Coocl (young stems, leaves and pods are eaten) and ~t.s soil
binders; produced ;is a cash cmp in SE Asia
Leucaena !lentil_ 1842
22 spp. ; Ml'xico (10 endemic spp. with 2 extending to S USA ['l'CX<lS :111cl Ne\\
Mexico] and 'f spp. to C America); Ii endemic spp. in C A111e1ic:1; 1 sp in S
Americ:1 (N and \'(! co;:ist;:il regions S to Pe ru); 1 S[J . pantropical
From leucnino (L: becoming white), prolx1bly in refe1ence to the devclopil1g
flowe1· colou1 of most species
Trees and shrubs; t1opical and subtropirnl se;:isonally dry fo1est, semi-aiid tllom
scruh forest, to w;:irm temperate open habitats
Rcfcrence(s). Zaute 0994); Hughes 0998)
Pl;:icecl in a well suppoited cl;icle sister to Scb/einilzfu, Kmwlon ;:incl 1Jes111d11tl1115
(Luckow el di., 2003; Hughes el al, 2003), in the Leucacn;J g1oup
Le11coe11a le11cucepba/d (Lam.) de \'V'it (leuc;:ienJ, koa haoJe, jumbie bean, guajc,
i[)il-ipil) is cultivated pantropically a11cl has become natu1alised and weedy ill
many areas; this and other species ;:ire used for livestock feed, green manu1c,
timber (for construction, firewood and charcoal), s111~1ll 'ivood pmclucts, soil
conserv;1ticm (ground cover ;:incl reCorestMion) and human food (un1i pe pods
ancl seecls)
Leucaeno trichodes Photograph by G. P. Lewis
174 LEGUMES OF THE WORLD
Schleinitzia insularum Drawing by
Desmanthus bicarnutus Drawing by B. King
H. Lamourdedieu Desmanthus o/igospermus Photograph by G. P. Lewis
Schleinitzia Wmb ex Nevling & Niezgoda Cl978)
4 spp.; rvlalesia and Papuasia (Phili[)pines [1 sp], Papua New Guinea, Ivioluccas,
Solomon Isis ., N V;rnuatu [1 sp .]); SW & SC l\icific (S Vanuatu, E to Society Isis_,
New CalecloniJ, Piji, Tonga, Niue and Cook Tsls. [1 sp.]); :C--,1\Xi' Pacific (SE Guam
:111d adjacent Mari<inas [1 sp.])
Named in honow· of Vice Admi ral George Schleinitz (1831-1910), governor of
German New Guinea (Papu<l New Guinea)
Trees ancl sh11.1bs; tropical secondary rain fore.st, woodland, wooded grassland,
coast~1l phtins and beaches
l(ei'erence(s), Nevling & Niezgoda 0978); Guinet & Nielsen (1980); Nielsen 0992•
190-193)
ln a well supported clacle with Kmw!oa and Desmanth11s (Luckow el al., 2003;
Hughes el al~ , 2003), this clade is sister lo Leucaena, in the Leucaen;:i g1 'o up
Usecl for wood (ceiemonial Ci1ewoocl for cremations, fishnet frnmcs and
h;:indic1afts), medicine (from the bark) and soil imp1ovement
Desmanthus w;11c1 . 1806
c. 24 spp ~; USA to Mexico ;:incl C Ameri ca (14 spp. in Mexico [7 endemic]; 3 spp-
endemic to USA); S America (2 spp ~ endemic to SE 13razil, Pzm1guay, N Argentina;
2 srp [and one info1speciCic va1iant] disjunct between SE USA-E Mexico aml the
SE Bfazil, P:11~1guay, N A1·gentin<1 region); 3 spp. more widespread in the New
\'V'orlcl including the Caribbe<lll, but absent l"rom the Amazon Basin
Fmrn des111e (Gk.: bundle) and a!ltbos (Gk: flower), in reference to the dense
c1pitate inflorescences
Shrubs ancl herbs; tropic;:il to subtropical sea.son<1lly cl1y forest, woodland,
gl°assland, thorn sci'uh ::ind thicket, along w::1te1courses, in wetlands, rocky or
gr·avelly areas and old pastures
Reference(s} Luckow 0993)
Strongly surpoitecl in a clacle with Scb/einilzin and Kmzaloa by Luckow et al.
(2003) and Hughes el al. (2003)
Desma11tbuspe1·na111b11c:a1111s (L ) Thell . (Jong misinteipretecl as D. vi1go/11s (L)
\Xi'illcl . [which is nevertheless still a good species]) is a wiclespre;:icl p:intropical
weed; genus ;:ilso used JS om;:imentals, cattle Fockler <md in erosion control; D ,
il/i1wensis (Michx.) IvlacMill. (Jllinois or piairie bunclleflmver, prairie mimosa,
spicler bean) is used for human food (lea\'es, cooked seeds), medicine and is a
potenti<1 l rulse crop
TRIBE MIMOSEAE 175
Kanaloa kahaolawensis Drawing by A. Asquith Calliandrops/s nervasus Photograph by c. E. Hughes Olchrostachys akataensis Photograph by G. P. Lewis Dichrostachys cinerea Photograph by 5. Collenette

Kanaloa Lorence & IC.R.\Vood 1994 Dichrostachys (Dc.J Wight & A1n . J834
1 sp.; Hawaii, ende mic to the S coast Mak::1wao District of the isl.and of 14 spp.; Madagasc.11 (centre of diversity wi th 9 endemic spp .); 1 sp. each in
Kaho'o la we Ethiopia- Sornali:-i and Socot1.a; l sp. in Tndi<1 ; J sp. in N Au:-;t1alia, 1 s p . (D.
Named for the Hawaiian deicy Kanaloa, w ho was closely associa ted w ith the cinerea (L.) Wighr & Arn.), wi<lespre~1d in Afdca and Asia bur namialise<l
Island of Kaho'olawe where he could rest and recoup his ene1g ies throughout rhe tmpic:-;
Shrubs; steep rocky talus slopes a nd coast~tl shrubl~mcl Prom di- (Gk .: two), cbroma (Gk .: colour) ~incl stac/Jys (Gk ,: spike), in refe1•e nce
Refe1ence(s): Lorence & \Voocl 0994) to the bi-colou 1ed inflorescences founcl in the genus
Kanaloa kahoolawensfs Lorence & K.R.\\.'ood is known from only two plants on Trees and sh rubs; tropical moist and se~1 son~1 1ly c.liy foi csr, woodland, wooded
the islan d of Kaho'olawei strongly supported in a clacle with Sch/einftzia and grnss lancl, bushkrnd, thicket and xerophytic scnib
Desmanlbus in the Leucaena group (Luckow et al., 2003; Hughes et al., 2003) Refe1-ence(s} 13rcn;rn & Il1urnmitt 0965); Thulin (1993, 366 - 368), Villie l's in Du
Puy el al. (2002, 206-218)
Jn a well supported clade with Callicmdropsis, Gagnebina <rncl Ala11tsilude11dron,
Calliandropsis H .M .Hern. & r .Guinet 1990 in the Dic hrostachys group (Luckow et al., 2003; Hughes el ed., 2003), and sister to
1 sp.; narrowly distributed in Mexico , in disjunct populations from Du1:mgo :1s for Gagne/Jina in the m01phological an~il ysis of J-lt1ghes (1998)
sout h as Oaxaca Dic/Jrostacbys cinerea (sickle bush) is a widespread ex tremely vari1:1blc species,
From opsis- (Gk.: appearance), rcferriflg to its resemblance to Calliandra, another wirh nume1ous subspecies and va1ieties; it i!'> ~1 111ulti-pu1pose tree, the timber is
g:eous of mimosoid legumes (in tribe lngeae) used for construction, fencing, ca rpentry, firewood ~rnd chartoa l; othcl" uses a1e <lS
Shrubs; tropical dry tho rn forest and ::11 id scrub lands, on 1ocky rnlcareous soils; c. ornamenw l:-;, medicine, livestock fodder (goats), fibre from lhe Lr.11 k (rope), bee
1450-2000 Ill fo1age, hunting bows and for ~oil improvement; it is some1imes an inva.-;ive and
Reference(s), lleinfodez & Guinet 0990) persislenr weed of cultivated land
\Vell supported as ra11 of the Dichrostachys g1oup in the molecular analysis of
Luckow et al, (2003)
Alantsilodendron Villieis 1994
10 spp.; N, \V & S MaclagasG11
Gagnebina Neck. ex DC. 1825 frorn alcmtsilf, the Malagasy name for 'd 1y foresl', ancl de11dron (Gk i: tree)
8 .')pp.; Madagascar (6 endemic spp ), 2 spp. extend ing to sun·ounding Comoros, Shn.1bs or small t1ees; seasonally c.11y t1opi c:1 I wooc\bncl, thorn scrub and thicket to
Seychelles (Akh1b1 a and Astove) and Mascarene Islands xc 1ophyti c s hrubland, on calc.:<1reou s so ils
Named for Abraham Gagnebin de la FeniC1e (1707-1800), Swiss physician ~ind Refe1ence(s): Villiers 0994); Villiers in Du Puy el a l (2002, 198-206)
botanist Genus segrega ted fl'om Dichroslacbys by Vi llie1s 099'1), and closely allied to
Trees and shrubs; se::isonally dry tropica l fo1est 1 woodland and thicket (1 sp in Dicbrostachys, CaRne/Jiua :rnd Cal/iaudropsis in the Oich1ost::1chys g1oup
moist forest and forest mmgins) (Luckow et al , 2000; 2003)
Reference(s} Lewis & Guinet (1986); Luckow (1995: 2002); Luckow & Du Pu y Timber used fo1 posrs in house <..:onslit1ccion
(2000); Villiers in Du Puy et al (2002, 218 - 222)
Gag11ebina is suppo11ed as a monophyl e tic group by Luckow et al. (2003) w ithin
the Dichroscachys group (see note under Dicbrosracbys); VUlie rs in Du Pu y et rrl.
(2002) tre:tt three Madagascan species of Gagnebi1w t1mler Dicbrostachys but chis
is not suppo11ecl in th e analysis of Luckow et al. (2003)
Used for wood (const1u ction and charcoal), also for p<.1pe11

Gagnebina pterocarpa Photograph by D. Du Puy Alantsilodendron ramosum Photograph by D. Du Puy

176 LEGUMES OF THE WORLD TRIBE MIMOSEAE 177

 Anodenanthera calubrina var. colubrina Phot ograph by G. P. Lewis Anadenanthera co/ubrina var. co/ubrina Photograph by G. P. Lewis
Parkia gigantocarpa Photograph by H. c. F. Hopkins Parkia ignei{lora Photograph by H. c. F. Hopkins
Anadenanthera speg. 1923
Parkia R.Br 1826
2 spp; G 1ea(er Antilles through northern S America to Pern, Boliv ia, Argenti m1
c . 34 spp; pantropica l, but w ith three disjunct centres of dive1sity in S Ame rica, and Paraguay; probably introduced in the \'ii Indies
Africa-Madagascar and the lndopacific region; 18 spp. centred in Amazonia (but From an- (Gk.: lacking), adeno- (Gk.: gland) and anthera (L.: anther), in
extending from Honduras in the north to SE coastal B1azil in the souih); 3 spp. reference to the lack o f a gland a1 1he :apex of the anther (although one of the
.
. in WC to SE Africa and 1 sp in Madagascar; c. 12 spp. in Asia from NE India species has anrhers glandular in bud)
(1 sp ), Inda-Ch ina and China (1 sp.), Malesia (c. 5 spp • some exicnd ing to Trees; seasonally dry uopical to subt1opic~ I Jiverine forest and forest margins to
Inda-China), Papuasia (I sp.), M icronesia (2 spp_) and Fiji (1 sp.); I sp. wood land and thicket (ce11i:tdo) nnd wooded gmssland (savanna), from a wide

'
widesp reCld i n SE Asi~ range of ha b itats; ofre11 planted near villages
' Named in honour of Mungo Park (1771 - 1806), Scollish surgeon & t1<1vellcr in
Africa
Reference(s): A ltschul (1964; 1972)
Fonneily treated as section Niopa of Piµtaclen.ia; nested w ith an element of the
Trees; tropica l lm:vland rain forest (ripalian, swamp and non-inundated), hill polyphyletic Piptadenia (i.e. P. virldijlora (Kun th) flenth .) in the analysis of
forest, seasonally d ry fores t, woodla nd (ce1 rndo), wooded g1ass land ~111d coastal Lu ckow el al. (2003)
dune fo1est (restinga) Anadenanlhera peregrina (L) Speg (cohoba, niopo or yopo) is one of the classic
Reference(s): Hopkins 0983; 1986; 1994; 2000a , 2000b); Vill iers (1989: 26 - 33); hallucinogens of the Arnc1icas, lttken as snuff made from the crushed seeds; A
Burki II 0995: 244 - 252); Hopkins in Beny el al (2001: 659 - 664) co/11bri11a (Veil .) B1-cnan (angico preto, cebil , curupay) is used for timber (heavy
Of the three sections in Parkia only section Parkia is pancropicnl, the 01hcr cwo COOSlruct io n , Oooring, l<tihvay sleepers, turnety) and leathe1- tanning
being 1estricted l o the Ncolropics~ All 1ecen1 phylogenelic analyses indicate tha(
Parkia is most closely relaced to members o f the Pipcadenia group, :rnd is no( a
basally branching genus in the Mimosoidcae Pseudopiptadenia Rauschen 19s2
Parkia speciosa Hassk. is used as human food in SE Asia; man1re and slighdy
Monoschisma Brenan 0955); Newtonia sect Neonewlonia Burkart (1979) (=
immature green seeds ate e<Hen as a vegetable (peta i, sa 1aw bea n, chou dou),
'American Newtonia' of Lewis & Elias (1981))
sold fresh as bunches of s11-ap-shaped pods in markers and as pods or loose
11 spp; S Amer ica, centred in NE to SE Bmzil (c. 7 spp, " t least one to Paraguay),
seeds, enten fresh or tin ned, from supermarkets; seeds are also ferm ented and 1he
4 spp. in northern S America (to Peiu), one wic.lespl'e:.id to C America
pulpy endocarp makes a refreshing drink; other species are used extensively ~s
From pseudo- (Gk : fa lse) and Piptadenia, a re lated genus in M imosea e
food in W Africa (e.g., P. b/gltl//o.'ifl Qacq.) G.Don, or ncere, African locus t bean);
Trees and shru bs; tropical coa st<ll iain forest, galle1y fores t, seconcla1y forest,
also used 8.S Cfl ltl e fockler, cordage, shade tree.s, med icine Hncl th e timbc t in
wood land , wooded grassland (cerrado) and rho1 n scrub (caa tinga)
plywood mant1facrure 1 construction, utensi ls and as firewood
Reference(s): Lewis (1991a, c); Lewis & Lima (1991); Grimes 0993)
Placed in the Piptaden ia group and considered to be allied with Pa rapiptadenia
by Lewis & Elias (1981: 157)
Pseudopiptadenia pittie1i (Hal'ms) G J>.Lewis is used for timber (carbone10) in
heavy consl ruction, furnitu1 e, flooring, posts, llll nery and ra ilway sleepers; tannins
are extiaclccl from the bark

Pseudop1ptadenia
· brenanii Photograph by G. P. Lewis
Parkia bicolor Drawing by I. Zewald

TRIBE MIMOSEAE 179

178 LEGUMESOFTHEWORLD
 Piptodenio flovo Photograph by G. P. Lewis Parapiptadenia ilheusana Drawing by A. J. Beaumont Parapiptadenia pterosperma Photograph by G. P. Lewis

Piptadenia Benih. 1840 Parapiptadenia lllen•n 1963

Pil1yoca1pa Britton & Rose (1928)
c , 24 spp.; S Ameii ca with centres in Amazonia linking to the Atlantic foiests of
Brazil (c. IO spp.), NE & E Brazil diybnds with extension to Venezuela (c. 1 spp.,
one exrencling to C Am e1icCJ and Mexico); sub-Andean drylancls in Pern , Ecu:.iclor
and Bolivi" (c. 2 spp .); SE Brnzil (c. 4 spp ) ; 2 spp. widespfead in drylands f1om
Mexico and C America to Argentina
From pipto- (GL fall) and ade110- (Gk .; gland), for the caducous glands on the
nnthers
Trees, shrubs (e1'ec t anci scanclcn t) and li ~ma s; rropical lowland rain fo1est, co;1st<1I
and ripJTi<m forest , 1ropical to subtropical season<1 lly cl1y forest, scco11cla1y forest,
woodland, thorn scwb (caatinga) and 1ocky wooded grassh1ncl
Reference(s} Barneby & Grimes Cl984b); Barneby (1986); Lewis 0 991 b); !3;11
in Be1ry et al (200 1; 665 - 667)
Genus much in need o r revision and new t ~1xa await description Pol yphylclic in
mosr recent molecula1 analyses (Luckow et at., 2003); the type was not included
in th e ~:m<ilysis so its position within the genus is nor clea1; present ly placed in the
Pip1<1dcnia group, wheie two species form a st1ongly .suppo1tecl clacle with
Parapiptadenia, SllJjJbnodendron and 1l1icro/obius, in the :m.:ilysis of Luckow et
al. (2003)
Various species u.')ed fonin 1be1 (surucu\'.u, icarape, soroc;:1, jurema prer:i, pau
jaca re, angico de bcze110) for constructio n, posr.s, tool handles, ca rv ing ;incl
firewood; tannin~ a1e extracccd from che lx:irk
6 spp.; S America (SE & E Brazil, Paraguay, Uruguay, and Argentina)
Prom para- (Gk.: near) ~incl Pipwdenia, a related genus in Mimoseac
T1ees ancl sh1ubs; tmpical coastal nncl dune forest (restinga), seco nda1y growth
forest, woodland and scnib
!leference(s); Lima & Lima (1984); Lewis 0994)
See note on relationships un<ler Piptndenia
All species are used as commercial timber (chari , angi co, anchico colora<lo ,
kurnpa'y ra) in high quality furniture and construction (e.g ., P. rigida (Denth.)
Bremm); also used for gums, a mucilage constituent of m edicines and in
agroforestry
neby
Microlobius c Pres! 1845
Golt/111a11ia Rose ex Micheli (1903), pro pa11e
1 sp.; disju nct in \Y/ ~ind S Mexico to Hondu ra s, and in S 13rnzil , Pa1aguay :-tncl
Argentina
Fmm micro- (Gk.; small) and /obion (G k_: pod)
Sh1ubs or· sma ll trees; tropical to subtropical seas o nally city forest, thorn scrub cmd
seasonally wet wooded grassland
lleference(s): Sous:i & And1ade (1992)
The N and S populat ions h~• ve previousl y been considered as l wo separate
species, llltl :11·e now treated as one species wit h two subspecies; strongl y
suppo1ted in a cladc with Parupiptadenia, Piptade11icr pro pa rte and
St1yph11ode11dro11 by Luckow et al. (2003), and placed in the Piptacleni" g1•oup
Used for traditional medicine; tl1 e plant has a stron g gmlic o r onion-like smell

Piptadenia viridiflora Photograph by G. P. Lewis Micro/obius foetidus Photograph by C. E. Hughes

180 LEGUMESOFTHEWORLD TRIBE MIMOSEAE 181

 Stryphnadendron ratundifo/ium Photograph by G. P. Lewis
Stryphnodendron Mall 1s37
c. 30 spp.; S America (1 4 spp. in Amazonia, w ith I sp. to C A111e1icn !Costa Rica];
13 spp. in C & SE llra zil, I sp to llolivia and P:uoguay; 3 spp NE B1 azil)
From sliJ'fJbno- (Gk.: ast1ingent) and -dendron (Gk .: lree), in reference co 1he ba1k
w hich contains tannins and ha:-; been used medicinall y
Trees and suffrmices; tropical rain forest and riparian forest, se;1sonally dry fo1es1,
wood l:ind (ce nado), wooded g1ass land and tho rn scru b ( ca:1ting;1)
Refe1ences: Occhioni Molli ns & Mollins (1 972); Occh ioni Ma1tins 0974; 198 1);
13arncby & Grimes (1 98<la) ; Rizzini & I-leringer 0 987); Ne ill & Occhio ni ~·1"rtins
(1989) ; Guinet & Cacca vari (\992); Occhioni in 13erry et al. (2001: 673 -667)
Stro ng ly suproned in ;1 cladc with Parapiptadenia, Piptadenic1 and J1;/icrolo!Jilfs in
the Piptodenia group (Lu cko w el al., 2003); eight species o f geox)'li c scdl1 uticcs
occt1r in th e ce rrado of Goi f1s <111tl Minas Ge r~ii s ( Braz il)
T he ba1k of S ads1r;11ge11s ( M ~ut ) Cov ille (barba timflo) is used fo r medicine
( possibl y effecti ve :ig~1inst l~ ishm:.miasis); o ther uses in ag1 o fo1e.st1y are fo1 timbc1
(cons1ruc1ion), ni11 ogen fix ing and reforestatio n (e.g ., S microstacbywn Puepp. &
Emll., o r v; millio); some species arc toxic tu li vestock
Adenopodia c Presl 1s5 1
Pseudoenlttda Bdtcon & Rose (1928); Eutada subge n. Act1n the111ada Brc n<l n
(1 966)
c 7 spp ; Affirn (I sp. in WC ond 3 spp. in SE Afri ca {2 sp1> in the Zan zib:ir-
lnhambane and I sp in th e Tongaland-l'oncio l:tm l regions]) ; Mex ico (3 spp . l
exte nding to C Am erirn)
From adeno· (Gk.: g l:.1ncl) :.ind pod ion (Gk.: fo ot), referring to th e conspicuo us
gland ab ove the base o f th e petiole
Lianas, shrubs 0 1 .'.'i mall trees; tropic::1l to subtr o pi ca l, seasonall y d1y fo res t, ripaiian
fo rest and forcs r m:ugins, 1hicke1 crnd bu shland
Rderence(s): ll1 enan 0 986)
The three Amazonian sr ccies included b y Bren ~ln (1986) in 1Ul 4!1tOjJodia arc
retained in Pip1ade11ia ( 13orneby, 1986); the genus (as 'Pse11//IJl!11/11tla') is pl:tced in
the l'iptadenia g1oup by Lewis & Elia' 0 981: 166)
Used for medicine
Adenopodia patens Drawing by E. Catherine
182 LEGUMES OF THE WORLD
Mimosa townsendii Photograph by G. P. Lewis Mimosa loxensis Photograph by G. P. Lewis
MimosaL 1753
Scbra11kia Wilk!. (1806); Scbranckiastmm Hnssl. (19 19)
c 490 - 510 spp .; 1110~1 in the Neotropics: centi ed in Mex ico (62 endemic spp , 15
s pp. ex tending co USA and 7 spp. to C Ame 1ica (3 and 2 spp .. endemic to each
regio n 1·csp ecti ve ly)); Calibbean (8 endemic s pp.); 7 s pp. disjuncl between
Mex ico - C Amc 1i ca- C~1ribbcan and S Amcl'ica; 20 spp. w idespread in New \'(1orld ,
3 o f w hich ai e pan1ropical w eed s; c. 350 spp. endemic in S America in a) Brazil (S
o f Amazon ia) and adjacent Pal'aguay, Arge ntina and Uru gu11y; b) Ecuado r, Peru
a nd Boliv ia; c) Orinoco lx1sin 35 spp endemic in the Palaeo trop ics (30 spr in
Macla g~1sc;-1r; 2 spp in SE tropic.il Afri c.:;1 ancl 3 spp . endemic to the Indian
subcontin ent)
F1om miino- (Gk.: a mimic or imitato r) , referring to the rcl<Hion between the
.c;e n.c;iti v it y o f rhe leaves of several species whic: h fold ll[J wh en touched , imitatin g
lhc hand rhat to uches them
T1 ees, shru bs o r herbs; seasona lly dt)' tropical and s ub tro pical foresc, \voodland ,
wood ed g1:.1ssland , thrn n fo1cst, tropical mo niane woodland , temperate g1assland
:ind d esert
Reference(s): llarneby 0984; 1991); Grethe1 el al. 0 996); 13arneby in Bc11y et al.
(2001 : 645 - 657); Vi llie1s in Du Pu y et al ( 2002: 170 - 194)
Placed in the Pipwdenia g1 o up, allied ro Parapipla d en.ia, Pipladen;a,
St1yp lm od en dro11, Micro/obi11s and Anad e11antbera ( Lu ckow el ul., 2003)
Used as o rn a me nt a ls , li\'ing fences, soil binders, fo dd e r-, green manure, shade
pl ants, fuelwood and medicine (e.g., ,H.. p 11dica L., the sensitive pbnt, sleeping
grass, humb le plant); many spec ies are weedy, ca using r mhl e ms in ag1i cultu ra \
lancl
Mimosa tenui{loro Photograph by G. P. Lewis
TRIBE MIMOSEAE 183
TRIBE
Mimozygantheae
Tribe Mi mozygantheae Burkart 1939
The monogeneric tribe Mimozygantheae was described
to accommo d ate the single sp ecies Mimozyganthus
carinatus (Griseb.) Bu rkart, orig inally described in the
genus Mimosa. Burkart comme nted that the genus
Mim ozyganth us was transitiona l be tween the
Mimosoideae and Caesalpinioideae esp ecially because
of its peltate stigma, imbricate sep als and essentially
valvate petals (the upper half of the corolla always w ith
the petals valvate but the lower half with some overlap
at the m a rgi n s). H e su gges ted, b ecau se of the
combina tio n o f p eta l a nd sepal aes ti vatio n, a
relatio nship w ith tribe Parkieae (the two genera of that
tri be cu rre ntly p lac ed e lsewhe re a nd the tribe
disba nde d) but p o inted o ut th at Mimozygan thus
diffe red in having spinescent stip ules, free sepals, free
stamens, eglandular anthers, no staminodes, a few-
FIG. 2 5 Diagram ofrelationships to tribe Mimozygantheae after Luckow et al. (submitted a)
184 LEGUMES OF THE WORLD
by R.H. Fortunato
cl cl ovary and a larg p ltate stigma . Burka rt (1952)
propo ed that the genu Dinizia Ducke shou ld also be
included in the Mimozygantheae b ut Hutchinson 0964)
placed Dinizia in the Mimoseae and Schul ze-Menz
(1964) placed it in the Parkieae. Guinet (1981) grouped
Mimozygantbus with Dinizia in his m imosoid pollen
stu dy, b ut Elias (1981b) repositi oned Dinizia in the
Mi1noseae, thus re-establishing the Mimozygantheae as
mo nogeneric. Luckow et al. (2003) did not include
Mimozygantbus in their mo lecula r study but recent
w o rk (Lu ckow et a l. , submi tted a) indica tes that
Mimozygantbus sho uld be placed in the Prosopis group
of Mimoseae, near Piptadeniopsis and Prosopidastrum.
While treated in its tradition al sense he re pend ing
publication of this analysis (Fig. 25) , there is no doubt
that tribe Mimozygantheae should be disba nded.
Mimozyganthus carinatus Photograph by M. Luckow
Mimozyganthus Buoi<art 1939
I sp .; Neotropics, from SE Bolivia and SW Paoaguay 10 NW and cc ntml W
Argenti na
From zygo- (Gk.: juga te, paired) and a n/hos (Gk .: flower), re fe rring lo the flowers
in heads 1hac arise in pa irs or foui s from betwee n a pair o f spi nescent slipules,
and the prenx m imo- (Gk .: mime or mimic); the th ree elements o f the name
Cae_salpinieae pro parte (see page127) together allude to inclusion o f the genus in the Mi1nosoideae
Sh rubs 01 sm all trees; ri opical and subtro pi ca l add rind semi-alid scrub bushlan d
(clw co vegeta tio n), often associctte<l w ith cacri
Refe rence(s): Burka rt 0939b, 1952); El ias (1981b); F011unato (1997); Lu ckow el al.
(subm itted a)
1Wi111ozyga nt/Jus has a unique combinati o n of mo1phologic<1 I fe<J turcs that suppoiiecl
Mimoseae (see page 163)
trea ting it in an isolated taxonomic position (see above). Moleculm analysis o f the
genus, however, has conftimed its posi tion within 11ibe Mi111ose:.1c (Luckow et a/_,
submiued a) and ri ibe Mimozyganlheae w ill soon pas.is inlo synonymy
Used locally for flrewood and tuinery; appa1e ncly a dye is exu-actcd fmm the stems
Mimozyganthus Mimozygantheae
Acacieae sens. strict. (see page 187)
lngeae (see page 193)
Mimozyganthus carinatus Photograph by M. Luckow
TRIBE MIMOZYGANTHEAE 185
TRI BE

A.cacieae b y G. P. Lewis

·be i\ c·icicae Dumorl . 1829

1'1'1 '

'frib Acaciea is idely attributed t Bentham 18 2),
;i sat 1 81) and Ma ·Jin et. a l. (2001 , bur Re v a l
e.. g,
(l 997) giv Dumorri r (1829 a th first pl ce of
ubli ·a ci n f th tribe a nd thi s is confi rme d by
~nim111ill pers. comm., 2 0 . Th enus Faidberbia
A.Che . wa ind ud cl in the Acaciea by VassaJ (1981
ii nd is sci.II r ra ined a parL f th u-ibe by Ma. !in et al.
(2003). The tribal p ' it io n f Fa idherbic1 r m a in.
qu iv al, althoug h l ewis & Rico Arc (th. volum
place 1h genus in rrib lngea following Polh ill (1 99 )
and Lu ·kow et a l . ( 2003) r ndering rhe caciea
monogeneric, w ith the single ge nu s Acacia . The
taxonomic status of Aca cia and its relationship to other
mimosoicl genera is, howev r, as yet u nresolved. At
present three subgenera a re recognised within Acacia
sens. lat.: Acacia, A culeiferum and Phyllodin eae.
Pedley (1986) proposed that these three subgenera be
given generic rank, namely A cacia, Senegalia Raf. and
Raco~perma Mart., respectively, but th is was not widely
<1ci optecl, alth o ugh the d e b a te s urro unding thes e
suggested nomenclatural changes con tin ues. Pedley
(2003) has recently published combinations (several
hu nci reci of w hich are n ew) in Racosperma for all
Australian phyllodinous acacias . What is clear is that the
genus Acacia , a s c urre n tly circumscribe d , is n o t
monophyletic (Maslin et al., 2003; Miller & Bayer, 2003;
Miller et al., 2003), and at le ast five genera shou ld be
resurrected or newly described fro m within it in clue
course. The five genera correspond to those recognised
by Ped ley except that Senegalia sens. lat. is regarded
as comprising three gen era : Sen egalia sens. strict.,
Acaciella Britton & Rose (syno nym Acacia su bgenus
A culeiferum section Filicinae (Benth.) Taub .), and an
undescribed genus based on Acacia coulteri Benth.
and a small group of related specie s. Acacia subgenus
Aca cia appears more closely related to tribe Mimoseae
and subgenus Phyllodineae nests w ithin the Ingeae in
most recent studies (Luckow et al., 2003). Wh ile these
two taxa are not moved o ut of the Acacieae in this
trea tment, this seems a likely future consequence of
recent research. Removal of Acacia sens. strict. from
the Acacieae wou ld leave the tribe w ithout its type
ge nu s so that Acacie ae could the n n o longer be
re tained . It is not appropriate here to re instate , change
o r d e scribe new gene ri c n a m e s, es pecially as
application of the names Acacia and Ra cosperma are
currently under review (Maslin et al. 2003, O rchard &
Maslin , 2003; Luckow et al., su bmitte d b) . A proposal
to retypify Acacia b ase d o n a n Au s tra lian taxon
(O rchard & Maslin , 2003) has recently been p assed by
the Committee for Sp erma tophy ta bu t this decisio n
awa its ra tifica tion . Th e p resent tre atment of the
Acac ieae thus recognises a single genus containing c.
1450 species (Fig. 26).

Caesalpinieae pro parte (see page 12 7)

Mimoseae pro parte (see page 163)

Mimoseae pro parte

Acacia subgen. Acacia (= Va chellia)

Acacia subgen. Aculeiferum sect. Aculeiferum sens. strict.(= Senegalia)

'Acacia coulteri Group'

Acacia subgen. Aculeiferum sect. Filicinae sens. strict.(= Acaciella)
Acacia subgen. Aculeiferum pro parte

lngeae (see page 193)

Acacia subgen. Phyllodineae (= Racosperm a)

FI G. 26 Diagram of relationships in tribe Acacieae largely based on Miller & Bayer (2003)

LEFT Acacia anthachaera Photograph by B. R. Maslin

TRI BE ACACIEAE 187

Acacia piptadenioides Drawing by 5. Wickison Acacia hopperiana Photograph by G. P. Lewis Acacia brandegeana Photograph by G. P. Lewis Acacia hindsif Photograph by G. P. Lewis

Acacia Mill. 1754 Acacia (continued)

Vachel/ia Wight & Arn. (1834); Racosper111a Mart (1835) ; Senegalia Raf. (1838); charcoal, medicine, tan ning leather (e g., A mearnsii De \Xfild., black watcle)
Acaciel/a Britton & Rose (1928); and several other, mostly obscure, names listed in and oils in arornachempy; nwny species impo1tant in agroforesrry systems (e.g .,
Maslin el al. (2003) A mangium \Villd., mangium or brown salwood, grown w ide ly JS a plantation
Cosmopoli trm w ith in excess of 1450 spp. dist ributed in all conti nen ts except species in Asia); some species planted as ornamentals, cut flowers of A .
Anta1ctica; Acacia sens. strict. (syn _: Acacia subg Acada; \lache/lia), c. 161 spp., clealbata Link and a few other species are sold as 'florists mimosa'; many
pantropical (73 in Africa and Madagascar of which c. 15 extencl to Asia , 21 s pecies are good bee forage ; ~owers of A.jarnesiana (L.) Willd are used in the
reslricted to Asia, 7 in Australia and the Pacifi c; c. 60 spp. in the New \'V'oild, of perfume industry; several species a major source of gum arable although A
which c. 35 in N and C America and c. 25 in S America); Sen egafia (syn.: Acacia senegal (L.) Wi\ld. is rhe 'trne gum arabic'; inany African species are multi·
subg Aculeifel'Ulll sens. sll1cl.), c. 207 spp., pantropical (69 in Africa and purpose trees with a wide range of local uses (see Burkill, 1995: 177-203); in
Maclagasca1, of which 7 extend to Asia, 36 restricted to Asia, 2 in Australi<t ancl the Australia used in rehabilitation and soil improve ment programmes, waterproof
Pacific [1 extending to Asia]; c. 100 New World of which c. 40 in N and C glue production, pulp, for tools, aborig inal weapons and musical instrumen ts;
America, c. 60 in S. America)i Acaclel/a (syn.: Acacia subg. Aculeifen1111 sccc seeds have minor use as human food
Filicinae), 15 spp restricted co the Neotropics (mainly Mexico and C Ame1 ica but
ex tending thinly to S America); Gen nov (the 'Acacia coulleri' g1oup), 13 sp p. (see also next folll figures on fo llowing page)
restricted Lo N and C Am erica; 'Racosperma' (sy n.: Acacia subg_Phyllodi11eae), c.
1045 spp. in Austialia (of which 941 endemic and 7 extending to Asia , and c 100
spp. new and yet to be desc ribedjide M<1s lin et al., 2003) , 7 in the Pacific, 3
confined to Asia, 2 in Madagascar and the Mascarenes
Derived from acacia, first used by Dioscorides (A~D 40- 90) in his Mate lia rv leclica
to desuibe a pre p:ll'ation (used in tanning) from leaves and pods of Acacia
nilotica (L.) Willd., the type species of the genus; deri1,ed from acis (Gk : sh:11 p
point, tip, thom 0 1• barb) in refere nce to the thotns of that species
Armed or unarmed trees, shrubs and Jianas; wide rangin g in habita l, from n1in
forest to a lpine co mmunilie.s, domi nant shmbs and trees in seasonally cJ1y
rrop ical ;:ind subrropical bus hland, woodl and, wooded grassla nd, co<1stal dunes
;1nd deserts
Reference(s): Britton & Rose (1928: 84 - 120, under severo 1 separate genera); Ross
(1979); Lock (1939, 62-79); Lock & Simpson (1991, 9-13); Lock & Hea ld (1994:
33-36); Nielse n (1992: 34 - 64); Pedley (1986, 1999b); Ebinge r et al, (2000);
Maslin el al. (2001, 2003 and refs, therein) ; Du Puy & Villiers in Du Puy el al
(2002 : 224-242); Kumar & Sane (2003: 79 -100); Rico-Arce (2003); Pagg &
All ison (2004)
Leaves, fruits, wood and bark of many species are used for li vestock fo<ldc1
(although pods and leaves of some yield toxic cya nogenic glycosic\es), 1imbc1
(cons truction, handicrafts, ule nsils, implements), famin e food, firewood,

(continued on next page)

Acacia pervitlei Photograph by D. Du Puy Acacia lanuginophylla Drawing by 5. Ross·Craig

188 LEGUMES OF THE WORLD TRIBEACACIEAE 189

Acacia willardiana Photograph by G. P. Lewis Acacia aphy/la Photograph by A. McRo bb Acacia daemon Photograph by G. P. Lewis

TRIBE ACACIEAE 191

190 LEGUMES OF THE WORLD
TRIBE
Jngeae by G. P. Lewis and L. Rico Arce
.... 0 Ingeae Benth. 1865
f!ILI'-
Niel n 19 la) recogni ed 21 nera in In ae (Tabl
) although were n~t gi~en gen~ric names, ~ut were
7 partly due to a lack of appropriate material for DNA
referred t a Gen. A to Gen. D . He recogms d the
genus Mc1rmaroxylon, although withom a generic
number, so that the trib , to the casual observ r
gpp ared ro contain only 20 genera. The genu
pu.njuba appended by Niel ·en 1981a) under "genera
and pedes of unknown affinity'' i h re Lr at d as a
synonym of Aharema, following Barneby & Grimes
(1996 , although we suggest that this may be reinstated
as a g cl genu in th future . Ni n J9 la) al o
included Pithecellobium incuriale (Yell.) Benth. as a
" pe ie1> of unknown affinity" but this is now placed in
Leucocbloron Barneby & Grimes (1996)
Polhill 0994) increased the number of genera of
Ingeae to 25 (Table 7). He recognised Nielsen's 'Gen.
A' as Paraserianthes LC.Nielsen, 'Gen. B' as Archiden-
dropsis LC.Nielsen, 'Gen. C' as Pararchidendron
LC.Nielsen, and 'Gen. D' as Macrosamanea Britton &
Rose ex Britton & Killip. He placed Faidherbia in
Ingeae for the first time, reinstated Cathormion,
Samanea and Chloroleucon, and recognised the
monospecific Obolinga Barneby (subsequently
subsumed into Cojoba by Barneby & Grimes, 1997).
Zapoteca, a segregate of Calliandra described by
Hernandez (1986), was also added by Polhill 0994).
Klugiodendron, recognised by Nielsen 098la), was
considered a synonym of Abarema by Polhill (1994),
and Ajfonsea was placed as a synonym of Inga, a
position later confirmed by Pennington 0997).
The present treatment of Ingeae recognises 36
genera (24 of which are New World endemics) and
(935)-951-(966) species (Fig. 27). We follow Barneby
& Grimes (1997) in placing Obolinga as a synonym of
Cojoba. Eight genera: Blan~hetiodendron, Ebenopsis,
Hesperalbizia, Hydrochorea, Leucochloron, Painteria,
Pseudosamanea, and Sphinga, have either been
reinstated or described as new since 1994 (Barneby &
Grimes, 1996). Paraserianthes section Falcataria was
raised to generic status as Falcataria (LC.Nielsen)
Barneby & Grimes (1996). Balizia Barneby & Grimes
0996) is considered a synonym of Albizia following
Rico Arce (1999). Guinetia L.Rico & M.Sousa was
described as new (Rico Arce et al., 1999, pub!. 2000),
and Viguieranthus Villiers in 2002.
LEFT Pithece/lobium excelsum Photograph by G. P.
Clarification of generic relationships within tribe
Ingeae still suffers from a paucity of molecular data,
extraction of the recently described and reinstated
genera. Luckow et al. (2000) included four ingoid
genera in their analysis of the basal genera of
Mimosoideae. These formed a group together with
Faidherbia (then still considered a member of tribe
Acacieae , although moved to Ingeae by Polhill
(1994)). Barneby & Grimes 0996) concentrating on
neotropical taxa, divided American ingoids into five
informal alliances: the Abarema-, Samanea-,
Chloroleucon-, Pithecellobium- and Inga- alliances.
Genera of uncertain position within their system
included Albizia, Enterolobium and Cedrelinga.
Lysiloma was considered as intermediate between
tribes Ingeae and Acacieae. Luckow et al. (2003)
carried out a phylogenetic analysis of the
Mimosoideae using chloroplast DNA sequence data.
They treated sixteen of th e 36 ingoid genera
recognised in this account, including Faidherbia, but
concluded that relationships within the Ingeae are
generally unresolved and that, with only a few
exceptions, clades within the ingoid part of their
topology were not strongly supported. Albizia
proved to be polyphyletic, supporting the findings of
Grimes (1999).
Any new classification of the Ingeae will require
sampling of all the genera not included by Luckow et
al. (2003) and more extensive sampling of the larger
and putatively non-monophyletic genera. Relationships
between ingoid genera and the various elements of a
polyphyletic Acacia have still to be resolved, although
Luckow et al. (2003) have an Acacia subgenus
Phyllodineae clade nested within the Ingeae ,
suggesting that at least part of Acacia sens. lat. (the
Australian phyllodinous acacias) might be included
within the Ingeae in the future, or that the Ingeae, as
currently circumscribed, may have to be broken up
into several distinct suprageneric taxa. Such suggestions
are premature as 20 ingoid genera, including Abarema,
Archidendron, Pithecellobium, Zygia and the largely
Madagascan Viguieranthus have not yet been included
in molecular analyses.
Lewis
TRIBE INGEAE 193
 TABLE 7 Additions and changes to tribe lngeae since 1981 Acacieae (Acacia subg.
Aculeiferum); see page 187
LEWIS & RICO ARCE (this volume) P,O LHl~L19 94 NIELSEN 1981a

Faidherbia Faldherbia Faidherbia

Zapoteca Zap0feaa Zapoteca
Guinetia
Calliandra Calliandra Ca\liandra
Viguieranthus Guinetia
Macrosamanea Macrosamanea Genus D Calliandra
Cojoba Cojoba·+ Obolinga Cojoba Viguieranthus
Inga Inga Inga+ Affonsea lngeae Macrosamanea
Cedrelinga €edrelinga Cedrelinga Cojoba
Inga alliance
Zygia Zygia Zygia Inga
Marmaroxylon Marmaroxylon Cedrelinga
Marmar0xylen
Zygia
Archidendron Archidendron Archidendron
Marmaroxylon
Falcataria
Archidendron
Serianthes Serianthes Serianthes
Paraserianthes Paraserianthes Genus A
Archidendropsis Archidendropsis Genus B Falcataria
Wa\laceodendron Wallaceodendron Serianthes
Pararchidendron Pararchidendron Genus C Paraserianthes Old World group
Hydrochorea Archidendropsis
Abare ma Abare ma Abarema + Klugiodendron + Punjuba Wallaceodendron
Blanchetiodendron z
Leucochloron [Pithecellobium incuriale] Pararchidendron G'>
Chloroleucon Chloroleucon Hydrochorea Abarema alliance rn
Cathormion Cathormion Abare ma )>
Thailentadopsis Havardia rn
Sphinga
Blanchetiodendron
Havardia Havardia Havardia
Leucoch\oron
Ebenopsis Chloroleucon
Chloroleucon
Painteria alliance
Cathormion
Pithecellobium Pithecellobium Pithecellobium Thailentadopsis
Hesperalbizia
Pseudosamanea
Samanea Samanea Sphinga
Havardia Pithecellobium
Albizia Albizia Albizia
Ebenopsis
Enterolobium Enterolobium Enterolobium alliance
Paintera
Lysiloma Lysiloma Lysilorna
Pithecellobium

Hesperalbizia
Pseudosamanea Samanea alliance
Samanea

Acacia subg.
Phyllodineae (see page 187)

UNPLACED TAXA
Albizia
Enterolobium
Lysiloma

FI G. 27 Diagram of relationships in tribe lngeae mainly after Barne by & Grimes (1996, 1997); Luckow et al. (2003)

194 LEGUMESOFTHEWORLD TRIBE INGEAE 195

 Faidherbia a/bida Photograph by P. Van Wyk Guinetia tehuantepecensis Drawing by E. Catherine Ca/liandra bahiana Photograph by G. P. Lewis

Faidherbia A.Che v. 1934
1 ~p . ; Africa (widespread in Suda nian, Zambezian and S<1 he lian regio ns f1om
Egypt, Senega l and the G;1mbia south to Botswana , ~md KwaZu lu Na1al in S
Africa) and in the Arabian Peninsula to \ Y,/ Asia (Ir~rn) ; iniroduced in India , Nepal,
P:Jkistan, Peru and some At lanric islands
Named for the Fr·ench general L.L C, Faiclhe1be (1819-1889), govern o1 of Senegal
in 1854 <tncl 1863, who pllblishe<l seve1al woJ'k s about Senega l and Algeria
Ti-ccs; seasona lly dry twpical to subt1opic:1I rive rin e and g1ounclw:He1 fotcs t,
woocll<incl, swamps and llooclplains
Rcfcrcn cc(s): B1c nan 0959: 78-79, ;is Awcia a/bida Delile); Wickens ( 1969, as
Acacia albida); Ross 0979: 83--'35); Burkill 0995: 233-236); VcntCI' & \'enter
0996: 22-23); Timhel'iake e/ a/, 0999: 24-26); Barnes & Fagg (2003)
Segrega ted from Acacia by A. Chevalier in 1931, but this w~1s ignore d unlil
recently; 111ole<.:ul;;11· studies of Luckow el al (2000, 2003) confirm rli~1t the genus
belongs to Ingeae and not ro Acacicac ; the slrn pe of the ripe pods inspi1cd the
common n ~ 1 rne 'apple-ring aG1cia'
Faidberbia nlbida (Delilc) A.Chev. (wi11te1 tho rn; see Barnes & f;igg (200.'\) for
1eview o f vemacula1 names) is used in ;ig1·oforeslry, for livestock fodde r,
rnedicine, famine food ( humans), shelter and shade trees (unwmal in bc in ~
leaness duting the rniny season <tn<l lea fy during the chy season), hedge!->, ;u1d
timber (foi light construction , utensils, tools , co nwiners <mcl fences); rh e lxuk,
roors <incl powdered pods have been used ::ls a fish poison; bark srrips a1c used
( like dental no."i.s) to cle;:m teeth; in some areas a popular o rn amental and is s:iid
to be an indicator of fc1'lilc sites
Zapoteca H.M.Hern. 1987 ('1986')
20 srp.; Neotr'op ics, 11 spp. f10111 S\V' USA ~mcl N Mexico th1·ough C Ame1 ict :ind
th e C;:i ribbea n (6 sp p. endemic in Mexico with a concen t r'~lli o n in 0:1x;1ca, 3
endemic in C America ;:ind 1 in Hispaniola); 6 s pp. restricted to S Americ:t ;111d
largely ci rcum-Amazonian (1 endemic in Colombi;:i, I in F.cuaclor, 1 in Peru and 1
in Brazil), 3 spp. widespread rh1ough N :rnd S Amcric.i ( J extending 10 Pa 1;1gu~ 1 Y
and N Argentina)
Named for the Zapotcc people of 0 ;1xaca, Mexico .
Shrubs; common in o pen sites derived from seasonally diy 1ropical fo 1'Cst, ~ind in
arid and semiarid scru bby vegetatio n and wet evergreen forest
Re fere nce(s): Herna nd ez 0986, 1989, 1990); He rn itnd ez & Campos (1994): Jlcisslcr
0998: 153 - 158)
All species o f 11enthain's Callim 1dm ser ies Laeteuirellles ~ir e inducled in Zopoteca;
the single species o f ZajJoteca included in the study of Lu ckow el al. (2003) is
sister to Fctidb erbia , basally bi;rnching in Ingeae
Used as o rn ~ 1111 e ntal s
Guinetia L.Rico & M.Sousa 1999, publ. 2000
1 sp ; Mexico (endemic to O:axaca)
Named for Philippe Guinet 0 925 - ), french sc hool teache r and palynologist
specialising in mimosoi<l legumes
Slm1bs; seasonally d1y tropical lowland forest, 0 - 150 m
Hefcrcnce(s): Hico Arce el nl, (1999, publ , 2000)
Calliandra llenrh. 1840
/lnn es/ia Salisb. (1807)
c. 135 sp p .; endemic to the Americ1 s, 1anging from S\V' USA to Uruguay, wa1m
tempera Le Argentin;J and N Chil e; 30 srp restricted to N A111e1 ica, 4 spp
extending from. N AmeriG1 into north ern S Am erica, 6 spp . endemic to the
Caribbean, a c.:0 11ccnt1ation of taxa (66 sr p.) from NE to SE 131'azil (\vith 31
end emic to I3ahia), 26 spp. restricted to N, NW and \Y/ S AmeriG1 ( from the
Gu i<inas to S Peru :ind \Y/ l3oli via)
Fro m calli- (G k : beautiful) and a11dros (Gk .: man); m osr species have brigh!l y
colour ed 0 1 showy slamen filaments
Trees :.rnd shrubs; moslly seasonally dry tropica l forest (man y 1 iparian), wood land,
thorn fof'est and sc1ub (cciatinga); wooded grassland (cena<lo) and rock}'
shrubla ncl (campo rupestre), some tropical su bmonwne, several acbptecl to clese1t
e nvi ronmen ts
Reference(s): Thulin el al. 0981, for Afri c~1n species, but .see notes below),
Barneby (1998); Bassler (1998: 158 - 168); Chamberlain (200 1)
lfarneby (1998) rest1ictecl Ca!lirmdra to the Ameri cas , rejecting Olcl \\lorkl species
named as Calliandra from the genus (see notes und er V(q11ieran1h11s); he implied
new ge ne ric status for two African species, referred Madagascan species to
\figu.ieranlbus and rejected 3 Tnclo-Bunnese species fro m Callirmdra sens. strict.
\'V'i dely culti va ted as ornamentals (calli;111dr:1, powder puff); used in agrofrn'Cst ty
(e .g., C. bousloniana (Mil L) StanclL va r. ca/othyrsus (Meis n.) I3ameby, as live.stock
fodder, fuclwood, green manure, pulp fo 1 pape1 production, erosion control,
filebreal<s, r efo1·es1ation, bee forage, le;1 f meal ( protein sou rce), medicine and as
shade frees
(see also nexc four figures on following page)

Zapoteca caracasana by G. P. Co//ia d . .

Photograph Lewis n ra tumbez1ana Photograph by G. P. Lewis

196 LEGUMES OF THE WORLD TRIBE INGEAE 197

 Ca/liondro coccineo Photograph by G. P. Lewis V/gu/eranthus ombongensis Photograph by D. Du Puy Mocrosomoneo mocracolyx Drawing by 8. Angel

Viguieranthus Villie1s 2002

c. 23 spp.; Macl.igascar (18 spp., 17 of which are endemic, with one extending to
the Comoro Islands) :and Asia
Named fc)I" Rene Viguier (1880-1931) whose unpublished account o f the legumes of
Madagasca r w;is the basis for The Leg11111i11osae oj'Madagasca1•by Du Puy el al, (2002)
Trees and sh rubs; tropical lowland to submoncane huiriid fores[ and seasonally
d1y woodland, thicket and scrubbnd
Reference(s): Villiers in Du Puy el al. (2002: 271 - 285)
Villiers in Du Puy el al (2002: 271) implied that the genus contains five species
add itional to those occurring in Madagascar; l1e did not, however, fo1~11ally
tl8nsfe1 them from Cal/iandra so no combinations exist in Viguieranthus. Villi ers
(l.c.) also stated that the genus occurs in Asia where only three Indo-Burmese
species (one from Myanmar [Burmal and two fl'Qm India) have been ascribed to
Ca/liandra (13arneby, 1998: 3). While it cannot be assumed that Villiers also
intended to include the two rejected African calliand ra s to make up his total of 23
species, this does highlight the poo1· .<Hate of know ledge of generic Jirnits within
the Old \'(<'orfd calliandras Relationships of this gmup to the three Asian species of
n1ai/entadopsis should also be invest ig~1led Vfguieranthus was segregated from
Calliandra, since it differs in having all tlowers fertile and similar to C<tch other,
sepals imb1icate in the flower buds, <ind seamen bases fused to the petJls
Used for timber (constmction ;md joine1y) and firewood

Macrosamanea Britton & Rose ex Britton & Killip 1936

11 spp~; S America, most diverse afld numerous in rhe Amazon basin extending N
into the O rinoco valley <111d the Guianas
From 111acro- (Gk.: g1eat) and Samanea (from the abo riginal sarnJn == rain tree)
Trees and sh1 ubs; tropic<il, mostly riparian and seasonally nooded f01 est, 2 spp. of
seasona lly wet sandy wooded giassland (savanm1)
Reference(s} Barneby & Grimes (1996: 182 - 203)
Largely equivale nt to 'Genus D' of Nielsen (1981a); placed by Bai neby & Grimes
(1996) in their 'Inga-alliance' Nectaries occUJ-on floral bracrs in most species and
are unique co rhe genus in neotropica\ Inge~1e

Cojoba Britton & Rose 1928

Calliandra leptopoda Photograph by G. P. Lewis
Co/ob 0
Calliondro /onota Photograph by G. P. Lewis arborea Photograph by c. E. Hughes
Obolinga Barneby (1989)
12 spp.; Caribbean (4 spp. endemic to the Greate r Antilles), NW Andean and
trans-Andean S America (4 spp.), SE Mexico and C America, 1 sp. (C arborea (L.)
Britton & Rose) nearly coextensive with rhe genus
Derived from a vernaculm name for Cojoba arborea in Pueno Rico
Trees and i:;hrubs; u·opical, mostly wet low land or subm o ntane forest, 3 Caribbean
spec ies in seasonally city scrub forest or in sclerophy llous shrnbland (chaparral)
Reference(s): Rico Arce 0991); Barneby & Grimes (1997: 36-60)
Grimes 0995) indicated that A1acrosamanea and Inga are the outgroups to Cojoba
CojohCJ arborea (arclilla, jolote beard) i.s used as timber, ornamentals and shade trees

198 LEGUMES OF THE WORLD TRIBE INGEAE 199

 Cedre//nga cateniformis Drawing by P. Holliday Zygia macrophylla Photograph by B. B. Klitgoard
Inga sapindoides Photograph by G. P. Lewis

Inga MilL 1751 Cedrelinga Duckc 1922

A/fonsea A.St .-Hil. (1833); Fe11il/em Kuntze (1891)
c. 300 spp (including 45 to 50 imperfecrl y known and undescribed spp.);
Neot1opics f1om 24°N in Mexico co 34°5 in Uruguay, including 2 spp restricted to
the Caribbe~tn. highest diversity in Peru , Ecuador, Colombia and sout hern C
Arne1ica. 40 spp. confined to Mexico and sou thern C America, 75 spp. essentially
Amazonicin, 33 spp. rest1 icred to coastal Brazil, c. 60 spp. exrra-Amazoni;rn within
an 8r'c f1om the Gui~11us through Colombia to Peru ~tnc.l 13olivia, the remain ing
spp. Cc. 40-45) extend ing across the Parn:tma isthmus (seve1al of these
widespread especially in S America)
Derived from the S Arne1 ican Tupi-Guarani vcrnacl1lar n<irne 'ing{t'
Trees; mostly of lowland ancl mont~me r'ain fore.<;t throughout the hurnicl t1opics,
0 - 3000 m, half in l'ain forest on non-flooded land, half in rip<1.1i~in habitats on
pe1 iodically flood~d land , 1arcly in seasonJlly dry areas
Hcference(s): Sm1sa (1993), Pennington 0997), Pennington & Revelo 0997),
Reynel & Pennington (1997), Pennington & Pern<1ndes (1998)
Richardson et al. (2001), bJsed on DNA .sequence da1a , indicate that /l(~a has
dive1sified 1elatively 1cccnt ly, and that speciation has been concc ncratcd i n the
past 10 million years, w ith rn~m y species arising ~ts recently as 2 million y<.:a rs <tgo
All Inga species ha ve sim ply pinnate (not bipinn ate) leaves; in the Mimosoideae,
Z)'gia (1 sp.) and Cojoba (o few spp.) a1e the only other gene10 with some
species simply pinnate
Used ~ts multi-pu1pose soil resto1ation and ag1oforestry t1 ees (e.g. ,/. ed11!is Matt.,
or ice cream bean, guava machete, inga cip6), for edible fruirs (the whice pulp is
eaten or- usec.l in flavouring desserts), shade fo1 crops, l e~if mulch, nitrogen fixing
propc 11ies, timber, fuelwoocl ~md medicine
1 sp.; Biazil, Ecuador, Pern, Surinam, French Guian<1, a nd Venezuekin Guayana
From Cedrela (in family Mcliaceae) <lnd !uga (anothe1 mimosoid legume) due to
its similarity co these cwo gene1a in ba1k and wood characters
T1ees; mainly Amazonian lowland tropical mi n forest to seasonally dry forest,
especially along streams, 50-250 (-800) 111
Reference(s} Ducke (1922); llarneby & Glimes (1996: 25 1-25/i)
Accoiding to Barneby & Glime.s (1996) the genus has no obvious relatives within
Lhe Jngeae but is closest ro Albizia and E11terolobi11m
Mmure trees c01n att01in 60 (-66) m; C cateniformis (Ducke) Ducke (cedrc)l'ana,
lOrnillo, huayl'acaspi) is used in agrofo 1es1ry for nitrogen fixing, fuclwoocl,
medicine and timber (for con.st1uction, boat bui\cling, frames ancl furnituie; a
substirute for 1rn1hog;:1ny)
Zygia P.l31owne 1756
c.. 45-50 spp.; Neotropics, from Mexico <1nd rhe Greate1' Antilles ro S America,
lllO:)t diverse in C Ame1ica and S Mexico (c. 10 spp_), Amazonia Cc.. 10 spp.), N ;md
no1thweste1TI S America Cc. 20 spp., most diverse in Colo mbi;:1 ~ind the Guian~1s)
P1om zygo- (GL yoke) alluding to the s ingle pair of (joined 01• yoked) pinnae
and also ro the p;:11tially fused stamen filaments
Trees and shrubs; t1opical, mostly ripa1'ian fore.st ~ind coastal habitats, generally
below 900 m, but a few species 1e;:1ch 2800 m
Reference(s} Rico Arce (1991); llarneby & Gl"imes (1997, 60-131)
B<imeby & Grimes (1997) adopt a b1oad defini1ion of the genus and include
MarmaroJ..y!on in synonymy; Marma rrL\)llD11 was co nsidered to be a distinct
genus by Rico A1'Cc (1991) and is 1'etain ecl as such in this work based on macro-
mo1phological evidence; Zygia has not yet been included in molecular studies
Used fo1 so il s t~1bilisatio11 and erosion co ntro l (e g. 1 Z . lon8 !fO!ill (\'(filkl) B1itton &
Rose, OJ SOLOC<lba llo)

Marmaroxylon Kil lip 1940

c. 9 -1 3 spp.; 1m1inly Venezuela and the Gu i01n as (6-7 spp.) , Amazonia (Peru ro
Ilrazil), also Bano Colorado Island
from marmaro- (Gk .: m::ll'ble) and J..y/011 (Gk.: wood) 1efe11·ing to the hard wood
Trees and sh 1ubs; tropical, mainly floo<led fotest
Reference(s), Nielsen Cl98h1), Rico Arce (199 1)
Rico A1cc (199 1) rccognisecl 1l1ar111aroxylo11 as a disti11Ct genus; Barneby &
Gtimcs (1997) plJcecl it as a synonym of Zyg ia . Pending molecubr analysis of
both raxa, 1hc posiLion of 1Harmaro.".,y/011, and th e exact 11umbe1· of species,
remains unclea1
Used frn· timber, e.g. , l11 race111os11111 (Ducke) Killip (marhlewood, angelim rajaclo ,
se 1pentwoocl, snakewood) fo1 fu1 nit111 e, floo1 ing, cab inct1y, rna1quetry, musi ca l
Nlormaroxy/on basijugum Photograph by J.·C. Svenning instrumen ts, rnrnery and constructi o n
Inga feuillei Photograph by G. P. Lewis

TRIBE INGEAE 201

200 LEGUMES OF THE WORLD
 Falcataria moluccana Drawing by K. Tind Paraserianthes /aphantha Photograph by G. P. Lewis Archldendropsis paivana subsp. tenulspica Photograph by J. c. Bradford

Archidendron EMuell 1865

Paraserianthes LC.Nielsen 1984 ('1983')
1 sp ; SE Asia (Ma lesi::1) ;md SW Australia , widely cu ltivated in tropical and warm
Cylindrokelupba KosternL 0954); Morolobium Kostenn. 0954); Para/bizzia
Kostcom. (1954) te mpera te regions of both hemispheres
F'rorn para- (Gk : beside) and Serianthes, resembling that genus (but this is now
94 spp.; Asia and Austo"l ia (India , Si·i Lanka [1 endemic sp.), lndo·China (a centre
unfortunate given the recircumscribed, monospecifc status of Paraserianthes; P
of diversiry in Thailand, L1os and Vietnam wirh 10 endem ic spp~l, rhrouglmut
Malesiil 14 spp. endem ic in the Philippines, 5 in Borneo, 3 in Sulawesi, 3 in the
lophantba (Willd .) l.C Nielsen is less like Serianthes than is Fa/cataiia
Moluccas, and 30 in New Guinea and the Solomon Islands), 7 endemic in NE
moluccana, which was origin<Jlly a lso included in Paraserianthes)
Australi~1, J sp. ex tending ro Micronesia)
Shn.1bs or tl'ees; coastal 01 near coasta l open e ucalypt forest, thicket, shrubland
Fro1n archon- (Gk: chief or chieftain) and dendron (Gk .: tree) in refe1cnce to the
and grass land (SW Australia) and tropical mo ntan e and e lfin forest and grassy
large St<Jture of many species, or archien- (Gk.: anci ent), or arche- (Gk: plains, 600 - 3265 m (in Malesia)
Reference(s): Nielsen et al 0983a); Nielsen (1992: 149-150), Bameby & Grimes
beginning) and dendron, suggesting <in early evolution
Trees ;me\ sh11Jbs; mostly of t1opical lowland prirna1y and second<try 1<1in forest, 0996: 256 - 257). Cowan 0998: 35-36)
Apparently taxonon1ically isolated in the Jnge<Je acc01ding to Barneby & Grimes
on sandy or lateritic soils, a few species in swampy h<1bit<Jts
Rderen ce(s} Nie lsen (1979a, 1982, 1992: 86 - 141 ); Nielsen et al (1984a); Cowan 0996); 'Genus A' of the lngeae in Nielsen (198Ja); Nielsen et al 0983a & b)
rlaeed the three species of Fa/catatta (LC Nielsen) Barneby & Go imes in
(1998: 40 - 48)
RelJted to Cojoba and Zygia of the Ameri cas
Parasen'cmtbes section Falcalaria IC.Nielsen; resolved in the molecular
Used for limber (construction), fencing (stems), hurn <J n food (f1uir, e.g , A jiringa phylogeny of Kajita et al. (2001) as sister to the type species of Albizia
Qack) l.C Nielse n, or jeo'ing, luknieng) and navouring (seeds). ta nning, fis h Paraserianthes lophantha (Cape Leeuwin wa ttle, crested wau le, brush wau le,
plume albizia) is used as an ornamental and for reforeslation, al though sometimes
poisons, dyes, soa ps and medicine
becoming naturalised and an invasive weed
Fafcataria (l.C.Nielsen) Barneby & J ,W.Grimes 1996
Parase1ianthes sect. Fa/ca/aria LC.Nielsen 0992)
3 spp~ ; Moluccas, New Guinea, Bismarck Archip elago, Solomon Islands, l sp Archidendropsis 1 C.Nielsen 1983
endemic to Aust1"a li a (Queensland), 1 sp. cullivated in the New \Xi'orlcl 14 srp.; SW Pacific (8 spp., New Ca ledonia), Australia (3 spp, Queensland,),
Frornfa/catus (L.: sickle-shaped), probably alluding to the lea Flet shape Papuasia (3 spp ., New Guinea-New llritain-Solornon Islands)
Trees; tropical rain forest, and coa.stal 'chy rain fores t', 0 - 2300 m Derived from its resemblance to the genus Archidend1-on
Reference(s): Nie lsen 0992: 150-154, as Parase1iantbes) , !la1neby & G1irncs Trees and shrubs; tropical mostly lowland rain forest 1 but 2 of the Australian
0996: 254-256), Cowan 0998: 36, as Paraseriantbes) species in drier eucalypt woodland, vine thicket and acac ia shrubland
s :-ti In refor'Csttoli<Jn fo r its ra pid g!Q\\1h. as sh:idc 1rees (C(X'O;I ,ond cuff<..:). Refcrence(s): Nielsen eta/. (1983b), Nielse n (1992: 141 - 142), Cowan 0998:
m:in1e111r1l<, for w00<l (pu Ip, er.lie :ind llgh1 fomillore), lltwc (l1:1rkl, h11m.111 food 37 - 39)
(green (>Qds), OrewOQCI and ch,,~o>I; f·: 111oluccn11fl (Miq.) !);irncl>)• &J . ~ .{;riones 'Genus B' of the Ingeoe in Nielsen (198la)
(balai, peacocks plume, white albizia, jeungjing), is an invasive weed Various species (salmon bean, red lancewood, d ead finish) used for timber
(cabinet making and turnery), whips and as ornamentals
Serianthes Be nth 1844
c. 18 spp.; Jndo-China (Tha iland), throughout SE Asia and Pacific (Male,ia,
Papuasia, Micronesia, Melanesia and W Polynesia; 6 spp. restricred to New
Caledonia whe re the genus shows most vari<Jlion)
From se1ico· (Gk.: silky) and antbos (Gk.: flower), referring to the dense silky
hairs on the outer surface of the petals
Trees and shrubs; tropical rain fo1est, scrub fo1est and coastal scrub, 1arel)' in
open wooded grass land
Reference(s): Fosberg (1960), Kanis (1979), Verdcourt 0979: 196-203), Nielsen et
al. (1984b), Nielsen (1992: 157-162)
Used for making canoes; S ne/sonii Merr. (firetree) is endangered in che Pacific
. flu/gens Drawing
Archide ndrops1s . by Turpm.
Serianthes grand/flora Drawing by K. Tind

TRIBE INGEAE 203

202 LEGUMESOFTHEWORLD
 Pararchldendran pruinosum Drawing by unkn own artist Abarema fi/amentosa Photograph by G. P. Lewis Abarema lehmannii Drawing by P. Halliday

Wallaceodendron Komd. 1898 Aba1·ema Pittier 1927

I sp.; endemic to Malesia (N Su lawesi [CelebesJ and che Philippines) }upunba Boilton & Rose (1928); P11nj11ba Boiuon & Rose 0928); Klugiodendron
Na med for Alfred Russel Wa llace (1823-1913), British ex plorer, zoologist :ind B1i uon & Killip (1936)
botanist, and dendron (G k : tree) 46 spp.; e ndemic to the Neotropics, l sp . exte nding N to Bahamas and 2 spp. S
Trees; tropical coasial and in land 1ain forest, 0 - 850 m ~long the coast of E Braz il to Santa Catarina ; 7 spp.. restricted to the Caribbean, 6
Refe1 ence(s): Nielsen el al. Cl983 b), Nielsen (1992: 162 - 165) Lo Mexico and C America, 10 to Amazonia, 6 to Atlan tic Brazil, c. 14 in N and NW
The seeds are in indivi dual e ndoca1p envelopes S A m eri ca (especially the Guayana Highland), <lncl 2 on both sides of tl1e Panama
\flal/aceodendron celebicu m Kooid . (banCtyo [=i1onwoocJ], cJcrham mahogany) isthmu s
timber is used for furniture, cabinet making and fl oo1ing Most p rob~bly derived from the vernacular nam e 'auaremo-temo', the basis of
Pifhecellobium section Abare111otemo11 Be nth., from whi ch the genus was erected
by Pitt ie r
Pararchidendron 1.c .Niclsen 1984 ('1983') Trees; in a wide range of habitats including (ropical lowland terra flrme and
1 sp~ ; Male.s ia (Javn) to NE Australi<1 (Queenslan d, N New South \Xlale.s) montane humid frnest, wooded slopes on sa ndstone t;:ible mountains, coastal
From pam· (Gk.: beside) and Archidendron, i"ese mbli ng that genus scrub wood land, often on white sands (both restinga a nd ca mpina), rarel y in
Tree or shnib; tropic;.1] hilly to mon tcme 1ain forest (M ;:desia), seasonally di y rarest, riparian nooded fo rest
and coastal scrub (Aust1alia), 0- 2250 111 Heference(s): [la rneby & Grimes 0996: 41 - 111), BUssleo· (1998: 116-127)
Re ference(s): Nie lsen et al. (1983a, 1984b), Nielsen (1992: 145 - 148), Cow<1n Toget her wi th Balizia Barneby & J.W.Grimes (effectively sunk into Albizia by
0998: 39-40) Ri<.:o Arce (1999) and not recogn ised in this wo1k) and Hydrochorea forming the
'Genus C' of the lngeae in Nie lsen 0981a) 'Abarema·alliance' of [larneb y & Grimes (1996)
Pararchidendron pn,inosum (Benth) LC.Nielsen (snow wood, tu lip siris,
sl inkwood) is used as ~in ornamental an d street rree
Blanchetiodendron uarneby &J.W.Grimes 1996
1 sp.; E Brazil (Bahia and Minas Geia is)
Hydrocborea Barneby &J.W.G1imcs 1996 Named fo r Jacques Samuel Blanchet (1807 - 1875), Swiss botanist, and dendron
3 spp.; S Ame1ica in the Orinoco and Am:::izon basins, th e Gu ianas and in B1azil (Gk.: tree)
extending inlo Maranh~o and Mato Grosso do Sul Tree; seasonally dly tropical forest, woodland (ccrracl o) , liana forest ('mata de
From hydro· (Gk.: water) ancl chorein (Gk.: to tra vel) in a llusion to the water- cip6') and wooded grassland, 400 - 1000 m
borne propag ul es Reference(s): Bameby & Grimes 0996: 127-130)
Trees <.Jnc.l shrubs; tropica l , ofte n inundated, ripa1 ian fo 1est and woodland Placed in the 'Chlomleucon-alliance ' of llal'J1eby & Grimes 0996); the single
Refe1ence(s): Barneby & G1imes 0996: 23-34), Rico Arce 0999) species B h/anchelii (Uenth.) Uameby & J W Grimes, has previously been placed
Placed by Baineby & Grimes 0996) in their 'Abarema alliance'; t\ico Arce (1999) in four different legume gene•~ . including Albizia
d e monslmted that Hydrocborea (?) acreana (J.F.Macbr.) Bt1rne by & J .\X',Giimes is
an Abarema

Hydrochorea carymbosa Drawing by H. E. Ireland Blanchetiodendron blanchetii Drawing by P. Halliday

204 LEGUMES OF THE WORLD TR IBE INGEAE 205

Leucochloron limoe Drawing by E. Catherine Leucochloron limae Photograph by G. P. Lewis Cathormlon umbellatum Drawing by P. Halliday Thailentadopsls vietnomensis Drawing by H. Lamourdedieu

Leucochloron 13emeby & J w Grimes 1996 Cathormion Hass k 1855

4--5 spp.; Aclanric and phinatine Brazil 1 sp.; S As ia (I n di~1 ::md S1i La nka), Inda-China, Malesia, Papuas ia a nd Australia
From leu co- (G k.: white) a nd cbloro- (Gk.: gree n ish yellow); ;rn ;inagr.1111 of [he From catborm ion (Gk.: neck lace) in reference to the moni li fo 1m fru i[ th at
ingoid genus Cblv roleuco11 to which it has affin itiCti resembles a string o f beads
Trees ( I sp. occasionally shru bby); seasonall y dry tropi cal, main ly wooded Shrubs; tro p ica l riverine fo rest (and mangrove), .seasonally d ry woodl a nd and
g rassland, woodland and thorn sc rub (incl uding 'e1ari nga') wooded grass la nd
Rcfere nce(s} llarneby & Grimes Cl996: 13G-136) Refe rence(s): Nie lsen (1992: 143 -145), Cow"n (1998: 49-50)
Pla cecl in the 'C hl orole uco n-a llia nce' of 13arne by & Grim es (1996); a pa 1ti a ll y- Eleve n African a nd American species fo rme rl y inclu ded in Cathormion ;:11 e now
known tree in Venezuel:J might rep1esent a fifth sp ecies re fe rred direc tly or in comment (Nie lse n, 1992; Ba rn e by & G1i mes 1 1996) to
Albiz ia, Chloroleucon and IJydrocborea; Ca tbormion is sister to Chloroleucon in
the a nalys is o f Luckow el al., 2003
Chloroleucon (Ben th .) 13, itton & Rose 1928
10 sp p.; Ame ri ca, fro m NW Mexico (1 sp ) to the Ant illes (2 spp.), remaining
species in S Ameri ca, S {0 Argentina ancl SE Brazil
Thailentadopsis Kosterm l977
From cbloro- (Gk.: greenish yellow) and leuco- (Gk.: wh ite), probably alludi ng to 3 spp ; Sri La n ka (I sp.) and Ind o-C hin a (I sp., Thai la nd; 1 sp . S Vietna m)
the p:Hchwork bark of it~ species Derived frnm Thaila nd (co unt ry o f origin of [he type species), Entada (a
Trees a nd shn.ibs; wa rm te rnpemte and tropical lowla nd ~ncl less often mi mosoid legum e ge nus, q .v.) and -opsil· (Gk: appeara nce), for the s uperficial
.<i ubm onrane se:-1so nally diy forest, xe1 om01·phic bmsh-woodl ::mcl, crn:1stal rhickct , resembla nce o f the fru i[ to chat genus
woo ded grass la nd, shmbland ancl clcseit Sh rubs or small trees; seasonally dry tropical fo rest ~d ong l'ivers or in inundated
Reference(s): Barneby & Grimes 0996: 136-157) areas, often on limesto ne
Characterised by axi ll t1ry spi nes ( modi fied sterile peduncles), st1iate rest ing buds Re fere nce(s): Koste rma ns 0980: 494 -495); Nielse n (198lb: 109 & 111); Nielsen
~in d fl owering: preceding leafi ng (1985b: 205 - 206); Lew is & Sc h1ire (2003b)
The 3 species (trea ted separ"ately in each of the fi rst three refere nces above) ha ve
bee n placed in several genera including Pilhecellohtum a nd Painteria, and
relatio nships have been suggested with Hava rdia (Nie lse n, 1992). All three ge ne ra
a r>e no w co ns idered to occur exclu sive ly in the Ne w \'\/oriel, and Kosterrnans'
genus Tha ilentadopsfs wcis 1'esrn rected to acco mm oclate these Asian species,
pending fu 1t her gene ric-level studi es (Lewis & Schrire, 2003b); the genus a ppea1 s
ma c romoipho logicall y close to Ca1bor111 io11, b ut 1e lat io n.':i hips to the 3 As ian
species o f Vi1.511iera nthus sho ul d also be in vestiga ted

Sphinga Ila rn e by &J .W.Gri mes (1996)

3 spp.; Cuba (1 s p.), Mex ico (2 spp., one ende mic), Guaterna\;:i, a nd north-western
S Ame ri ca (Colombia , Venezuela and extending in to the Caribbea n is land of
Aruba)
From Sphinx (Sp hing idae - haw kmot hs), the pu tative poll inatofs a nd th e Tu pi
Amerindia n 'ingfl', ttie vernacul ar na me for several Mi mosoideae
Trees a nd shru bs; seasonally d ry ti o pica l to alid lowla nd a nd submontane fores t,
thi cke t, wooded grassland and thorn sc ru b
Refe rc nce(s): 13arneby & Grimes (1996: 160 - 165)
Orig ina ll y desc ribed in Havardia by Britton & Rose (19 28), these 3 species were
placed in Sphlnga by 13arneby & G1imes (1996), form ing p a1t of their
Chloroleucon mangense Photograph by G. P. Lewis Sph·
mga acatlensis Photograph by c. E. Hugh es 'Pirhecellobi um-alliance'

206 LEGUMESOFTHEWORLD TRIBE INGEAE 207

Havardia campy/acantha Drawing by M. E. Rickards Ebenopsls con(inis Photograph by c. E. Hughes Plthecellabium histrix Pho tograph by G. P. Lewis Pithecellobium excelsum Photograph by G. P. Lewis

Havardia small 1901
5 s pp.; Mexico, Texas and C A1ne1ica
Named fo r Dr Va leiy 1-JavJrcl, Assistant Surgeon-Genera l of rhc US Army , ;.1
student of the N American floi'a
Trees; wa1m temperate and seasona lly c11y t1opica l woodland, wooded g1ass land
and d esert th o rn scn.1b, mostly below 450 m
Referen ce(s): 13arncby & Grimes (1996: 165 - 173)
Placecl in th e 'Pithecellobium-alliance' of Bai neby & G 1imes (1996)
Hauardia pa/lens (I3cnth.) Britton & Rose (tenaza, huaji\Jo) is used for wood
(turnery, urc nsil.s), livestock fodder, revegetatlon, bee forage and as an orrn1mental
Ebenopsis B1itton & Rose 1928
Sideropsis Small (1901)
3 spp.; Mexico (including the Yuca tan) and S Texas
From 'eba no', the comon name of the type species, perhaps because of its hard
\vood .similar to Ebenus
T1ees and sh111bsi warm temperate and seasonally dry tropicfll woodland, thicket,
thorn forest and deseit shrubland
Reference(s): Barneby & Grimes 0996: 173-178)
Placed in the 'l'ithecellobium-alliance' of Barneby & Grirnes (1996)
Ebenops1:s ebano (13erl ancl.) Bai neby & ] W.Grirnes (Texas ebony) is used for
wood (fence posts, charcoa l) and as ~m 01 m1rnental
Pitbecellobium Ma1t. 1829 (110111. and orth_ conserved 1837)
J8 spp.; N and S America , including the Caribbean, 12 spp. restricted to Mexico, C
diligent America and the Gre~He1' Antilles (3 exrencling N into subt1opica\ Mexico, the
lfahJmas and Florida, 1 circum-Ca1ibbean) 1 3 on ly in no11hern S America and 3
more wiclespreJd both sides of the Pananrn isthmus
From pitheco- (Gk.: monkey) and e/lobion (Gk.: eal'J"ing) in allusion to the
pendulous, contoitecl fruits
Trees and shrubs; lowland seasonally d1y tropirnl woodland and thorn .scrub
below 500 m, some to 1550 111, a few on co<1s tc1l dunes or in 1ipa1 ian woodland
Reference(s): Barneby & Grimes 0997: 2 - 36), Basslc1• (1998: 168-182)
Prior to 1981, Pithecellobium se1vcd as a dumping ground for many ingoid
species of unknown affinity but the genus now excludes all Old \\forld taxa <it one
tim e placed within it
Pithecellobiwn dulce (Roxb.) IJenth. (Madras thorn , Manila tamarind,
guayarnochil), is a common ornamenta l, hedging and s hade tree, where it is also
grown for the edible pulp in the seed aril (also made into beverages); other uses
include timber (constructio n and po.sts), li vestock fodder, firewood, bee forage
(honey). seed oils (soap) and ba1k for tanning; plants can be tenacious weeds

Painteria Britton & Rose 1928

3 spp.; Mexico (Mexican Pl~teau, with 1 sp~ disjunct in lowland T<1maulipas)
N arnecl fol' the Ame1 ica n Joseph I-L Painter 0879-1 908) who co llec ted in Mexico
and d rowned in rlle Potomac river near \'V'ashington DC
Shnibs and treelets; deser1-g1asslan<l ;ind shrubland ('rnatorrnl')
Refcrence(s): 13al'neby & Grimes 0 996: 178 - 182)
Poorly distinguished from Havardia; part of the 'Pithecello bh.1n-1-alliance' of
I3a1neby & Grimes 0996)

Painteria /eptaphylla Drawing by P. Halliday Pithecel/obium dulce Photograph by D. Du Puy

208 LEGUMES OFTHE WORLD TRIBE INGEAE 209

Hesperolbizio occidentolis Photograph by c. E. Hughes Pseudosomaneo guochopele Photograph by c. E. Hugh es Albizlo multif/oro var. multif/oro Photograph by G. P. Lewis Albizia mainaea Photograph by D. Du Puy

Hesperalhizia Ba1 neby & J .W.Grim cs (1996) Afhizia Durazz. 1772

1 sp.; SW Mexico Pseudoalbizzia B1itcon & Rose (1928); Jt11hrosamanea Brillon & Rose ex Brillon
From bespero- (Gk : rhe evening star, and by ex tension the \Xlesc) and 1hc gcnc1ic & Kill ip 0936); Ba/izia llarneby & J.W.Grimes 0996)
name Albizia (q.v.) c. 120 - 140 spp ..; discontinuously circumtropica l, most di ve1.se in tropica l Ame1iG1
Trees; m:ainly seasonally chy tropical woodland, from nea1 seJ level to !500 m (22 spp.), c. 14 sp p. in S Ameiica, 6 spp. in C Ame rica, Mexico and the Ca1 ibbean
Refe1ence(s)o Ilarncby & Giimes 0996' 112 - 113) and 2 spp. w idespread in lhc Neotropics; Africa (c. 36 ende mi c .<; pp~), MadagasG11°
A member• of the 'Sama nc;J-alliance' o f lfa1 neby & G1imcs (1996) (30 spp., 24 endemic), SE Asia (cent1 ecl in Malesia [20 spp.l, India, Inclo~C hina to
Used for livestock fodde1 Chin a le. 15 spp,, the type species of the genus, .1J julihrissin Du1azz. is
w idespread in subt1opical to ternpeiate \Y/, C and E Asia]); 1 sp., endemic in N
Austra lia
Pseudosamanea Hanns 1930 Named fo 1 Filippo degli Albizzi, 18rh centllly It<11i~m naturalist, who brought seecls
2 SPf1·i one endemic to Cuba, the other wid espread from SW Mexico to NE Peru from Constantinople to Florence in 1719
From pseudo- (G k,: f;:ilse) ~incl the gcnu.'i Sc1111 cmeo (q.v.); in tlowe r easil y Trees, shrubs, <1ncl li~mas; tropical , mostly lowloncl (so metimes inundated) rn· low-
co nfused with Sa manea sc1man (J;:icq .) Men'. monta ne se;1sonall y chy ripa1fan fores t, woocll:rnd, wooclccl grnsslancl, bushland
Trees; seasonally dry t1opics: P. guacbapele (Ku nth) Harms occ u1 s in tl1ough t- and 1hicket; some restricted to J'ain fo1est; othe1s ex rrarfopical , ra1el y in desen
cleciduous woodla nd Lo 1000 m , P cuhana (Br ilton & Rose) ll~tr nc:by & j .\V Gri mes footh ill s, often in seconda ry vegetation
in wooded grassland (palm-savanna) and Jiang warcr cou 1sc.:s below 50 m Reference(s} Nielsen 0979b, 1985a, 19920 64 - 86); Huang 0983); Ila1 neby &
Hcference(s)o Ilarncby & Grimes 0996' 11 3 - ll 7) Glimes 09%0 203 - 245); Vi llie1:-; in Du Puy el al. (20020 243 - 271)
ConsidcJ'ed a good genus by B;11"11eb y & Grimes (1996) and placed in 1liei1 Albizia has become a dumping g1o und for uru e lated species in the lngeae and
'Sa manea -alliancc' needs to be monographed. Kajita el a(. (2001)~ in the k molecular Jnalysis, have
Pseuc/osamcmea guachapele (gtwch;1pele, frijol illo) is used for timhc1' in 1easonable support linking A . julibrissin (the type specie.s of Albizia) w ith
shipbuilding (planking, ri bs, decking), railw.:1y sleepers, gencn.1 1 construc1 ion, Paraserianlhes, thus suggesting a more basally brnnching posirion in the tii bc for
flooring , clecomtive veneers and furniture , ;rnd as li vestock foc.klcJ at leas t one elemen t of Alhizia (cl usleling with the O ld \Vo1l d Group in fig 27).
Luckow et al. (2003) include 8 species of !llbizia sens lat in thcir 111olecula1·
~ma lysis of the Mimosoideae and the genus, as currenrly circumsclibecl, is sh own to
Sarnanea (Be nth .) Merr. 1916 be polyphyletic; three neotropical species form a well-suppo1tecl group but th e Oki
3 spp.; mostly circum-Amazonian S Ame1ica (Venezue la, Colombia , Pei u, NE Wadel species appear as disparate elem enrs sca ttered amongst other genera of rh e
llolivia, Parnguai•, S, E and N ll1'azil) to C Ame 1irn (El Sa lvador) Ingc;.ie. The spelling 'A/bizzia' , adopted by Benth<tm in 1844 ~md much copied, is
Frnni rh e native S American name 'sa man' derived from the French C11 ihlJc~in incoi rect. Rico Arce (1999) reduced Batiz/a co a synonym of Alhizia
vein:acuh1r 'zam<1ng' or 'ra in tree' because 1he leaves fold at dusk ~ind a t the V~1dou.s sp ecies, e.g., A (ebbeck (I..) l3entll . (East Indian wa lnu t, sif'is t1ee, woman's

approach o f stornt~ lt.'t.ling the rain fa ll through ro the ground tongue), arc used for timber (constr1.1ct ion, furni ture, cabinet w01k, ve neers,
Trees; seasonall y dry 1ropic.al deciduous to moist evergreen ()1es1, woodla nd and general ca rpentry), livestock fodde1·, human food (fruit p ul p and seeds), bark (fibre
wooded grassland and pulp fo r pa per), medici ne, firewood, gums, tannins, <l yes, ink, soaps, fish
II ·rcrencc(s)1 Darn •by & Grimes (19960 117- 126) poisons, ornamentals, street and plantatio n shade tree!-i a nd for reforesca tion
111c genus w:is rcin.st:ued hy llarncby I!< G1imes ( 1996). who coiwin i1q;tly :ur;uc<l
for the recognition of th ree species
Samanea saman (Jacq.) Me l'r. ( 1ain tiee, sc1111an, rnonkeypod, coco 01 cow
1mnnrlnd) is wl<l<:ly pl:m1cd In the Id ood New W.,rld 1rople< for shad•·· :is on
15
ornamuntnl :md for le< 1m1ri1ie1\1, pod• (for """'Moc k fodder nnd chc i;wc<• pulp
a lluincrn food , also mJde into a beve1age); also used for meclicint:, ti111IJc:1
(furniture, genern l co nstruction, interior trim , box es and crares, panelling,
plywood and venee r), hats (made from w oocl shavings) ;ind bee forage (honey)

Somoneo somon Photograph by c. E. Hugh es A/biz/a vm

· 'd'ts var. zygo1des
· Photograph by D. Du Puy

210 LEGUMES OF THE WORLD TRIBE INGEAE 211

 '
I
I

Entero/obium schomburgkii Drawing by B. Angell

Enterolobium Mart. 1837

11 spp~ ; centred in S Ame1ica (4 .spp Amazonian Brazil lO neighbouring counrrie:S,
1 of which more widespread to C America and Mexico; 5 spp. in All<mtic fo1cs1 10
seasonally d1y vegeta rian in E Brazil, Parngua y , N Argemina , Uru guay and Bolivia;
1 sp. in d1yland Venezuela co Colombia and 1 sp. widesp1ead in C Ame1 ica,
Caribbea n and Mexico extending to Venezuela and Co l ombi~1)
F1'om entero- (Gk.: intestine) and fobion (Gk.: pod) in rel'eience ro eithe1 the
shape or the contenrs of the fiuit
T1·ees; tropic;:il lowland <mcl submontane rain forest (teria fi1·me), often ;dong
rivers , ancl seasona lly cJ1y low forest, woodland (ce11ado), wooded gr<oissland,
thorn scrub (ca::11inga), co<1stal fo1est (restinga) ~incl inunclntecl g1assbnd C1x1ntanal)
Reference(s): Mesqu ita (1990), Barneby & Grimes 0996: 245-251)
Apr eLJring related 10 Albizia in its bro<1dest sense on rnorphologic<1I g1ou11cls, but
its t<1 xonornic affinities arc still to be resolved
The ti111he1 of se\'e1·al species (guanacaste, baribatra, l<Jmboril, corotu, c;.11 pod
tfee) is valued for high quality furnitrne, cabinet wo1k, joine1y, panelling, 'enee1s
~ind water-resis tanr construction (canal sides and troughs for example); ~tl so used
as sh<tde trees, 0111amen1~ib :a nd for p<1stu re imp1ovem enr, livestock fodde1
(although the fruils o f some species a1c tox ic to cattle), fib1c (fo1 papc1), crnk,
so;1p substitutes, gums and medicine

Lysiloma Be nrh. 1844

8-9 spp.; centred in Mexico to C America (1 sp , wicJespre;.1d to FJ01·ida ancl
Cai ibbcan Hnd l sp. to SE Arizom1); 1 sp.. endemic to the G1e8te1 Ami lies
riom (ysis- (Gk.: loosening or separation) and loma (Gk.: border 01• m<irgin); the
fruit valves of most species .sepm<1te from the pe rs istent s urures
Trees and s l1rubs; seasonally dry tropical ancl .s11 btropica l woodland and descit
veget;:1tion at low <incl submo nt<me elevations
Reference(s): T homrson (1980), Barneby & Grimes 0996: 257-263); Gale &
Pennington (2004)
The permanent fu.o;ion of the cwo parallel sutura l ribs around the whole periphery
o f the pod distinguishes Lysiloma from c1 1l other New World Jngeae; some. bur 1101
all, sp ecies have crasped ial dehiscence; records from S America are b;,1sed on
misidemified Acacia species; gene1ic affinities of Lysiloma aie nol known but <1

possible relu r lorL~hlp wnh ll<"{ximlblzln I wo11hy of lnvt:,<tlgt11ion .

Voriou~ pcdc. ~abicu, l'7:th1111, :ibcy. fomher tree) :ire 11scd for 111111><« (in
construction, l'untllure, .1bfnc11y, noorlng, wh • I~, roles, fine • ipcnlry, .r.1f1< :ind
musical in.<trum •ms), as shndc u ecs. ornnment;ils, for :11fcy cropping, ll\' ·' fomi ng,
1rcw1xxl. fodd.,r, medicine oncl erosion o mrol: h stonc:nlfy 1hc 1imbcr of/., m/Jicli
Be nth. was one of Cuba 's most valued exports

Lysilomo tergemlno Photograph by c. E. Hughes Inga Sp. aff. densifloro Photograph by G. P. Lewis

212 LEGUMES OF THE WORLD TRIBE INGEAE 213

TRIBE
swartzieae by H. E. Ireland
'ffl. ' ,varrzieae
• . DC. 1825
.
li·ibc Tounateeae Ba1ll . (1870)
111
2
,s
e wartzieae sen . lat. , compri in 17 ene.ra a nd .
sp cie · (Fig. 28 , i largely tr picaJ and
disrributed from Mexico to Argentina and the Caribbean
ith Bobgunnia, Cordy/a, Mildbraediodendton and
Bcipbiop Is re tricl d ro tropica l · frica and Madagascar.
co~ an (198l a) included 11 g n ca in the wartz.ieae,
d1en lacer (Polhill 1,994) Iran fi rred four genera from the
phor a Ambu.rana , Ateleia, yc1tbo t.eg ta and
Holoca~)V). Bobgunnia (K.irkbrid & Wiers ma, 1997)
and '/i"iscbidiwn Ireland , submitted) w re add cl
subsequently.
The flowers of Swartzieae genera are unusual and
varied, and do not totally conform to the typical
'papilion.oid' structure, resulting in much d bal over
the ysrematic placement of the tribe. Dispariti · with
the rest of the family, of some but not all Swartzieae
taxa, include a closed calyx in bud, non-papilionaceous
corollas (often with a single petal, or these lacking
altogethe r clue to comple te loss of some petal
primordia) and polystemony (often numerous stamens
resulting from an innovative developmental feature,
the ring me ristem) (Tucker, 2003) . Although now
generally accepted to be papilionoid, the tribe has
frequently been shifted between the Papilionoideae
and the Caesalpinioideae, and is even recognised by
some as a fourth subfamily (De Candolle, 1825;
Bartling, 1830; Endlicher, 1840; Corner, 1951).
Research based on pollen (Ferguson & Sehr.ire,
1994), macromorphology (Herendeen, 1995), wood
anatomy (Gasson, 1996) and DNA sequences (Doyle et
lEFT Cordylo madogoscoriensis Photograph by G. P. Lewis
al., 1996; Ireland et al., 2000; Pennington et al., 2001)
has shown the Swartzieae to be polyphyletic, with
many members of the tribe more closely related to
genera in the Sophoreae, Dipterygeae and Dalbergieae
than they are to each other (Fig. 28). In a phylogenetic
analysis of DNA sequence data (Ireland et al., 2000;
Pennington et al., 2001), Swartzia emerges in a
monophyletic group with Bobgunnia, Bocoa,
Trischidium, Cyathostegia and Ateleia. This group of
genera, with the addition of Candol!eodendron, are
likely to constitute a redefined Swartzieae sens. strict.,
with the remaining swartzioid genera being moved to
other tribes (Fig. 28). Wojciechowski et al. (2004) find
moderate support for including Swartzieae sens. strict.
in a monophyletic clade together with basally
branching genera lacking the 50kb inversion in
Sophoreae, and Dipterygeae.
The reclassification of Swartzieae sens. strict., and
realignment of the remaining swartzioicl genera in other
tribes, needs to be corroborated by further evidence.
For the present, Swartzieae sens. lat. is retained in a
basally branching position within the Papilionoideae
following Polhill Cl981a).
[Author's postscript: Mansano et al. (2004a) recently
undertook a molecular-morphological analysis of the
Lecointea clade of Herendeen (1995) and found strong
support for the inclusion of Harleyodendron and
Exostyles within this clacle, rather than in the Vataireoid
clade as reported here]
TRIBE SWARTZIEAE 215
 Bobgunnia
Bocoa
Swartzia
Candolleodendron
Swartzieae sens. strict.
Trischidium
Cysthostegia
A Ateleia

Sophoreae sens. lat.

(non-so kb inversion
/ genera); see page 228
Dipterygeae (see page 250) Aldinoid clade pro parte;
basally branching non-
Amburana so kb inversion genera
Mlldbraediodendron
Cordyla
Aldin a

Sophoreae sens. lat.

(see page 228)

Cali a
Uribe a
(Sophoreae sens. lat.,
see page 228)
Lecointeoid clade pro parte;
basally branching so kb ~
Zollernia inversion genera '--J
Holocalyx
B Lecointea Bobgunnia madagascariensis Drawing by E. M. Stones Bobgunnia madagascariensis Photograph by a. D. Schrire

Bobgunnia J .H.Kirkbr. & Wiersema 1997

Sophoreae sens. lat. 2 spp ; Africa (1 sp, Gu ineo -Congolian WC region; 1 sp Zambezian to Sudania n
(see page 2 28) regions); no t occu rri ng in Madagascar as implied by one of the species names
Named in honour of Dr Charles Robert 'Bob' Gunn (1929-), botanist at the
Agricultural Resea rch Centre, Beltsville , USA
Sweetia Trees; trop ical rain forest (B. fist u/oides (Harms) j .H.Kirkbr. & Wiersema) and
Luetzelburgia seasonally dry tropical woodland, wooded grassland, bushland and thicket
(B. madagascariensis (Desv.) J.H.Kirkbr. & Wierse ma, or snake bean)
(Sophoreae sens. lat., Reference(s): Aubreville 0970: 298-304, as Srucll1zia); Kirkbride & Wiersema
see page 228) (1997)
and vataireoids Previously included in Swarlzia section Fistuloides, as the African species of
(see page 310) Swa11zia, and not accepted universally as a genus distinct from Swartzia
Used as tim ber (construction, furniture , cabinetry, tu rnery, carving), firewood
Vataireoid clade (c harcoal), medicine, anti-fungal agen ts, fish [JOison, insecticide, fibre (ba rk clorb)
Harleyodendron? and a dye; leaves and pods are poisono us to li vestock
c Exostyles?
Bocoa Aubl. 1775
Sophoreae sens. lat. 3 spp ; S America (I sp. Fre nch Guiana and Surinam, 2 spp Brazil, one fro m Para
and Amazonas extending to French Gu iana and Surinam, the other recorded o nly
(see page 228) from Marnnhao)
Derived from the vernacula r name 'llois !loco' (French Gu ia na) fo r the type
species B prouacensis Aubl.
Baphiopsis Trees; tropi cal Amazonian (non-inundared) rain forest (2 spp.), woodland
(ce rrado) and bushland (I sp.)
Baphioid clade Re feren ce(s): Cowan (1974); 11eland (submitted)
D Based o n mo lecular data Bocoa is closely re lated to Bobgunnia and Swmtzia
Sophoreae sens. lat. (I 1e land et al, 2000)
Used for timbe1 (furnirure, musical insuume nts) 1 medi cine, cosmetics and
(see page 228)
considered to have magical powers

? =doubtful placement of genera in this group

FIG. 28 Diagram of relationships in tribe Swartzieae after Ireland et al. (2000); Pennington et al. (2001); Wojciechowski et al. (2004). Genera traditlonallY
placed in Swartzieae now occur in at least four different positions within the Papillonoideae. Clade A comprises two separate elements, the basallY
branching tribe Swartzieae sens. strict., as well as a group of basally branching non-5okb Inversion genera sister to elements of the Sophoreae sens.
lat. and Dipterygeae (Aldinoid clade of Ireland et al., 2000, pro parte). Clade B represents elements of Sophoreae and Swartzieae sens. lat. that com prise
the basally branching genera of the 5okb inversion clade (Lecointeoid clade of Ireland et al., 2000, pro porte). Clade C comprises geneta of SwartzJeae
sens. lat. possibly sister to the Vataireoid clade, but see author's postscript on previous page, where a recent analysis (Mansano et al., 2004a) finds
Harleyodendron and Exostyles are strongly supported together with Zollernia, Holocalyx and Lecointea in the Lecointeoid clade (Clade B). Clade D contains
Bocoapr . .
Baphiopsis sister to the Baphioid clade of Sophoreae sens. lat. ouacens1s Drawing by H. Rypkema

216 LEGUMES OF THE WORLD TRIBE SWARTZIEAE 217

 Swartz/a myrtifolia var. e/egans Photograph by G. P. Lewis Candolleodendron brachystochyum Drawing by P. Halliday Trischidlum mo/le (combination in press) Drawing by P. Halliday

Swartzia Schreb. 1791 Candolleodendron R.s.cowa11 1966

Possira Aubl. (1775); To11nalea Au bl, 0775) 1 sp.; S Arnelica (French Gu iana, N and NE B1azil)
c. 180 spp.; Mexico, C America , Ca ribbea n (c 10 spp.); S Amel'ica (to SE Brazi l, Named in ho no ur o f August ine Py1amus De Candolle (1778 - 1841)
c . 170 spp) Trees; trop ical Amazo nian (non-inundated) ra in fores t
Named in honour of the Swed is h botani't O lo f Peter Swa11z (1760-1818) Heference(s): Cowan (1966); Ba 111eby & Heald (2002: 301-302)
Sh111bs, trees and lianas; tropical ra in forest and seasona ll y dry fo rest, wooded Candolleodendron brachystachyum (DC.) RS.Cowan (saboneteira) was p1eviously
grassland, often riverine, to high mo un rai n slopes, d1y scmcly plains and 10cky included in Swa rtzia; now considered to be closest to Bocoa an d Swcu1zia
beac hes (Ireland, u np ub l. data)
Refere nce(s): Cowan (1968); Cuello & Cowan in Be1•1y el al. 0999: 391i - 415);
Toi ke (2004)
Ove r 150 species desc ribed, with c. 30 species awaiting desc 1ipti on ; Cow<rn's Trischidium Tu t. 1843
(1968) r11onog1aph does no t clivkle Szva1tzia in to natu ra l g rm 1pi ngs and gh en the 5 srp; S Ame ri ca (all spr . in Bia zil , l sp . exle ncling to Paragrn1y. 1 sp . to Guy:rn a,
lack of a species level phylogeny of the genus, it is qui te p1obable rhat Su·o rlzia Bolivia and Pe ru)
wi ll be furt her segregated ::it the generic level (as in the cases of Bocoa, From lri· (Gk.: th ree) and schidio (Gk ,: fragment), 1"Cferring to the cal yx, which
Cando//eodendron , 'Jhschidi11m, G)ia /h oslegia and Bobg11 n nia) splits into thiee seg men ts
Used for rirnber (Swm1zia species vm iously known as bastard rosewood, \\'am:ira, T1ees; tropic::1 I Amazonia n ( non-inundated) m in forest (2 sppJ, coas tal forest (1
ba nnia, COJ'a'!;?l.O de neg10, <lfe used for in la y, parquet florning, turnc ry, furnitute, sp,) and seasonally diy woodland and bushland (caatinga and ceirado; 2 sp r J
c::ihinetwork, violin bows, specialcy items; suggesred ''s a subsrit ure for ebo ny); Refe1ence(s): h e land (submitted)
also used in med ici ne On the basis o f mo lecular data, Triscbidiu m is closely re lated to Ate/eia and
Cyathostegia (Ireland et al., 2000); rrev iously included in Bocoa
Used fo r med ic ine and fuel

Cyathostegia (Benth .l sche1y 1950

Swarlzia Schre b. sec t. Cyathostegia I3en th.
1 sp.; \'ifcs tern S America (inlcr-Ancl ea n valleys of S Ec uaclo1 and N Peiu)
From cyath o- (Gk: cup-) and -stegia (Gk_: shelter, covering ), referring to th e
sliape of the calyx
Trees und shru bs; se:tsona lly dry trop ictil bus hland and sh rubl a ncl, often o n 1ocky
hillsides
Reference(s): Rudel (1968a); !Jela nd (2001: 190 - 198); Lewis et al . (2003b)
This genus was previously included in the Soplloreae; see co mment o n
relationshi ps unde r Trischidi11 111

Swartzia simplex Pho tograph by G. P. Lewis Cyathostegia mathewsii Photograph by G. P. Lewis

218 LEGUMES OF THE WORLD TRIBE SWARTZIEAE 219

 Amburana cearensis Photograph by G. P. Lewis
Ateleia (Moq. & Sesse ex DC.) Ilenth. 1837
20 spp.; Mexico, C Arnerica & Caribbean (c 17 s pp.); S America (Ve nezuela to N
Argentina ; 2 spp .); J sp. widespread
From ate/eta (Gk : t:ncompleteness, impe1jeclion), referring to the tlowers lacking
four of the five petals
Trees and shrubs; tropical to subtropical seasonally d1y forest, woodland,
bushlaf'ld and xerophytic shrubland, often in rocky areas or riverin e; 1 sp. of high
humid forest
Refere nce(s): Mohlenbrock (1962b); Rudd (1968a); Ireland (2001: 199-270)
This genus was previously included in the Sopho1eae; see comment on
relationships under Triscbidium
Used for timber, insecticides and wildlife-food
Amburana Schwacke & Taub. 1894
Torresea Alle mao (1862)
3 spp.; S America (non-Amazonian Brazil, Peru, Bolivia, Parnguay, N Argentina)
Derived from 1he vernacular name 'imburana de cheiro' (Brazil), fo1 ics
resemblance to 1he imbu Frui1 (Spondias - Anacardiaceae)
Trees; seasonally thy trnpict l to subtropical forest (often on 1ocky hillsides or in
ravines), bushland and thicket (caatinga)
Reference(s): Smith (1940); Burka1t 0952: 206); Bernardi (1984); Uliballi (1997: 5)
Closely related to the following 3 geneia based on molecular ev idence (Ireland el
aL, 2000; Wojciechowski , 2003); this group is allied to a number of traditionally
sophoroid genera lack ing the 50kb inversion (Doyle et al., 1996), e.g. Dussia,
Myrospermwn1 Myroxylon, Castanospermum, Angyloca(yx and Xanthocercis,
togethe1 with the Dipterygeae (11eland et al., 2000; Penn ington et al., 2001)
Used for timber, e.g., A cearensis (Allemao) A.C.Sm. (cheny wood, ambu,.111a,
cerejeirn, ishpingo), for furn iture, construction, frames, cabinet1y, veneers and
carpentry; also L1sed as medicine
Mildbraediodendron Harms 1911
I sp.; WC Africa (Guineo-Congolian and L'ke Vicloria regions)
Named in honour of the German botanist and explorer of 1ropical Africa, Goufried
Wilhelm jolrnnnes Mildbraed (1879-1954)
Tree; tropical rai n fores t and seasonally d1y forest
Reference(s): Ilrenan (1967: 223-225); Aubreville 0970: 304 - 306)
Closely related to Cordy/a and Aldina (Pennington et al., 2001)
Mildbraediodendron excelsum Harms (muyati) is used for tirnbe1 and as wildlife-
food (especia lly elephants)
Mildbraedlodendron excelsum Drawing by E. M. Stones
220 LEGUMES OF THE WORLD
Cordy/a africana Photograph by P. Van Wyk Cordy/a africana Photograph by P. Van Wyk
Cordyla Lour. 1790
c. 7 spp.; c , 5 spp. in Africa in the Za mbezian, Sudan ian and Somalia-Masai
phytochoria of White 0983); 2 spp in Madagascar
From cordyle (Gk .: club), referring to the club-shaped Fruit
T1ees and shrubs; seasonally dry tropical fores! (often riverine), woodland,
bushland and scl'll b, often on rocky hillsides
Heference(s): Milne-Redhead 0937); \:lrern1n 0967: 221-223); Thulin 0993: 361);
Du Puy & Labat in Du Puy et al. (2002: 294 - 297)
The genus is being worked on by Ki1 kbrid e who suggests tlut two generic
elements may be involved
Used as ornamentals and shade trees, timber, medicine and human food
Aldina Encil. 1840
c 14 spp. (currently under revision by B. Sccrgios, who suggests that there are
at least 22 species); S America (Colombia, VenezL1ela [most species}, Guyana, N\V
B1azil)
Named in honour of Tobias Aldinus, an Italian medical doctor of the 17th centu1y,
auchor of Ho11us Farnesianus
Trees; rropical momane and fowl.and Ama zonian (ei ther inundated or riverine)
forest, and seasona ll y dry wooded grnssland (whi te sand savannas) and thicket
(caatinga)
Reference(s): Maguire et al. 0953); Stergios & Cowan 0999: 245-253)
Used for 1irnbe1, e g., A heterophyl/a Spruce ex Ilenth. (mac ucu de paca, angelim
eta Campina) for heavy construction, sleepers, flooring, exte1ior panelling, joine1y
a nd fencing
Aldina Ia t'' . Drawing
110 110 . by P. Halliday
.
TRIBE SWARTZIEAE 221
 Zollernio g/obro Photograph by G. P. Lewis Lecointeo omozonico Drawing by P. Halliday Horleyodendron unifo/iolotum Photograph by G. P. Lewis

Zotlernia Wie<l-Neuw. & Nees 1827 Lecointea Ducke i 922

10 spp.; S America (1 sp '1 Venez uela, Guianas; 9 spp NE to SE Biazll from Beliceodendron Lundell 0975)
Arm1zon lx1sin south to Santa Catarina State) 4 spp .; C Ame ri ca (1 sp.) and S America (Colombia, Ve nez uela, N 13razil, ECllador
Named in honou1• of Frekl1ich-\X'ilhelm Guilelmo IH of the Hollenzollc1n family and Peru )
en. 1797-1840), named after a GlStle on the hill of Zollern, in !:iOLl(hern Ge1many Named in honou r o f Paul Le Cointe (1870 - ?), Pr-ench-13.azilian botanist and director
Trees and shrubs; tropical Amazonian rain fo1esr and seasonally dry foresc, of the Museo commercial do Para
woodbnd (cerrado) ;rnd lhorn shrubland (c<ialinga) o n rocky coastal hillsides Trees; trop ica l A mazonia n inundated forest to seasonall y dry, liverine and hills ide
(restin ga) forest
Reference(s): Fil ho & Andrade 0967); Yakovlev Cl 976b); Carva lho & !3a1neby Refc1 ence(s): Yakov lev Cl976b); 13arneby 0989a)
0993); Mansano et al. (2004b, publ 2005) Pa r rebtionsh ips see under Zol/ern ia
Closely related to the fo llowing 2 genern based on molecula1 ev idence (l1cbncl el Used for timber
al., 2000) and fo I'ms a group with Urihea (Sop llo reae sens lat.). This ancl the
fo llowing 4 gene1a arc members of a well suppoitecl Lecointcoid cl;.1cle (i\fans<mo
et a l, 200/i::i), with in a broad sophoroid clade possc~si ng the SO kb in ve1sion Harleyodendron R.S Cowan 1979
(Doyle el al., 1996) I sp.; NE 13iazil (13a hia co;10l)
Used as l imber; potential as orna men tals Named in honour of Dr Raymond M. Harley 0936-), Kew botan ist specialising in

'
the B1:1 zilian flora
Small trees o r shrubs; tropical coasta l fores t on steep rocky slopes
Holocalyx Miche li 1883 Refe1 ence(s): Cowan 0979)
I sp.; NE Argentina, S Brazil anJ Paraguay Sister to Exoslyles based on molecu lar evidence (Ireland et al, 2000); well suppo1ted
Prom bolo- (Gk~ : entire) and calyx (Gk.: c11p) 1 referring to the entire c:i lrx wit h in the Lecoirneoid clade (Mansano et al., 2004a)
T1 ecs; se.a!'ionally d1y tropica l to s ubtropical riverine forest and wood land
Reference(s} !3u1ka11 0952: 188); Ulilrn•ri 0997: 6 - 7)
The tribal placemen t of /Joloca(lix is un certain, and it has often been pb rcd in the
Sopho1·eac; analysis of DNA sequence data r:d accs it with Lecointea and Zoller 11ia
( wuri~I ~le) and Urtbea (Sophoreae) (Ireland el al., 2000~ Wojcl~'('hmvsk i , 2003)
/Jaloct1/jn: l1nlf1n$.1.Ul Micl1eli (wood rosemary, rosem aty-of-c:unpina:-.-. ibir;1pepC) is
used fo r its resistant and durable timber (for constru crio n, be.:1rns, fur11iw1e •ind
carpentry), ::1s otnamenrnls and shade trees, but the leaves :i re toxic (ronwi ning
cya nogenic glycos ides)

Ho/ocolyx bolonsoe Drawing by P. Halliday Harteyodendron unifoliolotum Photograph by V. de F. Mansana

222 LEGUMES OF THE WORLD TRIBE SWARTZIEAE 223

 6

Exostyles venusta (1-4), E. amazonico (5 , 6) Drawing by P. Halliday Exostyles venusta Photograph by v. de F. Man sano

Exostyles Scho tt 1827

ft spp.; J sp Surinam to Ama zonian Brazil, 1 sp. EC Bw zil, 2 s pp SE B1 ~11_il
from exo- (Gk.: protruding ) and st)>lis (Gk.: pillai'), refen·ing to the prot1 uding style
T1 ccs or sh1ubs; tropical 1ain forest, non-inundated Aclantic coaswl fo 1c:-.1 and
resringa, and se<1sonally dry 1ropical sub1nomane forest
Jtc fc1c ncc(s): Hutchinson 0 964 : 249); Ya kovlev (1 979a); Soares·Silva & ~ Jansano
(2004); Man sano & Lewis (2004, publ. 2005)
Sister to Harleyode11ch o11 in I i eland et al. (2000)
Used fo 1 timber

Baphiopsis llenth. ex !laker 187 l

1 sp.; \VIC Africa (Gttin co-Congolian and Lake Victoria regions)
Fro m Hapbia <lnd - oµs 1~'\ (Gk.: appearance), 1efe 1ring ro irs silnil:.1rity to the genus
Bapbia (Sophoreae)
Trees 0 1 (sometimes ~candent) shru bs; tropic.ll lowland win fo1es1 0 1 ::i\\:11n p frncsl
Reference(s ), Aubreville (1970, 297-298); Y:ikovlev CI977a)
13ascd on evidence f1om pollen morphology (Fe1guso11 & Skvaila, 198 1).
macromorphology ( I-Jcrc ndeen, 1995) and D NA (Pennington el al . 2001) this
genus is beu~r placed w ith Bapbia in rhe Sophoreac seus~ fat., tlrnn wit h the
above assemblage o f genera
Us~d fo r \YOoden implemems

Baphiopsis parviflora Drawing by E. M. Stones Swartzio cf. fl oemmgu

· ·· Photograph by J. Ratter

224 LEGUMES OF THE WORLD TRIBE SWARTZIEAE 225

TRIBE
soph0 re a e by R. T. Pennington, C. H. Stirton and B. D. Schrire
tribe sophoreae Spreng. ex DC. 1825
polhill (1994) treatm nt the following informal
10
oups w r r ogni d: the Iyroxylon group 11
~ en1 ; 10 eocropic , on Africa); Orm sia group (3
11
genera; eotrop.ic , Africa , A ia); Angyl alyx group (4
!enera; eotropics Africa Au tralia); Baph.ia group (6
genera ; Africa to Asia)· ussia group 9 genera·
Neotr pies and ophora group (14 genera; Africa ,
Asia, Neotropics).
The only formal change made to the classification of
sophorea e since Polhill 0994) is the tra nsfer of
Bowringic1and Baphiastrum to Leucomphalo (Breteler,
1994b). Jn this account we maintain Bowringia and
Baphiastrum, not because we disagree with Breteler
(1994b), but in the spirit of this volume, to encourage
future workers to verify the monophyly of
J,eacomphalos sens. lat. with new data. We also do not
follow Polhill's 0994) suggestion that Riedeliella,
Etaballia and Inocarpus belong in Sophoreae. It has
been generally accepted (e.g., Polhill, 1981b) that these
belong in Dalbergieae, which is confirmed by the recent
study of Lavin et al. (2001a) that places them in the
Dalbergioid clade. They are therefore treated as
Dalbergieae in this volume (see page 307).
Cladistic analyses of overall morphology (Chappill,
1995; Herendeen, 1995) and pollen data (Ferguson et al.,
1994) showed Sophoreae to be non-monophyletic
because Swartzieae genera were mixed in the same
monophyletic groups as Sophoreae. These results have
been corroborated by molecular studies. Doyle et al.
0996) showed Sophoreae to be heterogeneous for a
large inversion in the chloroplast genome. This suggests
that Sophoreae is non-monophyletic if it is assumed that
the inversion arose only once. Doyle et al.'s 0997) DNA
sequencing study of the chloroplast gene rbcL included
18 genera of Sophoreae. Cladistic analysis showed these
to be scattered widely across the papilionoid tree. More
recently, these results have been conoborated by another
chloroplast locus, the trnL intron (Pennington et al.,
2001). This study sa mpled more putatively basal genera
of Papilionoidea (26 of 41 ophorea ; 14 of 15
Wartzj ,1e and all Dalbergieae and Dipte1ygeae). The
trnL tree (summarised in Fig. 29) is also largely congrnent
With other molecular studies that include some taxa of
ba al PapiJionoidea .g. , Hu et al. , 20 O; Ireland et al,
2000; Lavin el al., 2001a; Kajita et al., 2001; Wojci chowski
et al., 2004). fl clearly shows g nera of oph r e to be
Ill.embers of disparate papilionoid clades.
Diverse data ets now indicate Sophoreae to be non-
lb.onophyletic a 1 olhill (1981b; 1994) predicted. If the
LEFTS
ophora mtcrophy//a
. Photograph by P. Gasson
trnL results are corroborated, it seems likely that
Sophoreae will be dismembered with its genera
scattered across several tribes. This would entail
extensive taxonomic changes. Yakovlev Cl972b; 1991)
split Sophoreae into five and nine tribes respectively.
These classifications have not been widely accepted,
and although they are not congruent with the most
recent molecular topologies, they will need to be
considered in any formalisation of new tribal names. In
any new scheme, Sophoreae sens. strict. will comprise
a group of genistoid clade genera from among Polhill's
0994) Sophora group (Fig. 29), but published
molecular phylogenetic studies have not yet sampled
sufficient genera to suggest its delimitation.
A new classification for Sophoreae requires sampling
of the genera not included by Pennington et al. (2001;
see Fig. 29) and other authors, in future molecular
systematic studies. Some of the clades discovered by
DNA sequence data (Fig. 29) are cryptic in that they are
not marked by obvious macro-morphological features ,
and it is therefore perilous to attempt to determine the
affinities of genera based upon macro-morphology
alone. It may be that these clades are defined by
anatomical or chemical characters. For example,
quinolizidine alkaloid accumulation may be a
synapomorphy for the Genistoid clade (Pennington et
al., 2001; Kite & Pennington, 2003), and lack of these
chemicals in Styphnolobium species supports the
segregation of this genus from Sophora sens. strict. The
presence of quinolizidine alkaloids in Calia, which is
not placed amongst the genistoids, suggests that this
genus is a strong candidate as sister group to the
Genistoid clade, a relationship that might be resolved
by more robust molecular phylogenies. Such
phylogenies should incorporate information from
nuclear genes (Lavin et al., 1998; Doyle & Doyle, 2000)
which would be particularly useful to test hypotheses
that are currently based solely upon evidence from
chloroplast DNA. Careful integration of morphology,
preferably as part of a simultaneous cladistic analysis,
is also critical. Such morphological study may be best
achieved by focusing on separate monophyletic groups
because assessment of homology of morphological
features across all Papilionoideae is difficult. The
monophyletic groups discovered in the trnL analysis
provide a framework for statting these future studies. In
all 45 genera and (393) - 396 - (398) species are treated
here (including c. 76 basally branching, c. 262 genistoid
and c. 58 baphioid species of Sophoreae; Fig. 29).
TRIBE SOPHOREAE 227
 Dalhousiea Baphia group
Airyantha Baphia group
Leucomphalos Baphia group
Baphioid
Bowringia Baphia group
Baphiastrum** Baphia group
Baphla Baphia group
Swartzieae p.p. (see page 216, clade D)

Ormosia Ormosia group

Haplormosia** Ormosia group
Perico psis* Ormosia group
Acosmium Myroxylon group
Bowdichla Dussia group
Diplotropis** Dussia group
Clathrotropis** Dussia group
Genistoid Dussia group
Petaladenium**
Sakoanala** Sophora group
Neoharmsia** Sophora group
5okb inversion Bolusanthus Sophora group
Platycelyphium Sophora group
Dicraeopetalum Sophora group
Cadia Sophora group
Podalyrieae (see page 267)
Ammodendron* Sophora group
Ammothamnus** Sophora group
Maackia* Sophora group
Sophora Sophora group
Salweenia Sophora group
Euchresteae (see page 260)
Camoensla* Sophora group
Thermopsideae (see page 263)
Brongniartieae (see page 253)

Sweetia Vataireoid Myroxylon group

Luetzelburgia Myroxylon group
Dalbergieae p.p. (see page 309) Alexa conorocunensis Drawing by A. J. Beaumont Alexa canoracunensis Photograph by W. Milliken

Cali a Sophora group Al.exa Moq 1849

Uribea Lecointeoid Myroxylon group Alexandra R.H.Schomb. 1845, non Alexandra Bunge 1843
9 spp.; S America (Venezuela, Guyana, Surinam, Fre nc h Guia n<1, Amazonia n
Swartzieae p.p. (see page 216, clade B, but see legend
Brazil)
to Fig. 28) Substitute name fo r Alexandra R.H.Schornb, non Bunge, which w£1s dedicated to
Dalbergieae p.p. (see page 309, Andira, Hymenolobium) Alexandra, Empress of Russia
Trees; tropical lowland (occasionally upland) r• in forest
Reference(s), Yakovlev (1977b); Ram irez (1995); Stirton & Aymard in Berry et al
Cladrastis Sophora group (1999, 254 - 258)
Styphnolobium Sophora group Morphology and DNA sequence data both indicate that Alexa is closely related to,
Pickeringia (see page 263) Thermopsideae and possibly congenelic with the Australian Castanospermu.m (Pennington et al,,
2001; Stirton, pers. obs.)
Dussia Dussia group Alexa imperatricis (R.H.Schomb.) Ba ill . (haiari, haiariballi) is used as a commercial
Myrocarpus Myroxylon group timber (constn.tction, crating and plywood); species' also used for fish poisons
Myroxylon Aldinoid Myroxylon group and medicine
Myrospermum Myroxylon group
Swartzieae p.p. (see page 216, clade A) Castanospermum A Cunn. ex Hook. 1830
Monopteryx** Dussia group 1 sp.; NE Australia, SW Pacific (New Ca ledonia, Vanuatu)
Dipterygeae (see page 250) From castano- (Gk., chestnu t) and sperma (Gk., seed), referring to the
appearance of the large seeds
Alexa Angylocalyx group Trees; tropical ra in forest, alo ng rivers, strea ms and coasrs
Castanospermum Angylocalyx group Reference(s), Boland et al. (1934, 586-589)
Angylocalyx Angylocalyx group Closely related to Alexa (Sophoreae); see note thereunder
Angylocalyx group Wood used for furniture making and ve neers; C. australe A.Cunn. & C.Fraser ex
Xanthocercis
Hook (Moreton Bay chestnut or black bean) is ofte n cultivated as an ornamental
Swartzieae p.p. (see page 216, clade A) Swartzieae sens. strict. (usually as street trees); also used for medicine

CAESALPINIOID
OUTGROUPS
Amplilmas Myroxylon group
GENERA FOR WHICH NO MOLECULAR DATA ARE Panurea Dussia group
AVAi LAB LE AND WHOSE PLACEMENT IS UNCERTAIN Splrq.tro'pls Dussia group
tlleanthus Dussia group

FIG. 29 Synopsis of relationships of Sophoreae, based on cladistic analysis of nucleotide sequences of the chloroplast trnL intron (Pennington et al., 2ooi;
and unpublished data [genera marked by an asterisk*)) and of the chloroplast gene rbcL (Kajita et al., 2001). Clades (e.g., 'Genistoid', 'Aldinoid') are well·
supported monophyletic groups that have been given Informal names. Dotted lines Indicate weakly supported ctades. The node congruent with the presen~e
of the 5okb inversion In the chloroplast genome is arrowed. Genera are arranged in groups (marked byverticalllnes) that correspond to smaller monophylellC
groups in the trnL analysis. Genera marked with two asterisks("*) are those for which no molecular data are available, but for which relationships have been
inferred from other features. The system of genera as assigned to the informal groups of Polhill (1994) is also provided in the right-hand column. Castanospermum oustrale Photograph by G. P. Lewis

228 LEGUMES OF THE WORLD TRIBE SOPHOREAE 229

 Xonthocercis modogoscoriensis Photograph by D. Du Puy Oussia tessmannii Drawing by A. J. Beaumont Dussio sp. Photograph by G. P. Lewis

Atigylocalyx Taub. 1896 Dussia Krug & Urb ex Taub , 1892

7 spp.; \VIC and \YI Africa (Liberia to C 1meroon, Congo (Kinshasa) and Angolai Vexillifera Ducke (1922); Casba/ia Stancil , (1923)
and 1 sp. in Za nz ibar-Inhambane E Africa (Kenya and Tanzania) c 9 spp.; c. 4 spp. in S Mexico, C AJT1c rica and Ca ribbean and c. 5 spp. in S
From angylo- (Gk.: curved) crnd ca lyx, because of the shape of the G1lyx America (Ve nezuela , Colombia and the Gui:rnJs to C Pe n.1 and Amazonian Brazil)
Trees and s hn.1bs; rropical lowland 1ain forest and seasonally dry everg1een forest Named nfrer Anmine (Peie) Duss (1840 - 1924), Swiss clergyman and botanist in
and rhickct, often by rivers the \'Vest Indies
Reference(s): Yakovlev el al, (1968) T1 ees; tropic:il lowl:ind and upland rain forest
Closely related to Xanthocercis; see note thefeuncler ReFe 1·e nce(s) : Rucki (1963); Zamora el al (1999)
Used as medicine and for timber (construction and furniture) DNA clata (!relancl el al, 2000; Pennington el al, 2001) ploce this genus in" clade
with some Sophoreac, S\vartzieae and Dipterygeae th<1t lack the 50 kilobase
inversion (Doyle el al., 1996)
Xanthocercis B;iill. 1s70 Usec.1 f<ll" timber (local construction); the 1Jlood-1<::tl sap is sornetitncs used as
Pseuducadia l-1<.irms (1902) med ic ine
3 spp.; \VC and Za mbesian southern Africa (Gabon [1 sp.li Za mbia, Ziinhahwe,
llotswana and S Africa [1 sp.]) and Madagascai (1 sp.)
from xantbo- (G k~ : yellow) and Cercis, a genus or legumes Myrocarpus Alle111ao 1817
Trees; 11opical wet and seasonally dry lowland foresc, woodland ~ind bu shbnd, 5 spp.; S Ame rica (Venezuela [l sp.l, S and E Brazil, Bolivia, Paraguay and N
along 1ive1s and sometimes associa ted with te1mite mounds A1gentina [4 spp.D
Refei:ence(s): Dumoz-le-Grand 0952); Peltier (1972); van dcr Maesen (1997); Du F1om myron- (Gk ~: sweet oil, pel'fume) and cmpos (Gk.: fruic), refe1ting to its
l'u y & L<1bat in Du Puy el al (2002: 299-301) amnrntic fruits
lvlolecu lar data indicate this genus to be close to Angyfoca~px at the base of the TJ'ees; wet to seasonc1\ly diy tropirnl lowbncl foie.sl and woodland
Papilionoideae (Doyle el al, 1997; Pennington el al., 2001) Hefe1ence(s): Rudel (1972c); S<11tori & Azevedo Tozzi (2001; 2002)
Usecl for timber, as shelter or sh::1Je trees (especially X zamhesiaca (Riker) Por re hnionships see notes under 111y1·ospermu111.
Durnaz-le-G1anc1, the Nyala tree), ornamentals ;incl for fu c lwood; the fruits are a Used for timber (durable, fragrant Jnd of good qualiLy for construction, furnitu1"e
famine food with the pt1lp around the seeds being edible but Gtrc is nceclccl as :incl wood c:irving); essential oils (balsam) from the b;i rk (e .g ., of 1l1.frondos11s
ot her pa11s are reported to be toxic All emf10 [cabieuva]) arc used in aromatllen1py, ~II.so as medicine, perfumes ~mcl
incense

Xonthocercis modogoscoriensis Photograph by D. Du Puy Myrocarpus frondosus Photograph by G. P. Lewis

230 LEGUMESOFTHEWORLD TRIBE SOPHOREAE 231

Myroxylon balsa mum Photograph by c. E. Hugh es Myraspermum frutescens Photograph by c. E. Hughes Cladrostis sinensis Photograph by R.B.G., Kew Styphnolobium joponicum Photograph by G. P. Lewis

Myroxylon Lr 1182 Cladrastis na t, 1824

Tol11i/ era L. (1 753)
2 spp.; Mexico, C America ; wesrern and noi thein S Amelica co N A rgent in;1
f1 om my r ou- (Gk.: swccr o il , pc1fume) and :!.:y/o ,, (Gk : wood), pc11ai11ing to its
aro1mllic wood , from which both the ·nalsarn of Peru' (from M per11ifer11m L f)
and Tolu bals<tm (f1om ;11 halsa11111111 (L.) Harms) of cornmeice are cxt 1-;1eted
Trees; tropica l lowland rai n Forest to seasonally city fore.st and womllancl
Refe rence(s): Rudd (l968b): Dillon (1980); Sartori & Azevedo Tozzi (2002)
For relat io nships see notes uncler Jl 1f1 11·ospermlfm
Used fo1 timhe1 (furnirure, tumcry, inte1io r t1i111 and co nstru crion); h~ils;1m
ex tracti on fo r lll<:!dicine, pe1 fumery, thickening agents, etc.:. (cultivated in e.g.) Sri
Lrnka a ncl \YI Africa)
MyrospermumJacq. 1760
3 .srp.; N & C Mexico, C America, Caribbe~m
:111d no11hern S Ame• ic:1
From myro11- (G k.: sweet oil, pe1fume) and sperm11111 (Gk.: .seed), 1e fc..:11ing toils
:uo rnaric seeds
T1 ccs :ind shrubs; seasonally d ry cropical to subtwpical woodland ;tnd bu shh111d,
on scony hil lsides and along rive1s
Hcfcrcnce(s): Rucld (1968b); Delga clo Sa linas &Johnston (1984); S;11tori &
Azeveclo Tozzi (2002)
Mo l ec.:u l ~11 cit.Ha ind icate 111yrospe1•111111n to be in :1 monophyleric group w ith
11'ryrO.\J!/011 and J'l-~)wocc11J;11s that is lx1sally bran ching in a clade com~lining some
Sw ~1rt zieae cmd Dipterygc ae (lrel~ind el al,, 2000; Pennington el ell, 2001)
Used (spmingly) for timber
Monopteryx Sp ru ce ex Bench. 1862
3-4 .spp.; northern S A m erica (Colombia, Venezueb , French G ui~ina and
Arnnzoni;1n I3r;1zil)
from 1110110- (Gk.: single) and p t e1_l'X (Gk.: wi ng), referring lo th e 1wo- lipped
ca lyx, o f w hich the uppe1' lip is much hlrger and cove1s the flower in bud
Trees; non-inunda ted tropic.al low land rain fo1esl , o n sand y soil
Hcfe rcnce(s): Ducke (1949: 158 -159); Dufour & Zarucchi 0979); Stirton &
Ayma rcl in 13eiry el al. (1999: 355 - 356)
Affinities :ire possibly wilh the Dipterygeae ( Lima, pers comm.)
l'c.ucnl ially use ful wood for furniture and construction; th e wood is resinous
h:1s ~1 lx d:o;amic smell ; seeds edible when r oasred
Monopteryx inpae Photograph by 5. A. Mori
7 s pp ; E Asia (6 spp .; China , _Japan, KOl"ea), eostern N America (l sp ; SE USA
[from southern Illinois and lndi a11~ 1 to western N Ca 1o linc1 and souch west through
Al;i.bama , A1kansa!-; to E Ok l:1ho ma))
from dado- (Gk .: br.rnch) ancl th1<111sto- (G k,: lwittlc, fo1g ilc)
Trees; w~1rm tempe1a tc montanc eve1g 1een and mixed deciduous foJ''C S I
Hefc1cncc(s): Takeda (1913); Robeibon (1977); Ma (1982): Spongber-g & Ma
0997); And rews (1997): Dulc y & Vincent (2003)
Ana lys is or DNA sequences shows Cladrastis to be closely related to
S1yphnolohi1111i (Doyle el al, 1997; Pennington el a l , 2001) and Pickeringia
(\V'ojcicchow ski el al., 2004), the btte 1• being ti ea tcd here in the The rmopside<i e
T he woocl is used for construction and is <1 sm11 ce of a yellow dye; the N
America n C ke11/11ckea (Dum ,Cou1s) Rudd (Amc 1ican o r Kentucky yellow-wood)
is cultivared as an o r m1111ental and for landscaping
Styphnolobium Schon 1830
Sophora sect. S1_111bnolohii11J1 (Schott) Y»kovlev 0964)
9 s pp ; N & C Amc1 ica (8
spp ; S USA (from Ok laho ma and Texcis to A1 kansas and
Louisiana]; Mexico and C A 111e1ica ; l sp _ to Colo mbic\) ; \Y/ & C China (1 ~p .;
1K1tw•a lised in Japan)
Fro1n s1ypb110-, s11ypb1to- (Gk.: sour, ~1sclingcnt) and lobion (Gk: pod), alluding to
the taste of rhe Aeshy pulp of rhc pods
Ti-ec:;; tempc1;:ire ro se1son<Jlly chy tropi c~tl wood land, and lowland to montane Forest
l(derence(s): Yakovlev 0991); Sousa & Hudcl (1993)
l'vlorphological ;:111d molecular studies sl1ow ~ 1 close re b:itionship with Cladraslis
ttncl Pickeri11gia (see note unde r Cladrastis); S1ypb 11 0/obium is one of the m~1ny
gene ri c splits of Sophom 111aclc by Yakovl cv (1967)
S1ypbnolobi11 111japo11ic11111 (L) Scho1t (= Sopborc1jc1po11ica L, the Japanese
pngoch1 tree) is widely pbnted as a11 o rm1111e11 w l; also used for timber (fu111iture),
medicine, hL1man food (leaves ancl flow ers) :ind :i yellow dye wh ich w;:1s
extl'<lctcd fro m che pods and used in the .sil k ~rnd ba tik indust1ies
Calia Ter:in & Be r-land . 1832
Bro11ssone1ia O nega (1799); Den11atop1Jyl/11m Scheele (1848); Sopbora sec1iun
Ca/ia (Ter:in & Berland.) !lucid (1971)
4 spp i S USA (New Mexico to Texa s) south to C Mexico
N<lfflCd after SD. A nto nio Cal, Professor of I3otttny in th e ci ty of Puebla, Mexico
Tiees ~m ci sh rubs; wa1m temreraLc woodland, wo od ed .sh11Jhl aml , thicket
~incl (chaparrn l) and deseit; rocky slopes, grrivel washes, gu llies
Refe re nce(s): Yakovlev (1968); Sousa & Rucld (1993); lsely (1998: 843-845, "' Sop/Jora)
Cnlia sew ndiflora (Ortega) Yakovl ev (- Sopbo ra sew ndijlora (Oitega) DC.; Texas
mount<1in laur'el, me.seal bean), is culti va tt:cl as an o rnamental and foi• bndscaping
(th~ l ~u ge fed seeds are used to make neckb ce.s ;1nd a1c c.:onsidcrecl to be highly
Calio se cun d"{I
I ora Drawing by P. Halliday. tox ic to hlllnans); ~1lso used for n1 cclicinc

232 LEGUMESOFTHEWORLD TRIBE SOPHOREAE 233

 0 :1
p~

Uribea tamarindoides Drawing by A./. Beaumont Sweetia fruticosa Drawing by M. Warwick Ormosio amazonica Drawing by A. J. Beaumont Ormosia smithii Photograph by G. P. Lewis

Uribea Dugand & Romero 1962 Ormosia Jacks. 1811

Placolobium Miq (1858); Arillarla Kurz (1873); Podopelalwn F.Muell . (1882);
I sp.; C & S America (Costa Rica, Panama and Colombia)
Ormosiopsis Ducke (1925); Fedorovla Yakovlev (1971 ); Ruddia Yakovlev 0971);
Named in honour of J oaq uin Antonio Uribe (1858-1935) and his son, Lorenzo
Trichocyam os Yakovlev 0972)
Uribe 0900-1980), Colombian botanists
c 130 s pp.j Mexico, C America, Caribbea n, north ern and western S America to
Trees; tropical lowland rai n forest
Brazil and Bolivia (c . 80 spp ., mosl diverse in northern S America); SE Asia (Ind ia,
Reference(s): Holdridge & Poveda (1975); Quesada el al 0997); Weber (2001)
lndo·China, China, Malesia, Micronesia, Solomon Is .) and Australia (c, 50 spp.),
On the basis of DNA sequence data, U. Jamarindoides Dugand & Romero is
closely related to Holocalyx, Lecointea and Zollernia (Swartzieae) (Ireland el al.,
most diverse in lndo-China and China
From hormo- (Gk.: necklace, chain); the seeds are used as beads
2000; Pennington el a l, 2001), and to Exostyles and Harleyodendro11 (Ma nsano el
Trees and shrubs; Lropical rain forest (sometimes riverine) o r seasonally dry forest,
al, 2004a)
bushland, shrubland and xerophyti c woodland, on sand over limestone
Reference(s): Ducke 0939); Me 11ill & Chan (1943); Rudd 0965); Yakovlev (1971);
Stirton & Aymard in Berry el al, 0999: 364-372); Faridah Hanum (2001)
Sweetia Spreng. 1825 Ormosia is placed in the Genistoid clade based on mo lecu lar (e g ,, Doyle el al .,
Ferreilia Allemiio 0851) 1997; Pen nington el al., 2001) and chemical data (e.g., Ricke r et al,, 1999)
I sp.; S America (Bolivia, Brazil and Paraguay) A number of species (variously known as ten to, kokriki, sirari and huayniru) are
Named in honour of Robert Sweet (1783-1835), British horti cu lturnlist, bota nist used for 1he i1· durable timber (construction, furniture, carpenlry and veneer); also
and ornlth<Jlogist used for medicine and orname ntatio n (bright red seeds used as necklaces and
Trees; uopici-JI rain forest (sometimes riverine) and seasonally dry forest, bushland charms; species variously kn own as lady bug tree, jumby bead or necklace tree)
and thicket to subtropical forest
Reference(s): Yakovlev (1969)
Moleculm studies place Sweetia as part of the Vataireoid clade, Logethe1 with
Valairea, Valaireopsis and Luetzelburgia (Penni ngco n et aL, 2001); th is is
Haplormosia Haims 1915
1 sp.; WC a nd \V/ Africa (Sierra Leone Lo S Nigeria and E Cameroon to Gabon)
supported by wood anatomy (Ga'5on, 1994) and largely supported by
From baplo- (Gk.: single) and Ormosia , because it resemb les the genus Ormos;a,
morphological studies (Lima , 1982b; 1990), bt1t is co ntrary to its tradition<il
but has simple leaves
placement close to Acosmium
Trees and sh iubsi tropical lowland evergreen forest, a long rive1s and in swampy
Sweetiafruticosa Spreng. (mani) is conside red a future crop cree for logging in
llolivia valleys
Refere nce(s) : Voorhoeve (1965); Burkill 0995: 361); Faridah Hanum (2001)
Morphology indicates a close rela tionship with Ormosia (Sopho reae) but no
Luetzelburgia Harms 1922 molecula r data are available
The timber of H. monophy//a (Harms) Harms (black gum) is used for
8 spp.; Brazil
Named in honour of Phillip von Luetzelburg (1880-1948), German botanist and constm ction, fu rniture and canoes
explorer, who collected widely in NW Brazil
Trees and shrubs; seasonally dry tropical, lowland woodland and wooded
grassland, occasionally lowland rain forest
Reference(s): Yakovlev 0976a); Rizzini & Mattos Filho 0 977); Lima (1983a)
Analysis of DNA data places this genus in the informal Vataireoid clade, close to
Valairea, Vataireopsis and Sweelia (Pennington el al., 2001)
Used for timber, fuelwood; roots milled for fiour as a famin e food

Luetze/burgia auricu/ata Drawing by P. Halliday Haplormosia monophylla Drawing by L. van der Riet

TRIBE SOPHOREAE 235

234 LEGUMES OF THE WORLD
Perlcopsis ongolensis Drawing by D. Bridson Acosmium brochystochyum Photograph by G. P. Lewis
Pericopsis Thwa ites 1864
Ajrormosia Harms (1906)
4 spp.; Africa (3 spp; Zambezian/ Sudanian [2 spp.] and Guinea-Congolian WC (1
sp.] regions), Sri Lanka, Malesia to Papuasia (1 sp .)
From pericopto- (Gk.: cur ~ II round), 1efe1Ting to the deep cuts in the Gilyx
Trees and shrubs; tropica l lowhmd forest ( riverine or mangrove in Asian spp.) or
seasonally dry woodland and wooded g1assla nd
Referencc(s} Knaap van Mecuwen (1962); Yakovlev (1978); Faridah Hanum (2001)
Tradirionally chis genus was considered to b e close to Ormosia (Sophorcae). This
is con fiimed by molecular data, which place both genera in the Gen isto id clade
(Doyle et al., 1997; Pennington et al., 2001)
Various species (satin wood; afrormosia, obang, nandu, nedun) provide excellent
timber which is a subsritute for teak in cabinet work and furniture rnuking; also
used as a popular medicine and as an arrow poison; P angolensis (13:-iker) va n
Meeuwen (nmvange, 1nbanga) is a multi-purpose species, being used for charcoal,
crafts, domestic uses (pest les, mo1tais), fencing (post.s), firewood, furniture, gum,
land improvement (nil1 ogen fixing), medicine, rituals (used to cha.se aw<1y
witchcraft) and timbe1 (e.g., 1<1ilway sleepers)
Acosmium Schott 1827
leptolobi11m Vogel (1837)
17 spp.; S Mexico, C America, S America (Brazil [majority of spp.l to Paraguay and
N Argentina)
From a- (Gk., without) and -cosmos (Gk,, beauty, rnder, elegance)
Trees and shrubs; seasonally dry tropical woodland and wooded grassland,
occasionally lowland forest and dunelands
Reference(s} Yakovlev (1969); Bridgewater & Stirton 0997); Aymard & Gonza lez
(2003)
Previously considered basally b1anching within the Sophoreae; analys is of DNA
data show it to be closer to ge ne1a in the Genistoid clade (Doyle el al, 1997;
Pennington et al., 2001)
Acosmium panamense (Ilenth.) Yakovlev (Ilil!y We b, ca1 boncillo) is an important
hardwood noted for its strength and du1ability; the inner bark (cascara amarga)
conrains an alkaloid used for medicine and as an anri-1m1larial; it is also the mai n
ingredient of 'Sweet Blood', a comn1ercial remedy sold for diabetes
Acosmium lentiscifo/ium Drawing by A. J. Beaumont
236 LEGUMES OF THE WORLD
Bowdich/a vlrgilioides Drawing by A. J. Beaumont Bowdich/a virgilioides Photograph by G. P. Lewis
Bowdichia Kunth 1824
2 spp.i northern S America (Colombia, Venezuela, Guyana, Su1inam to C Brazil)
Named in honourof1110mas Edward Bowdich 0791-1824), early plant collector
in \Y/ Africa, for his many scientific achievements
Trees and shrubs; tropical, seasonally dry lowland wooded grassland, grassland
and bushland, occasionally lowland fo rest
Reference(s} Yakovlev (l 972a); Ferguson & Stilton 0 993)
Molecular data indicate that Bowdicbia is a member of the Genistoid clade
(Pennington et al., 2001)
Bowdicbia nitida Benth~ (sucupi ra) is an important timber used in construction
(flooring, beams and f1ames), also used for charcoal and as fish poisons
Diplotropis Benth . 1837
Dibracbion Tul (1843)
12 spp.; S America, mostly Amazonian basin (Colombia, Ve nezuela, Surinam,
Guyana, French Guiana, Brazil, Ecuador, Pem and Bolivia)
From diplo- (Gk , double) and rropis (GL keel), the wing and keel petals are
similar in size and shape
Trees and shrubs; tropical rain forest (riverine, immdaled and non-inundated) to
seasonally dry forest, wooded grassland and bushland
Reference(s), Yakovlev (1972a); Lima 0985a); Lima & Ayrnard in Berry et al.
(1999, 315 - 318)
T1aditionally this genus has been placed with Pmmrea, Bowdicbia and
Clathrotropis (Sophoreae), for which Yakovlev 0991) e 1ec ted the tribe
Diplotropoideae; Diplotropis has not been included in any morphological or
molecular daclistic analyses, however, and its 1elationships remain uncertain
Used for timber, e.g, D pwpurect (Ricl1) Amshoff (suct1pira; tatabt1; boto nallare)
in heavy const ruction, shipbuilding, flooring, turn el)' and manufacture of
furniture, tool handles and railway sleepers); also used as a mammal poison
Diplotrap/s racemosa Drawing by A. J. Beaumont
TRIBE SOPHOREAE 237
 Petolodenium urceo/iferum Drawing by A./. Beaumont Neoharmsia baronii Photograph by G. P. Lewis Neohormsia baron ii Photograph by J. -N. Labat

Clathrotropis (Be n th ) Hai ms 1901 Neoharmsia R.Vig. 1951

DijJlolropis sect. Clalb m lmpis llenth. (1862) 2 spp.; N and \VI Madagascar
From neo- (Gk.: new, pertaining to) and honouring He rm ann August Theodor
6 spp.; Ca ribbean, western and nmthe1n S America (Pell\, Colombia, Venez uela,
Hanns (1870 - 1942), Professor o f Botany in Berlin
Suri nam, Gu yana , French G uia na :ind I3ra zil)
Trees or sh n1bs; seasonally dry tropical woodland and succulent vegetation, o n
f'ro m clathron- (G k.: latti ces o r pierced w ich o p enings) and tropis (Gk : keel) ,
refe1ring to rhe opening between the incompletely joined petals
1i mestone or coasta I sands
Reference(s): Peltier 0972); Du Puy & L'bat in Du Puy el al. (2002: 307- 311)
Trees; rropical lowbncl (ofte n riverine :and seasomdly inundated) 1t1in fo rest,
Both species rare; no molecular cl~lta are available for this genus but Du Puy &
occasionally upland or monrane Forest; al so se~1 so nally city woodland :incl
Labat in Du Puy et al, (2002) note that it is closely re lated to Sakoanala
shrubland
Wood of N. baronii (Drake) R,Vi g. ex M,Peltier (manango na) is used for
Rcfe rence(s): Ducke (1949: 151-152); Stirton & Ay rnard in Berry el al (1999:
278- 280) construction, carpemry, firewood and cha rcoal
Gcneially considered to be close to Dip/ot1•opis; mo lecular datJ suppo 11 ;i n ~1ffinity
w ith lhe genistoids ;.m cl possibl y basally branching to Tirongnia11ieac (Pe nnington
el at., 2001) Bolusanthus Harms 1906
Used as timber, e g., C. brachypeta/a (Tul.) Kl e inhoonte (aromata; ;.1lnw. 11cg1a) fo1 1 sp.; Zambezian southern Afri ca (Za mbia, Malawi, Mozambique, E Zimba bwe, E
constnJCtion, boat building, ful'n iture, flooring ;ind railway sleepe is); <1lso used for Botswana, Swaziland and NE South Africa)
medicine and a mammal po iso n Named in hono ur of Prnfessor Hany Bolus (1834 - 1911) , distinguished South
African botanist, and anthos (Gk,: flowe r)
Treesi seasonall y <lty lmpical to subt ro pical woodland and wooded grnssland o n
Petaladenium Ducke 1938 clay soils
Reference(s): Harms (1906); Yakovlev (1972a); Este rhu yse (1994); Venter & Vente r
1 sp.; Biazil (Rio Negro river)
(1996); Brummitt in Brummitt el at. (submitted)
Frompelalon- (Gk.: pet;i l) " '"' adell (Gk.: gland) , refeiring to tbe gland ulai pecals
Molecular dara place this genu s close to Platycelypbium and Dicraeopetalwn in
Trees; tropical lowland 1·ain forest
Referencc(s): Ducke (1949: 152) the Genistoicl clade (Pennington el al., 2001)
Bolusarzthus speciosus (Bolus) Harms (t ree wisteria, Vanwyks hout) is cultivated as
Morphologica lly, this genus appears close to, and is rossibly congeneric with,
an ornamen ttil and for landscaping; th e wood is used for furniture, househ old
Clalbrotropis (Sophoreae) (St i1to n, pers. obs); thel'e are ve ry fe,v collections of
Pelaladenhtm 11rceo/iferum Ducke, none o f them made s ince 1966 articles and fence posts; the roots and bti rk are used for medicine

Sakoanala R.Vi g. 1951

2 spp.; Macbgascal' (1 sp in coastal E Madagascar; the other in I\l\X' & \X1
Madagascar)
Derived from the Malag'1sy ve rnacular name sa koanala for s. madagascariensis
RVig
Trees; t1·opic~l lowland humid coastal frn'est (1 .sp.); other species in season:illy
cliy wood land o n limestone
llefcrun< -ls)! Y. k<Wlc1• (11)8()~): Du l'uy & l;ih:u in Ou 1'\1y l!I nl, (l!l()l: 3 11 ~ ;.l• >
N(l lllf)k cul"r UHi I ore ~v:111,,1:>1e ror 1hi~ gcnu:s. hut Polhill\ 198 lb: 19')-1) ph•~"'d It
in the Sopho ra group o f Soplloreae
Wood of S madagascariensis is a goocl constru ct io n timber

Sakoonala vil/osa subsp. vil/oso (A-L), S. villosa subsp. menabeensis

(M), S. modogascoriensis (N- S). Drawing by A./. Beaumont Botusanthus speciosus Photograph by P. Van Wyk

238 LEGUMES OF THE WORLD TRIBE SOPHOREAE 239

Plotycelyphium voense Drawing by c. Grey-Wilson Plotyce/yphium voense Photograph by B. Hucks Cad/a purpurea Drawing by P. Halliday Cod/a purpureo Photograph by S. Collenette

Platycelyphium Harms 1905 Cadia Forssk 1775

7 spp.; Madagascar (6 spp.), NE Africa (montane Ethiopia, Somalia and Kenya) ro
1 sp.; NE and E Africa (Tanzania, Kenya, E Ethiopia and S Somalia)
SW A1abia (I sp.; C. p111purea (G.Piccioli) Aiton)
From plaly- (Gk.: bl'Oad) and celyphos (Gk.: pod), re ferring to the wide pod
From kadi or badie (arabic: judge or inspi1 acional guide); Forssk<il refers to 1he
Trees and shrubs; seasonally dry tropical bushland
Arab belie f that young leaves, when pressed on the body, are said co lift the
Reference(s): van der Maesen 0970); Polhill in Milne-Redhead & Polhill (1971:
38-40); Van Wyk et al 0993 b); Thulin 0993: 391) spiri ts
Trees and shrubs; seasonally dry tropical plateau woodland to submontane forest,
Molecular data indicate a close relationship with Bolusanrbus ~incl Dicraeopetalmn
1 sp. in humid eve rgreen co deciduous co;:1srn l forest
within th e Genistoid clade (Pennington el al., 2001)
Re ference(s): van cler Maesen (1970); Gillett in Milne-Redhead & Polhill 0971:
Platycelypbiwn uoense (Engl.) Wild (saman samando) is a potential forage and
33 - 35); Du Puy & Labat in Du Puy et al. (2002 : 314-318)
browse crop for livestock in arid and semi-arid areas
A number of Madagascan species are r~ue; fl owers of Cadia are morphologically
imermediare between the Caesalpinioideae and Papilionoideae (Tucker, 2002c);
the genus has sometimes been placed in the Cae:;~ lpinioicleae, but molecular data
Dicraeopetalum Harms 1902 clearly place it in the Genistoid clade o f the Papilionoicleae (Doyle et al., 1997)
Lovanafia M.Peltier 0972) and basally branching to the Pocbly1ieae (e,g., Pe nnington el al., 2001; Kajita et
3 spp.; NE Africa (S Somalia, E Ethiopia and N Kenya (1 spJ) and S & SW al., 2001 ; Wink & Mohamed, 2003)
Madagascar (2 spp.) The bark and roots are used as fish poisonsi C. pedicellata Baker is recorded <ls
From dicraeo- (Gk.: forked) and pelalon (Gk.: petal), referring to the slender petal
havi ng medicinal uses
claws th<it point fo1ward or are somewhat spreading
Trees and shmbs; seasonally dry tropical bushland, s hrubland and xerophyric
woodland, on sa nd over limestone
Reference(s): van der Maesen (1970); Yakovlev 0977a); Thul in 0993: 390-391);
Du Puy & Labat in Du Puy et al. (2002: 305-307)
Morphologically this genus is similar to Acosmium (Sop horeae) from S Amelica1
but molecular data shows a closer relationship with Bolusantbus and
Platyce(ypbium (Sophoreae) (Pennington et al., 2001)
Wood used for tools and construction; the bark of D. stipulare Harms is used for
tanning

Dicroeopetolum copuronionum (A-J), D. mohofo/iense (K-U). Cad/

Drawing by M. Tebbs. ,o sp. cf. pubescens Photograph by D. Du Puy

240 LEGUMES OF THE WORLD TRIBE SOPHOREAE 241

 Sophora a/opecuroides Photograph by G. P. Lewis Sophora mlcraphylla Photograph by G. P. Lewis
Ammodendron conollyi Drawing by P. Halliday Ammothamnus gibbasus Drawing by L. M. Ripley

Ammodendron Fisch. ex DC. 1825 Sopbora L. 1753

Edwan/sia Salish. (1808); Vexibia Ra f. (1825); Keyserlingia 13unge (1872);
c, 4 - 5 spp.; C & W Asia (Iran, Afghanistan to Uzbekistan and Ka zakhstan) to NW
Echinosopbora Nakai 0923)
China CXinjiang Prov.)
c, 50 spp.; SE Eu1·ope to W, C & E Asia and south th1ough tropical regions to
From ammo- (Gk,: sand) and dendron (Gk-: tree), referring to th e species' habitat
Australasia and the Pacific; c . . 3-4 spp. in sect. Edwardsia in western S America
Trees and shrubs; dry continental temperate habitats and desert
(Chile, Arge ntina and Juan Fernandez Is.); largely introduced in Africa; 1 sp
Reference(s): Yakovlev et al. (1982; 1996)
endemic from coastal Kenya south ro S Africa and Mad aga sca1~ S. tomentosa L.
The genus (based on a sample of A . argenteunl Kuntze) is sister to Sophora in the
widespread in the coastal Palaeotropics and also in coasral E Brazil
analysis of Wojciechowski et al. (2004)
Derived from rhe Arabic sufayra, th e name for a legumino us tree
Used as sand stabilise rs in desert regions, dyes, ornamentals and bee crops
Trees, shn1bs and he1 bs; seasonally chy tropical to warm te mperate lowland a nd
upland forest or d1y vegetation types and sand dunes
Reference(s): Yakovlev (1967; 1980b); Yakovlev & Malov (1981); Tsoong & Ma
Ammothamnus Bunge 1847 (1981); Hulme (1995); Hurr el al (1999); Heenen et al. (2001); Mitchell & Hee nen
c. 2 spp.; C Asia (Kazakhstan, Tadzhikistan, Uzbekistan and Turkmenistan) (2002); Brummitt in Brummitt et al (submitted)
From ammo- (Gk.: sand) and tbamnos (Gk.: shrub), referring to the habitat in Some authors maintain a broad circumscription of Sophora; Y:akovlev (1967),
which it grows however, split it into a number o f ge nera and the1 e is n tend ency to recognise
Shrubs (dwarf); dry continental temperate grassland and desert some of these segregates, e g., C'11ia and Stypbnolobiwn have been shown by
Reference(s): Yakovlev (1975); Yakovlev et al. 0996) molecular and morphological data to be distinct genera; sect. Edwardsia (c. 16-18
Some authors consider this genus to be a section within Sophora (e g., S. gibbosa spp. with various hybrids in New Zealand) is most distinctive biogeographically,
(DC.) Yakovlev and S. pacbycmpa C.A.Mey. from W Asia and the Middle East), comprising the Australasian, Pacific, Mascarenes and S A1ne rican element of th e
but within Hs area of distribution it is regarded as a distinct genus genus; the centre of the remaining species is China, India and Indo-China . On
Used for medicine, dyes, forage, orname ntals, preventing erosion and as bee crops the basis of DNA sequence data Sopbora sens. stn'cl. be longs with the Genistoid
1
for honey dade (Doyle el al, 1997; Pennington et al., 2001; Kajita et al.. 2001;
Wojciechows ki et al., 2004)
Cultivated as ornament<ils (in sect. Edwardsia, with ye llow flowers, e .g., S.
Maackia Rupr. & Maxim. 1856 tetraptera J.F.Mill [kowhai] and S microphylla Aiton [small-leaved kowhai]; and in
c, 8 spp.; E Asia (S Russian Far East, China, Korea and Japan south to Taiwan) sect. Pseudosophora, with blue to white flowers, e .g, S. davidii (fornch .) Skeels);
Named in honour of Rkhard Maack (1825- 1886), an Estonian-bo rn Russian also used for its durable timber (for bearings, turnery and cabinet work), as
na turalisl based in St. Petersburg who introduced many plants into cultivation medicine (e g , S jlauescens Aito n [ku s hen], S. tonkinensis Gagnep. [shan do u
Trees and shrubs; warm (from humid subtropical) to continenral temperate gen) and S. micropbylla); some species are toxic and used as insecticides
deciduous broad-leaved forest and open woodland, on plains and uplands
Refere nce(s): Takeda (1913); Hoshi & Ohashi (1987); Andrews (1997)
On the bnsl s of ONA Wllll, M(lnckfa is . member of th!! Cenistold ( /ucJc, d <>.<C ro
!Wp/.)(lrt1 (e.g., J)Qylc (!{al .. 19')7; K:1jl1:i ct al., 200 I; \\'link & Mo hanu.-tl, 2003) :ind
M. t111111ru11sls b sl~1cr to ·numnop.~ldeae (e .g., Kallt;1 "'ol,, 2001; Wo j<;:l1:d1owsld <if
"'·· 2004)
Used :\S o rn.1men1als, e .g., M. amurensis Rupr. & Maxim. (Chinese yellow-wood;
Amur m;m kla); wood used for interior finish of houses, fumicure, utensils ~nd
handles of implements; plants also used for forage, to prevent e rosio n an<l as a
bee crop

50
Maackia hupehensis Drawing by P. Halliday Phora velutlna subsp. zimbabweensis Photograph by R. Zabeau

TRIBE SOPHOREAE 243

242 LEGUMES OFTHEWORLD
 i
J•,
.< l
;I
;}

'~

0
Salweenia ward/I Drawing by A. J. Beaumont Salweenia wardii Photograph by x. Zhu Comaensia scandens Photograph by G. P. Lewis Dolhousiea bracteata Illustration from Roxburgh, Pl. Coromondel

Salweenia Bake1 r. 1935 Camoensia welw. ex Benth. 1865

1 sp.; N\V China (Tibet, W Sichu;rn) 2 spp.; \VIC Africa to Angola
Na med afrer chc Sal wee n gorge in the Chando Disrrict of T ibet, w here ir was first Nmncd in ho nour o f Luis de Ca moens (1 524 - 1580), Portug uese e pic poel and
collected soldier; he sailed w ith Vasco da G<l ma o n his voy<lges of discove1y
Shn.1bs (evergreen); diy conti nen tal temperate monlane regions, th ickets o n dry Shrubs and lianas; rropical lowland 1ain forest
stony slope!;, o n gravel rerraces and in sand ~long streams Reference(s): Toussain t (1953); Hepper 0958: 510)
Referen ce(s): Bakc1 0935); Ying el al. 0 993) Some authors h<tvc suggested that this genus might belong in the
Polhill 098lb; 1994) placed this genus in th e Sopbora gro up of Sophorc"c; Caesalp inioicleae, but recent mo lecu lar evidence places it within the Geni.sto id
molecu lar data indica te that it belongs to the Gen istoicJ clade, with Sa/11•ee11ia clade (Penn ington, unpub l.)
wardii Baker f. sister to 1'.1aackia amurensis in th e ana lys is of Kajita et of. (2001) Camoens/a scandens (\Velw.) J~B .Gill e lt has been w ide ly introduced in the t1opics
as an o rn ame ntal because of its 10 cm w ide, w hite, butterf1y-shapcd frag1 ~mt
flowers w ith c1·inkled, l'eddish- to goklen-margined peta ls

Dalhousiea wall. ex !lenth. 1837

2 - 3 S[lp.; \VIC Africa O sp. a1ound Gulf of Guinea), NE India (t ropical E
Himalaya) and Bangladesh (1-2 sp p.)
Named in honour o f James Andrew Ramsey (1812 - 1860), Ma1quis of Dalho usie,
and Govc1 nor Gene 101l o f India (1848- 1856)
Trees 1 shrubs and lianas; tropica l lowland ra in fo rest
Refeience(s): Panig1ahi 0975, rubL 1976)
See note under Baphia; placed in the 13ttphia group of Sophore:1 e by Polhill
(1981bi 1991); molecula1· Jata contlrm its 1·ela1ionship w ith Bapbia 1:md 1elatives
(Ire land el al., 2000; Pennington el al., 2001)

Airyantha Ilrummitt 1968

2 spp.; WC Africa (1 sp.) and Malesia (Borneo and Philippines, 1 sp .)
Named in ho nou 1· of Herbert Kenneth Airy Shaw 0902 - 1985), botanist
specialising in Malesian flora and nomenclatl1rc, and anthos (Gk.: tlowe1•)
Trees, shrubs and li ~ n tts; trop ical lowland min forest
Rcference(s): Bn11nmitt (l968b)
See note u nder BajJhia; molecular data place Ai1ya11 l bt1 in a strongly su pprnted
dadc w ith Baphia, Dalbousiea, Leucompbalos (Sophoreae) and BajJhiojJsis
(Swal1Zieae) (Pennington el al.. 200 1)
Used for medicine

Sa/weenia wardii Photograph by x. Zhu Airyantho schweinfurthii Photograph ex Wageningen, Netherlands

244 LEGUMES OF THE WORLD TRIBE SOPHOREAE 245

Leucompha/os capparideus Drawing by H. de Vries Bowring/a discolor Drawing by H. de Vries Baphiastrum brochycorpum
Leucomphalos Benth. ex Planch 1848
1 sp.; \Y/C Africa (E Nigeria, Cameroon, Gabon and Equatorial Gu inea)
From /euco- (Gk.: w hite) and ompha/os (GL navel), referring to the w hite aril on
the seed
Lianas or small trees; tropical lowland rain frn est
Reference(s): !3reteler Cl994b)
llreteler (l994b) included Bowringia and Baphiastnmz (Sophoreae) under
Leucompbafos, increasing the number of species in this genus to six; [he traditional
view is maintained here, however, pending review of the entire Baphioicl clade;
see also under Bapbia for discussion on systematic re l~ti onsh ips
Bowringia Champ. ex llenth. 1852
4 spp.; \Y/C Africa (\YI Liberia to Angola) and Madagascar (3 spp., B mildbraedii
Harms disjunct between \Y/C Africa and Madagascar); SE Asia (SE China, Inclo-
China and Malaysia [!3omeo], l sp.)
Named in honour of Sir j . Bowring 0792 - 1892) and l1is son, C !3owring (1821-
1893) plant collecto rs in Hong Kong and China
Shrubs and lianasi tropical lowhmd rain forest and seasonal forest, often ;dong rivers
Refeo·ence(s} !3reteler 0994b)
!3reteler Cl994b) included Bowringia unde r Leucompha/os (see note under that
genus) and also described a new African species wh ich there fore does not have a
co mbination in Bown·ngia
Baphia Afzel ex Lodd , 1820
47 spp.; tropical Africa (most spp. in \Y/C Africa), E South Africa (! sp.), \YI
Madagascar (2 spp., one of which is disjunct to W Africa)
Fmm hapto- (Gk.: to dye); the dye obtained from the wood is bright reel
Trees, shrubs and lianas; tropical lowland min forest and :seasona lly dry forest,
woodland, wooded grassland, bushland and thicket
Reference(s} Soladoye (1985); Du Puy & Labat in Du Puy et al (2002' 301-303);
Brummitt in Brummitt et al. (submitted)
This genus is not monophyletic (Pennington el al, 2001) and requires revision .
Analysis of DNA sequence data places Baphia in a clade with Dalhousiea,
AllJv11111Ja and IJtllCQmfJhfllO> CSophore:icl ond lk1/J/1fO(>S£< (Sw:111zk,,.c) .
(l'ennfng1on ut al.. 2001), 1hls dadc is pl:ia'<.I well :•P"t1 from 1hc gcnbwicls, sister
:ind 1>osslhly lx1s:1lly hmn hing to tl1c pr<:doonlna11tl)• Old \'(lorl(( cladc comprising
the mirl.>eliolc.L<, nlllleuioicls, plmscolol ls :ind llnlog:ilcgin:i (I' ~mlngion et al.,
2001; Wojciechowski et al., 2004); this relationship is suppo1ted by pollen
(Ferguson et al., 1994) and morphology, since all 111<'111hcrs of 1his group have
unifoliolale leaves. Yakovlev 0991) recognised this group of gener<1 as a distinct
tribe !3aph <tac (excluding /Ji1p/1lupsts) .
Used as orn:11rn:n1:1ls Cshncl.: or Street rrccs), for medicine , timber (e.g., construcuon,
boat building, 111rn l)', (':t binetoy :ind t<>ol handles), a dye (e.g., B. nitida Lacki.,
Amph'1
Baphio modogoscariensis Photograph by G. P. Lewis camwood) is sometimes used in cosmetics, firewood, charcoal and forage
246 LEGUMESOFTHEWORLD
Photograph ex Wageningen, Netherlands Baphiostrum brochycorpum Photograph ex Wageningen, Netherlands
Baphiastrum Hao ms 191 3
I s p.; \Y/C Africa (Ca meroon, Gabon, Congo [!3razzaville], Congo [Kinshasa] and
Central Afoican Republic)
From Bapbia and -astrum (Gk .: simila1 co); the genus is morphologically similar
to Baphia
Lianas; tropica l lowland gallery and semi-deciduous forest
Reference(s): !3reteler Cl994b)
13reteler (1994b) included Baphiastrum under Leucompha/os (see note under that
genus)
Genera of uncertain affinity
Amphimas Pie11e ex Hanns 1906
3 - 4 spp.; \Y/C Africa
From amphi- (Gk, all oound) and himas (Gk, strap, l;ish), referring to the strap-
like Jobes of the petals surrounding the central part of the flower
Trees; dense tropical low land rain forest and seasona ll y dry frnest and bushland
References' \Y/ilczek 0952); Aubreville (1970, 29 - 32)
Placed by Polhill (1994) with Riedelie/la, /noca1pus and Etabal/ia, these three now
in the Dalbergieae sens. lat_, and on the basis of noral morphology Amphimas
does appear to group with these, however molecular ev idence is lacking and the
relalionships of this genus <ire still to be resolved Evidence from wood anatomy
(llaretta-Kuipers, 198L 686), chemistry and leaf morphology (Polhill, 198lb, 217)
indicate an affinity to Myrocmpus (Sophoreae) and Mildbraediodendron and
Cordy/a (Swartzieae)
Ampbimaspterocarpoides Harms (bokanga, lat i, yaya) is used as medicine (the
bark contH ins a red sticky resin); toxic alkaloids t.1re also present; the timbe1• is
used in construction, planking, joinery and veneers
mas pterocorpoides Drawing by M. Boutique
TRIBE SOPHOREAE 247
Ponureo /ongifolio Drawing by A. J. Beaumont Spirotropis longifolio Drawing by A. J. Beaumont

Panurea Sprnce ex 13enth. 1865

2 spp.; S America (Colombi:i :in cl N 13r:izil)
Named after the typ e loca lity, Panure, in N Brazil
Tiecs; t1opical lowl ~md fo 1est (Amazonian caa tingas) on white s:incl
J(eference(s} Yakovlcv Cl977a)
The systematic rc\;n ionships o f Panurea are uncc1tain and one of the two species
is ~ 1 s yet unde!<iclibecl; the distinct ive pollen resembles that of Howdicbia
(ferguson & Sti r1on, 1993)

Spirotropis Tul . 1844

2 or 3 spr.; northc 1n S A111erie<1 (Yenezu eli.1 , Guya na, Surinam and Fr<::nch Guiana)
Fmm speiro- (Gk.: coil) and tropis (Gk ,: keel), 1efen·ing to the twisted keel
Trees and li:inas; tropic<1 l low land to m ontane, ri ve1ine foresc
J(cferencc(s} Sti r1on & Aymarcl in lleny et al (1999, 391- 392)

Uleanthus Hanns 1905

1 sp.; S A1ncriG1 (Arnazoni:rn Brazil)
N<imcd in honour of Ernst Heinrich Georg Ulc (1854-1915), 13r:.tzilian bo ta nist
Trees; rr opical lowland fo1esr, along margins of rivers :.ind waterfalls
Rcfcrence(s} Du cke (1 949: 158)
This genus has nevc1· been included in any comprehensive molccld<tr 01
morphological :.1nrilyscs; it has some intlorescence and florn1 features in crn rnnon
with A11gylocalyx (Sti 1tori, pers. obs.), but sy napo morphies to assc1t re l;1 tion ships
w ith :.111r other genera are :.1s yet Licking

50
Uleonthus erythrinoides Drawing by A. J. Beaumont Phoro tomentoso subsp. tomentoso Photograph by D. Du Puy

248 LEGUMESOFTHEWORLD TRIBE SOPHOREAE 249

 TRIBE
/ Dipterygeae byJ.Barham
Tribe Diptetygeae Polhill 1981, as 'Dipteryxeae'
The ea rli e r affinitie s of this grou p of gen era are
discussed by Po lhill (1 98 1c). The a n alysis o f
Pennington et al. (2001) confirms that the three genera
of Dipterygeae form a monophyletic group which is
sister to eleme nts within Swartzieae sens. lat. and
Sophoreae sens. lat. (Fig. 30). The Amazonian genus
Monopteryx (treated here in trib e Sophoreae) may
belong with this group (Lima, p ers. comm .). Two of the
three genera in Dipte1ygeae are essentially Amazonian
-
---
!
I Mildbraediodendron, Cordyla, Aldina
FIG. 30 Diagram of relationships of tribe Dipterygeae to neighbouring tribes after Pennington
FIG. 31 Diagram of relationships within tribe Dipterygeae after Pennington
250 LEGUMES OF THE WORLD
wet forest i.n cti tl'ibocion (Dipte1yx a lso oc urring i .
C ncral America w ith a sin le pecies to drier ar as i~
central Brazil, Bolivia and !)araguay , whi le Pterodon
inhabits the d rier areas of a t and central Brazil an,d
ea t Bolivia . Wood anaLomy ha prov d diagno lie
parc icL1larly in di. tingu i bing berw n Dipte1y.'I: anci
Tttralea (Gasson, 1999), and Lhis has u nderpinn d u1e
concl usions of Lewi & Gas on 2000). Th tribe as
tre Led here comprise c. 22 species.
Swartzieae sens. strict. (see page 216)
Sophoreae sens. lat., e.g. Alexa,
Angylocalyx, Castanospermum,
Xanthocercis, Styphnolobium,
Cladrastis (see page 228)
Taro/ea steyermarkii Drawing by E. Fro eschner Pteradan abruptus Photograph by G. P. Lewis
Dipterygeae Tara"lea Aubl. 1775
c. 7 spp.; Amazo nian S America (Venezuela (4 spp.), the Guianas, Amazonia n
Brazil, Columbia and Peru)
Sophoreae sens. lat., e.g. Dussia, From tara la, the vernacular na me given by the Galibis in the Guianas
Trees and sh ru bs; tropical lowland rain forest (rive rine and non-inu ndated o r
Myroxylon, Monopteryx (see page 228) inundated pla ins) to seasonally d ry forest (sometimes on white sand), woodland
and marshes; also 1- 2 spp. in montane forest and open moist wo odland to scrub
on sandstone
Swartzieae sens. lat., e.g. Amburana, Reference(s): Ducke 0934a; 1940); Lima & Ayma rd in Berry et al. (1999: 416 - 419)
The very hard wood is suitable fo r construction and fi rewood; T oppositifolia
Aubl. (cuma ru-da-praia) contains an un scemed o il e xtracted for industrial use
(see page 216)
Pterodon Vogel 1837
Remainder of Papilionoideae c . 3 spp.; S America (EC states of Braz il and E Bol ivia)
(diagnosed with 5okb Inversion) From ptero- (Gk : winged) and odonto (Gk : tooth), re ferring to the two large
wing-like (petalo id) teeth of the calyx
Trees; seasonally d ry trop ical forest, woodland (ce rrado) and thorn shrnbla nd
(caatinga), often associa ted wi th rocky outcrops
Refe rence(s): Pedersoli et al. 0976)
et al. (2001); Wojciechowski et al. (2004) The very hard wood is suita ble for items requiri ng durnb ility; also used fo r
medicine, firewood , fodde r, as ornamenta ls and for refo1estat ion
Dipteryx Schreb. 1791
Coumarou na Aubl. (1775); O/eiocarpus Dwyer (1965)
c. 12 sp p.; S America (mostly Amazon ian [4 spp . in Ve nezuela]; 2 spp. to C
America (Pa nama to Hondu1as]; 1 sp. to drier a reas of C Bra zil , E Bolivia and
Paraguay)
From di- (Gk,: two) and pteryx (Gk ,: w ing), referring to the two large w ing-like
Tara lea 0 (petaloid) lobes of the calyx
"'O Trees; tropical rain forest (riverine and usua ll y non-inundated) ro seasonally dry
--l fores t (some o n w hite sand) and woodland (cerrado)
m Reference(s): Ducke (1940); Flores (1992); Lima in Beny el al. (1999: 318-320);
Pterodon ;:o
Lewis & Gasson (2000)
-<
G) A subgroup of c. 3 species produce fragrant coumarin-yield ing seeds, e .g., D .
m odorata (Au bl) Willd, (cumaru, tonka bean, sarrapia, almendro) is a major source
)> of coumarin whose vani lla-like fragra nce is used fo r scenting tobacco, snuff and
Dipteryx m confecLion ery, and is an ingredien t of perfumes a nd cosmetics; the beans also
yie ld a h igh percentage of solid fa t o r ton ka butter wh ich is used to flavour food ;
plan rs produce balsam resins (oleoresin) and red g ums from the leaves, ste ms and
bark; 1 species bears pods with a sweet, ed ible pericarp; the hard wood is
gene1all y used for heavy constru ctio n, fl ooring, shafts, bearin gs, veneers and in
et al. (2001) Oipteryx a/ata Photograph by G. P. Lewis craft objects; also used for cha1 coal and med icine
TRIBE DIPTERYGEAE 251
TRIBE
srongniartieae by J. H. Ross and M. D. Crisp

. 'be Bron niartieae 13cnth . Hutch. 1964

1rl .ribe ;ilcg •'1 subuibe Bro ng n.iartiinae Benth. (1865), as 'Brongniartieae'
1

The tribe is geographically disjunct, compnsmg a
araphyletic group of tropical American genera
rcyclolobiun·1, Poecilanthe, Harpalyce and Brongniartia),
and nested within it, a monophyletic group of Australian
genera (Templetonia , Hovea, Cristonia, Tbinicola and
umprolohium). The Australian genus Plagiocarpus
apperu: to be more closely related to the neotropical
Brongniartitt than to the other Australian genera.
Arroyo (1981) considered the tribe to comprise the
two genera Brongniartia and Harpalyce, both from
the Neotropics, so that the concept of Brongniartieae
has expanded considerably . The Australian
Templetonia group was transferred on the basis of a
cladistic analysis of morphology (Crisp & Weston,
1987), and recently two new genera have been
described in this group, Cristonia and Tbinicola (Ross,
ZOOla). Two more neotropical genera (Cyclolobium
and Poecilanthe) have been transferred into the tribe
on the evidence of DNA sequences and phytochemistry
(Crisp et al., 2000; Hu et al., 2002). A putative new
genus from Bahia, Brazil is under study by Queiroz,
Lewis and Wojciechowski. Current molecular data
indicate its sister relationships are to Harpalyce and
Poecilanthe, but formal description of the new genus
awaits further analysis.
Polhill (1994) placed Brongniartieae next to
Bossiaeeae, partly because Crisp & Weston (1987)
removed the Australian Templetonia group from the
latter and placed it in Brongniartieae. Polhill (I.e.),
however, noted that their very different alkaloid
profiles suggested Brongniartieae and the Templetonia
group had an affinity to the genistoid tribes, while
tribe Bossiaeeae was more closely related to Mirbelieae,
Hypocalyptus (now in Hypocalypteae), a group of Old
World tropical tribes (including Indigofereae, Millettieae
and Phaseoleae) and the Hologalegina group of tribes.
Recent phylogenetic analyses, especially those using
DNA sequences (e.g., Crisp et al., 2000; Doyle et al.,
2000; Hu, 2000; Pennington et al., 2000a; Hu et al.,
2000; 2002; Wojciechowski et al., 2004) support a
placement of Brongniartieae either next to, or within,
the main clade of genistoid tribes, which includes
Sophoreae sens. lat., Euchresteae, Thermopsideae,
Podalyrieae, Crotalarieae and Genisteae. Pennington et
al. (2001), Kajita et al. (2001) and Wink & Mohamed
(2003) place Brongniartieae sister to a clade including
Sophoreae sens. strict. , Thermopsideae, Podalyrieae,
Crotalarieae and Genisteae. The tribe as treated here
comprises 10 genera (not including the putative new
genus) and c. 152 species (Fig. 32).

Sophoreae sens. strict., Euchresteae,

Thermopsideae, Podalyrieae,
Crotalarieae, Genisteae

Cyclolobium
z
Poecllanthe fT1
~
Genus nov.? ~
0 OJ
;:o ;::c
Harpalyce r
0
Cl
z
Brongniartia G'l
z
Plaglocarpus
)>
;::c
Templetonia )> -i
c IT1
i'-lovea l/l
-l )>
;:o IT1
)>
Cristonia r
)>
ThinicaJa

Lamprolobium

FIG. 32 Diagram of relationships in tribe Brongniartleae after Crisp et al. (2000); Thompson et al. (2001); Hu et al. (2002)
LEFT Hovea pungens Photograph by M, D. Crisp

TRIBE BRONGNIARTIEAE 253

{
I

Cyc/olobium brasiliense Photograph by G. P. Lewis Harpalyce arborescens Photograph by C. E. Hughes Harpalyce sp. (from Cuba) Photograph by G. P. Lewis

Cyclolobium 13cnth 1837 Harpalyce M0<;. & Sesse ex DC. 1825

l sp.; S AmeriGl ancl :m me wbat disjunc t (I3r~1 z il [from H.ond6ni<1 ;111cl Go i;ls cast to 2/i spp_; amphitropi cal and somewh at disjunct: 7 spp. in S M exico, C Ameri ca,
I3ahi;i ;incl south to Sft o P;.\Lllo, M~lto Gros.so do Sul ~111d Par;Jnfll, P ~1ra g ut1y Cai ibbean (Cuba); c. 8 spp. in E Brn zil
[Co rdillera de Altos and Ybycur National Pai kJ , Bolivia [Santa Cru z]) Named after a G reek hunting goddess of great beauty, vario usly considered Lo be
Fl o m cyclo- (Gk,: circle) and lobion (Gk.: pod), refer r ing to tile distinct l\'e circular, the claug l1te r o f Cly menus or Harpcdyc11s; coincidentall y the G reek haipy rnet1 ns
fbttened, jndehiscent , na 1r owly w inged p ods twisted 0 1 co nto rted and could possib ly l'efer to the tw iste<l keels present in the
Small trees; seasonally dry lropical forest, o fcen along ri ver banks nowc rs or most species
Refe1encc(s), Hoe hne (194 1a); Warwick & Pe nningto n (2002); K ite (2005) Shrubs o r sm:tll trees; sea sonally d1y l 1o pical lO w arm-tempe rate humid forest,
Diagnosed by unifolio l<lte leaves (shared wi th some other gencrn in the u·ibe) and woodland , bu shl:tnd and thick et, shrublancJ and grassland; m ost species are
distinguished from the cl osely related Poecila nlhe by the orbic:ul ;.1r, i mlc hiscent, evergre en and fl ower during th e dry sea.son
stipitate, membranous pod , with nJrrowly winged sutu1es; placed in 1\Jillettieae Refeiencc (s), Arroyo 0976); G1 e inwald et al. (1996)
until r'ecently, but DNA :-;equenccs cle:nly indicate a position in B1 o ngnia1ticac A rnorphol ogically well ci1tumsclibeci genus with no evid ent close extant 1elative
(Crisp eJ al., 2000; Hu el al., 2002); re ce ntly redlJCCd from five 10 o ne species, C. and w hose :.1ffinities arc not cle;-t r; disting uished fr'Orn o ther gene1a of
brasiliense Denth (\Xl<1rw ick & Penningto n , 2002) llrong nia1cie:1e by rhe large pelt:He glands on many parts o f the plant, largely
Used as 01 namcntals resupinat c flowers, strongly bilabiate calyx and helicall y com o rtecl keel
(scconda1 il y .stra ightened in 3 to 'i species), Species fall in ru ughl y equal numbers
within three regio ns, Mexico-C Am e1ic:1, Cuba and E Brn zil
Poecilanthe Ilcnth 1860 The wood is used for small tL11 'n t ry ite ms
c. 10 - 12 spp.; S Am erirn (mostly l3razil, Sul'inam)
Fr o m poecilo- (Gk_: vmi-coloured) and antbos (G k ,: flower)
Trees untl shrubs: U'Oi>iC!ll Jnwlnn I min forest •\m:iwnfnn basin • i><'Cle> it""'p);
Si."US<ln:tlly dry 1ropl1::1l for= I w<ll'.l<llancl and thlcl<ct, nr rh·crlnc fo1cM (P. llrnzil
10 Uru guay and Argentina species group)
Refere ncc(s} Amshoff 0939); Owen (1 989) : Lewis (1989b); Lav in & Sousa 0995);
G te imvalcl el al. 0 995)
Diagnosed hy the vex illary stamen conmlte with the others zind fndt s thickly
w oody zincl clehi sce nt, o ft en explosively; pl aced in va1ious t1ibes including
Millenie<1e but D NA .<;equenccs cleaf'ly inc.li c~He a position in B1 o ngni;11licae (Ciisr
et al., 2000; Hu et al., 2000; see also La v in & Sousa, 1995); Poecila11J/w r oinprise.s
two distinct groups o f species
Used as timber, medici ne, anti~fungal agents ~md ornament~1 ls

Poecilanthe fa/cata Photograph by G. P. Lewis Harpa/yce brasiliana Photograph by G. P. Lewis

254 LEGUMES OF THE WORLD TRIBE BRONGNIARTIEAE 255

 Hovea /ongifolio var. montana Photograph by G. P. Lewis Hovea trlspermo Photograph by B. Fuhrer
Plogiacarpus axi/laris Drawing by P. Halliday

Hovea R,Br. ex W.T.Aiton 1812

Brongniartia Kunih 1824
c. 63 spp.; amphitropical and somewhat disjunct: SC USA (Texas), N and C
Mexico (c 98% of genus occur.i in Mexico) and C Am erica, w it h c. 2 spp. in
western (Andean) S America (llolivia and Chile)
Named for Adolphe 111 'od re Brongniart (1801-1876), a distinguisl1ed French
botanist and palaeontologist
Shrubs or small trees; tropical to warm-temperate (often montane) forest,
woodland, bushland, thicket, shrubland and grassland, mostly in doy aoeos
Re ference(s): l\ydberg 0923); McVaugh (1987: 297 - 321); Greinwakl el al. (1996)
Brongniarlla is distinguished readily from Hmpatyce as the vexillary st:11nen is
free (except in B. ulbricbiana Harms from Bolivia), th e flowers :.ire not
resupinate, the keel is not helica ll y contorted and plants lack peltate i;bnds;
closely relc1ted to the Australian genern Plagiocmpus, Lamprolobium, Cristonia
and T1Jinicola, and possibly paraphyletic if Plagiocc11p11s is incl uded CJ'hompson
el al., 2001); the genus is being revised by O.Dorado in Mexico
Used as orrnunentals
Plagiocarpus Benth. 1873
1 sp_; N Auslralia (the Kimberley region of \V/ Australia lO western Queensland)
From plagio- (Gk.: o blique) and ca1pos (Gk frnit) , in reference to the obliquely
ellipsoid pods
Shrubs or subshrubs; seasonally dry tropical to subtropical woodland, bushland and
thicket. shrubland ~ind g1:tssland, associated usually w ith sandstone or s[l ndy soils
Reference(s): Ross (1992)
Distlngmshcd by the sessile mostly cligitately 3-foliolate leaves, suhsessi le solitary
axi11:1ry nowers With linear bracteoies, pods less than twice '1.S long <IS broad, and
seeds with nearly circular arils; closely related to, and possibly ne.sted within,
Brongniartia (111o rnpson el al., 2001)
37 spp.; Australia (widespread, but absent fro m the centoal Eremaean Zone)
Named fo r the Polish collector An ton Pantaleon Hoveau (la te o· Anthony Hove) fro m
\V/arsaw, who was sent to India via Namibia in 1795, ro collect plants for Kew
Subshrubs, shrubs and smnll trees; tropical to warm temperate, coastal to monlane
foresl, woodbncl, bush land , lhicket and m ed iterra n e~tn shrubland
Reference(s): Stanley & Ross (1983: 270-272); Weber (1986: 692-693); Wheeler
in Marchant elal (1987: 268-269); Elliot &Jones (1990, 5: 378-384); Ross in
Wa lsh & Entwisle (1996: 804 -808); Thompson (2001); James in Harden (2002:
487 -496)
Distingu ished by having usually blue or mauve (except for markings) flowers,
simple leaves, pods not or scarcely Jonge1 than broad, and the aril 3 or more
times as Jong as broad (except in H /ong1jJes Benth.); see note under
Templetonia ; the greate.st species divers ity occurs in E Australia with ~1 second
cenr1 e in the SW coiner of \'V Australia
Used as 01 namentals (e g., H . ell1plica (Smith) DC. or 'tree hovea' has attractive
blue flowers); seeds and pods used as human food; the foliage of some species is
tox ic to livestock

Tempktonia R.Br. ex W.T.Aiton 1812

10 spp.; Auslralia (widespread on the 1nainland, and o n some islands off rhc W
Australian coast; absent from NE Queensland)
Named after the Irish botanist John Templeton (1776-1825)
Shrubs or subshrubs; mainly subtropical ro warm t emper~ue forest, wood land,
bushland and thicket, ;ind mediterrn nean shrubland
Reference(s): Ross in Jessop 0981: 150-151); Ross (]982); Stanley & Ross 09 83 '
268); Weber 0986: 696-699); Wheeler in Marchant el al. (1987: 298-299); RoSS
in Walsh and Entwisle (1996: 802-804); James in Harden (2002: 186 -lo87)
Distinguished by the usually ovate papery l.11'1•"1t.'Oles and the ]J(ICl.s being 1nore
than twice as long as broad; Thompson el al. 2001) found this gi:11us lO be sister
to Hovea; flowers of T. re/Ilsa (page 259) are adapted for biod pollinatio n
Used as o rnamentals. particularly T.. retusa (Vent.) R Dr (parrot-bush. c:ock ies
Templeton/a oculeota Photograph by M. D. Crisp Hovea chorizemifolia Photograph by G. P. Lewis
tongue), which has brgc mtrnctive red fl owers; also used for medicine

TRIBE BRONGNIARTIEAE 257

256 LEGUMES OF THE WORLD
Cristonio bi/obo Drawing by A. M. Podwyszynski Thinico/o incona Drawing by A. M. Podwyszynski

Cristonia J.H .Ross 2001

1 sp .i \Y./ Aust1 ~1lia (coasl<ll phiin tmd ranges N o f Pc 11h)
The narne is <l conrrnction and acknowledges the cool! ibu1io ns of bo1anis1.s
Michael D Crisp (Canbeua) and Pete1 H Weston (Sydney)
Sh1uh; 1nediten«rnean fore:->t, woodland and shruhland) fo vouling s:indy soil and
hiterite
Hefc1 ence(s): Ross (1982 , as Temp/e/011ia bi/ohu (Benth,) Polhill); !loss (200la)
Distinguished from all species of 1"empletonia, in which genus it was f"u1meil)1
[Jl:lc.:ed, by the simple leaves that are LJSLJally distinctly biJolJeci tlpinilly, lhe 2
upper calyx lobes unirecl into ;J tl'uoc:1te limb, and the linc;:ir he1baccous
b1act eoles; cli sringuishccl from othe1 Australian genera in B1ongni~11tic:1c by the
combination of simple leaves, yellow and purplish-bi own co 1ollas, c1lyces that
a1~ not lx1so.lly circum.':icissile, ancl oblong pods; ~1ffinities <lre with lr1111/;ro/obi11m

Thinicola J H.Ross 2001

l sp. , \VI Ausrrn lia (NC area of tile S1<1tc)
F1·om lbini11m (L.: dune) and -cola (L: dweller), in 1cfe1ence 10 1he p1x.>fe1rcd
hahi1a1 of T incmw (JJ·I Ross) J.I-LHoss on 1he ci esr of sa nd dunes
Shiub; tropical to subtropical shruhland and sand y dese1ts; occurs in deep icd
sand on the crest of sa nd dunes
Refercnce(s): Ross (1980, as Te111p/e/011ia inca11a .J !'I .Ros>); Ross (2001a)
Distinguished by the dense spreading silve1y ioclumc nrum on th e vcgdati\'C p:iris,
rccliccls and ex rern;:1J su1face of the calyx ; large. persist ent , foliaceous stipulcs;
l~rge , p encluluu.i.; fl owers with basicllly J'ed pe1als, s1and<.1rcls with conspicuous
au1 ides m the apex of the claw, ancJ linear h erbaceous IJrncteoles; ~dfinitic~ :ii'c
with Lamprolohium; the flowers are adapted for bird polli11 ;,1tion

Lamprolobium !Jenrh. 1864

2 spp ; Au stralia (NE Queensl:rnd)
from /ampro - (Gk.: shiny) and /o/Jion (Gk*: pod), in referenc.:e to th~ shi ny
g labrous pods
1
Shrubs; seasonally dry u opic.:al forest , woodland, bushbnd 1 ch icket, shruhl<ind an(
grassland
lkfcrcnc :{s): Jlcndc~n (199 1); mum &Jone.< ( 1993, (,, •l7- 8)
D~inj(ulltht'tl ,,_.,,c1ny by 1hc cln:um;I lie • lyx; cl scly rcbtc<l 10 Cri:J!olllfl ond
71Jinicola (Thompson el al ., 2001)

Lomprolobium fruticosum Drawing by P. Halliday Temp/etonia retusa Photograph by G. P. Lewis

258 LEGUMES OF THE WORLD TRIBE BRONGNIARTIEAE 259

TRIBE
Euchresteae byH. Ohashi
Tribe Euchresteae (Nakai) H.Ohashi 1973
Tribe Dalbergieae subtribe Euchrestinae Nakai (1940)
Polhill Cl981r) followed Ohashi (1973b) in recognising
the monogeneric tribe Euchresteae based on the genus
Euchresta (c. 4 species).
Euchresta was previously included in subtribe
Geoffroeeae of tribe Dalbergieae and was believed to
be intermediate between this and subtribe
Lonchocarpinae (Bentham, 1860, as 'Lonchocarpeae').
Recent molecular evidence places Euchresta in the
Sophora group of the core genistoids (Doyle et al.,
1997; 2000; Crisp et al., 2000), supporting previous
hypotheses of a close affinity with Sophora (Ohashi &
Sohma, 1970; Ohashi, 1973b; Polhill (198lr: 402);
Polhill, 1994; Chen et al., 1992; Matsuura et al., 1994).
Crisp et al. (2000) note evidence that strongly
FIG. 33 Diagram of relationships of tribe Euchresteae to neighbouring tribes mainly after Crisp et al. (2000); Kajita et al. (2001)
260 LEGUMES OF THE WORLD
upports th groupi~g f Euchresta and ·ophora, and
the ana ly i c f Kapta et al. (2001) ven s LLggests
Eucbresta may be mb dded within opbora. As
traditionaJJy circumscribed, ophora is polyphyletic
(Kass & Wink, 1997; Crisp et al., 2000; Pennington et
al., 2001) with certain elements, e.g., Styphnolobium
and Calia probably falling outside the Genistoid clade
(Wojciechowski et al., 2004).
A new, more narrowly circumscribed Sophoreae
sens. strict. will thus have to include Euchresteae
(providing the type species of Sophora, S. tomentosa
L., belongs with the taxa so far sampled from subgenus
Sophora (sensu Tsoong & Ma, 1981).
Euchresta japonica Photograph by H. Ohashi Euchresta formosana Photograph by H. Ohashi
Euchresta Benn. 1840
4 spp.; Indian subcontinen t (Bhutan, Assam), Jndo-China (Thailand, Vietnam);
China (3 spp ., centie of distribu tion), E Asia (Taiwan, Ko1ea [Cheju Doi, Japan)
and Malesia (Sumatra, Java, Bali, Philippines)
From euchresto- (Gk: usefu l), with reference to the medi cinal use of the seeds
Shrubs; seasonally dry tropic-JI rnonrane to warm tempera te evergreen forest, in
Brongniartieae (see page 253) undergrowth on damp humic slopes
Reference(s), Ohashi & Sohma (1970); Ohashi & Kurosawa (1979); Chen et al.
(1992); Matsuura et al (1994); Ohashi el al. (1999a)
The rhizomes of E. japonica Hook.f. ex Regel (shan dou geog) are used for
treating diseases of the mouth and throat in China and japan and seed extracts of
Sophoreae sens. strict. (see page 228) E ho!!fieldii (Lesch.) Benn. a re a tonic in Indonesia
Euchresta EUCH RESTEAE
Thermopsideae (see page 263)
Podalyrieae (see page 267)
Crotalarieae (see page 273)
Genisteae (see page 283)
Euchresta hors{ieldii Drawing by P. Halliday
TRIBE EUCHRESTEAE 261
TRIBE
Tbermopsideae by J.M. Lock

. ·rJicrniopsiclcae Yakovlev 1972

T111be

'f ditionally, the mall tril ' Tb rm psideae includ
: gen era with a toraJ. o'. (43)-45- (46) spe ·ies
r cl rhrougb th med1rerran an and temp rate
sea lre
regions of America, tbe Meditemrnean Basin and C
ro NE ASia (Turner, 1981). Recent molecular data,
nQwever no w places Pickeringia Wilhin the Cladrasc:is-
s1yphnolo bi ~1 m clade in the basal Papili noideae
(Wojciech w kl el al., 2004). 17.1ermopsis .it. el f o
in borh N America and E A ia, and Bc1pt'isic1 is
exclusively N American. The two species of Anagyris
occur around the Mediterranean Basin and in
Macaronesia. Piptanthus and Ammopiptant!nts occur in
E Asia. The tribe as a whole is absent from the S
Hemisphere. The free stamens have always marked
the tribe ;1s distinct, and have led some to associate it
with Sophoreae. Its ea rliest placement (e.g., Bentham,
1865) wcis in his Podalyrieae, in which he included
C)'clopia ancl Podalyria , as well as many genera now
placed in Mirbelieae. However, anatomical and
phytochcmical work (summarised by Turner, 1981)
points to a closer relationship with Genisteae.
Molecular analyses (e.g., Pennington et al., 2001;
Kajita et al., 2001; Wojciechowski, 2003) and studies
combining molecular and phytochemical data (e .g .,
Kass & Wink, 1995; Wink & Mohamed , 2003), all
associated Thermopsicleae with Genisteae, and placed
them as part of the 'core genistoids' (Thermopsideae,
Sophoreae sens . strict., Euchresteae, Podalyrieae,
Crotalarieae and Genisteae). The Thermopsideae form
a clacle sister to Sophoreae sens. strict. in some analyses
ur (e.g., Crisp et al., 2000 [where Thermopsideae is
stro ngly supported as monophyletic] and Wink &
Mohamed, 2003). In other analyses, however, a
monophyletic Thermopsicleae is not supported (e.g.,
Kass & Wink, 1995; Kajita et al., 2001; Wojciechowski
et al., 2004). In the latter analysis elements of the tribe
are nested separately within a paraphyletic Sophoreae
sens. strict. The combined Sophoreae sens. strict.-
Thermopsic.leae clade is sister to a Podalyri eae-
Crotalarieae-Genisteae clacle. Ammopiptanthus does
not appear to have been sequenced to elate and its
placement here is based on comments by Turner
(1981). A recircurnscribecl Thermopsicleae, excluding
Pickeringia (Fig. 34), thus reqt1 ires further study to
ascertain its tribal composition and validity.

Pickeringia

Sophoreae sens. strict. (see page 228)

Euchresteae (see page 260)

-i
Ammopiptanthus? :r:
rn
Anagyris ;;o
Piptanthus $
0
-0
Vl
Thermopsis 0
rn
Baptisia )>
rn

Podalyrieae (see page 267)

Crotalarieae (see page 273)
Genisteae (see page 283)
? ~Placement in tribe uncertain

FIG, 34 Diagram of relationships in tribe Thermopsideae after Turner (1981); Crisp et al. (2000); Wink & Mohamed (2003); Wojciechowski et al. (2004)
LEFT Th
ermapsis barbata Photograph by P. J. Cribb

TRIBE THERMOPSIDEAE 263

 Ammopiptonthus mongolicus Drawing by unknown artist Plptanthus nepalensis Photograph by G. P. Lewis Thermopsis rhombifo/ia var. montono Photograph by M. Wojciechowski

Pickerlngia Nutt. ex Torr. & A.G 1~1y 1840 Piptanthus Sweet 1828
I sp; N America (W USA [California to Baja Ca lifornia! and adjacent ~ lexico)
2 spp.; S Hirnabyas (India , I3huw n, Ne pal, Myanmar (13urma] and \Y/ Chin a)
Named after Charles Pickering (1805-1878), American hot<mist, zoologist and F1'om piplo- (Gk.: fa ll) and an/hos (Gk.: flower); the flo1al parts foll togethe1 after
anthropologist anthesis so rh e1'e is no calyx on the young pod
Shru bs; mecliterranean bushland (chaparral), open wootlland :md thicket, often Shrubs; montane grassl~md, thicket and forest margins
on dry slopes and in w:Jshes Reference(s): Tu mer (1980)
Refe rence(s): Hickman 0993: 641); lsely (1998: 801 - 802) Peng & Yuan (1992) recognise thre e species
Two varieries are recognised; Tu1 ne1 ' (1981) considered this robe ;111 isobtcd Piptanlhus neprilensis (Hook ,) Sweet (evergreen laburnum) is grown as an
genus with ri base chromosome number of x = 7 co mpared to x = 9 or 10 in the 0 1rnimc ntal

rest of the rribe; lupine alkaloids are .:dso reportedly absent in Pfr;ke1·f11gia ancl
pie.sent in othe r gene ra which have been tested; Wojciechowski el rJ/ , (2004) have
Pickeringia nested, with strong .<;upport, in rhe Cl::iclrnstis -Styphnolobium cla<le in
Thermopsis lLBr. ex W.T.Aiton 1811
rhe ba.sal Papilio noideae c. 23 spp. [but see below]; C ancl E Asia (Uzbekistan and Kazakhstan to Mongo lia,
Pickeringin m o11Jana NurL ex Torr. & A.Gray (chaparrnl pea) is a wildli fe (deer) Russia [Siberia and Far Eastl, Chim, Japan and south to the Himakiyas [13 spp D
foiage and N Amclic1 (\Y/ Canada to S Dakora, New Mexico and C\lifornia; anll N
Carolina and Tennessee to Alabama and Georgia k . 10 spp.)
Fmm thermos (Gk.: Jupin) and -apsis (Gk.: appearance), refeffing to the
Ammopiptanthus s .H,Che ng 1959 1escmbl::tnce of rhe yellow-flowere d inflore.scenccs to lupins
Herbs; ma inly n-1ontane warm ;md contincnral tempe1':ite to suha1cric woodland ,
1-2 spp; C Asia (Kyrgyzstan and W China 10 Mongol ia)
foiest margins, shmbland, grassland, desert orncl nmdi"a (in As ia) and
Fro1n ammo- (Gk.: sa nd) a nd Piptanlbus (q.v.)
mediterr~mean shrubland and thi cke t (charparal), forest, woodland and grassland
Shrubs; conrine nrnl rempcra rc shrublancl, montan e dese1t ;ind open com1nuni tie5
of stony c.lry pl ~ 1 ces and rocks (in N America)
Refere nce(s): Peng & Yuan 0992); Che n el al. 0994); Ya kovlev el al. 0996); Iscly
Refe re nce(s): Che ng (1959); Peng & Yuan 0992); Yakovlev el al. (1996)
Used as ~in o rnamental and A ~ mongolicus (Maxim. ex Kom.) SJ-I.C heng is a 0998: 863-87 1)
lsc ly Cl 998) considers the genus to comprise 8-10 srecics ancl treats the
so urce of antifreeze proteins for reducing the freezing point of solutions
Ame1ican taxa as Ii species (2 in \YI USA [ancl adjace nt Canada! and 2 in SE USA)
Used as ornamentals (e.g., T rho111hifo/ia (Pursh) Ric hal'<lson va r. monlana (Nutt.)
Isley, golden pea or golden banner); other 'fol.'ie lupin' species are so metimes
Anagyrls L 1753
weeds an<l a1e used in medicine and for e1osion control
2 spp.; Meditenane~in and \Y/ Asia (Cana1y Is., S Europe, N Africa , Tu rkey. Middle
E:tst to Iinn, Sa udi Arabia a nd N Yemen); introduced in Malll itius
Fwm anagyros the Greek name for A Joelie/a L. (stinking bea n tJ'efoH); ,ii.so
Baptisia VenL 1808
a11a- (Gk.: again) a nd gyro (Gk.: round), refe rring to 1he curved r ods
15 - 17 s pr.; E (SE coasta l rlain) a nd C USA (SC Si;Ites to slightl y adjacent Canada)
Shrubs; mediterranean bushland, deciduous woo<lkrnd and thicker, on rocky
From baplo-, haplein (Gk.: to dip, to dip in dye , colour); the plants have been
slopes
used as a n jndigo substirurc
Reference(s): Perez clc Paz (1975)
Herbs; con rincncal a nd wa1m tempe1me grassland and open woodla nd, along
Toxic ~ind thus unpalatable to stock, lea ves and seeds used for medicine; al.so
st1 e<1ms ;;incl in sandy or JOcky areas
used as ornamen tals and inclicatol's of degraded Janel
l\ercrencc(s): L>uisey (1940); Jsely 0998: 454 - 464)
fn1 e1'.s pecific hyb ridisation is comm o n
Used as 0 1 narnentals (e.g., B . austrafis (L.) R Br,, or blue wilcl indigo), fo1 · ground
cove!', me dicine ancl dyes (e.g~, B linctoria (L) RJJ 1•., oi· false indigo); unpal~tab l e
to s1ock

Bapt' ·
Anogyris foetido Photograph by J. Hooper isia austro/is Photograph by I. K. Ferguson

264 LEGUMES OF THE WORLD TRIBE THERMOPSIDEAE 265

TR IBE
podalyrieae byB.-E.vanwyk
====~"tt·•- -- ·
. . p da lyrie.1 e l3e r'lth. 1837, e mend. A. L. Schutte 1998
Tr1o;rl:e Loteiic . .subtribe Lipariinae Benth . (1843) , as 'Liparieae'
'frfbe Upa ricac 13e nth .) Harv . (1862)
dlUll & Vai \'X~yk c1?98'1) u~nn.1ari ~d recent ha~ges
generic and m bal circwn cripoons m th Podalyn ae
~o d Liparieae. T he Liparieae has b en formally placed
1 hydroxylated anthocyanins in purple-flowered taxa. A
'. n synonymy und r Poclalyrieae, a wa also sugge ted
~y Polhill (19 l o : 396-397 39 and Kas & Wink
(l997), but th g nu. Hypoa1/ypt.u was excluded a nd
.given triba l ta tu a the Hypocalyptea ( 'chmt &
VaJ1 wyk, l998b). The g nu Coelidium Veg I ex Walp.
was ubseq ue otly re d uc d to s yno n ymy wiLh.in
Ampl.litbcilea Ec kl. Z yh. chutt , 1998) . T h i
treaunenl ~ Uow Va n Wyk & chutte 1995a but
reflects several more recent modifications and revisions
(referen es give n below . p ci in Lhe tri e are Ii ted
b)' konki et ctl. 2003 . As pr s ntly circumscribed, the
tribe includ s 8 e nera and 125 p i (Fig. 35 .
It is interesting to note that taxonomic modifications
based on detailed analyses of morp hology, cytology
and chemist1y (Schutte & Van Wyk, 1998a & b) have
subsequently been su pported by molecular evidence
(e.g., Crisp et al., 2000; Van der Bank et al., 2002; Wink
& Mohammed, 2003). Secondary metabolites have
contribu ted substantially to the curre nt generic and
tribal concepts. These include, amongst others, the
FIG. 35 Diagram of relationships in tribe Podalyrleae after Van Wyk & Schutte (1995a); Van der Bank et al. (2002)
LEFT 0YeIop1a
.
maculata Ph otograph by B.-E. Van Wyk
presence o f uniqu e quinolizidin e a nd piperidyl
alkaloids, the absence of canavanine, the absence of
proa nthocyanidins an d the presence o f esters of
clear-cut dichotomy (sprouting versus non-sprouting)
exists in the fire-survival strategy of many mediterranean
shrubla nd legumes an d this feature is particula rly
relevant in the Podalyrieae (Schutte et al., 1995).
The Podalyrieae belongs to a mo nophyletic clade,
the 'core genistoids ' which includes Crotalarieae,
Genisteae, Podalyrieae, Thermopsideae, Euchresteae
and Sop h o reae sen s . strict. (Crisp et al., 2000;
Pennington et al. , 2001 ; Wojciechowski et al. , 2004). In
Crisp et a l. (2000) a nd Wink & Mo hamed (2003) ,
Podalyrieae is sister to a clade comprising the Old
World Crotalariea e a nd the Genisteae , the latter
incorporating the 'African Genisteae', i.e. Argyrolobium,
Dichilus, Melolobium and Polhillia (see Van Wyk &
Schutte, 1995a) . Cadia (in Sophoreae sens. lat.) is
b asally branching to Podalyrieae in the analysis of
Kajita et al. (2001) and Wink & Mohamed (2003), and
the Podalyrieae-Crotalarieae-Genisteae clade is sister
to a Thermopsideae - Sophoreae sens. strict. clade.
Crotalarieae (see page 273)
Genisteae (see page 283)
Cyclopia
Xiphotheca
Amphithalea
""O
0
Stirtonanthus 0
;r::.
~
;;c
Podalyria rn
;r::.
rn
Lipari a
Virgili a
Calpurnia
TRIBE PODALYRIEAE 267
 Xiphotheca phylicoides Photograph by 8.-E. Van Wyk Amphithalea ericifolia Photograph by B. -E. Van Wyk Stirtonanthus taylorianus Photograph by 8..f. Van Wyk

Cyclopia Vent. 1808 Amphithalea Eckl. & zeyh 1836

23 spp.; S Africa (endemic to the Cape region) Lalh1iogy11e Eckl. & Zeyb. (1836); Coelidi11111 Vogel ex Walp. (1840)
from eye/a- (Gk.: circle) and podion (Gk.: foot) , refe rring to che circular 42 spp,; S AFrica (endemic to che Ca pe region)
d ep ressio n m the base o f rhe calyx From amph i- (Gk.: around) and lhaleia (Gk: b looming), refcning to cbe o fte n
Shmbs; medilerrnne~m shrubland (fynhos and forest margins), often localised at copious blossoming o f the plants
high a ltitudes, in rocky and sandy are as Shrtibs ~rnd sh1 ublets (often e ricoid); ineditcrra nea n shru bhmd (fy nbos), usually at
Refere nce(s): Sch utte (1997a); Schutte in Golclblall & Manning (2000: ·183-485) medi um o r low alt itudes, often localised
A distin ct genus wit h inrric'<tte infmgene1ic rch1tionships; it appears to he an Reference(s): Granby (1980, 1985); Schutte 0998); Schutte in Goldblatt & Manning
iso lated basa ll y branchi ng group within the Poclalyiieae (see V~in clc1 fkmk el al.1 (2000: 462 - 465)
2002) Despite infrageneric diversity, Ampbitba/ea sens. lat. i.s a coherent genus with
C)ic/opia i nlermedia E.Mey, and C subternata Vogel ~ire cLwrently under severn l sy napomorphics; species with fu sed sw mens (formerly Coe/idium) foJ'tn a
commercial develop me nt as a traditional hcrlx1l tea ( ho neybush tea), which also clacle that is nested within the rest of th e genus (see Schutte, 1998)
has hea Ith benefits
Stirtonanthus Il .-E.van Wyk & A.L.Schutce 1995
Xiphotheca Eckl. & Zeyh , 1836 Sli11011ia Il.-E.van Wyk & A.L.Schutte (1994)
3 spp; S Afric;1 (e ndemic to the cencral patt of che Cape region)
Prieslleya DC sect, A11eisolhea DC, (1825)
Named for Charles Howa rd Stilton 0946 - ), a SouLh African botanist and the first
9 spp.; S Aflica (endemic to the Cape region)
Directo1· of the National Botanic Garden of \X'a les
from xipho- (Gk.: a swrnd) and -lheca (Gk .: a cont aine r), 1efen'ing lO the sword·
Shrubs; medite rranea n shrubland (fynbos), al medium to high al titudes , all highly
s haped pods of some species
Shrubs; mcdite1ranean shrubhtnd (fynbos), usun lly m mediu m or low ;11tiwdes ,
lornliscd
Ilefe tence(s} Van Wyk & Schutte 0 994 , 1995b); Schutte in Goldblatt & Manning
often locolised
(2000, 510)
Refe re nce(s): Schutte 0997b); Schutte in Goldblatt & Manning (2000' 511-512)
Superfi cially similal' to Podalyria but tl1e }e llow fl owe1s ~mcl unique alk:iloicls,
1

Related to Amphilhalea, which togethe r' form sub11•ibe Xipbo lbeci11ae A LSchutte
rogerhe r with DNA sequence data, .show that tl1 e genus is monophyletic (see Van
(sec Schutte & Van Wyk, 1998a)
t!e 1· !lank el al., 2002); Sli11011ia A.L.Srn , (1926) is a name cunently in use for a
genus of lichens

Podalyria wi1k1. 1799

19 spp i S Africa (sube ndernic to the Cape 1egion, ex tending slightly eastwards to
KwaZulu-Nata\)
Named for Podaliiius, a G1eek physicia n, son o f lhe Greco-Roman god o f
rnedic.:ine, Aesculapius
Small trees o r s hrubs; mediterrnnean shrnbland (fynbos and forest ma rgins), at
low, mediu m and high altitudes , mostly in rocky and sandy places
l\efcre nce(s): H~rvey (1862); Schutte (1995); Schu ti e in Goldblatt & Mann ing
(2000: 503 -504)
Despite morphological and chemica l uniformity , the ge nus appears to be
gene tically diverse; the ITS arrnlysis o f Vr111 der I3ank et al. (2002) indic1tes that
Podalyria is paraphyletic, compri sing three line;:iges co rresponding to three of
four sections of the genus proposed by Schutte 0995)
Podalyria ca~vptrata (Retz.) \Vil\cl. (bush sweerpea, ke urblom) and othe1· species a re
Xiphotheca canescens Photograph by 8.-E. Van Wyk Poda/yria ca/yptrata Photograph by 8. ·E. Van Wyk atlwct ive 01·namental shrubs; alkaloids in so me species have potential as inseccic ides

268 LEGUMES OF THE WORLD TRIBE PODALYRIEAE 269

 Liparia sp/endens Illustration by Mills
Liparia L. m1
Pries/I~)'" DC. sect p,•iesll~)'a ( ~ sect. Eisotbea DC.) (1825)
20 ~pp . ; S Af1ica (endemic to the Cape 1egion)
From liparo- (Gk: blight, glossy); the sta lks aod leaves arc smooth, che b11er also
silky at times or w ith a sil vet)' .sheen
Shrubs; mcditet'1'anean sh n.ibbnd (fynbos and forest margins), al low, medium rn
high altitudes, often high ly Joc;dised, mostly in 1ocky or .s;mdy pl::ices
Refe rence(s)o Schutte Cl997c); Schutte in Goldb latt & Ma1111i11g (2000 0 49j-196)
L1parict appe:irs to have some genetic affinity wirh Podalyric1 but the suggestion
that Uparia is not monophyletic (Crisp el al, 2000), or is nested within :t
paraphyletic Podalyria (Va11 der B~ink el al,, 2002) needs verification th1ough
furthe r study
Ltparia splendens (I3urm.f.) Bos & de \X'it, the spcctac.:ul~1r 'mountain d;1hlia' and
some other species ~ ire om~11nentals
Liparia hirsuta Photograph by 8.-E. Von Wyk
270 LEGUMES OF THE WORLD
Virgilio oroboides Photograph by P. Van Wyk Ca/purnia aurea Drawing by C. Grey-Wilson
Virgilia Poil'. rnos
2 spp.; South Africa (endemic to the Cape 1egion From the C;ipe Peninsula to
Grahamscown)
Nmned frn the Roman poet Vil'g il (70 to 10 13.C .)
Trees; s ublropical Afromonwne forest m~irgins , o flen along .'i tre11ns and 1·ive1·
banks
Rcfc rcnce(s)o V"n Wyk (1986); Schutte in Goklbla 1t & Manning (20000 511)
Claclistk analyses all confir·m a close rclationshi[1 with Calp11rn;a (see Schutte &
V;t11 Wyk, J998a and Ci isp el al., 2000)
Anractive :incl populJr gaiclen trees in many patt.'i of the worlcl (known as
keu1boom in South Af1 ica); of ecological i111porrance as a forest mal'gin pioneer,
the wood is so metirncs used foi furnitl11 c and has othet general use.s
Calpurnia E.Mey 1836
7 spp.; Afric::1 (Afromontane to Somalia-Masai regions) <llld India (ma inly the
easte1n palls or southern Af1 ic.1 , with one s pecies extending through cc1ste10
tropical Africa to S India)
N::unecl for 1hc Rom~m poel Ca lpurnius, an imitator of Virgil, alluding to a
similarity wirh Virgilia
Trees and shrubs; sub1 1opical or tropical, lowland ancl monrane, .seasonall y dry
forest (mostly Jlong m~irgins) , wooclbncl, bush land , xerophytic shrubland and
grnssbnd, ~dong livers, or on s~ind or rocky ou tcl'ops
Referencc(s)o Ileaumont et al. (1999)
CalpHr11ia was tr•ansfem~cl to Pocblyri e;1e from Sophoreae by Van \\!yk &
Schutte (1995a); molecub1 data supports this as siste1· to Virgilia (Van de1 Bank
el al., 2002)
Used as ornamentals and shacle plants, for timber and medicine
Catpurn1a
• aurea Photograph by P. Van Wyk
TR IB E PODALYRIEAE 271
 otalarieae byB.-E. VanWyk
. (Oenth.) Hui It. '196
·tlnoe:le (B -n) Dumori . subtri be Crotalariinae Benth. (1865), as 'Crotalarieae'
(jeni.'>1C:t ror.
a JiX>niea Hut l. (1964)
i:1ooonic!e;1e Hut ch. (196 )
nt state of knowledge of the Crotalarieae was
by Van Wyk (199la) and by Van Wyk &
a) . The most conspicuous .recent clun~e
0995
the exclusion of the Argyrolobzum group (six
i.e. Argyrolobium , Dichilus, Melo/obium,
.,' Anartbrophyllum and Sellocharis) which
,a, 1 . C 1 .
in tribe Genisteae rather t 1an m rota aneae,
they were previously placed (Polhill, 198lq: 399
. New insights into relationships within the tribe
come mainly from chernosystematic studies of
cir (summarised in Van Wyk & Verdoorn, 1990)
em! recent generic monographs (see below).
Crotalarieae forms part of a monophyletic clade,
'G©re genistoicls' (Fig. 36) which also includes
e, Poclalyrieae, Thermopsideae, Brongnia1tieae,
teae and Sophoreae sens. strict. (Crisp et al.,
Pennington et al., 2000a; Kajita et al., 2001) .
·eae appears to be sister to the Genisteae and
are sister to the Poclalyrieae (Crisp et al., 2000;
howski et al., 2004). This clade is in turn sister
'Fhermopsicleae and Sophoreae sens. strict.
g Euchresteae).
Crotalarieae shares with the Podalyrieae the
t"C! of a-pyriclone alkaloids such as cytisine and
ine that are a typical feature of all other 'core
oid' tribes. Despite a lack of defining characters,
onophyly of the tribe as circumscribed here is
Ctritolar/a m.
icons Photograph by D. Du Puy
well supported by molecular evidence (Crisp et al.,
2000; Wink & Mohamed, 2003) and by cladistic
analyses of morphological, cytological and chemical
characters (Van Wyk & Schutte, 1995a). The latter study
suggested an early diversification of the genera with
uniform anthers and lupanine-type esters of
quinolizidine alkaloids (Pearsonia, Rothia and
Robynsiopbyton) followed by the poorly known
Spartidium and then the so-called 'Cape group of
genera' (Polhill, 198lq: 399- 402), which now includes
Lotononis and Crotalaria. Relationships between the
seven genera of the 'Cape group' remains unresolved
despite several recent molecular studies because
sampling is still relatively poor. However, a basally
branching position in the tribe of the 'Cape group',
notably Leheckia and Wihorgia - as considered by
Polhill (1976, 1981q) - is now accepted here. The
exclusion of the Argyrolobium group, based on
morphological and chemical characters, is also strongly
supported by DNA sequence data. Due to reticulate
and overlapping patterns of character state distribution
in the Crotalarieae sens. strict., generic delimitations are
intricate and subject to misinterpretation. Several of
the large and diverse genera appear to be either
monophyletic or paraphyletic depending on the choice
of characters. As currently circumscribed the tribe
includes 11 genera and c. 1204 species (Fig. 37).
TRIBE CROTALARIEAE 273
 Sophoreae p.p. (see page 228)

Sophoreae p.p.

-
Brongniartieae (see page 253)

Sophoreae sens. strict., (see page i 28)

Euchresteae (see page 260)

---
II Thermopsideae (see page 263)
I

Podalyrieae (see page 267)

Crotalarieae
I
I Genisteae (see page 283)

FIG . 36 Diagram of relationships of tribes in the Genistoid clade (i.e., core-genistoids), after Crisp et al. (2000); Pennington et al. (2oooa); WoJ Lebeck/a macrantha Photograph by B.-E. Van Wyk
et al. (2004)

Spartidium Pomel 1874

1 sp.; N Africa (Morocco to Li bya), in the Sahara Regional Tra nsition Zone (White,
1983)
Derived fro m the superficially s imilar genus Spa1tit1m L. ('Spanish broom') and
-idlum (Gk. suffix for d iminutive), thus meaning 'little Spa1tium'
Shrub; desert on sand d unes
Reference(s} Polhill 0 976)
This is a poorly known genus in need of further study; mo rphologically S,
saha..ae (Coss. & Durieu) Pomel, is practically inseparable from Lebeckia (Polhill ,

- Spartidium ?
1976) but the genelic concept is supported by the accumulation of ammodend rine
(a bipiperidyl alkaloid) and the virt<1al absence of the characteristic quinoli zidine
alka loids of Lebeckia (Van Wyk et al., 1989); the genus may thus be better placed
in Gen isteae

L~pecki a Lebeckia Th unb. rnoo

Wib0rgia c. 35 spp; Africa (mostly in the Cape region of South Africa, extending
northwards into the dry interior of South Africa, Botswana a nd Namibia)
Rafnia The generic name commemorates HJ. Lebeck (?-1800), a merchant and collector
Aspalathus in Inda-Ma laya who stud ied in Sweden under Thunberg
Shrubs o r sh ru blets, rarely he rbs Cflreweeds); med iterranean shrubland (mainly
fynbos and re nosterveld) or less often in dese1t, xerophytic scrubland o r
seasonally dry tropical to subtropical woodland, often in rocky and sandy p laces
Lotonon is Re ference(s): Harvey (1862); Polhill (1976); Schutte in Goldblatt & Manning (2000:
494 - 495)
Belusia Lebeck/a is closely related to Wiborg ia (q.v.) . The genus is morphologically
Cretalaria d iverse and d istinct infrageneric groups (sections) can be recognised (Harvey
1862; Van Wyk, unpublished data); a rev ision and phylogene tic analysis at
sectional level are in progress. Lebeckta s pecies are remarkably simila1· to some
shrubby Lalananis b ut the loss of stipules appears to be a convergent character
Pearsonia and the accumulation of large amounts of quinolizid ine alkaloids and the absence
of cyanogenesis in Lebeckia a1e significant d iffere nces (shrubby species of
Rothia Lotononis have macrocyclic pyrrolizid ine alkalo ids and are cyanogenic)
Robynsiophyton Some species of Lebeckia a re locally dominant and ecologically important; some
are used as ornamen tals and for fuelwood

? =doubtful position of Spartidium in Crotalariea e

• k & Mohamed
FIG . 37 Diagram of relationships of genera in tribe Crotalarieae after Polhill (1976; 1981q); Van Wyk & Schutte (1995a); Wm
esooJ>

274 LEGUMESOFTHEWORLD TRIBE CROTALARIEAE 275

 Rofnio meyeri Photograph by a. -E. Van Wyk Aspolothus lineoris Photograph by B.-E. Van Wyk

Wiborgia Thunb 1800 Aspalathus i . 1753

JO spp.; Africa (endemic to the Cape region of South Africa) Borbonia L. (1753)
278 spp .; Africa (sub-endemic to the Ca pe region of South Africa, extending
Named after El'ik Yibmg 0759 - 1822), a Danish botanist
slightly eastwards to KwaZulu -Natal Province)
Shrubs; mediterrnnean shrublan<l (fynbos and 1cnosterve ld), mostly in
From the Gree k aspalathos, an ancient name for a plant, dating from biblical
and sandy flats
times; over the cen turies it has been appl ied to several differe nt legume genera
Reference(s): Dahlgren 0975); Schutte in Goldblat t & Manning (2000: ~Ill
Shrubs or shmblers, rarely small trees; mcditerrane'1n s hrubland (mainly fynbos
A morphologicall y uniform and dearly monophyletic group with Jllrikinl!
and renostervekl) and grassland, often in rocky and sandy places
to some species of Lebeckia, but generic affinities have yet to be
Reference(s): Dahlgren 0960; 1961a; 196Jb; 1963a; 1963b; 1988); Schutte in
depth; the indehiscent, winged, sa ma ra-Jike fruits a1c <l unique dfill(
cha meter Goldblatt & Manning (2000: 466 - 483)
Earlier attempts at subdividing th e ge nus into naturnl groups were '1bandoned by
Dahlgren (1988) for practical reasons; no phylogenetic analysis of infragenel'ic
1elationships has yet been published although a molecular study is cunenrly in
Rafnia Thunb. 1800
progress by J. Hawkins and co-workers
19 spp.; Africa (sub-endemic to the Cape Region of South Africa, t!Xt
The chopped and 'fermented' stems <rnd leaves of A linearis (Burm. f.)
slightly eastwards to KwaZulu-Natal Province) RJ)ahlgren are widely known as rooibos tea, an important commercial crop plant
Named after Carl Gottlob Rafn (1769-1808), a D:rnish botanist
cultivated on a large scale in the western parts of the W Cape Province of Sourh
Shrubs or suffrutices; rnedite1rnnean sluubland (m<1inly fynbos) and ~rill
Afiica (annual harvests of up to 9 million kilogrnrns have been recorded and
mostly in rocky and sanely places . exports llave increased in recent yeais); recent resea 1ch has shown that
Reference(s): Campbell (1998); Campbell & V:m Wyk (2001); Campbell Ill
consumption of the tea has a range of he<1lth benefits; many species are loe<1lly
Goldblatt & Manning (2000: 507 -509) dominant and of tmijor ecological imprntance as pioneers, especially of fin~-rrone
Seve1al st udies, based on morphological and molecular drn1:•1ctcrs, ha~
vegeration; also used as soil binders
Rafnia to be close ly related to Aspalathus The genus is easily tt'tQBand
infragene1 ic rehuionships ~1re not yer resolved; species idcnrifiaitJOn
circumscriprioi: have hitherto been co111plic.1ted by.su_perfi:::il s_IJ~~;~
unrelated species and by extreme geogrriph1cal va11~H10 n, icsultmg
regional fo1ms within species (Campbell & Van Wyk, 2001) . herbal It'll
Leaves of some species have been used For making a ranrnn -nch
1111
*
the roots of others (notably R amplexicaulis (L) Tliunb. or ·socd• 1so ....,.r
a hitter-sweer taste and were an early Cape substitute for liquorice; a
ornamentals, for medicine, soil binders and ground cove•

Rofnio per{olioto Photograph by B. ·E. Van Wyk Dahlgren

276 LEGUMESOFTHEWORLD TRIBE CROTALARIEAE 277

Lotononls corymbosa Photograph by 8.-E. Van Wyk Lotanonls magnlfica Photograph by 8. -E. van Wyk
Lotononis CDC.) Eckl. & Zeyh. 1s36
Buchenroedera Eckl. & Zeyh. (1836); Listia E.Mey (1836)
c. 150 spp.; Africa, mainly in southern Africa with a few extending U> (
monlane) tropical Africa (c. 145 spp.); c. 5 spp. Mediterranean .(S. p:i1A.
Macaronesia and N Africa [Morocco)) to W Asia, Arabia and Pnklslan
The name is derived from Lotus and Ononis (q.v.), suggesting a sim~
these genera
Shrubs, shrublets, suffrutices (rarely geophytic), perennial or :trul\lul hcsblt,
mediterranean shrubland (dry fynbos and renosterveld), desert, . d!llflll)'llc
scrubland and grassland; also seasonally dry tropical to 011l)1roplct1l fl
wood land, usually on sand and in rocky places, more rarely in liell•)' or
calcareous soils
Reference(s): Van Wyk 0991b); Schutte in Goldblatt & Manning l:ZOOO:
Polhill in Pope et al. (2003: 233-245)
Considerable infrageneric variation has been accommodated in 15 " " -
of which have convincing apomorphies 10 support their m nopllyl • W.:
relationships between sections have been explored in a series r Llcfi,tdic
of morphological, cytological and chemical characters (Van \Vy!{, 1!)9lbl
Lo/ononis bainesii Baker is :i widely cultivated livestock fodder and ~
due to their prominence in dry regions and disturbed places, 60\'l:l':t\
of ecological importance as soil improverS and in erosion conitoh ut IL."f
are used for medicine and have potential ~$ ornamentals; some~
prunasin and other cyanogenic glycosides and have caused Stodt "'"'
Bolusia Benth. 1873
5 spp,; SC and southern Africa . ~(l1'1tcut
Named for Harry Bolus (1834-1911), :t businessman, plo nc<!n ~<;olul•
11
and founder of the Bolus Herbarium, University of Cape To7~d p,,.J
Perennial herbs; seasonally dry tropical to subtropica l "~ a ('I
bushland, shrubland and grassland, usually in rocky or_san Y p~ /r(,
Reference(s): Polhill (1976); Lisowski (1995); Van Wyk m Pope
228-23.~) I! nd Jl.'llral~
Ensily rc'COJlni.<ctl hy the str:mgc ltdirnll)' w1l~>d k1.-cl flC"' s • l<Clll'I Ot"'
b ul the genus seems do.<cl)• rchuud to C1'1J1t1/mim the Mntlug; • (()(ti"'
comrH 1111moirls ll.Vi.!1· np pc.1!ll to he :r. 1110rpholo1: 1<"tl 'pr'OIO!~.: l)UI ;Ill
chamete~ (o und In '1Jol1t.<f(I: th<: gencrl rot11.'<!pl ~hv~lcl be ""
1 (see also next four figures on following page)
• :1m1 lc 11r Crotulmi" ;pcd.;,. would n :cd 10 be ind11th:<l
.5. R-C.
Bolus/a resupinata Drawing bys. Ross-Craig
278 LEGUMES OF THE WORLD
Cratalarla grevei Photograph by o. Ou Puy
Crotalaria L 1753
Goniogyna DC. (1825); Hey landia DC. (1825); Priotropis Wight & Arn (1834)
c. 690 spp.; especially in the southern hemisphere: most spp. in Africa and
Madagascar (c. 510 spp., mostly E and southern Africa, c 34 of wh ich e ndemic to
Madagascar and 5 widespread in Old World); c. 18 spp. widespread in tropical
Asia; c, 60 spp. endemic to the Indian subcontinent; c 5 spp. to W Asia, c. 15
spp. to Inda-China, c. 12 spp. to China, c. 6 spp. to Malesia and 9 spp. to
Australia; 59 spp. endemic (and a further 15 introduced) in the New World (fide
Windler & Skinner in ms.), with c. 35 spp. in S America (mainly Brazil), 20 spp. in
N and C America (mainly Mexico) and c. 5 spp. widespread in New World
From crotalon (Gk ,: castanet; rattle), referring to Lhe rattling of the seeds in the
inflated pods
Shrubs, shrublets or perennial herbs, rare ly small trees or annual herbs; seasonally
dry tropical, subtropical to warm temperate forest, woodland, xerophytic
shrubland and grassland, often in disturbed places, on sand or rocky outcrops
Reference(s): Polhill (1968b); Bisby (1973); Bisby & Polhill 0973); Windler 0974);
Matos (1978); Polhill (1982); G6mez-Sosa (2000); Flores & Miotto (2001); Du Puy
& Labat in Du Puy el al. (2002: 672-708); Holland (2002); Polhill in Pope et al.
(2003: 68-228)
Crotalaria appea rs to be closely related 10 Lotononis (q.v.); the two genera are
unique in rhe family in their accumulation of severa l macrocyclic pyrrolizidine
alka loids (Van Wyk & Verdoorn, 1990); pyrrolizidine bases, however, are also
known (Kinghorn & Smolenski, 1981: 592-593) from Ammodendron
(Sophoreae) and Adenoca11Jus and Laburnum (Genisleae); infrageneric
relationships are poorly known but the genus seems to have originated in Aflica,
wi th more recent diversification into other regions of the world; the Asian species
are in need of revision
Several species are of commercial importance as fibre crops, fodder and g1een
manure; others are used as medicine, ornamentals, nurse crops, heavy metal
indicators, human food and for soil improvement (intercropping); some cause
croralism in li vestock and humans - acute bul more often chronic poisoning
affecting the lungs and liver - clue to the ingestion of pyrrolizidine alkaloids with
an unsaturated necine base; C. jtmcea L (Sunn hemp), is a major crop for its bark
(phloem fibres) which provides a high quality bast fibre for cordage, fishing nets
and fine paper
TRIBE CROTALARIEAE 279
 Rothia hirsuta Drawing by E. M. Stones

Pearsonia Dummer 1912

Pbaenohoffmannia Kuntze (1891); Gamwel/ia Baker f. 0935)
13 spp.; Africa and Madagascar (12 spp . in Africa S of the equator, mainly in the
southern Afromoma ne Region [D1 akensberg and Inyanga ni Cemres], and l sp. in
Madagasca r)
Named for H.H.W. Pearson (1870-1916), a pionee ring botanist of S Africa and
rirsr Dii'ector of the Na tional Botanic Gardens, Kirsrenbosch
Shrublets ~nd herbs; seasonally clry tropical to subtrop ica l montane xerophytic
bushl and, shrubland ancl grassland, sometimes in fo rest or woodland; on sand or
rocky o utcmps
Refc1e nce(s): Polhill (1974); Campbell-Young & Dalkwill (2001); Du Puy & Labat
in Du Puy el al, (2002: 669-671); Polhill in Pope el al. (2003: 56-63)
Morphol ogica lly similar to Lotononis (pa1t icu larly in the lorononoid calyx) but
distinctive in th e gullet-type pollination m echan ism ancl the accumulation o f
unique esters of quinolizidine alk~loids, two features shared only by Rothia ancl
Robynsiophylon (Va n Wyk & Verdoorn , 1991)

Rotbia Pers. 1807

2 spp.; Africa and As ia to Austra lia (1 sp. widesp1•ead in the drier parts of Africa,
the o th er from India to N Australia)
Named for Albrecht Wilhelm Roth 0757 - 1834), a German physician and bota nist
Annual herbs; mainly in seasonally dry tropical xerop hytic s hrubland and
grassland
Refere nce(s): Polhill 0976); Polhill in Pope et al (2003: 64 - 66)
Re lated to Robynsiopbylon and Pearsonia (q.v.)
Used as famine food , livestock foclcler and green 111anu1'e

Robynsiophyton R.Wilczek 1953

1 sp .; SC Africa (Ango la, Zambia and Congo (Kinshasall
Named for F.H,E.A.W, Robyns 0901 - 1986), a Belgian botanist ;ind phyton (G k.:
plant)
Herb; seasonally d ry tropical grnssl:md and deciduous woodland
Reference(s): Polhill 0 976); Polh ill in Pope el al (2003: 66 - 67)
Closely re lated to Rotbia bm the flower has nine .sta mens of which o nly five are
fertile

Crota/aria flavicarinata Drawing by J. A. Lang horne Crotalaria variegata

280 LEGUMES OF THE WORLD TRIBE CROTALARIEAE 281

TR IBE
Genisteae byR.M.PolhillandB.-E. vanwyk
. Genistcae ( Bi:oon) Dumo rt . 1827
nibe t iribe Genistinae Br 11 11 (1822), as 'Gcnisteae'
,.°1 ~c Cyti eae H or<m. (1847), as 'Cytlsinae'
'f~Jbe Uliceac Webb (1.852), a 'Uli inea '
TrlbC taburn a 0\ouy) Hutch . 0 964)
'fribe 1.uriine;ie (Rouy) Hu tch . (1964)
Bi by (1981: 409-425) divided the Genisteae into
ubtribes Genistinae and Lupininae, a principal
dh'ision taking both morphological and chemical
ev.idenc into account. Polhill (1976) drew attention to
rh. similarities between woody Thermopsideae
(Anagyris and allies) and the main part of Genisteae,
and between the herbaceous genera of that tribe,
Baplisitl and Thermopsis, and Lupinus. Crisp et al.
(2000) emphasise the lability of staminal fusion in
genistoic.l tribes as a whole, and it might be supposed
dull the free-stamened Thermopsideae would be
better included in Genisteae. In recent molecular
analyses (Kass & Wink, 1997; Crisp et al., 2000; Wink
&. Mohamed, 2003; Wojciechowski et al., 2004),
howeve r, all Thermopsideae grouped with Sophora
<µld close allies, and Lupinus was relatively basally
bran I ing in Genisteae, above the southern African
genera Melolobium and Dichilus (previously placed in
Crotalarieae). The similarities between Baptisia,
Therm op is and Lupinus are likely due to strong
~cologkal convergence in temperate habitats.
The Genisteae forms part of a monophyletic clade,
the 'core genistoids' which also includes Crotalarieae,
Podalyrieae, Thermopsideae, Brongniartieae,
Euchresteae and Sophoreae sens. strict. (Crisp et al.,
2000; Pennington et al., 2000a; Kajita et al., 2001).
Genisteae appears to be sister to the Crotalarieae and
both are sister to the Podalyrieae (Crisp et al., 2000;
Wojciechowski et al., 2004). There is now convincing
evidence that Melolobium, Dichilus, Argyrolobium and
Polhillia (the Argyrolobium clade of Van Wyk &
Schutte, 1995a), together with Anarthrophyllum (and
probably Sellocharis), are part of Genisteae and that the
similarities with the Crotalarieae (the remarkable
parallels between Dichilus and Lebeckia for example)
are superficial only (e.g., Wink & Mohamed, 2003;
Wojciechowski et al., 2004). Crisp et al. (2000)
suggested that this group of genera may be sister to
Crotalarieae but their analysis did not include critical
genera such as Adenocarpus and Lupinus. Less
emphasis is now given to stamen fusion, and these
genera have been .form~ Jly transferred to Genisteae
based on the shared presence of a trifid lower lip of the
LEFT Lupmus
· weberboueri. Photograph by c. E. Hughes
calyx and the distinctive quinolizidine alkaloids of the
a-pyridone type (Van Wyk & Verdoorn, 1990; Van Wyk
& Schutte, 1995a; Wink & Mohamed, 2003). Van Wyk
et al. (1989) suggest that Spartidium, currently in the
Crotalarieae, may be better placed in Genisteae.
More detailed molecular studies are necessary to
fully assess relationships of the genera. Kass & Wink
(1997) undertook the first detailed molecular analysis
of Genisteae using rbcL and ITS sequences, and
limited ITS sampling by Crisp et al. (2000) is largely
congruent with this, also supporting a paraphyletic
Argyrolobium. The two southern African species of
Argyrolobium analysed by Crisp et al. (2000) are
dispersed within the Genista group Clinked to
Spartium and Retama). Kass & Wink (1997) sampled
four species of Argyrolobium, two from southern
Africa and two from the Mediterranean Region, and
these are more basally branching in the tribe (Fig.
38), although the Mediterranean species A. zanonii
(Turra) P.W.Ball groups elsewhere near the base of
the Cytisus group. These results are largely supported
in the chloroplast rbcL analyses of Wink & Mohamed
(2003), with more species of Argyrolobium sampled
and a placement of A. zanonii as sister to the
combined Genista and Cytisus complexes. The ITS
analysis of mainly Spanish Genisteae (Pyne, 1999) is
also largely congruent with Kass & Wink 0997),
except for Argyrolobium where one southern African
species groups within the Genista complex and A.
zanonii is basally branching to it. The sequence for
Lembotropis is acknowledged as suspect in the Pyne
analysis, and its position allied to Cytisus (as C.
nigricans) (Kass & Wink, 1997; Wink & Mohamed,
2003) is more in agreement with morphological
evidence. The biggest problem areas remaining,
besides resolving relationships within Argyrolobium,
are generic delimitation within the Cytisus and
Genista groups and the decision whether to recognise
a few large, or many small genera. Morphologically,
Argyrolobium is more coherent than the current
molecular evidence seems to suggest and a broader
sampling and reanalysis would be informative. For the
main part of Genisteae, the segregation of the groups
TRIBE GENISTEAE 283
 of genera around Cytisus and Genista is evident (Kass
Lupininae Rouy - lupinu -, Anartb1·opby1t
& Wink, 1997; Cubas et al., 2002; Wink & Mohamed,
ell chttri Am rica , parri ularly montan r gio u,~,
2003; Pardo et al. , 2004), but the placement of many the w t; M dit rran an regi n u·opi al Afrl a · ns 1n
genera is not well supported. Kass & Wink 0997)
conclude that the position of many of these genera Cytisinae (Hora n .) Benth . - Cytisopb if/.
which lie between the Cytisus and Genista complexes
.
A rgyrocytisus, P etteria,
. c·
ytisus, Lembotr im1.•
cannot b e esta blished with certainty, probably Calicotome (Europe , Mediterranean region· op1s,
becaus e of the small numb er of nucleotide Macaronesia) and
substitutions available (due to rapid and recent Laburninae Rouy - Laburnum, Podo cyu
dive rsifica tion) and homoplasy. So as not to obscure la b urnum (Me diterranean,
' . . Stts
u-
nespero pnnc1pally Balka~
the morphologica l relationships prior to further and Atlas Mts); these genera are treated here as bei.Q
mol ecular s tudies, the subtribal classification of nested within Cytisinae g
Talavera & Salgueiro 0999), slightly extended and
modifi e d, is ou tlined here . The Genisteae here Erinaceinae Talavera - Erinacea (SW Europe and
comprises 25 genera and (551) -562 - (572) species Africa); genera in the Erinaceinae and in Span iin~
(Fig. 38). below, are treated here as bemg. nested within '"'e
the
Genistinae
Adenocarpinae Rouy - Melolobium, Dichilus,
Po!hillia, Argyrolobium and Adenocarpus (South Africa, Spartiinae Benth. - Gonocytisus, Retama, Spartiurn
tropical Africa (mostly montane) and Mediterranean (Mediterranan region)
region, thinly to the Indian subcontinent); this group Genistinae Bronn Genista, Ecl:1in sparturn,
largely comprises a basal grade on molecular evidence, Stc11traccmthus, Ulex (Europe, mostly w ·tern, and
but all genera need further analysis Meclilerranean region

Crotalarieae (see page 273)

Melolobium
Dichilus Melolobium macrocalyx Drawing by V. C. Gordon Melolobium wilmsii Photograph by A. E. Van Wyk & 5. Malan
Polhillia
Melolobium Eck l & Zey h 1836
JS spp .; Afri ca (sourhern Africa, including drier areas of Namibia, Botswana a nd S
Argyrolobium pro maj. parte Africa)
From me/o- (Gk : li mb) and -/obion (Gk: a legume o r pod), re fening to the
constrictions in the pods of some species
Lupin us Shrubs and h erbs; mediterran ean shrubl a nd (fy nbos and renosterveld), dese1t,
xerophytic scrubland, bushla nd and thi cket or grass la nd , often on sand,
Anarthrophyllum sometimes in rocky places
Sellocharis? Reference(s): Harvey (1862); Mo teetee & Van Wyk (2001); Schutte in Goldblatt
& Manning (2000: 500); Moteetee el al. (2002); Polhill in Pope el al. (2003:
250-253); Moteetee (2003)
Allied to Dichilus as basally b 1a nching genera in the tribe (Van Wyk & Schutte,
Argyrolobium pro min. parte 1995a; Hu et al, 2002; Wink & Mo hamed, 2003); Melolobium species are. eas ily
recog nised by the auriculate stip ules, gla ndu lar t richomes and (often) spmy
inflorescences
Adenocarpus Used as pasture p la nts a nd as ma terials (e.g ., in bas ketry)

Dichilus De. 1825

Cytisophyllum, Argyrocytisus, 5 spp.; Africa (E Na mi bia, Botswana and w ides pread in the eastern a nd centra l
Petteria, Laburnum, Q pa11s o f southern Africa to Zimbabwe) .
-i From di- (G k: two) and -chi/us (Gk: li p), re fe rring to the two-lipped calyx
Podocytisus, Hesperolaburnum, l/l Shnibs o r suffrutices; grassland 01 xe1ophytic s hrubl a nd , in sandy or 10cky places
Cytisus, Lembotropis, z Reference(s): Schutte & Van Wyk 0988; 1990); Polhill in Pope el al. (2003:
)> 248-250); Moteetee (2003) ..
Calicotome rn A d isc rete genus, allied to Melolobium, forme rly in Crotalarieae and superf1c1ally
si mil ar to Lebeckia, but now pla ced iT1. Ge ni sleae neLJr Argyrolobtum (Cr_1sp et_ al.,
2000); the a lkaloids and DNA profile provide the best evidence foi relat1o nsh 1ps,
but the two-lipped calyx is a usefu l mo rph ological marker
Gl
Echinospartum, Erinacea?, rn
z
Retama, Gonocytisus, l/l
Genista, Spartium, -i

Stauracanthus, Ulex z
)>
rn
? = position in group uncertain

FIG. 38 Diagram of relationships in tribe Genlsteae after Kliss & Wink (t997); Pyne (i999); Crisp et al. (2000); Cubas et al. (2002); Wink & Mohamed
(2003); Pardo et ol. (2004)
Dichilus reflexus Photograph by B.·E. Van Wyk

284 LEGUMESOFTHEWORLD
TRIBE GENISTEAE 285
Po/hiI/lo conescens Photograph by B. -E. van Wyk Polhi/lio pollens Photograph by c. J. Burgers
Polhillia c .H.Stirt. 1986
7 spp .; W and SW parts of the Cape region of South Africa
Named for Roger Polhill (1937- ), a botanist al Royal Botanic Gardens, Kew, Jn
honour of his major contributions to legume systematics [approbation added by
ed itors]
Shrubs and herbs; mediterranean shmbland (re nosterveld) and scrub-grassland,
mostly on heavy soils
Reference(s): Stirton (1986b); Van Wyk & Schutte (1989); Van Wyk (1992); Schutte
in Goldblatt & Manning (2000: 504 -505); Moteetee (2003)
Allied to Dichilus and Melolobium, but the species are mostly rare and highly
localised in the SW Cape region; it can easily be distinguished from the closely
related Argyro/obium by the absence of true peduncles, the conduplicate leaflets,
virtual absence of bracts and bracteoles, the sheathing stipules and the
chromosome number
Polhi/llo obsoleto Photograph by B.-E. Van Wyk
286 LEGUMES OF THE WORLD
Argyroloblum ltremoense Photograph by D. Du Puy Lupinus p/losus Photograph by R.B.G., Kew
Argyrolobium Eckl. & Zeyh. 1836
Chasmone E.Mey. (1836); Calispepla Vved. (1952)
c. 80 spp.; principally southern Afri ,, ( • 50 spp. in S Africa), c. 10 spp. in the
highlands of tropical Africa to Ethlopi", 3 spp. in Madagascar and c. 15 spp. in the
Mediterranean (N Africa and S Europe) to Arabia, W Asia and thinly to India
From argyro- (Gk.: silvery) and loblot1 (Gk.: pod), for the often sericeous pods
Shrubs and herbs; seasonally dry tropical to warrn temperate grassland, woodland
and forest edges, more rarely in mediterranean shrubland; also extending into
xerophyti scrubland, desert and coastal arens. t.L<uully on sand and in rocky places
Refore11re{S): Polhill (1968a); Van Wyk & Schutte ( 1989); Edwards (1994): Chaudhary
(1997); Edwards & Schutte in Goldblatt & Manning (2000: 465-466); Du Puy &
Labat in Du Puy et al. (2002: 709-711); Martins in Pope et al. (2003: 253-262)
Formerly placed, with Dicbilus and Melolobium, with some uncertainty (Polhill,
1976; 1981q) in Crotalarieae, but, with more inforrnation on the phytochemistry,
now confidently attributed to Genisteae (Van Wyk & Schutte, 1989; 1995a);
molecular evidence (Kass & Wink, 1997; Pyne, 1999; Crisp et al., 2000; Wink &
Mohamed, 2003) suggests that the genus is not monophyletic and there is a need
for a comprehensive study; some species are morphologically and chemically
closely related to the genus Polhillia; relationships among the South African
species were explored by Edwards (1994) and Moteetee (2003)
Used for ground cover, erosion con trol, human food (roots) and medicine
Lupinus L. 1753
c. 220-230 spp.; Mediterranean basin in Europe to Turkey, Middle East and N
Africa (c. 13 spp.), plus montane E tropical Africa (2 spp.); disjunctly to N
America (c. 120-130 spp.; mainly California and W Rockies to Mexico and C
America) and S America (c. 80-90 spp.; mainly Andes and thinly to Brazil and NE
Argentina), widely introduced elsewhere
Classical name in Latin, possibly derived from lupus, a wolf, on the erroneous
supposition 1h:u these plants impoverish soils
Shrubs and h<-'flJS; mostly open h:ibirnts, in disturbed places, in poor (often acid) soils
Reference(s} Smith 0938-1953); Gladstones (1974); Monteiro & Gibbs (1986);
Barneby (1989b); Hickman (1993); Isely (1998); Kurlovich (2002); Ree et al. (2004)
Isolated in Genisteae (apart from Anarthrophytlum and Sellocharis) and given
separate subtribal status (Bisby, 1981; Talavera & Salgueiro, 1999); species
delimitation is very problematic and no workable infrageneric classification is yet
available; New World species are currently being revised by C.K Hughes
Important as a domesticated human food crop; L. a/bus L. (white lupine, lupini
bean) and L tuteus L. (yellow lupine) From the Mediterranean and L. mutabilis Sweet
(pearl lupine) from the New World are grown for edible, high protein seeds (also for
flour); several species widely grown for livestock fodder, green manure, cover crops,
fish poisons, medicine, oils and as attractive ornamentals (e.g., Russell hybrids of L.
polyphyllus); a number of species are variably toxic and lupinosis causes death in
LuPmus
· touris Photograph by G. P. Lewis animals, due largely to ingestion of Foliage containing quinolizidine alkaloids
TRIBE GENISTEAE 287
 Lupinus mutabilis, pink variant Photograph by G. P. Lewis Anorthropliy/lum desideratum Photograph by M. F. Gardner Sel/ocharis paradoxa Drawing by R. M. Polhill

Anarthrophyllum Benth. 1865

15 spp.; S America (Andes in Argentina and Chile)
From anarthro- (Gk.: unjointed) andphyllos- (Gk.: leaf), referring to the highly
modified leaves lacking pulvini
Shrubs; dry shrubland, bushland and grassland in cold parts of the Andes, on
sandy soils and in rocky valleys
Reference(s): Burkart (1952); Sarani 0974); Van Wyk et al. 0993a)
The alkaloid pattern, circumcauline stipules and chromosome number provide
evidence for a relationship with Argyrolobium and Lupinus (Van Wyk et al., 1993a)
Used for forage, windbreaks and firewood

Sellocharis Taub. 1889

1 sp.; S America (SE Brazil)
Named in honour of Frederich Sellow 0789-1830), a German naturalist, and
charis (Gk,: graceful), referring to the attractive habit of the plant
Shrub or herb; temperate grassland (pampas)
Reference(s): Taubert (1889); Polhill 0976)
The unusual arrangement of the leaves, with 5-7 leaflet-like structures in a whorl
at the nodes, is unmatched in legumes; known only from the type collection until
re-collected recently in the Rio Grande do Sul region; Anarthrophyllum and
Lupinus would seem to be the most proximate genera

Adenocarpus Dc. 1315

c. 15 spp.; Mediterranean Region CS Europe, Middle East, N Africa [many in
Morocco], Macaronesia) and montane tropical Africa; a further 10 minor
segregates (Castroviejo, 1999) sometimes recognjsed in Europe
From adeno- (Gk: gland) and -carpos (Gk: fruit), referring to the sticky glandular
pods
Shrubs; edges of montane forest, woodland, bushland, shrubland and grassland;
colonisers of disturbed places
Reference(s): Gibbs (1967); Rivas-Martinez & Belmonte (1989); Castroviejo (1999);
Castroviejo in Talavera et al. (1999: 189-205); Percy & Cronk (2002); Cubas et al.
(2002)
A distinctive genus, relatively basally branching in the tribe and probably closest
to Argyro/obium; placed as sister to the Cytisinae and Genistinae clades by Cubas
et al. (2002), but as sister to Cytisus sens. lat in Wink & Mohamed (2003); placed
in a separate subtribe Adenocarpinae Rouy by Talavera & Salgueiro (1999); see
also note under Argyrocytisus
Used as ornamentals

Lupinus conicus Photograph by C. E. Hughes Lupinus sp. 'paniculatus alliance' Photograph by c. E. Hughes Meno carpus comp/icatus Photograph by M. F. Gardner

288 LEGUMESOFTHEWORLD TRIBE GENISTEAE 289

 Argyrocytisus bottondieri Photograph by G. P. Lewis Laburnum onogyroides Photograph by G. P. Lewis Laburnum x wotereri Photograph by 5. D. Barfoot

Cytisophyllum o ,Lang 1843 Laburnum Fabr. 1759

Cytisus ,;eel. Phyllocylisus W, D.J.Koch (1836) 2 spp ; montane SC and SE Europe
1 sp.; Meditcrrnnean area of S Europe from Spain ro Jta ly Derived f1om the ancient Latin name for a broad-leaved trefoil , mentioned by
From cytisos (Gk.: ve rmtcular for several kinds o f woody Legumino:-;ae) and Pliny
-pbyl!os (Gk.: leaved) Small rrecs or shrubs; edges of forest and in woodland and associated scrub
Shrub; me<lite nanean sh ruhhmd and montan e woodland ReFerence(s): Frodin & Heywood in Tutin et al. (1968: 86); Ern in Cullen el al.
Refere ncc(s): Huxley el al. (1992: 229); Talave1a in Ta1'1vera el al (1999: 216-248) 0995: 537); Talavera in Talavera et al. (1999: 248 - 251)
A bnsr11ly brnnching clement of Cytisinac (Cub::i s et al., 2002; \'\/ink & Mohamed, llasally b1anching genus in the Cytisinae (Wink & Mohamed, 2003; Pardo et al.,
2003) 2001)
G)ilisopbyllwn sessilifolium (L..) O .Lang is used as an ornamental \Xlicl ely grown as an ornamental and 1'eacl ily natu1-alisecl; L. anagyroides Medik.
(golden rain or golden chain tree); L alpinum (Mill.) 13ercht. & J .P1esl, and 1he
hybrid L. x watereri (Kirchner) Dippel, with cu ltiva1 'Vossii' most well known;
Argyrocytisus (Maire) Raynaud 1975 + Laburnocytisus adamii (Poit.) C.K.Schneid ., the cu ltivated graft chimer::i
co mprisi ng Cylisus pu,pureus Scop. grafted on to l. auagyroides; also used for
l sp.; N Africa (Morocco, Atlas Mts)
wood (a substiture for ebony); all species poisonous, especi"lly the pods and
Prom argyro· (Gk: silvery) and -cylisos (Gk.: vcrnacul:.ir fo r se,'e1:1I kinds of
seeds, due to the presence of quinolizicline alkaloids
woody Legllm in osae)
Shtub; 1nontane ceda r-o;ik fore!'it edges cmd associaced bushland
Refere nce(s): Huxley el al. 0992: 813)
Relativ •ly bas;illy branching in the C.)~l<ft/(/e ( ub:is (I/"'" 2001; Wink &
Podocytisus Iloiss. & He ldr. 1849
Mohalll{.'<I, 2003); Talave"' & Salgueim 0999: 216) observed snmll i;fa nds under 1 sp.; SE Europe (Albania, Greece, Yugosla via) and Turkey
the intlumcmum of the pod ::md tr.rnsf~rrcd the: species to .11do110<·m1111s, but rht: Promj>ado- (Gk: foot), alluding to a Cy1is11s-like plant with stipitate pods
llower.; ore d iscorda tlt nncl lhc glnn<~ :trc nOI l:l\':1tc Shrub; edges of rnontane forest and woodlancJ, ri verine
AtJV•mc li.<11.( /Jollrmtlierf [Maire) lt1y11aud (• Cytf$11Sballandieri ~faire; pineapple Refcrence(s): Chamberlain in Davis 0970: 33); Prodin & Heywood in Tutin el al.
broom, Moro<' .H\ brornn, or ydlow mll) is ~in nHrm.1ive ornamen tal shrub or small (1968: 86); Mayer (1981)
tree for tempe1a1c ga1·dens Podocytisus caramanicus Boiss. & Hcldr. is a relictual species with a disjunct
clis1ribution, ::1llied to Laburnum, Hesperolaburnum and Cytisus

Petteria C.Prcsl 1845

Cylisus sect. Pe/leria (C. 1'1csl) Polhill 0976), in pa1t
l .<p.; SE Elll'ope (A lbania , Yugoslavia)
Named for Frnnz Petter (1798-1853), <rn Austrian schoolteache1 who wrote about
Balkan plants
Shrub; moncane m cdi te1ranean shrubl:rnd and scrub
Refercncc(s): Heywood in Tutin el al. (1968: JOI); Ern in Culle n el al. (1995:
537 - 538)
l<clic1ual :incl b:is:illy lmmching in Cy1 sm:ic Wink & Moh11mcd, ZOO.~ ) 1 ~1
l'ellona IYllllL' ll/tlC(}{I (Sleb<!J') C. l'resl (Allx111ian broom) is i:rown us n11 .,rnonien

Petter/a ramentoceo Illustration by H. G. Reichenbach Podoc•"'

T•tsus coromanicus Photograph by R.B.G., Kew

290 LEGUMES OF THE WORLD TRIBE GENISTEAE 291

 Cytisus scoparius Photograph by G. P. Lewis
HesperolaburnumMai re 1949
1 sp ; Morocco ( Anri-Aclas Mounrains)
from bespero- (Gk .: west) and Laburnum (q.v.), referring ro its weste1n dis11ibulion
Shrub; Arga nia woodland on alluvium
Reference(,): G1einwakl el al, 0992); Lewalle & Monlforl (1997: 150); P:i rdo el al.
(2004)
Reli ctual and basa ll y branching genus in lhe Cytisinae (P~udo el a/_, 2001), <1lli<:'.d
to Podocytisus, with <:1 genei;JI ;:1ffinity to Labr11·11u111 (Gl'cinwJlcl et al , 1992)
Cytisus D cs f, 1798
Cbai1iaecy1is11s Link (1831); Sorol/Jamnus \V'imm. (1832); Spa rlocyfr:;us \Vebb &
Gertlicl, (1840); Coro1ha1111111s C.P1esl (1845); Cbro11m11b11s K.Kocli ( lH'i.1)
<.:. 65 spp.; Mac:aronesia, N Af1ic1, Europe, Turk ey, m~1rgin<illy into the /\liclcllc East
~rnd Caucas us northeast to Russi~1, widely inc1oduced elscwlie1 e
fr'om cytisos (Gk.: vcrmicul~11 for scve1al kinds of woody Leguminos: tc)
Shrubs and he1bs; edges of forest or· woodland, mon1anc: and coa:;1al ~CJ uh cind
grassl::i.nd , ex tending into medi1e11anean sh1ubland (m::i.quis) and di~tui bcd rl~JCeS
Hcference(s): Frodin &. Heywood in Tut in el al 0968: 86 - 93, including
Cba11iaeC)'lisus) ; Cri slofolini (1991); de V1ies in Cullen el al. (1995: 538- 51'1);
Y:ikovlev. el al. 0996); K;iss & \Vink 0997); Cub"s el al (2002); \Vink & Moh:11ncd
(2003); Prn·do el al, (2004)
Di stin ctive genus and, with Lhe other min01~ seg1cg;11es in the suh11ibl' mostly
included by Polhill (1976), e:isily dislinguishecl from the Gen isti n<Je b\' the
chara cteristic.; folcme rather than oblong keel, th ough compo nenrs or the two
subtribes ha ve often been intermixed in the 1x1sr. Th ese -=scgregales persist in loc'.11
flori sti cs. The sc heme of Bisby (1981) is adopt~d here, with rhe mostly 1nonotypic
basally brnnching elements, Podocylis1t"<i~ Cytisopby/111111 Pelleria and A1p,yrocyli5 U5
1 Cytis11s
excluded from Cyl isus as well as rhe minor .-;egrega tes 1 Calicolome and
Lumbo!ropls (which probahly should be re-inclt1de d now, see be low). Bi,..,by also
cxd udecl 11\e h1rge S<:8rc8'11e C/H1111mKJ11.<11.<, which has ne3rly 30 ·fl""' . wilh "
tubular cal yx, but is re-inc.ludcd here ollo' ing Cristofolmi (1991); K:i<S ·"Wink.
( 1997); Cubas q111/. (2002) and \Vink & Mohan!C(I (2003). l'lorisi ic n oun1 fol" E
Eu1·ope, the Ibe ri~1n Peninsul:a and Macarones ia, ho wcve1', still rend to seg1eg:ite
01a111aecylisus (e g., Y<1kovlev, 1996, but nol Tal:i ve ra & Solgueiro, 1999); 1he
.:ir u111S<."rip'1on rem..iln II • AAmc whc1hct :u gt ncrl~ or :o.t.~C onftl I '\1 •I
1\ considc1:oble number or p<.'ci • "'" 11.IC<l n~ omamcnials; pro/Jf<t111S. f. ti~
lu~crnc) Jnd ttllie• from th• C:m •ry Jsl:inds :ire US<."<I for w111dhtc:1k., hl!dW'-" :in
for rorage, ~1lthough the presence of alk1!oicls c:in cau se poisoning; C. sco/111';/{s.
(L.) Link (broom, scotch broom) is used as ;rn ornam ental, for soil sW l)ili~:UifJI\ (it
.:11lonL<es cn•lly ond h:ls lx:com~ w idely n• iurnllscd), b:iskcsry, hrooms. set ,._'llS,
(lyes, J1bre (forP"Jl"'" clorh ond ncrsJ, c:rnnlns, csscn1 fn l ofls (from no""-'"'· or
J><:rf11111cf)•), medicine, :md ured :1< n coffee .<ul ,;i 1t.u1c n.ntl condi ment 11hhu<1i:h
Cytisus proliferus Photograph by G. P. Lewis Giution is need ed ~ 1s plants ~ire toxic
292 LEGUMESOFTHEWORLD
Lembotropis nigricans Photograph by J. Hooper Cal/cotame spinosa Photograph by J. Hooper
Lembotropis Griseb, 1843
Cy1is11s secl , Le111bo1ropis (G 1·iseb.) Bcnlh. (1865)
2 spp.; C & SE Eu1'0pe n011heast to C Russia
from le111bo- (Gk .: a li1tle boat) and l1'0pis (Gk.: keel of a ship), alluding 10 1hc
shape of the keel-petals
Shrubs; mesophyric shruhland and forest
Reforence(s) : Sk:ilicka 0969); Heywood in Ttllin el al. 0968: 86)
Uns~tisfactory minor segregate of C)iffsus, Lo whi ch Calicotome might be allied;
probably beue1 t1eatcd as a sec1ion of Cy1is11s (Bisby, 1981; K;iss & Wink, 1997;
Cubo s el al., 2002; \Vink & Moh:irnccl, 2003)
Lembonupis 11igricans (L) Griseb. (bbck b1oom) is used as <in ornamental. but
ca n become a weecl
Calicotome Link 1808
2 - 3 spp. (Greulcr, 1986); Mediterranean region of S Europe, N AfriG1 to Middle
East and Tu1 key
From calycos- (Gk.: ca lyx) and -tome (Gk.: cut , division), alluding to the media ll y
circu11"1scissile cal yx
Shn1bsi medilen:mean shrubland ( maqui~) o f co:ists and inland 1nountains, in dry
l'ocky a1 c:1s
Heference(s): Gibbs (1968); Polhill 0975); G1c ut er (1986); Ga1t:ia Murillo in
Talovera el al. (1999: 182-189)
Close Lo C)'tisus sect Tuhocylisus (C/Jm1wecytis11s) but with ;i distinctive calyx;
cun·em 111~lecula1· analyses <ill pL1ce Ca/icolom e ;is nested within C) 1tisus (Kfiss &
\'\link , 1997; Pyne, 1999; Cubas el al., 2002; \'<link & Ivlohamed, 2003) and it seems
;1ppropriate to now include both Lembotl'Opis and Cnlicotome ~1s sections of
Rare ly culcivatecl a.s an ornamental
Echinospartum (Spach) Rothm 1941
5 spp.; SW Europe (S F1~nce , Spain , Portu gal)
From ecbi110- (Gk,: hedgehog) and spc111011 (Gk .: foe· Stipn te11acissi111a, a kin<l o f
g 1·ass used in weaving <ind cordage) , alluding to the spiny leafless habit
Shl'ubs ot suffiutices; exposed mo111ane slopes and rocky areas
Reference(s): Talave10 in Tabve'"' el al (1999: 119 - 127)
Segregole o f Genis/a (see Gibbs, 1966); apparently ba sall\' branching in the
Ge ni s tin ~1 e :md rolyphyletic in the an;:ilyses of Pnrclo el n/, (2004), with two clacles
(the c:ilcicolous and silicolous species groups) .se rx 11'::1tely nested in or nea1 to
Cen ista
T he authority is sometimes att1ibuted to Fm11reau, but the n:imc is only
m entio ned by Fourreau in Ann . Soc , Linn Lyon 16: 358 (1868)
Echinospartum algibicum Drawing by A. Jordan
TRIBE GENISTEAE 293
Erinacea anthyl/is Photograph by G. P. Lewis Retama raetam Photograph by 5. Collenette Genlsta sp. (= Rivasgodaya nervosa) Photograph by RBG, Kew Genista tinctoria Photograph by G. P. Lewis

Erinacea Adans 1763 Genista L 1753

I sr; S\Y/ Europe (Spain, fl<mce) and N Africa Cbamaespai tiwn Adans, (1763); Genisle/la Oitega (1773); Teline Medik. (1787);

Q
From erinaceus (L.: rc:;embling a hedgehog), alluding to the spiny tufted habit Pterospw111111 (Spach) K.Koch (1853); Riuasgodaya Esteve 0973)
Subshn.tb; mc<lite rranean shrubland and scrub on limestone and dolomi[e; rocky c..: . 90 spp.; Macaro nesia, N Africa and Europe co Turkey, Midd le East, Caucasus
moun1ain slopes and northeast to Russia, introduced elsewhere
Reference(s): Haynaucl (!989); Ern in Cullen et al (1995: 511); Tala\'CJa in Talaverd From genes/a, genisla (L.: for a broom plant); the name from which che
et al 0999: 208-211) Plantage net kings and queens of England lOok their name Cplanta genisla)
D istinct genus possibly Jllied to Cylisus (on th e ba.si.s of morphological l'viclence) Shrubs and he rbs; forest edges, grassland, medltenanean shrubland (often in
~incl raised to s ubtribal 1ank by Talavcr::i & Salgueiro (1999); molecular evidence rnaquis), rocky mountain slopes, coasts ancl disturbed places
currently inclicares a position in the Genisrinne (or in Ge11isla sens /C1/J near Reference(s): Gibbs (1966; in Tutin et al., 1968: 94 - 100); Gibbs & Dingwell
Retallla or SjJm1iJ1111 (Pyne, 1999; Wink & Mohamed, 2003; Pardo el of, 2004) (1971); Arco (1983); Ern in Cullen et al (1995: 546-549); Talavera in T"lavera et
Erinacea a111byllis Link (hedgehog b1oom) is an at1mctive orn:rn1enta l for 1ock al. (1999: 45-119; 128-137); Percy & Cl'Onk (2002); Pardo et al, (2004)
ga1clens Main genus of the Genistinae but its circumscription is difficult, app~rently
cont<l ining a numbe1 of other genera in the Genistinae in its broadest sense, and
with va1ious segregates commonly recog nised (nm;:ibly Chamaespa11i11111,
Retama H:rf 1838 Pterospar/11111 ancl Teline; e.g ., Yakovlev, 1996 :r nd Talavera el al., 1999) in its
f.ygos Adans (1763) o;:inowest sense The analyses of Pardo el al. (2004) suggest recognising Genis/a
4 spp.; I\'lacamnesia, S Europe, N Af1ica 10 Midd le r:asr, Saudi A1abia and Yemen in the broad sense (i.e ., including the gene ia Retama , Stauracanthus, Ulex and
From the Arabic relem rn• re/am for this broom-li ke hush
Ecbinospmtum ~11nong othe rs) with at le~1 st three subgenera. Teline (Rivasgodayt1)
Shrubs; high mountain slopes, bushland, thic ke t rind grassland ro desc1 1s and Jacks any evident apomorphy and is polyphyleric in the analyses of Cuba:; et al.
sandy coasts, in arid places (2002) and Pardo el al. (2004); it is often conside red intermediate to Cytisus, but
Referen cc(s): Zoha ry (1959) can be clearly distinguished by the oblong keel, ~n apomorphy unique and
\V'cll-clefin ecl genus, p 1obably bas~dly branching in the Genistin:1e 01 within consiste nt within the Genistinae (Gibbs, 1974 ; Polhill, 1976; Bisby, 1981). Talavera
& Salgueiro (1999) revert to an aitificial classification of the subt1'ibe, separating
Gemsta se11s. lat (Zohary, 1959; Polhill, 1976; Crisp et al., 2000; Wink &
Moh"mcd. 20\U: 1'11rclo et"'·· 20011; "°""' :rn:rl)':'<..,. (e.g., Pynl'., 19?9• Ka. •"'
Teline from Genista by the millatc ('strophiolate') seeds (a notably unstable feature
in the tribe, found "lso in sect. Genis/a, see Gibbs, 1974 and Polhill, 1976).
Wink. 1997: Pttrtlo ut al., 2004) pl.ice l?Oltw111 :~ :r m re tlcri•'t'<I cl.,tm:nl "11hln
Genis/a TalaveGJ & Salgueiro 0999) 81so include sect. Chronanlhus from the middle of
Retama mo11ospei·111a (L.) Boiss. (white broom) is grown as ;111 orn;1111cntal C)'tlsus in Teline, beG1use it bas a deeply bifid uppe1 calyx lobe; this is rejected in
the ana lyses of Cubas el al. (2002) and Pardo el al. (2004), who place Cytisus
fontanesii Spach ex Ilall (~ Chronantb11s bijloms (Desf.) Frodin & Heywood)
Gonocytisus spact1 1845 within Cytisinae
A number of species are grown as 0 1namentals; G tinc1011·a L. (clye1"s gree nweed,
3 s pp.; E Mcditerrane~111 region in SE Europe, Tul'key and Middle E.asl
dyer\ broom) was used frnrnerl y as a d ye plant in Eu1'0pe, the yellow d ye from
from .'<onia
. (Gk.: angled), alludi ng to a Cytisus-like pl:tnt w ich angular-,,·tngcd
rhe flowe rs was mixed wit h the blue dye of woad to produce Kendal g1een, a fine
stems
green textile dye; also used (as are ot he r species) for fibre, medicine, ground
Shrubs; woodland and mediten·anean shrubland (maquis), cliffs and d1 y hi llsides;
covci~ low hedging, fo1· perfume (from essenli:ll oils in rhe flowers) and as a
lowland to submOl'Umu~
coffee substitute and condiment
lll.ference(s): Heywood in Tutin el al. (1968: JOI); Gibbs in Davis (1970: 21 - 23 );
Polhill (1976)
Discrele genus allied ro G'enista and Rela111a, but remains to be sampled in
molec..: ul:i r am1lyscs
Alk81oicls in G'o11ocylisus show 8nti-microbial activily

Gonocytisus pteroc/adus Drawing by P. Halliday

Gen·1
sta microphyl/a Photograph by G. P. Lewis

294 LEGUMESOFTHEWORLD TR IB E GENISTEAE 295

 Stouroconthus genistoides Photograph by c. E. Hughes

Spartium i 1753
I sp; around th e Me dite minean CS Europe, Tu1key, Middle East a nd N Africa), but
widely cultivated and naturalised (including N Andes in S Am e rica)
From Greek spanon, a name given to a numbe1 of plants like the broom and Slipa
lenacissima L. (a kind o f grass used in weaving and cordage), a llud ing to their habtl
Shrub; mediterranea n sh rubland (maq uis); cliffs, ra vines and d ist u1 bed places
Re fe rence(s} Ake royd in Cullen el al. (1995, 545); Tala vera in Tala ve ra el al.
(1 999, 206-208)
\Xlell known ~me! possib ly basally branching ge nus in the Ge nistinae near
Erinacea (e.g, K{iss & Wink, 1997) or e mbedded within Genislc1sens lat. (Cubas
el al., 2002; Wink & Mo hamed, 2003)
Spmtium junceuni L. (S panish broom, weave rs broom) is wid e ly g1own as an
orna mental and so il stabiliser bur it is read il y in v<isive be co min g <1 nox io us weed;
also used frn its fi bre (for cordage com para ble co jute, made from rh e b r;;mches),
baskc ny , as a dye, perfume (from essential o ils in the flowers), a nd fol' medicine
but the plant is h ig hl y roxic, es peciall y the seeds

Stauracanthus Link 1807

3 spp .; SW Eumpe a nd NW Africa (P o 11uga l, Spain, Algeria and Mo rocco)
Fro m s/auro- (G k.: cross) a ncl -acantbos (G k,: spiny), refe1ring to rhe spines on
the angular br<inches
Shrubs and suffrutices; mediteiranean shrubla nd ( maquis) and woodl<md; heaths,
sat'lti)' r -~rony nllul'la, coo.i:1l .scrub :mt! dunes
Hcfofl)nC\.'( ~: lJinlz (1 ftl. (1990): P.Jh';I & ornlnho in T:1!:11>c1:1 el ' " · ( 1999, 240-245)
: •grcg:11 >or UI<'" :mcJ n•,,;tcd w irhfn G<'llf.<ffl In the • nn l)'se of Pan lo el al. (200 4)

Ulex 1.. 1753

10-20 spp.; Euro pe (m os tl y in th e wes t) a nd N Africa; U. <'llllJ/XIQttS widely of
introduced and natura lised in many montaoe (including u·op1cnl monwric) partS
the world
Derived from the a ncient Latin name ulex, ulicis used by Pliny fo r a shrub
resembling rosern ciry
Shrubs; mediterrane an s hrublancl (rnaquis) , bushland and woodl and; exposed
places, heaths and d unes ,
Re ference(s} Maxwe ll in Cullen et al. (1995, 550); Cubas in Talavera el al. (l999
212-~9): .'\ino1rchc et al (2003) _ ~ue
\X'cll-knnwn 1(<'111•< In liu! Gcnlslinae; species <lclimil:ulon is 11Jl!>l~hle. "Ith rcucu
char.i•ier '~uiuri<.>11 nffocu:~I by hybridls uion ~nd p0l1•11loldy (Cub:is In 'l'•i,ivcn; c<f
al., 199'): ll2- 239); riesled withi11 c,.,,1.11a in 1hc ~nolyseis of l'nrdo el 111 t.lOO ).
' e11rofN1(!t_1s L. (~orsc, 1'l.trze
Chex • or w 1· . to 01 Ilt.'I =-1""'c:fe<)•<
u n , nmncs :1 tso applied ·- l IU
w rnc1im"' gmwn for h•>dges I<.> e nclose ll\·cs1ock mos u .soil st:11Ji1iS<·1, Jmr ~' '
1

to become ::1 w eed a nd highly inflammab le in dry areas; <ilso L1sed ::is a tea
Lab
Ulex europaeus Photograph by G. P. Lewis (infused from rhe fl o we rs), as orn<1me ntals, d yes a nd fo ewood urnum anogyroides Photograph by B. D. Schrire

296 LEGUMESOFTHEWORLD TRIBE GENISTEAE 297

TRIBE
j\Jilorpheae byJ.M.Lock
. AJUOrpheae Boriss. 1964
1~ 'be Galegeae subtribe Psorale inae A.Gray (1863), in part, as 'Psoralieae'
1
; be Daieeae Hutch. 0964), as 'Daleae'
:r:be Psoraleeae sensu Hutch. (1964) in small part, as 'Psoralieae'
The A.morpheae were expanded by Bentham (1865) as
ubtribe 'Psoralieae ' of his ~ery broadly drawn
(i~le.geae. The latter also contained the genera now
,1aced in Indigofereae, Brongniartieae, Millettieae, and
kobinieae, as well as in the present restricted (but still
hererog n Ott Galegeae sens. lat. Both Gray's and
Bentham's concepts of Psoralieae included genera here
placed in Amorpheae, as well as a broadly
.ircums ri bed Psoralea . Borissova (Nov. 1964)
propo eel a tribe Amorpheae containing the single
genu Amorpha. Almost simultaneously, Hutchinson
(De . 1964) introduced a tribe Daleeae (as 'Daleae')
in luding four genera : Dalea, Thornbera,
Petafoste1non, and Kuhnistera, the last three placed
as synonyms of Dalea by Barneby (1977).
Hutchinson attempted to use petal insertion as the
character separating Daleeae and Psoraleeae, but
Barneby (1977) showed that Hutchinson's definition
and interpretation of this character were unsatisfactory.
Barneby (1977, 1981) added Apoplanesia, Eysenhardtia,
Panyella, Amorpha, Errazurizia, Psorothamnus, and
Marina, all formerly placed in Psoraleeae, to
Hutchinson's (1964) Daleeae to produce the concept
of Amorpheae as accepted here.
The main character distinguishing Amorpheae lies
in the inflorescence position, which is terminal in
Amorpheae and axillary in Psoraleeae. Additionally,
the leaves of Amorpheae are basically pinnate
(although sometimes trifoliolate or unifoliolate by
reduction), while those of Psoraleeae are basically
trifoliolate (although sometimes palmately >3-foliolate
or even pinnate). The basic chromosome number
appears to be 1O in Amorpheae (reduced in some
species of Dalea) but 11 in Psoraleeae. Stirton (1981a)
added further characters that reinforce the distinction:
the form of the cotyledons, the positions of the embryo
and radicle in the seed, seed shape, fruit structure,
and the pollen. Ferguson & Skvarla (1981) also pointed
out that the pollens of Psoraleeae and Am rpheae
have very little in common. Barneby (1977) ug ted
LEFT p
sorothamnus spinosus Photograph by R. Spellenberg
that the papilionoid flower of the more derived species
of Dalea may have evolved independently from that of
other papilionoids, but this hypothesis is rejected by
McMahon & Hufford (2004), based on studies of floral
development in Dalea, Marina and Psorothamnus
(McMahon & Hufford, 2002) .
Molecular work (Doyle, 1995) initially placed
Amorpheae among the New World tropical tribes,
as sister to a clade comprising Lotus and members of
the IRLC clade. More recently (e.g., Doyle et al.,
2000; Pennington et al., 2001 ; Kajita et al., 2001),
Amorpheae emerges as one of a number of equally-
ranked groups with the Dalbergioid and Genistoid
clades. Lavin et al. (2001a) included sequences from
four genera of Amorpheae (Apoplanesia,
Eysenhardtia, Amorpha and Marina) as part of their
study of the Dalbergioid clade . Three analyses, each
based on a different gene, a nd one combined
analysis using both molecular and morphological
data, all agreed firstly in placing Amorpheae as sister
to the whole Dalbergioid clade, and secondly in
supporting the monophyly of Amorpheae. This is
confirmed in subs equent analyses (e .g.,
Wojciechowski, 2003; Wojciechowski et al., 2004;
McMahon & Hufford, 2004) . All genera in the tribe
were sequenced in the analysis of McMahon &
Hufford (2004) and two well supported clades were
recognised: the Amorphoid clade comprising
Parryella, Amorpha, Apoplanesia, Errazurizia and
Eysenhardtia; and the Daleoid clade with
Psorothamnus , Marina and Dalea (Fig . 39) .
Psorothamnus is paraphyletic in the latter study with
two monophyletic clades dividing neatly along
taxonomic lines. One of these, the Psorodendron
clade, is basally branching in the Daleoid clade and
McMahon & Hufford (2004) state that this will be
resurrected at generic level, thus recognising 9
genera in the tribe. For this account, however,
Amorpheae is treated as comprising 8 genera and
(245) - 247 - (248) species (Fig. 39) .
TRIBE AMORPHEAE 299
 Andira
Hymenolobium (see page 309)

Dalbergioid clade (see page 309)

Apoplanesia )>
s::
· Parryella 0
;;u
"O
Amorpha :::c
0 )>

Errazurizia
0
n
s Apoplonesio poniculoto
cu
0..
0 Apop/ones/o poniculato Photograph by C. E. Hughes Photograph by G. P. Lewis

(1)
;:;o
Eysenhardtia "lJ
Apopla11esia c.Frest 1832
1 sp.; S Mexico, Nicaragua to Ve nezuela
I From apoplanesis (Gk~: aberration. leading astray), referring to the structure oF che
Psorodendron clade 0 rn flowers w hich is unusual w ithin Papilionoideae, hav ing 5 free petals o n the 1im of
a hypa nthium
)> )>
I
Sma ll trees; seasona lly d1y tropica l fo1est and woodland, hillsides, river valleys
Psorothamnus sens. strict. m rn and roadsides
0 Refe1e nce(s). McVaugh (1987: 280-281); McMa hon & Hufford (2004)
In the ana lyses of McMahon & Hufford (2004), th e ge nu s is either basally
0 brnnching in the Amorphoid clacle (based on ITS sequence dara only) or sister to
Marina n the Errazurizia-Eysenhmdtia subc\ade (in the trnK-matK analysis and the
cu
0..
combined analysis with ITS data)
The wood of A. paniculata C.Presl (cl1ulul) is veiy hard and used fo r cabine t1y,
Dalea (1)
cr<i fts and bows; the bark yields a yellow dye ; also used as ornamentals and in
agroforestiy

Parryella To rr. & A.Gray ex A.Gray 1868

1 sp .; SW USA
Named afte r C.C. Parry (1823-1890), English-born Ame rica n botan ist and exploreo
-- = Psorathamnus sens. lat. known as the 'King of Colorado botany'
Shrub; warm and co ntinental temperate dese11s; open sandy plains and hills,
dunes and washes
Heference(s): Isely 0998: 779); McMahon & Hufford (2004)
Pcmyella is either basally branching in the Amorphoid clade (in the analysis of
11·nK- matK dat~1 only, and the combined analys is with ITS data), oi· sister to
Amo1pba in the ITS analysis only (McMahon & Hufford, 2004). Tl1e flowers of P
filifolia Torr. & A.Gray lack a corolla and in som e trees in McMahon & Hufford
(2004) this is sister to Errazurizia rotu ndata (Wooton) l3arneby, which unlike the
other species of the genus also lacks a co ro lla; E. rotunda/a may thL1s be better
placed in Panyella (in which it already has a combinarion)
Used as soil binders and in erosion control, for medicine, insecticides and sterns
used for baskets and brooms

10

FIG. 39 Diagram of relationships of the tribe Amorpheae after Lavin et al. (2001a); McMahon & Hufford (2004) Parryel/a filifolio Drawing by P. Holliday

300 LEGUMESOFTHEWORLD TRIBE AMORPHEAE 301

!
I
Amorpho fruticoso Photograph by G. P. Lewis Amorpho nano Photograph by G. P. Lewis
Amorpha L. 1753
c. 15 spp.; N America from S Ca nacl:-i co N Mexico, most species in SE USA
From ammpho- (Gk ~ : shapeless, deformed), refe11 ing ro the corolla which IJcks
wings and keel
Shmbs or subshrubs; subu·opical to co nti nent~1I ;md cool rempc1atc woodhrnd,
wooded g 1-::1ssl<mc1, thicket crnd grassland, often in sandy a1eas and on hillsides or
along .5t1eams and in other moist areas
Reference(s} Wilbur 0975); Jscly (199g, 132 - 141); McMahon & Hufford (2004)
Very litrle sequence divergence is evident between species of A11101pba in eithe1
ITS or 1rnK-111a/K datasets (McMahon & Hufford, 2004); the A . ji-11/icosa complex
is ve iy diverse mo1 phologically
Use d <15 windbreaks, soil cover, in erosion co ntrol and as ornamcnt~1ls, e ~g . , A
CC1J1esce11s Nutt ex Pursb (lead pl'1nt, shoest ring) and A,J h11icosa L. (h:1st:ird
indigo, false indigo) are cultivated; th e latter occurs widely in lcmpcr;Ht: regions
of the N Hemisphere (and S Ame ri ca) and can become n;Huralised; other uses are
leaves as :.l tea and tobJcco substitl1te, medicine, insect repellents and dyes
Errazurizia Phil , 1872
Psorobal11s Rydb. (1919)
4 spp.; SW USA ;rncl NW Mexico (3 spp.); coasta l Chile (1 sp.)
The name commemorates the prominent Chilea n fomily Errftzutiz of Basq ue
ex tra ction, who were poets, ·writers, bishops nn<l statesmen (Barneby, 1977)
Xcrophytic shrubs; subtropical and mcdit crrnn ean fog-cleseits (Baja California,
Sonoran and At;:1cama deserts), on 1ock or sa ncl
J(e fere nce(s} Ilameby (1977, 13 - 21); McMahon & Huff0t•cl (2004)
See note under Panyella about relationships or E~ roflmdola (\V'ooton) U:1111eby
(1 ou nd leaf broom)
A co mmon name fol" species is dune b1oom
Errozurizio megocorpa Photograph by G. P. Lewis
302 LEGUMES OFTHEWORLD
Eysenhardtia orthocorpo Photograph by c. E. Hughes
Marina parryi Drawing by P. Halliday
Psorothomnus schottii Photograph by R. Spellenberg
Eysenhardtia Kunrh 1824
c. 12-15 spp.; mostly Mex.ico exte nding to SW USA and C America (one possibly
confined to Guaternala)
Named after C.G. Eysenharclt (1794-1825), botanist and zoologist, Professor at
Konigsberg (now Kaliningracl)
Shrubs or uces; tropical to warm lemperate seasonally city forest, woodland nnd
xerophilous scrub and thicket (matorrn.I), often on rocky slopes
Reference(s} Lang & lsely (1982); McVaugh (1937, 519-523); Isely (1993,
561 - 562); McM;ihon & Huffot'd (2004)
Eysenhardlia 1exa11a Scheele (Texas kiclneywood, bee brush, vara duke) is used
fo1 wood, medicine, livestock fodde r and wild-life food (including bees), el'osion
control, ornamentals, reforesta1ion rmd as a source of a yellow-brown dye
Psorothamnus Ryclb. 1919
Psorode11dron Hydb (1919)
c. 9 spp., some with varieties; SW' USA and NW Mexico (Baja California, Sonor;m,
Mohave and Chihuahuan <lese11s, and desert basins within the Colo1aclo Plmcm1
and intermountain USA)
From psoro· (Gk , scab) and 1bamnos (Gk., shrnb), referring to the gland clots o n
the plant
Xe1ophytic sh11.1bs and subshrubs; subtropical <ind Medi[erranean deserts, on
outwash Fans and by tempoi ::ity wateicourses
Reference(s), Ilarneby (1977, 21-54); lsely (1993, 805-815); McMahon & Httffot d
(2004)
Psorothamnus section Psorotbamnus (Barneb y, 1977) is based on 4 species with
sessile or nearly sessile flowers, non-exsc1tecl pods and non~spiny inflorescences
and= Psorotbamnus sens sl!'lct . (McMahon & Huffoid, 2004); the Psoroclendron
clade (Ba1neby's sections Xylodafea [- Rydberg's genus Psorodendronl,
Capnodendron and Winn emuca 1), however, comprises 5 specie!-i of stout s hrubs
or trees with spiny inflorescences and/or exsertecl pods; at present Psorolhn mnus
sens Jar. is a par<Jphyletic basally brnnching group in the daleoids (see
inrroduccion to the tribe above) and McMahon & Hufford (2004) state that in a
futrne publiGttion, the Psomdendron cla<le is to be resurrected at geneiic level
Psorotbammts spi11osus (A.Gray) Harneby (smoke tree, page 298), is <1n attractive
desen bndmark shrub; the wood is used for fuel and gives off an aiom~ti c
smoke; olher species, e.g ., P. sco/xwius (A.Gray) RyJb. (broom dalea), a1e used
for basket weaving, medicine, dyes and reforestation
Marina Liebm. 1854 (see next page)
TRIBE AMORPHEAE 303
Daleo c/iffartiana Photograph by D. Bailey Daleo carthagenensis var. brevis Photograph by G. P. Lewis

Marina uebm. 1854

38 spp.; all narive and mostly endemic to Mexico (although almost lacking on the
Mexica n Plateau), extending to SW USA a nd C Amelica ( highla nds of G uatema la,
with 1 sp. a weed through C America, to Cuba and Venezuela)
Named in honour of Ma rina, or Ma linche, the cacique's [chief's) cla ughte1 who
interpreted for Hernan Cortes du ring the conquest of Mexico (llarneby, 1977)
Pe renn ial herbs or shnibs; medite rra nea n and subtropical d1y gr~ssbnd, bushland
and woocJ lnnd to a rid tropica l deserts, on dunes, rocky or sandy ouLwas h fans,
hillsides and stream ba nks
Reference(s): Barneby (1977: 55-135); McVaug h (1987: 609 - 626); McMahon &
Hu ffo rd (2004)
A we ll-supported mo no phyletic genus siste r to Da/ea (McMa hon & Hufford, 2004)
Used for brooms; the roots of some species produce a ye llow dye

Dalea Lucanus 1758

Kubni:stera Lam, (1792); Petalostemon Mic hx. (1803); 7bornbera Rydi>. (19 19)
c. 165 spp.; SW Canada, USA, C Ame rica, Ca1·ibbea n and sourh to Argencina
Mostly Mexico (c. 11 5 sp p .. c. 75 e nde mic) a nd USA to Canada (c 23 e ndemic
spp.), and a cenrre of ra dia tion in the Andean region from Colombi;1, Peru,
Ecuador (and Galapagos Is.), Bolivia, to NW Argentina and coastal N Chi le (c. 27
spp.); l sp. nat uralised in the Phili p pines
Named for Dr Samuel Dale (1659 -1739), English botanist, apothecary an d
p hys ician at Braintree, Essex
Xerophy ric herbs o r shrnbs; temperate, continen ta l Lempernre, mecl itc rianean,
subtropical and montane tropical forest, woodla nd, thorn-t hicke t, gr~1 ssbncl,
shru bla nd and clese1t, o fte n in gull ~ and o n mcky hillsides
Refe rence(s): Wemp le (1970); 13arneby 0 977: 135 - 589); Jsely (1998: 493 -53 2);
McMahon & Hufford (2004)
Barneby 0977) trears Dalea as comprising 5 subgenern, most of which :u e
di\•ided further into sections; Dalea is a well-supported monophyletic genus
except for D.filiciformis B.L.Rob. & Greenm. which is sister to Marina (McMa 1 n
10

& Hufford, 2004)

Used ;ts soil stabilis\!rs, green ma nure, stock forage , rnedici ne, o rnaincncals,
brooms and in making baskets; D. pwp urea Vent. (purple prairie clO\'t.' r) mocs are
chewed for the ir p leasa nt taste and the dried leaves are a tea subscirurc

Daleo batteril Photograph by D. Bailey .\!or'

tna porryi Photograph by R. Spellenberg

304 LEGUMES OF THE WORLD TRIBEAMORPHEAE 305

rR tBE
p albergieae sens. lat. byB. B. KlitgaardandM. Lavin
.tJ!)ergic:ic sens. lat. including:
f)~be D:il bergieiie Bronn ex DC. 1825
T Tribe Aeschynomeneae (Benth.) Hutch. (1964)
Tribe Hedysareae subtnbe Aeschynomenmae Benth. (1865), as 'Aeschynomeneae'
Tribe Coronilleae subtribe Aeschynomeninae (Benth.) Schulze-Menz (1964)
Tribe Adesmieae (Benth.) Hutch. (1964)
Tribe Hedysareae subtribe Adesmiinae Benth. (1865), as 'Adesmieae'
Tribe Hedysareae subtribe Patagoniinae Taub. (1894)
Tribe Coronilleae subtribe Patagoniinae (Taub.) Schulze-Menz (1964)
tribe Desmodieae (Benth.) Hutch. subtribe B1yinae B.G.Schub.
'fhe pr · nr circum. cription of Dalbergieae sens. hit.
cenrnin radical hang ~ to Dalbe rgieae sen u. Polhill
(1981d : 233 - 2 2). In th fir ·r in ta n e it take a
rraditional view of the trib by in lu ling genera uch a
vatairea and Vataireopsis (now in the Vataireoid clade,
see Figs. 1 & 40) and Andira and Hymenolobium,
which in Wojciechowski et al. (2004) are sister to the
combined Dalbergioid clade of Lavin et al. (2001a),
plus Amorpheae. The bulk of the treatment, however,
recognises the cryptic Dalbergioid clade (Lavin et al.,
200la) as comprising tribe Dalbergieae sens. lat.,
diagnosed by the synapomorphy of aeschynomenoid
coot nodules. This clade includes all the genera placed
in the Dalbergieae sensu Polhill (198ld: 233- 242), the
Aeschynomeneae sensu Rudd 0981) and the
Adesmieae sensu Polhill 0981g: 355- 356), plus
subtribe Btyinae of the Desmodieae sensu Ohashi et al.
(1981) as well as the genus Diphysa (tribe Robinieae
sensu Polhill & Sousa (1981)). The placements of all
members of the Dalbergioid clade within the
classification presented here and in those of Polhill
Cl981cl: 233-242; 198lg: 355- 356), Polhill & Sousa
0981), Ohashi et al. (1981) and Rudel (1981) are listed
in Fig. 40.
Dalbergieae sensu Polhill (198ld) contained 19
genera defined by woody habit, supposedly
plesiomorphic flowers, pods with a specialised seed
chamber and seeds that accumulated alkaloids. Polhill
Cl981cl) noted that there seemed to be two centres
Within the Dalbergieae: one around Andira with
Hymenolobium, Vatairea, Vataireopsis, Dalbergia, and
MC1che1erium and o n around Pter caipu . He , 1 o
highlighted evidence from wo d a natomy (Ba r ua-
Kuip rs 19 1) bi h showed that An.dira,
Hymenolobium, Vatairea, and Vataireopsis have coarser
Wood structur more typical of members of the
Ophorea than the r maining mem be rs of the
DaJbe rgieae. The study of fruit and edling
lllorphology by Lima 0990) further supported these
LEFT Etaballla dubia Photograph by L. Y. Th. Westra
two centres within the Dalbergieae: one including
Andira, Hymenolobium, Vatairea, and Vataireopsis,
and the second the remaining genera. Most recently
several molecular and morphological studies (e.g., Lavin
eta/., 200la; Pennington et al., 2001; Wojciechowski et
al., 2004) confirm that these four genera do not belong
in the Dalbergioid clade. Since there is, however, still
much work to be done to resolve the phylogenetic
relationships of these four genera, they have been kept
in tribe Dalbergieae sens. lat. in this treatment to avoid
tentative placements, which might be treated by users
as formal.
The tribe Aeschynomeneae sensu Rudd (1981)
contained 25 genera characterised by lomentaceous
pods, although some members lack laments (e.g.,
Arachis, Ormocarpopsis, Diphysa spp., Ormocarpum
spp., and Pictetia spp .). None of the Aeschynomeneae
had previously been considered closely related to the
Dalbergieae, but the work of Lavin et al. (2001a) has
resolved all the 'aeschynomenoid genera' within the
Dalbergioid clade.
The taxonomic history of the monogeneric tribe
Adesmieae sensu Polhill 0981g) is ve1y different from
that of the Dalbergieae. The Adesmieae combines the
presumed plesiomorphic trait of free stamen filaments,
with presence of lomentaceous pods that are
supposedly derived . This combination of features
suggested a taxonomically isolated position far
removed from the Dalbergieae. The analyses of Lavin
et al. (2001a) based on molecular sequence and
morphological data, however, suppo1tAdesmia (nested
together with five genera of Rudd's Aeschynomeneae)
being sister to the Pterocarpus and Dalbergia clades.
The neotropical genera Brya and Cranocarpus
were placed together in a new subtribe Bryinae of
Desmodieae in the classification of Ohashi et al. (1981).
The features common to these genera are periporate
pollen and glochidiate hairs or glandular trichomes.
In the molecular studies of Doyle et al. 0995) and
TRIBE DALBERGIEAE 307
Bailey et al. 0997), Brya and Cranocarpus did not Peru (Hughes et al., 2004) . Three genera hav PLACEMENT IN ALS 1
been place? .in sy~1on:my since 1981 : the Cari~~~
Isolated genera
lack the intron for the chloroplast gene rpl2 nor for the
open reading frame ORF184, which are characteristic genus Belama which 1s a synonym of Pictetia (B ean Vatairea
Vataireoid clade
Dalbergieae
of the other desmodioid genera studied . Bailey et al. Matos c Lav in 1999 , a nd th genera Pacbecoa~t-d­ Vataireopsis Da lbergieae

0997), therefore, suggested that Brya and Cranocarpus A1tbroca1pum which Thulin 1999 synonynii ed nd Hymenolobium Dalbergieae
should be removed from the Desmodieae. Their Chapmannict. In this treatm nt 9 gen rn and ~~der Andira Dalbergi eae

findings were strongly corroborated by the three gene - 1325-(133 1 sped : a r recogni5ed in I alb r . 19)
analyses of Lavin et al. (200la) which place the two 1
·ens. kt!. In lucl ing 4. ba ally branching da ll ~ ~ • Amorpheae (see page 299)

genera in the Pterocarpus clade (Fig. 40) .

One further transfer has been made in the
genenl ompri ·ing . 8 s pecies, and (126J)-i ~~
(1273) pecies in tbe 45 g n era of the Datb ro· .-
26 Adesmia Adesmia Adesmieae
Poiretiinae
clade Amici a
. o~~
Dalbergioid clade since Rudd (1981) and Polhill & clad LLavm et ct/.., 20 la]). Zornia Poiretiinae
Poiretia Poiretiin ae
Sousa (1981). Lavin (1987) transferred Diphysa from the Ormocarpinae
Nissolia
Robinieae to tribe Aeschynomeneae based o n the The following informal groupings of genera are based Chaetocalyx Ormocarpinae
absence of ca navanine in the seeds, a feature on the work of Lavin et al. (200la):
consistently present in the Robinieae (Lavin, 1986). Adesmia clade: 6 genera; c. 360 species; neotropicaJ Riedeliella Dalbergieae
Lavin (1987) also listed 14 morphological characters except Zornia, which is pantropical. Discolobium Discolobiinae

that placed Diphysa with the Aeschynomeneae rather Pterocarpus clade: _2 genera; c. 200 ·p cics ntred Cranocarpus Desmodieae
Pterocarpus subtribe Bryinae
than the Robinieae. In the phylogenetic analyses of in the Neotr pi ·s, Pterocmpw and lylosu11tbe·a re clade
Brya
Lavin et al. (200la), Diphysa is reso lve d in the paniropi al, lnocarpu A. ian, and bapmannfa Platymiscium Dalbergiea e
Dalbergioid clade nested within the 'transatlantic clade' transatlantic.
Platypodium Dalbergieae
(Fig. 40), first identified by Lavin et al. (2000). Dalbergia clade: 17 genera; c. 706 species which are lnocarpus Dalbergieae
Finally, since 1981 four new genera have been pantropical, but nn· d in Africa ; lfleberl auerella Maraniona Dalbergieae
Tipuana Hughes et al. (2004)
published: the Brazilian monotypic Grazielodendron Soemmeringia, Pictetia and Diphy ct a r ne tr pica!'. Dalbergieae
Ramorinoa
(Lima, 1983b), the possibly exti n ct Madagascan Machaerium transatlanti c, Dalbergia and Centrolobium Dalbergieae
endemic Peltiera (Labat & Du Puy, 1997), Zygocarpum, Aeschynomene are pantropical, and Geissaspis Asian. Dalbergioid Paramachaerium Dalbergieae
cl ade Etaballia Dalbergieae
the Horn of Africa - Arabian segregate of Ormocarpum Isolated genera: 4 genera ; 58 species, neotropical Dalbergieae
Pterocarpus
(Thulin & Lavin, 2001) and Maraniona from northern except Andira, which has one amphiatlantic species.
Cascaronia Dalbergieae
Geoffroea Dalbergieae

Fissicalyx Dalbergieae
Fiebrigiella Ormocarpin ae
Chapmannia Stylosanthinae
Stylosanthes Stylosanthinae
Ara chis Stylosanthinae

Isolated position

Grazielodendron Lima (1983b)

Dalbergia Dalbergieae
Machaerium Dalbergieae
Aeschynomene subgen . Ochopodium Aeschynomeninae

) Subtribe Aeschynomeninae sensu Rudd

Aeschynomene subgen. Aeschynomene Aeschynomeninae
Cyclocarpa Aeschynomeninae
Soemmeringia Aeschynomeninae
Smithia Aeschynomeninae
Kotschya Aeschynomeninae
Humularia Aeschynomeninae
Bryaspis Aeschynomeninae
Geissaspis Aeschynomeninae
Dalbergia
clade Transatlantic clade

Pictetia Ormocarpinae
Diphysa Robinie ae
Zygocarpum Th ulin & Lavin (2001)
Ormocarpum Ormocarpinae
Ormocarpopsis Ormocarpinae
Peltiera Labat & Ou Puy (1997)

Isolated position

Weberbauerella Poiretiinae

FIG. 40 Diagram of relationships and Informal groups in tribe Dalbergieae sens. lat. after Lavin et al. (2001a) and placement in Polhill & Raven (1981:
2
33-242; 283-288; 292-296; 347-356) where genera formerly placed in tribe Aeschynomeneae are here referred to by subtribe

308 LEGUMESOFTHEWORLD TRIBE DALBERGIEAE 309

Votoireo mocrocarpa Drawing by P. Halliday Vataireopsis araroba Drawing by c. A. Sobel Andira fraxinifolia Photograph by G. P. Lewis Andira inermis Photograph by R.8.G., Kew

VataireaAubl. 1775 Andira Lam . 1783

Hymenolobium o/agoonum var. a/agoanum Photograph by G. P. Lewis
8 spp"; Neotropics (ce ntre of Jiversity in Airia zonia; 7 srp. confinccl to nonhcrn s L11111bricidia Veil . (1829)
29 spp.; Neo{ ro pics (most cliveise in wet forest Amazonia [c. 12 .spp .] and in th e
America, 1 sp~ in C Am erica, from south e1n Mexico to Pana nm)
Vtllairea is the v e rnacular name fo l' Fre n ch G uiana Brazilian Atlanti c forest [8 spp.J); c. S spp. in seasonally cl1y S America; 3 spp. in C
Tall e mergent trees; the majo rity of spec ies inh abit [fOpical lo wl a nd 1; 1in forest , America, Ca 1ibbe an anJ Mexi co; 11. i 11e1·mis (\'V,\Vrig l1t) DC. widespread in th e
bo th ig~1p6 and vatzea (inundated forests), 1 sp (V. macroccupa Ducke) occurs in Neat 1o pics and in \~C A fl'ica
seasonally dry fo1 est, w oodland (cerrado) and scrub forest (caatinga) Deri ved from lhe name for a bar in che Tupi Amerindian language; the fruits of
Refcrc nce(s), Lima (1982b) 1he majorit y o f Andira species ~ll'e dispersed by fruit ba1s
Polhill Cl981cL 233 -242) placecl Valairea in t1ibe Dolbergicac closest to VaWireopsis, T1ees and s hrubs; tropical rain fo 1est (most spp.), seasonnll y d1y forest, woodl and
(C Brazilian ce1rado), dune nnd sc11.1h forest (coastal 131azilian rcstinga); ofi en o n
Lima (1980, 1990) noted that Vatairea, Vataireop sis, /Jndim ~111d lfy111 e11olobi111n
seem more closely re lated to members of th e t1ihe Sophoreae tlwn to rite remaining st1'eam-banks an<l in swampy areas
members of the tribe Dalbcrgieae . Tn molecul<u· ;rn ;_1lyses these ge n c 1~1 :w.. : un1esolved Rcfcrcn ce(s), Pennington 0995 ; 2003)
within rhe Vatairco id cl ad e (Ireland el al , 2000), which also conwins clements of the Molec ulaf d .: 1W and the unusual rool nodule morpho logy indicate th::1t Anclira is
Sopho reae sens lat. (sec page 227), and is well removed fro m the Dalbcrgio id dadc sister to Hymenolobium (Penningto n et al., 2001 ; \'V'ojdechowski et al., 2004)
( Pennington el al. , 2001 ; Wojciechowski et al., 2004) These two gene1a ~11e not dosely related to rnhcr genera currently placed in the
The majrnity of the species (know n variou sly as fa veira a1rnirgosa, :111gel1111 amargos~. Dalbergicac, ancl rheir wider relationships are clllren tly u ncle ar, <tlthough rhe
am;:irgo, bitter angelim 01· d~mto) produ ce good quality wocxl used fo 1 co nstruction, matK data of \'V'ojciechowski et ul, (2004) shows modera te suppmt fo r tb em being
!'urnitu1e, G1binet w o rk , flooring, joinery ancl veneers; also used fo1• mccltcine sister to a combined Amorphcae a nd Dalbergioicl clacle
Used fo1· timb er· (e .g., A ine1'mis suhsp . i1ie1 ·111is [cabba ge tree , pa1tridge w ood o r
Vataireopsis D uckc 1932 bh1ck plum], fo r construction, fu1-11i1ure, cabinet work, milway sleepers ::inc.I fence
~ spp.; NE to Am <1zonian S America (Surinan\ Fre nch Guiana, and I3 1:1zil) posts), fish poi sons, ornam entals, shade plants for co ffee pl<lllt<llions ancl m edicine
Ft o m -opsis (Gk .: resembhmce), referring co i1s likeness to the ge nus \'a wiit!n (anti-h elminthic drugs nrc obtained fro m the bark [known as w orm ba1 kl and
Small to emergent trees; ff'Opical low land Amazonian and coa stal Atlant ic r:1in forest seeds, but these are poisono us in high doses); also used as fungicides (Goa o r
Rcfc rcnce(s), Limo (1980): Lima in Aynrn i<I in Beny el a/. (1999, 425 -·126) Bahia powde r)
Po r rekuionsbips see notes uncler Valairea
Usecl medicinally in pso riasis treatment; in 1876 chrysarobine wris extr<.1ctecl frrnll
I~ t 11r1ro/xl (Agu lur) l)llckc nnd In 1?16 n dcri,·:nlw, dlthr.inol, " "'' Oe st
syntht:SIS<:d; other s pec! • • loo used !'or trc;11mcm r I isl11m1niusis untl J ·nnnlill>
Hymenolobium llenth. 1860
c .. l7 spp.; con cent1:ued in I31azil, rhe Guianas, .ancl Venezu ela; bul ! sp . exrends
int o Pei u, 1 sp. into Ecuado r, and 1 sp. occurs only in c A111eri G1 (1' ;1nama, Cosrn
Rica, and Nicar~1gu n)
from hy111e11 (Gk.: 111e 111b1~rne) and lohion (Gk .. : pod), referring to th e thin papery,
m embrnnaceous fruit whi ch is a one-seecl ecl s am ~1ra
T"-es; very rnU. usua ll y cmergcn1 in troplcnl h11111ld low L1ncl r:iin rorcst
Refort:n<'e(s), M:itttis ( 1979); Limo Cl 982:t): Z!1n1<'dl ( 1!)93}, Neill ul al ( 19'.)9)
10
Pol hill (l98 1<J, 233 - 2·12) pl:I eel //y mcmcilolJ/11111 ro irlhc D.1lll<!<gicJ • ck'"-'SI <l
1l11rllrt1 1 I.Ima 1980. 19\XJ) hl1thligh1s th:u A!/tlim, f(1mrenol11lm1111, l~rt11irtvt, " "
\l((f(l/n.-0psls occm lllOI • do.w ly n<hlt<."<l to m<:111lx:r or 1rll>e Soph r<'.IC th:io t<l
genera of tribe OalbCrJ:ll<.."!lct see notes under Andira
All species procJ,1ce h:utlw{)ocls (e.g., H . ex cels11111 Ducke [pa1a-an gclill1. :111gelitll
ammelo , angelim rosa o r ~ingelim cla rn atal, is used for heavy :incl light
co nstruction , furnitw e, fl ooring, panelling, jo inc1y .and ven eers) Andira legalis Photograph by G. P. Lewis

310 LEGUMES OF THE WORLD TRIBE DALBERGIEAE 311

 .
515 : 516

Adesmia graci/is (515 a-g), A. guttu/ifera (516 a-d) Drawing by J. E. Lacour Adesmia sp. Photograph by P. J. M. Moos Zornio - various spec ies Drawing by J. M. Fothergill Zornio sp. Photograph by G. P. Lewis

Adesmia oc. 1825 Zornia J.F.Gmel 1792

c. 240 spp.; S Amcrie<t, from N Anclcs to Tien~1 clel Fuego (P~ru, Bolivi:l, S 13razil
c. 75 spp.; pantropical; centrccl in S America with c , 35 spp ., brn also c . 14 spp. in
and Ul'L 1gu~1y); centred in Ande< 111 Chil e and Arge11t i11~1 USA, Mexico and C America; c 30 s pp . in the Olcl Woolcl (c. 13 spp . in Africa, l
f"mm n- (Gk.: \Vitl10ut) ~ incl desme (Gk. : bund le), in refcicncc to til l' f1 ee sUtllll'.llS sp. end emic in Maclaga.sc:11, c. 8 spp ~ in J\si<l Jnd c . 8 .spp . c nclemic in Aust1':i li a)
Annu<t l ;rncl pe r'ennird herbs <mcl shrubs; subrropica l to ternper~lte scrniarkl :lllcl Named arte1 the Gcrm;Jn pha1111:1cisl and bot:mist Joh~1nnes Zol'n (J739-1799);
~ 1rid rnontane grassland ~ind shru bJand (c hap:u ial) author of Jco11espla11/t1n1m 111erticinalium
Refercncc(s)o Bu1•kart 0967); Ulibarii (1986, 1999); Miotto Sfoggia &. Lc i1ao Filho Usua ll y erect 0 1• p1os1rclte he1hs, sorne1imes s hrubs; seasonally clry tropica l to
(1993); Davyt & Izagu irre (1996); Ulibarri & llurkaot (2000); Ilianrn (7.002) ,-varm tempcnuc woodland, bushland, wooded grassland, g1-rissbnd and
Po1\1ill (198lg: 355-356) placed Adesmia in the monogeneric tribt: ,\dc.:smieac, .shrubland, often in 1ocky are~ts <:tnd weedy in cultiv<Hed lands
Lavin el ol. (200la), IJased on DNA sequen ce cl:1ca, resolved r ldesmio in the Refe1ence(s), Mohlenb 1ock 0961, 1%2a); Verdcou1 t (1974, 2000' 159-165);
Acl esmic1 clade with tile fo ll ow ing genera: !lmicia, C/J{fe/ocr1~i1x, Nis.~·o/f({, Poirl'lia, Reynolcl s & Hollancl 0989); Vanni 0995); AyrnaJCI in Ile11y el al (1999, ii31 - ii33)
and Zcu-11ia (Fig '10); / lc/esmia aclclitionally cont:1ins two distinct 1110110phyletic ltudd (1981) plcicecl Xornia in t1il1e Aeschynomene~1e, s ubcribe Poircliinae . Ii::1v i11
el al, (200 Ja), based on D NA seqt1ence cb t~1 , resolved 7on1io i n the Ad csm ia
subgrou ps (Lovin et nl., 2001'1)
The dead, spiny src rn.s of some species arc piled and stacked co rn:1kc clade, most close l ~1 1ebted to PoireJia :.ind Amicia ( fig . IJO); chi s ielationship is
impe netrable fen ces; used as grou nd cover, erosio n control, l n11n~111 food (e.g., :ilso discussed by Ohashi (1999)
roots of A . lotoides HookJ.), 0 1namcntals and medicine Used 8s fo1agc , a covei crop (g1een manu re), for medicine and soa p; some
species m ay be toxic

Amicia Kun1h 182ti

c. 7 spp.; clisiunct in Mexico (1 sp~) and And ean S America (T'.Cl1:1do1, l'c 1u ,
I3olivia , and Argentina)
Na med afrer tlil.' Iw lian :-.srronomcr Giovanni TialCisla A Lllici (1 786- 18().) ), Dircctoi'
of the Florence observ;l!ory, horanist and opt i ci~1 n w ho made impo11 :111 1
discoveries in th e construclion of mi1'ro1·.s fat telescopes ancl otl1e1 opric:d
instruments
Perenn ial be1b.s or sp indly Lo multiple-stemmed s hrubs; ·wcccly :1 long l'Oacls. :incl
in seaso nzilly clry l1opic;il rnontane grassland to 3000 m
llefercncc(s)o llu.-kal'I Cl939a, 170-175); l(uckl (1981)
Rudd ( 19al) plac~-<l ll1111clo in 1rlbc Aescl1ynomcnene, subiribe 1'rnretll11:1c LJivln
rl nl. (200ta). h:>$C(I on D A sequence dam. rcsolvc. Am; la in th · Ad .,0111i:I , L;clc.
masc closclr related to Poiretia and Zon1ia (fig. 40); this rel;Hion ship i:i abo
cliscussecl by Oha sh i (1999)
Occ1sionally usecl as ca ttle fo(klcr in Argen tina; /1 zygomel"is DC. is an ;ttti<icti''C
orn:1111e ntal
Poiretia vent. 1807
11 spp~; Ncotropics, cent1·cd in E Brazil to Pciragua y ;rnd N Argentimt; l sp., P.
p1mclata Desv , ex tending into no1the1n S Amc1ica, C Amelica, Ciribbe:io and
1'vlexico
N~1111ed after the P1ench bot;inist and clergym:m , Jc~m Louis Marie Poiret ( 1755 -
183fi), who comp leted L<m1ar·ck's R11cyc/opc!die 111et/Jodic111e
Twining, scandent 01 e1ect, p e re nnial herbs 01• shru bs; seriso nally dry tropicri l lo
subt1"0pi ca l rive line forest, woodland (ceri ~1do), gra ss land ;111.cl shru bland
J(cfe1ence(s} [Jmka1t 0939a, 1987); Rudd Cl972b); Mlillc1 (1986); Vanni 0999)
Pbced by Rudd (1981) in t1'ibe Aeschy nomene:ac , s ubtribe Poin:tiin.1e. La vi n el al
(200la), based o n DNA seque nce dma, re:->okcd Poiretia in the Adesm ia clade, in
a subcbde together with Zo1•11fa :ind Amicia ( fig 40)
One species is s~1id to be toxic to ca tr\ e ancl may b e a sou fte o! rotenones

Amicia zygomeris Photograph by R.B.G .. Kew Poiretia punctota Photograph by G. P. Lewis

312 LEGUMES OF THE WORLD TR I BE DALBERGI EAE 313

 Nissa/la fruticosa Photograph by G. P. Lewis Rledeliel/a magalhaesii Drawing by P. Halliday Disco/ob/um psoraleifolium Photograph by R. M. Harley

Nissolia Jacq. n6o RiedelieUa Harms 1903

13 spp.; Neotropics; 1 sp.• N.frnlicosa Jacq., is widespread f1om Mexico to Itaobimia Rizzini (1977)
Argentina along the Andes and in Paraguay, the remaining spp have 1estricted 3 spp.; endem ic to E and SE Brazil and Paraguay
Named after Ludwig Riedel (1790-1861), German plant collector and traveller; his
ranges in Mexico, but also extend into SW USA
Named after the French botanist and physician, William Nissote (1647- 1735) collection f10111 S Brazil was the basis of the generic desciipcion
Perennial twining herbs and shrubs; seasonally dry uopical to warm temperate Lanky, scandent shrubs; seasonally dry tropical forest , wood land (ce rrado) and
forest, shrubland and grassland, often in mesic s ites along forest margins, on lhorn shrubland (caatinga)
stream banks, hillsides and in ravines Reference(s), Lima & Fonseca Va z (1984)
Reference(s): Rudd (1956, 1975, 1991) Placed by Polhill Cl98Jd, 233 - 242) in tribe Dalbergieae; but moved to the
Rudd (1981) placed Nissolia in tribe Aeschynomeneae, subtribe Ormoca rpinac. Sophoreae by Polhill (1994). Lav in el al. (2001a), based on DNA sequence data,
Lavin et al. (2001a) resolved Nissolia in the Adesrnia clade, whe1e it is either sl:<l<Jf resolved Riedelie/la in the Pterocarpus clade as sister to Disco/obium with which it
to or derived within Chaetocalyx (Fig. 40); Pennington el al. (2004, 526-527) shares a disc-like fruit (Fig. 40)
show Chaetocalyx to be paraphyletic with respect to Nissolia Used locally as firewood and as ornamenrals
Used as a fish poison and as an antidote co snakebite

Discolobium Benth. 1837

Chaetocalyx De. 1325 8 spp.; NE Argentina, Brazil (Mato Grosso to Bahia) and Paraguay
13 spp .. ; Neotropics 1 centred in S America; so uth to N Argentina, S Brazil and From disco- (Gk,, disc) and /obion (Gk: lobe, frui t), referring to the disc-shaped
Uruguay; north through C America and the Antilles to S Mexico fruit
From chaite (Gk ' long flowing hair), referring to the calyces of most species Spindly shrubs or perennial herbs; seasonally dry tropical to subtropical wooded
which are covered in long glandular-based trichomes grassland or grassland, often hydrophytes in permanently inundated and
Twining herbs; seasonally dry tropical to warm temperate forest , and a few seasonally Oooded areas, in swamps and along rivers
species in Andean or Amazonian fo rest; orten in mesic habitats, C1long stream Reference(s): Burkart (1939a, 1952), Va nni 0999)
banks, forest margins and on hillsides Rudd (1981) placed Discolobium in tribe Aeschynomeneae, subtribe
Reference(s): Rudd (1958, 1972a); Zarucchi (1993); Neill el al (1999); Vanni ( J9'J')) Discolobiinae Lavin et al, (2001a) resolved Discolobium in the Pterocarpus clade,
Rudd (198 1) rln cd Cbt101oca(1•x in lribc Aeschynornc nc:w, sub1nl>c . as sister to Riedeliella (Fig. 40) with which it shares the disc-like fruit
Orm0<.~1ipinnc. l;l\'m el al (200 1n), hO\\'<!V<.'T, rcsolvL'<l lhe ~cnus '" 1hc Adesnua Used as forage and medicine
clade, whGrc i1 1s pnmph lc1ic wi1h respect 1u fl'ix!Olfa lFi!I- 40; P<'l1n111gton et al.,
2004, 526-527)

Chaetocalyx /ongiloba Photograph by G. P. Lewis

o·tscolobium psaraleifo/lum Drawing by P. Halliday

314 LEGUMESOFTHEWORLD TRIBE DALBERGIEAE 315

 ....
5 ~
~-

Platymiscium pubescens su bsp. pubescens Photograph by G. P. Lewis

Brya ebenus Photograph by G. P. Lewis Platymiscium pubescens subsp. pubescens Photograph by G. P. Lewis

Cranocarpus Benth 1859

Platymiscium Voge l 1837
19 spp~; Neotropics from Mexico and C America to tropical S America; 4 spp.
3 spp; e nde mic to NE Brazil variab le and w idespread, and 15 spp. occupying more restricted distriburions
From cranio11 (Gk,, skull) and cwpos (Gk' fruit) , 1eferring to the he lmet-shaped (with two centres of diversity, one in Mexico 16 spp-1 and one in Brazil [5 spp.]) .
fruit 3 e nde mic spp. in C Ame rica; 7 spp. in Amazonian Guianas, Brazil, Bolivia, Pern ,
Sle nder e rect subshmbs; seasonally dry tropical humid forest (2 spp.); the other Ecuado1, Colombia and Venezuela and 3 spp in E Biazil lo Bolivia
species in dry forest on sand (restinga) From platy- (Gk , compressed, nat) and 111/scos (GL stipe), referring to the flat
Reference(s} Halley 0978); Fernandes & 13ezet'ra 0979); Ohashi et al. 0981);
frui t stipe
Ohashi 0999) Tall uees; tropical 1ain forest to seasonally dry forest, woodland (ce rrado),
See notes under B1ya wooded grassland and thorn scni b (caalinga); 10 spp. in min forest and 9 sp p, in
d1y forest
lleference(s} Enrech & Agostini (1987} Kli tgaard (1995, 1999, in press)
Brya P.Browne 1756 Placed by Polhill (1981& 233-242) in tribe Dalbergieae related lo Pteroca1p11s
4 s pp; Caribbean (endemic to the Greater Antilles) L:ivin el al. (2001a) resolved Pla1ymiscl11111 in a polytomy in th e Pterocarp us clade
Named after J.T. de 13ry (1564 - 1617) born in Liillich (Liege, Belgium), co ppe1 (Pig 40)
engrave r and natura l historian Used as ornamentals; mosl species (va 1iously Gilled trebol, macawood,
Shrubs or small spreading trees; seasona ll y dry cropical forest and sh11.1bland macacauba, grnnadillo, coyote, rarara colomdo) are highly va lued regionally for
Reference(s), Ohashi er al 098 1); Lewis (1988) th eir hardwood ti mber, used in house construction and for fine furn iture, cabinet
Ohushi et al. ( 19KJ) pl:iced B t}W in •rlbc l'.>t.-snKKliea , sublribe Bryinae. Jn the making, decorative veneers, joinery and musical instniments
molecular phylog<:nie:1 of D yl · t:I t1/ ( 1995) an I B:uk-y et t1/ (19'J7), it was
su8JlCSted tha t Br;vt should he n:mov~'tl f1 111 tribe Dc.<moclie~c:. Tiiis was
Mrongly corrob i;u~'<.l by the Sludy by uwm or nl (.200tu), in wind> B1ya is
r~.,;oJved In the P1croc11rpus clode, slslcr to <:m11cx:mv11s (Fig . .fO) with whic h it
hn '' capitate i;l:mdulnr trichomes that llf microscopically glochidia te, and
penpomte pollen
Bov1 <>IX'.1111s OC- (cocuswood, J1muie:m c r \l;'cst Indian ebony) •~one of 1hc 11>0.<1
'"-'lu<.~I •:~xxl~ in Cuba: th" t1lmos1 hl-t~k hL1u1 1VrK1<l 1$ 11 _.,d for nrnking t\lOl' .ind
nm~ l."lll 1nsrrumcllts (c.g_, woodwinds ~ n J b•Qplpc ), nnd for lnl:i)' work on I
~~1b111 'L mokmg, 11lkln~ u •·cry ~mOOlh hli:h poli~h. II 15 s:ild to be th~ wood fron1
which the truncheons of London policemen were once m<.1de. Rich in res ins, the
wood has been used loca lly as torches. Allergic contact derma ti tis has been
reco1ded from the wood

Brya ebenus Drawing by A. /. Beaumont Piatymiscium /asiocarpum Drawing by s. Ross-Craig

TRIBE DALBERGIEAE 317

316 LEGUMESOFTHEWORLD
 Platypodium e/egans Photograph by G. P. Lewis Maraniorra /ovinii Drawing by R. Wise
Moraniono /ovinii Photograph by C. E. Hughes

Platypodium Voge l 1837 Maraniona C.E.Hughes, G .1'.Lewis, Daza & Reyne! 2004
1-2 spp; C ~incl S Am eri ca ( P~nama, Col o mbia, Venezuela, I31':1zil, 1'uli via t111d 1 sp ..; N Pel'u (Cajam:oin::a, Ama zo nas)
Pmagu"y) Namecl after th e dry inter-Andean Mnr:iilon Vc1lley (N Peru) where this ~ind a
Fm 111 platy- (Gk.: brot1cl, flat) anclpodfo11 (Gk.: fo o t), re ferrin g to th e ~: 1111a1oid number of o th er locnlised legume endernics haYc bee n discovered
Small tii:!e ; tro pi Gil seasonally dry t'orest ;rn<l thorn sc rub o n steep rocky slopes, in
fr tlit with a broad win g :tt th e base (the s ~ed c ha mber is at the ;1pcx)
Trees 20-30 m; seasonall y chy cropkal 01• humid gallcty or rive1in e fo1t~ sr , thi cket a na nu w eleva tionc1l zone between 1400 and 1600 m
and woodbnd Refe1en ce( s), Hughes el al. ( 2001)
ltcfe1·encc(s), Ayma1d in llen)' et al. (1999, 377 - 378) 1Harcmio11a /avin ii C.E.Hughes, G .P.Lcwis, 0~1z::1 & Hcy nel is considered most

Placed by Polhill (1 98 !d, 233 - 212) in lribe Dolbc rgieae; Lavin el al (20ll lco ) . closely related to Tip11a11a in a combined c hlo1oplast 11wtK-1r11K DNA and n o n-
b11se d o n DNA sequ e nce c lat~1, resolved Pfal)jJodi11111 in the l)lcroG llTJU S cladc rnolccular ~1nal ys i s (nlthougb with out b o otst1 ':1p supp o rt)
(rig, ,10)
Used fo r timbe r, e.g., P. eleg m1s Vogel (carcuc ia, coslilla, t1rbol soga , t; 111 ;1lua,
carwl c to) in house constru ccion, fur•niturc, tools ; 111d for decor'ation

Inocarpus J R.rorsL & G.Forst. 1775

2 -3 spp. ; SE Asi~1 (i\fal ~s i a
and Papl1<1sia)
F10111 i110 - (Gk .: Jibre, strength) and cmjJOS (Gk . : rruit), 1efe1'ring lO llH~ li;11·d
di·upaceous fruit
T1 ee.s trsually with buttresses; lowland r<lln fo res t
RelC rcnce(s} Ve1dcourt (1 979)
Pla ced h )' Polhill (198 1d' 233 - 242) in tribe D;ilbcrgieae, but rno\'ed 10 1hc
Sopho1eac by Polhill 0994). bvin et al (200 la ), based on DNA sequt:nce d:1t1 ,
i eso\\'cc\ I11oca1p11s in th e Pteroca1pus cbd e 1el;llcd to Etabo/lia ( fi g. 40). w ith
whi ch i[ s lla1·es radially s ymmetl°ic flowe1s
ln ocmpusfagifer (Parkinson) Posherg (P olyn es ian or't:1hitian chestnur, ; 1il ~1. Jal;r)
is cultivtHcd as a food CJOj1 fo 1 its seeds, which :ire roastecl or IJOilccl and c;1tcn in
~falcs ia ; the woocl is use cl fo r mouldings :incl inter-Lo r finishing ; th e ffces :dso ha\•C
m ecl icin;.il uses

lnocarpus edulis Photog raph by G. o. Carr Maraniona /ovinii Photograph by C. E. Hughes

318 LEGUMES OF THE WORLD TRIBE DALBERGIEAE 319


Tipuana tlpu Photograph by G. P. Lewis
Ramorinaa girolae Drawing by P. Halliday
320 LEGUMES OF THE WORLD
Ramorlnoa g/rolae Photograph by R. H. Fortunato
Tipuana (Benth.) Ile nth, 1860
Macbaerium sect. Tipuana Ilenth. (1853)
1 sp.; S Bolivia, NW Argentina and SE Bra zil
_'ttlllcll after the "lipunna va lley in l'•r:tn. (llr:11i1) where the best gold was found
f.n,'1.'I or spreodlng I :<!•1 characteri5tf .:omponcnt o f seasonally dry subtropical
(gollcry to monronc) fo<C'St and open ~ hrnhln nd
Reference(s): Burkart 0 952), Rudd (1974)
Po lhill (198 1d: 233-2 2) pl:icccl Tl[111<11w In tribe Dal bergieae, most closely
rela ted to l'tcrocmfi11 • Lavin N al (200 1:1). based o n DNA sequ ence d ata,
resolved 71p"""" in the PtcrOCll'pll< elude (Fig. 40)
Tipuww 1/p11 (Benth.) Kuntze (tipu, rQS(wOod, racehorse tree, pride of Bolivia,
palo morrcro) is widely used as an <1rnnmcn1nl and st ree t tree in sL1blmpical to
warm temperare areas; also in reforestation , as wi ndbreaks and for erosion
contro l; for fodder, timber (furniture and cabinet making), firewood and cha1coal
Ramorinoa speg 1924
1 sp; endemic to \Y/ Argentina (La llioja, Sanjuan, and San Lu is p1ovinces)
Named for the Italian lectu re r J. Ramori no, who we nt to Argencina in 1866 and
taugh1 botany at the University of Buenos Aires
Xelcphy1I ~httth or tn:c; uhtrop· .11 lowl•ml or lowcr mc1n1:me 1lry fo1 " t or
~hnthlanu In rocky or sa111ly ""'· (~an<l~ of R. glro/11q ·pt.~ :on: <':lllcd 'fhicales')
llcfet<:n..-.:(s): pc11nz!nl ( 192 ); llurk:m 0952): C 111cz-SOS,q 19'> , 19')'))
l'law<l I l'olhlll 0981d: 233-242) in 1rilx: D•llx:rgico " t~avin w al (lOOlal, based
on DNA seq uence data, resolved Ramorinoa in the Pterocarpus clade (Pig. 40)
The bea utiful and durable wood (chica) is used for musica l instnime nts erncl
furniturei seeds are eaten and also g1 ound up and used as a co ffee Sl1bs[itute
Paramachaerlum ormosioides Drawing by P. Halliday
Centrolobium ochroxylum Photograph by G. P. Lewis
Centrolobium Mart, ex Benth. 1837
7 spp.; S America G spp in Ecuador, Colombia , Ve nezue la, Guyana and N Brazil,
extending in to Panama; 4 spp. in SE Brazil a nd E Bol ivia)
Fao m centron (G k.: sp ur, pric kle) and lobion. (Gk .: lobe, fruit), refe rring ro the
spiny fruit
Trees; tropical rain forest and seasoT1a lly dry fores t at low e levations, thorn scrub
and woodland (caatinga), dune fo rest an d shrubland (restinga)
Reference(s): Rudd (1954); Lima (1985b); Pi rie (2000)
Placed by Polhill (1981d: 233-242) in tribe Dalbergieae; Lavin et al. (2001a),
based on DNA seq uence data, resolved Centrolobium in the Pte rocarpus dade
sister to Geoffroea
Valious s pecies (po1cupine w ood , ca na1y wood, zebra wood, ama lillo guayaquil,
arariba) used for timbe1 in house construction, Oooring, ship components,
decorative veneers, fine furn iture and cabinet work; also p lanted as hedge1ows, <ts
ornamentals and shade trees in p lantations; known as porcupine lrees in some
areas because of their large sa maroid fruirs with prickly seed chambers
Paramachaerium Ducke 1925
5 spp.j Amazon ian S Amcric:. (Peru, Um.ii, Guianas, exte nding to Panama)
From para (Gk.: near. ~;mil:)r) and Mc1chm11"i1.mi, referring lo its resemblance to
lhe genus Machaerium
Trees; tropical periodically inundated or non-inundated rain forest and seasonally
dry woodland 1 o fte n along rive rs
Reference(s): Rudd (1981)
Placed by Polhill (198ld: 233 - 242) in tribe Dalbergieae; Lavin el al. (200la),
based on DNA sequence data, resolved Pc1ramacberium in the Pterocarpus clade
(Pig. 40), w hile Macbaerit1111 is resolved in the Dalbergia clade
Etaballia Benth. 1840
1 sp.; no rthe rn S America (Ve nezuela, Guyana and Brazil)
Derived from etabally, the veinacu la1 na me fo r the species in Guyana
Trees; tro p ica l lowland ra in fo rest, o[ten a long riverbanks
Reference(s): Rudd (1970); Funch & Santos 0998); Cuello in De ny el al. (1999:
325)
Placed by Po lhill (1981d: 233-242) in tribe Dalbe rgi eae, but moved to the
Sopilorc:t · hy Polhill (1 994). Lavin el al. (200la), based o n DNA seque nce data,
resol\'cd Eltl/Jttlllrl in the Pterocarpus clade re lated to Inocaipus with which it
shares radially symmetric flow ers (Fig 40)
Etaball/a dubia Photograph by L. Y. Th. Westra
TRIBE DALBERGIEAE 321
 Pterocarpus ango/ensis Drawing by H. Wood
Pterocarpus rohril Photograph by G. P. Lewis
Geoffroea decorticans by M. F.
Cascaranio astraga/ina Photograph by G. P. Lewis Photograph Gordner

Pterocarpus ;acq. 1763 Cascaronia G1iseb, 1879

c. 35-40 spp.; pantropical, w ith the greatest diversity in Africa; c. 20 spp. 1 sp~ ; endemic to S Boliv ia and N\'Y Argentina
endem ic to Africa, 1 sp. (P. santahnoides L'Her. ex DC.) also native ro the from cascara (Spa nish: bark), refel'fi ng to the g rey, ci;:1cked ba1k of the trunk
l'k"011op cs, , nu I sp. ( I' indfc" ' Willcl.) n:uive 10 ss;i; 5 >pp. tl-strirti:cl to S Asia, Spindly shrnbs or rnultiple-stemmed trees; seaso nally dry subtropical fores t, frnest
lndo-Chimo and M~cla~ar; c. .11 spp. cndl!ni1 to the Neotropl (,\I tl< ico and c margins and alo ng rivers
Amctkn wllh 5 SJIJI· and S Ametic:i with 6 spp.) Referencc(s} llurkart (1952); G6111ez-Sosa (1999)
f'rom Plll>'011 k.: wong) and ctrrpos (Gk.• rnolt), rcrcrrlng 10 1hc wln!ft-'<I fruit of Placed by Polhill (198ld, 233-242) in tribe Dalbe rgieae, most closely re lated to
m:1ny spcties
Pterocatpus; Lavin et al. (200la), based on DNA sequence data, reso lved
Tall trees, sonic S(l(!ciC.-S :JJC buttressed rzi in forest emergents; tropic<i l lo w!Jncl Cascaronia in th e Pteroca rpu s clade (Fig_ 40). The l e~ves are characteristicall y
evergree n r:tin ror l (less than 10 spp.) to seasonal! )' ci ty forest, woocllancl, gland-do t1cd
thi cker and ' t><Xlc d (!no '!l:1 ncl
The wood is use d for fuel and tu rn ery
Rcfo~nt·~'(.~): llcppttr In ll111c hinson O:tl>.lcl (1958• 517-5110; Polhill in M1lne-
R~ lh<':lcl & Polhill (1971 : 81 - 91); llo)o ( l?n): 'lho1h:uhri CJ?Sn: Gii <'I ttl. ( 1987):
Enr<'Ch & Gii 1993): Enrceh of al. in llcrry ~I al. 0??9: 38 1- 384>; Wck in Geoffroea Jacq . !760
lln1111111i11 "' al ( ·ubmlned)
Gottrliea Gi ll ies ex Hook. & Arn . 0833)
l11c gcnu m:ty be sulxl" ·idcd into 1wo groups b3S<.-d on wing~'<!""""'' unw inged
2 spp.; S Amciica and Caribbean (1 sp . in Ch ile, A rgemina, Bolivia, P;:11aguay
f1uits. Polhill (198 1d: ljj-242) pl:!~'() PUJroc:t11p11.< In 1ribc l),1lbcrgiC:1 • most
and Urugu ay; the ot her sp _ also from Argentina ;rnd Paraguay to Bolivia, Peru,
closely rcl:ucd 10 71p11m10 on<! l'latypodi11111. This rcl.1iorL,h1r 15 o 110n11cd by the
Ecuador, N Colombia, Vene?.ue la, E Br;:izil and th e Antilles)
<~>mbincd nnalyS<.-s <>f I.: \'in Ct al. C-OOln), which rcsol\' !() P1111'fiaup11 on the
a med after the French chemist and botanist Cla ude Joseph Geoffroy, who
Ptcroc:1 rpus d:idc (Fig. 0), togc1hcr whit lhc majoriry of Jhc gcncm in
Dolbel'glcnc S()1w1 Polhill Cl981d) wo rked with j;>cquin from 1750 to 1752
T1ees or shrubs; chJracte 1iscic component o f dry open chaco vegetation in
f'tcrocmpu lttdlcus (1urrJ, Solomon's p:idnuk, l':tl""' New uill " rosewood.
southern S A111e1ica, and in disjunct areas of tl'opic::d seasonally dty fo rest in
:unh<rynn ) may be 1hc 1110>1 t'<."Onomicnlly impo11nn1 legume timber sped~"· Tue
nort he rn S Ame1 ica
Wood o r SC\'Crn l "J>l'CIC., ind uding f~ t111guf1!1tStS DC. v\rrit•:tn IC..~rk , bloodwood,
lleference(s} llul'ka1t (1952); Ire land & Pennington 0999)
kflrru, munlnlJ:o) mid I' so v111.,'fi Taub. (p:>dauk) is higbl)• , 11 fucd nod u.~..J ror nnc
Placed by Polhill (198ld• 233 -242) in tribe Da lbe ,.g ieae, most closely rela te d to
r11rnl111ro, l~1hlnc1 mnkinJ!. p:onelling. j<iincry, muslc:il in.'ln11nctll ', lmp lcuwnrs :rnJ
Pleroca1pus; Lavin et a l. (200 la), based o n DNA sequence d ata , J'e solve cl
curio.~. l11c re in ouppllc. d •e> (e.g., f~ t111cto1~11.< Wd, .). Mod in ~1 • Amaxun is
Geoffroea in the Pte rocarpu s clade o ften as s iste r to Casca ron;a (Fig. 40)
known "" 's.1n11rc de clmgo' ( • dmgun. hlood); 1hc ri: In ts nlso :1 wid ·ly u5t.'CI folk
Use d Fo r human Food ( the ed ible fnJ its are made into jam or used to flavour
mcdlnne ( the source of Kino guon.l; otl1cr US<.'5 .ore In tc1'<'i;<:l:rlio11 1111J "'111
wine); the timber is used fo r ca1pcn11y and furnin1re making; the bark and leaves
lmprO\'cmcn1, us s hade u·c.:.<, omamcnlal. , t-U.:111C1k :ond lxiske1. (from 11><: Inner
lurk) have medi c..:inal pro perties

I
I I Pterocorpus rohrll, sens. lat. Photograph by c. E. Hughes
Geo{froea splnosa Photog raph by G. P. Lewis

322 LEGUMES OF THE WORLD

TRIBE DALBERGIEAE 323
Fissicolyx fendleri Drawing by c. A. So bel Flebrigiel/o grocllis Photograph by G. P. Lewis
Fissicalyx Benth. 1860
1 sp.; S America (Venezuela, Guyana, Brazil and Paraguay) to Panama
FromjtSSus CL deeply deft), refening to the spathe-like calyx, split to half or
more its length on one side only
Trees; tropical lowland rain forest (on terra firme) to lower-level montane forest
(eve rgreen ro semi-deciduous), fo rest margins and wooded grassland
Re ference(s): Lewis & Owen (1989); Aymard in Berry el al. Cl 999: 326)
Placed by Polhill Cl98ld: 233 - 242) in tribe Da lbergl ·'"· most closely re lated to
Pterocaipm; Lavin et al. (2001a), based on DNA SL'<jUcnce da ta, resolved
Fissicalyx in the Pterocarpus clade as sister to Fiebrlgielfa to which it be<us no
obvious resemblance (Fig. 40)
Fiebrigiella Harms 1908
1 sp.; Andean S America (confined to S Ecuador, Peru and Bolivia)
Named after K Fiebrig (1879 - 1951); his collections from S Bolivia formed the
basis of Harms' description
Perennial herbs; tropical monta ne grassland, shrubland and thicket co 3500 m,
often in rocky areas
l!eference(s): Burkart & Vilchez 0971); Zarucch i 0993); Neill et al (1999)
Placed by Rue.Id I' I) in tribe Aesd1yn meneae, subtribe Ormocarpinae; Lavin
al. (2001a), however, resolved f'fcbti11fc/ln in the Pteroc-Jrpus clade as sister to
Fissicalyx (Fig. 40)
Chapmannia Torr. & A.Gray 1838
Arthrocatpum Balf.f. (1882); Pachecoa Stand!. & Steyenn. 0943)
7 spp.; New World (I sp. in Mexico, Guatemala and Venezuela and J sp. in
Florida), Old World (I sp. in Somalia and 4 spp. in Socotra)
Named after the American botanist Alvan Wentwonh Chapman (1809-1 899),
who cont ributed much to the knowledge of botany in Florida
Perennial hcrm, small shrubs o r trees; seasona ll y dry tropical to subtropical
woodland. bushland, grassland and open scmbland, often in rocky or sandy areas
and on roac.lslclcs
Reference(s): Gi llett (1966); Gu nn el al. (1980); Norman & Gunn (1985); Thulin
0993, 1999)
Rudd 0981) placed Chapmannia, Pach•'CtHt, and Arthroca1pwn in tribe
Aeschynomeneae, s ubtribe Stylosanthinae; l.:lvin et al. (2001a), based on DNA
sequence data, rcsolv~'(] Chapmannia sens. lat. in the Pterocaopus clacle, and
Chapma11nla is odclitlona lly sister to Arachis (Lavin el al., 2000; 2001a) (Fig. 40)
Used as an ornamenta l and for fodder in Somalia
Chopmonnio prismotico (1-6), C. reghidensis (7, 8) Drawing by P. Halliday
324 LEGUMES OF THE WORLD
Stylosonthes - various species
el
Stylosanthes copitoto Photograph by G. P. Lewis
Sty/osonthes guionensis Photograph by G. P. Lewis
Drawing by J. M. Fothergill
Stylosanthes sw. 1788
c. 25 spp., but generally poorly delimited and estimates of up to 50 species exist
in the literature; native lo New and Old World, ma inly neotropical (c 23 spp .;
centred in S America [13 spp.] with 4 spp. in N & C Ame rica and 6 spp
widespread in the Neotropics) and 2 spp. widespread in Africa, Madagascar, Jndia
and Sri Lanka (S fmlicosa (Hetz.) Alston [wild Jucernel and S erecta P.Beauv).
Several New Woo Id spp have been cultivated and are naturalised in Africa , Asia
and Austoalia (e g., S guianensis (Aubl,) Sw . sens. lat ., S. bu.mi/is Kunth and S.
vlscosa Sw.)
From stylo- (Gk.: style, pillar) and anlbos (Gk.: flower), oe ferring to the long,
slender style, often persistent in f11.1it
Subshrubs or perennial herbs; seasonally dry tropical co warm temperate
wood land, wooded grassland, thicket, shrubland and grassland, on sandy or rocky
soils, along streams and sornelimes weedy in old cultivated lands and on roadsides
Reference(s): Mohlenbrock 0957, 1960, 1963); Verdcourt 0974, 2000: 165-169);
Reynolds (1990); Sanjappa 0992); Du Puy & Labat in Du Puy el al. (2002:
662-665)
Mohlenbrock (1957) placed Sty/osanthes in tribe Hedysareae subtribe
Stylosa nthinae, related to Zamia, Chapmann ia and Arachis; Rudd (1981) placed
it in tribe Aeschynomeneae, subtribe Stylosanthinae related to Attbrocmpu.m and
Pachecoa (both synonyms of Cbapmann ia in this treatment), Cbapmtmnia and
Arachis; Lav in et al, (2001a), based on DNA sequences, placed Stylosauthes in the
Pterocarpus clade, sister to Aracbis, and nested with C1Japmannia (Fig. 40)
Major livestock fodder plants in warm temperate and tropical areas of the world
(e.g., S. gufanensis or Bra zilian lu cerne); several species are plan ted as soil
stabi lisers and improve1s, and as ground cover, in e.g., coffee plantations; also
used for medidne
TRIBE DALBERGIEAE 325
 Grazielodendron rio-docense Photograph by G. P. Lewis Oa/bergia ecastaphyllum (A-E), D. monetaria (H) Drawing by B. Angell Dalbergia cearensis Photograph by G. P. Lewis

Aracbis L. 1753 -, .. .
-
---- Dalbergia LJ. 1782

. Q ti . ·. . . . ~ · '"~
~ ., - -.
.
69 ~pp.; S AmeriC'".1,from SC & E Brazil (most species) west ro Bolivia ~nd south to ·' ' . I " 1!111eri11111011 P.Browne 0756); Ecastap/Jy/111111 P.l3rmvne (1756); Hecastopbyl/111n
PnrnAu:1y, Ur\lguray ;1.11<! N & E Argentina; but Cll ltiv::tted rxmtropic;Jll)' ancl in Kunth (1824); TriojJlo/emea Ma1t, ex Benth (1837)
. . I
wurrn tempemtc :in:n c, 250 spp.; pantmpical, cent red in the Old World with 60 - 70 species in Africa (1
Prom a- (Gk.: w ithout) a nd rbachis (Gk,: central ~1.xis), 1eferri ng to the absence of

.. \ . . .k . ~ ,.. "· ... ··

sp., D ecaswphyllwu (L.) Taub , reaches rndia), 43 spp. in Madagascar, of whic h
erect bra nches

.· 42 aie ende mi c; about 80 species in Asia w i1h 33 species in India (19 endemic.:),

· ~'
Herbs; ~e;isonal\y dry t ropical to subtropical , ciry ~ind well-c.lrainecl wooclecl 44 in Jndo-China, ancl eight in New Gu inea. Two of the Asia11 species (D .
gr<1 land and grassland
Rcforence(s), I3mko 1t (1?39:1); lludcl 198]'>: Kmpovl~kas,. Grt!l(Ory 099· I
-i' ' : • . ; .•·. ! ,f· ·. t' ca11den a le11sis (Dennst.) Prain and D denser Benth.) 1each Australia; c . 60-70
spp . occ ur in the neotropics with 45 -50 spp. in S America (centred in Amazonia)

~
Rudd 0981) placed Ami;h•.< ln iribe AC.<;ehynomcncac. sl 1l)lrlh • Stylosanihlnac. I ..•
and c. 15 - 20 spp in tropical Mexico to C Ame rica and the Caribbean
l.:1vln e/ al. (200 1"), b">ed on ONA S<lqucncc dnrn, rcsol•«:d An11; /Jf.< In 111<· Named after the Swedish bmthers Nils E. Oolberg 0736-1819) and Carl Gustav
l'oeroc:irpu.• cln<ic ns sislcr 10 Stylou1111/JN (rig. O)
Am~/Jl< IJJ'fJ08a(!{1 L (peonut, groumlnuO l " mnfc•r huin.111 food und :«>llr<" of
. .I . . . •J. 4
'
Oa lberg (// 1753-1775), both botanists and friends of Linnaeus
\ '
~'·~
, . Shru bs, trees <incl climbing li;rna.'), some species va1y in habit from scanclent

· . t ~ / ., ,\
.
''~'!!Cl•blc oil (,..C :ond only I<> soyb.!on in lmpounnre •mong legum ·s), uncl "ni111:ll ·,. . sh ru bs in dry hab itats to rnbust lia nas in humid a!'eas; tropic..:al l'ai n forest to
fodder (m;iinly lt';l.vcs): •lso use I "". Mlil rcnlll-;cr .ind ground CQ\'Cr: oth~"< seasonally dry tropical to subt1opical humid :.m d clry forest, woodla nd, bushland,
•p<..oci"" "'" '.1..""d slmll.1rly, but on n inore ICJC.1tl or rcglon:ol b.1S1sC•.g•. A.plmol thicket ;111cl wooded grassland
Kmpov. & W. . n:g.). l'unhcr USC!! nre ns medicine nnd focls; •he oil i> Uo: f).1.<C Reference(s), Prain 0901), Hepre1 in Hutchi ns on & Dalziel (1953, 513 - 516);
of numerous lnduStrlnl products (e.g.. polishes, paino.s, Jubrk~'""'· inscctlridl'S, Polhill in Mil ne-lledhcacl & Po lhill 0971' 95 - 112); Tliothat hli 0987); 1lo.'5e1 &
so:ips :ind t'O.~n1<;1ks) , ~· ,/ •
',., ~
!
' )
ltabevo hitra 0996); Carvalho 0997); Aymarcl in lleny el al. (1999, 295 - 301);
' . . ~· Bo.'5el" & R"bevohitra in Ou Puy e/ al (2002, 321 - 361); Niyonclham (2002); Lock

Grazielodendron H.CLima 1983 t' ' {·~' .\' .. ~. ·.., ..

in Brummitt el al (submitted)
Thothathd (1987) .subdivided Da/hergia fro m th e Inclian subcontinent into four

' · ~· ,
~ ~·t; '(. .~ ~
1 sp.; NE Brazi l (endemic to the state of Espirito Santo; rnre) sections based on anclroccium and rruit typ e; Ca rvalho 0997) treated the Bn1zi lian
Named after the Bra:t.ilian botanist Dr. Giazicl::t Maciel Barroso (1912-2003) . " Ii . .d . species in ftve sections based on inf101 esce nce and fruit types . The combined
Tall trees; tl'Opic~I Atlantic rain forest ana lyses of Lavin el al (200la) place Dalbergia in the Dalbergia clacle siste1 to
Reference(s), Lima Cl983b) Nlacbaerium and Aescby110111e11e subgen . Ochupodi11111 (Pig. 40)
Lwin et al. (200la) resolved Grazielodendron in the Pterocarplis cladc (fig . c}O) 1\fany fndian, Africm, ancl B1azilian species of" Dalbergia, including D. nigra
(Ye ll ) AJ\cmao ex I3enth -, D sissoo Roxb-, D. /atifolic1 Roxb. and D. 111e/ano;..:y/on

. ,: ~ · c$·
Guill. & Perr. a re known to produce high qua lit y timbei ' [h;1t is used for
construction, fine fl11niture, cabinet wo1k, marquetiy and inl;:1y, ri<inos ;md other
musical instruments , tool and cutle1y handle.s, turneiy, carYi ng and vario us
.specirtlty items. Common names include rosewoocl, blackwoocJ, tu li pwoocl,
kingwoo<l, ebonywood (not true ebony), cocobo lu, nambar, palis:.rndro and sisam.
The tenn rosewood most p robably refers to the pleasa nt smell of tbe wood. Sinc.:e

.· ~
the woo<l burns well, i[ is also often the preferred wood for cooking and
charcoal-making (e.g ., D _sissoo in Asi:.1) Sevc1'i.1 I srecies are cultivated in the Old
\Vorld tropics as multipurpose trees for timber, fibre, fodder, fuelwoocl ;:111d
medicine, or as ornamentals; contact c.leimatitis is reported from the wood

Grazie/odendron rio-docense Photograph by G. P. Lewis

Datbergia lemurica Photograph by D. Du Puy

326 LEGUMESOFTHEWORLD
TRIBE DALBERGIEAE 327

Machaerium hirtum Photograph by G. P. lewis
Machaerium Pers 1807
Drepa11oca1pus G,Mey. (1818)
c . 130 spp.j Neotropics, cen[led in Am:izonian S America , w ith c. 1S spp. from
Mexico (c. 2 endemic) and C America (2 spp., M isadelph11m CE.Mey .) A1nshoff
and 1l1. IHnalwn (LJ.) Ducke 1•eacb the Caiib bcan); 111 lunal111n is also ;1mphi-
atlantic, extending to th e west coast of Africa
Prom macbaira (Gk.: dagger, lnrgc knife), referring to the shape of th<..: fruit
and/or to the prickly stipulcs of some species
Shrubs, t 1ees and clirnbing Hanas; tro11i cal inundated nnd non-inund:ttt:d 1ain
forest, seasonally dry forest, d une foJ'esr, low woodland, thicket, tho1 n scrub,
sh n.1bland Hnd remnant shade trees in cocoa plantations
lteference(s): Hoehne (1911b); Rudd 0977, 1987a, 1987b, 1987c, and in Berry el
al., 1999, 339-352); Lozano & Klilgaard (submiued)
Hudd 0987b) di vides Macbaerium inlo four sections; Polhill (198Jd, 233 - 242)
placccl kfo cbaerium in Drilbergicae , mos t closely related ro Dalhergia, a
1cbtion.ship coi roboratecl by Doyle et al. (1997) and the combined :111alyscs of
Lavi n el al. (200 la); 1l1acbaeri.um is additionally re.solved close.".it to A esc/JJ'JIOllWfU!
•ect. Ochopodiu m (fig. 10) . into th ree subge nera : subgen~ Aeschynomene, subgen. Rueppellia, and subgen.
~lachaerium speci es are widely used in forest mancigemcnt as _.,;hadc plants and
for the re coveiy of clegrnc.led zireas; th e wood of 1rn1ny species (caviun ;1 , pau feno,
santos rosewood, moraclo), is h.:1rd ;rnd used for fine fu1 niture, c:1binetr~ 1 ,
decorative veneers, panelling, piano.s, flwes, axe handles and fence roses (the
wood is resisrnnt to decay); the reddish S<tp of some species has been used by
nati ve t1ibe~ to treat sn~1kebitc and the leaves of othe t species arc a cocai ne
substitute; con tact de1 matitis is reprntetl from lhe wood
Machaerium hirtum Photograph by G. P. lewis
328 LEGUMESOFTHEWORLD
Aeschynomene mediacris Drawing by V. C. Gordon
Aeschynomene indica Photograph
Aeschynamene martii Photograph by G. P. lewis
Aeschynomene L. 1753
Herminiera Guill. & Perr. (1832); R11eppellia A.Rich. (1847); Balisaea Taub. (1895);
Bakerophylon Q. Leonard) Hu!ch. 0964)
c 175-180 spp •• sometimes wi1h estimates as high as 250 spp~; c. 84 spp occur in
the Neotwpics flncl subtropics, centred in Mexico ro C Am elirn ; in lhe OlcJ Wodd
rhe distribufion is principally African-Madagascan (c . 90 - 95 spp), wifh 1 sp., A .
aspera L., endemic co Asia and Australia; c. 3 -4 .spp. are widely introd uced in the
Pah\eotropics from rhe New World
from aischynomeno- (Latinisc<l to aesch ynome no) an anc ien1 name for a sensitive
plant (derived from Gk.: modest, sensitive) , in refe1ence to the sensitivity of the
leaves to tou ch and temperatu 1'e
Sh1't1bs 01 herbs, in very few cases climbei s; seasonJlly chy tropical woodl;tnd,
wooded grassland, bushbnd and grassbn<l (sometimes mont;rne), often in 1ocky
or sand y areas; many species a re hydrophytes occurring in marshes, at the edges
of water holes, in swampy areas and flood p101ins
Re ference(s), Rudd 0955, in Berry et al, 1999, 241-245); Lock (1989, 101-110);
Reynolds (1990); Pe1nandes (1996); Verdcourt (1971a, 2000' 58- 117)
Verdcourt (2000) in his tre~1 1111e nt for Flora Za mbesiac~ subd ivided Aescbyno mene
Ochopodium . Rudel 0955), Fernandes (1996, treating !he 131azil ian species) and
b1vin et al (2001a) studying th e evolutiom1 ry iclationships of the genus, subdivide
Aeschynomene into Lwo subgene ra based on stipule placeme nt: subgen.
Aeschyn01nene w it h 1nedifixed stipules, and subgenus Ocbopodium with basi tlxcd
stipules. Rucki (1981) placed Aeschynomene in tribe Aeschynomc neae, subrribe
Aeschynorneninae rogethe1 w ith Soemmeringia, Kolscbyu, Smilbia, Geissaspis,
B1yaspis, Hu11mlaria, :mcl Cyclocc11pa, and Lavin et al (2001a) resolved subtribe
Aeschynomen inae as a well-supported monophylefic group (Fig, 40) , Ex!rnpolaf ing
from the small sampling, however, it is suggesled that 111embe1s of subgen
Ochopodium sensu Rudd, are mo1e closely 1elatecl to Machaerium (tribe
Dalbe1gicae sensu Polhill, 1994) [Jian to species in subgen. Aescbynomene; ii is for
a fut u1e revision of Aeschynomene to 1esolve these rebtionships; some species of
t1eschynomene produce stem nod ules
Used ecologically in management o f inundated a1eas, as o rnamenrals, fo1• fodder
and g reen ,-nanure; A . aspera (sola, s hola, a mbatch wood, joinc vecch) and
A. alt1pbro.xylo11 Guill. & Perr.) Taub. ;:ire major sources of pi rh , a white spongy
wc')()d used for pi1per, fibre, he lmets (solar 1opi), art work, handicrafts and art ificial
flowers (e.g., sola rosario tlowe rs); the wood is al.so used for floats, rafts and
ca noes
ex U.S.O.A. collection
TRIBE DALBERGIEAE 329
Cyclocarpa stellarls Drawing by unknown artist Soemmeringia semperf/orens Photograph by G. P. Lewis Kotschya strlgosa Photograph by J. Anton-Smith Kotschya a{rlcana var. bequaertll Photograph by D. Du Puy

Cycl.ocarpa Afzel. ex Urb. 1884 Kotschya Endl. 1839

1 sp.; widely distributed in the Paleotropics, through Zambezian and Sudanian 31 spp.; Africa (Zarnbezian-Sudanian, Afromontane and Somalia-Masai regions)
Africa, Indo-China 1 Malesia to N Australia and Madagascar (1 endemic sp.)
From cyclo- (Gk: circle) and catpos (Gk.: fruit), referring to the circular fruit Named after the Austrian botanist Theodor Kotschy (1813-1866)
Erect annual herb; seasonally dry tropical grassland, in damp places, rock fissures, Herbs, shrubs, rarely small trees; seasonally dry tropical riverine forest, woodland,
seepage zones and pond edges, often on cultivated land thicket, bushland and grassland (so metimes montane), especially on margins of
Re ference(s): Rudd (1981); Verdcourt (1971a, 1974, 2000: 157-159) rivers, lakes and forest, in swamps, floodplains and rocky areas
Traditionally cited as Cyclocmpa Afzel. ex Baker (1871) but this is considered as a Reference(s): Verdcourt Cl970a, 197la, 1974, 2000: 117-138), Du Puy & Labat in
provisional description (i.e., made in observation). Placed by Rudd (1981) in tribe Du Puy et al. (2002: 652-655)
Aeschynomeneae, subtribe Aeschynomeninae; Lavin et al (2001a) resolved Related to Smitbia with which it shares helicoid cymose inflorescences, Rudd
Cyclocatpa in the Dalbergia clade together with the remaining members of (1981) placed Kotscbya in tribe Aeschynomeneae, subtribe Aeschynomeninae;
Aeschynomeninae sensu Rudd (1981) (Fig. 40) Lavin et al (2001a), based on DNA sequence data, resolved Kotscbya in the
Dalbergia clade (Fig. 40)
Soemmeringia Mart_ 1828 Used for riniber, poles, firewood, forage, brooms and medicine

1 sp.; S America (Amazonian and NE Brazil, Venezuela and Bolivia)

Named after the German naturalist Samuel Thomas von Soemmering (1755-1830)
Prostrate shrub or perennial herb; seasonally dry tropical forest, woodland
(cerrado), wooded grassland and scrub, often along rivers, in floodplains and
disturbed areas
Reference(s): Ducke 0958); Lewis (1987); Lewis & Owen (1989); Cuello in Berry
et al. 0999: 390)
Rudd (1981) placed Soemmeringia in tribe Aeschynomeneae, subtribe
Aeschynomcnfnae; Lavin et al. (2001a), based on DNA sequenc resoh·cd
Soemmerl119ia in the Dalbergia clade in a subclade with most mcmhci:< of Rudd's
subtribe Aeschynomeninae (Fig. 40)

SmitbiaAiton 1789
Damapana Adans. (1763)
c. 20 spp.; Old World, mainly native to the Indian subcontinent (c. 11 spp.
endemic); a further 6 spp. widespread in Asia, cwo of which, S sensitiva Aiton
and S, conferta Sm. extend into N Australia; 2 spp. endemic in Africa and of the
two Malagasy species, one (S elliotii Baker f) also occurs in Africa and the other
(S. sensitiva) in SE Asia
'~med :ifter Sir James Sf11ith (1729 - 1828), round;,r o 1he Linne:i n Society in l.<>ndon
Herbs 10 subshn1bs: ~sonolly dry 1ropic:il gnc.<Sl:rnd, marshy ureas nnd sue 1111; ides
tlei'ftlnce(s): Vcrdcoun 097Ckt. 197 10, 1974, 1979, 2000: 138- 1 O); ll11dd (1!)111);
Thuan et al. (1987); Du Puy & Labat in Du Puy et al. (2002: 655 -657)
Genus in need of revision; related to Ko1schya with which it shares helicoid
cymose inflorescences. Rudd (1981) placed Smitbia in tribe Aeschynomeneae,
subtribe Aeschynomeninae. Lavin et al. (2001a), based on DNA sequence data,
resolved Smithia in the Dalbergia clade with Aeschynomene sect. Aeschynomette
(Fig. 40); both these taxa and Geissaspis share medifixed stipules
Used for forage, human food, medicine and as soap substitutes
Smithia elliotil Drawing by E. M. Stones l<otschya perrierl Photograph by D. Du Puy

330 LEGUMES OF THE WORLD TRIBE DALBERGIEAE 331

 Humulorio roseo var. reptons Drawing by M. Grierson Bryospls humu/arloides Drawing by P. Halliday Diphyso sp. Photograph by C. £. Hughes
Plctetia marglnato Photograph by G. P. Lewis

Humularia P.A.Duvign. 1954 Pictetia Dc. 1825

c. 3S s1 1>.; • Afrle:1 (Zalllhe>.lan and Afromom..'lnc regions)
Nam •d for •lie l'C.'«.'lTiblnncc of the inOorcsccncc. to those of the hop plant,
N1111111/11s /11p11ltl$ L.
Wcxxly·b,1setl he rbs or occ sion ully sma ll shru~ S(.11,;onnlly dry 1r •pl .,1
woodland, wooded gms.land, scrub nnd i:ra. ·land (so111<.'llmc,. 111Qnh11k!), often
3.lo.'iOCi:ucd with &n::tm b:inks, swamp mmgl~ lloodpl:iin • nd s:mdy ;11c:>s
llefcrcncc(s): Duvignc:iud (195-1); Cillcll "'(I/ In Mlln"· R~'<lhc:Jd l'u lhtll 0971:
29-435): Gl<-'<lhlll (1968); V'<tkoun (1971, 20001 140-1 ~)
Rudd 0981) placed l/1111111/arfri In 1ribc Ae><:hynomcncuc, sub1ribc
Ac.~ci1ynonh.'11 nae; l.1\•in ct ti/. (200.la). h!IS<.-c;I on D A sequence d:ita, resolved
//1111111/r1rl11 In 1he Dalbcrgl:i cl:u.le, sfsicr to /Jry•~<f>ls ond c;e1wapis (Fig. 40)
·d for lll~'<llcine :ind :ts a toni
Bryaspis P.A.Duvign. 1954
2 spp.; W Africa
From brya- (Gk.: be full of) and aspis (Gk : shield), referring to numerous bractS
that shield the flowers
Erect to spindly herbs; seasonally dry tropical grassland and open areas, often in
wet, swampy, rocky and sandy areas
Reference(s): Duvigneaud (1954); Gledhill (1968); Hepper in Hutchinson & Dalziel
Cl958: 582); Rudd (1981)
!lucid {1\)81) plac•"<l IJ1)"t1S/Jls in lrihc ACS<"hrnumcnc:1~. ~ublrlbc
Aeschynomcninnc: Llwin ~J 11/. (200 1:1), haS<.>d on L)NA ,.,...,.1,.,, 11 _.c d:t ta, resolved
l113wspl.< in 1he Dalhcrgin dude, si$1cr 1 N1111111/m'/11 <111d (,"a1$S11Sp s (fig. 40)
Sometin1 · t1 w•'<:cl In <:uhi .11c..l land; u~d u: med cin •
Geissaspis Wight & Arn 1834
c. 2 spp.; Asia, including the Indian subcontinent, extending to China, Jndo-Chin•
and Malesia
From l,'<!fsso. G k.: hem, borde r) and aspis (Gk.: shield), refening to th e large
bracts 1hn1 borde r and shield the flowers
Pl'Olit .uc to~ indent, woody-bo:;cd hcrhs; :;casonally dry tropical forest (in open
:11.-ns), gr.~ · land and In wc1 habiw~. weedy along edges of rice fields
Refcn;:nce(s): Ouvignenud (1954): Gledhill (1968)
Rudd (198 1) plaa.>d C11issas/Jis in 11ibc Ae."'h)'110111<:nc:1<:, suhtrilx:
Acschyno111.,111m1c; l~win q/ ti/. (200 1:1), b:lscd un ONi\ .cqu 'IK'C ,~ 1 1a, resolved
1he !;.'Cnus In rh ' P:ilbergfo dude, sister 10 /11111111/nrla :ind IJT)~l.<J!ls (Fig. 40)
Belai1ia A.Rich. (1845)
8 spp.; Caribbean in the Greater Antilles (Cuba, Hispaniola, Puerto Rico and the
Virgin Islands)
Named after the Swiss zoologist and palaeontologist F.j . Pictet (1809 -1872), a
friend of A.P de Candolle
Trees and shrubs; aggressive colonisers of seasonally dry tropical forest, in thicket
and thorn scrub, often on steep slopes, near streams and in disturbed areas
Reference(s): Beyra Matos & Lavin (1999)
In the molecular phylogenies of Beyra Matos & Lavin Cl999) and Lavin et al
(2001a), Pictetia is resolved in the Dalbergia clade, in a subclade together with the
Asian-African genus Onnocatpum, the Madagascan genus Ormocatpopsis, and the
C American genus D1physa (Fig 40)
Used as ornamentals 1 timber for fence posts and planted spiny hedgerows around
cultivated fields
Di.physaJacq. 1760
S/einbachiella Harms (1928)
15 spp.; Mexico and C America, with J sp. extending into S America, and another
into S USA (Arizona)
From di- (Gk .: two) and physa (Gk.: bladde r), referring 10 1he inflated exocarp
which forms two fused bladders
Shrubs and trees; seasonal tropical dry to humid forest and thorn scrub forest,
often a long rivers
Reference(s): Polhill & Sousa (1981); Sousa & Antonio (2001)
Polhill & Sousa (1981) placed Diphysa in tribe Robinieae; the phylogenetic studies
o f Lavin (1987, 1995), Beyra Matos & Lavin (1999) and Lavin et al. (2001a),
however, confirm the monophyly of Diphysa and resolve it in the Dalbergia clade
(Fig. 40)
Used as living fences and occasiona lly as ornamentals in Mexico; the wood is
strong and has been used as clubs in battle

Geissospls crlstota Drawing by P. Halliday

Oiphysa amerlcono Photograph by c. E. Hughes

332 LEGUMESOFTHEWORLD TRIBE DALBERGIEAE 333

Zygocarpum somalense Photograph by M. Thulin Ormocarpum sennoides Photograph by G. P. Lewis Peltiera nitido Drawing by f. Lemeux
ormocorpopsis parvifo/ia Photograph by D. Du Puy

Zygocarpum Thulin & Lavin 2001 Ormocarpopsis R.Vig. 19so

6 spp.; NE Africa (3 spp. in Somalia) ; 2 spp_ in Arabia (Yemen and Oman) and 1 6 spp.; endemic to Madagascar (throughout M.adagascar, exce~t in ~he e~sr~ r~
sp. in Socotra lowland rainforest); a numbe r of species, pa111cularly O~ aspe1'Cl H .~1~., exh1b1t
l'<Qm :Z)'gO- (Gk .: joined), aod cmpos Gk .. fruit). refctTing to the m;u~ lly two
complex variatio n pauerns; this group appears to be active ly spec1atmg
Prom -opsis (Gk.: appe arance), denoting a resemblance to Ormocmpum .
tLl11cles o f the legu me that ""' yokL"ll tO fOl'lll II ;cgrnentlld rl\111
Shrubs or small trees; seasonally dry tmpica l fo1est, woodland an d xerop hyt1c
Shru l>s Or small &Ices: S<".l>onall y dry 1ropic1 I wO<.xll:md, 1hickc1 o r bushla nd, often
o n ll1m~11c, l>u1 :11$() ,.....,..,.UL-d frool gr.mim or sn ndy ~u b>U111cs, 10 l OOO 111 shrubland, somelimes on rocky outcrops, sandstone or limestone
l<cfore ncL'(s): ·mulln & i.i•in (200 I) l(eferencc(s): Labat & Du Puy (1996); Du Puy & Labat in Du Puy et al, (2002:
Zygocarp11111 I a sesreg:llc genus or Ormoca1pum. Thu lin & Lavin (2001) , based 633- 639) I ·1
on ONA S<:qL1en<:es, rosol,•ed Zyy<x:wpum as monophyletic and sister to a clacie
Omwcmpopsis was placed by Rudd (1981) in tribe Aeschynomeneae, su >In J C
containing Ormocaipum, Pelliera and Ormocaipopsis (Fig_ 40) Onnoca rpimi e; Lavin et al. (2000; 2001a), basecl o n DNA seque nce analyses,
resolve Ormoctapopsis as sister to Ormocmpum in the Dalberg1a clade (Pig. 40);
see also note~ unde r Pictetia
Ormocarpum P.Beauv_ 1806 The hard wood is used in house and ca1t construction and for charcoal
D1phaca Lour. (1790)
c. 18 spp.; Africa (c. 15 spp., one widespread also in Asia; c. 3 spp. in Guineo-
Congolian, 4 spp. in Zambezian-Sudani an, c. 3 s pp. in Somalia-Masai and c. 5 spp.
Peltiera Labat & Du Puy 1997
in Swahelian regions) , Madagascar (2 endemic spp.) and Asia (1 sp. from the 2 spp.; endemic to C Madagasc.u (possibly extin~t? . . .
Indian subcontine nt to the Pacific islands (Fiji) and N Australia) Nnmed arter Maurice Peltier (1920- ?), in recogn1t1on of his outstandmg
From hormo- (Gk.: necklace, chain), and carpos (Gk .: fruit), referring to the conrributions to the knowledge of Legumi nosae in Madagascar
beaded appearance of the fn.1it Shrnbs· seasonally dry tropi ca l evergreen forest, forest margins and wood land
Shrubs and small Crees; seasonally dry tropica l forest (also mangrove or sw~llll(J
Refc rc~ce(s): Labat & Du Puy (1997); Du Puy & Labat in Du Puy et al (2002:
forest), woodland, wooded grassland, bushland and thicket 639 -642) . . -
Reference(s): Gillet t (1966); Verdcourt 0979, 2000: 51-58); Thulin (1990); Dupuy Placed by Lavin et a l. (2001a) as sister to Ormocaipopsis with w hich 1t sha res a
& Labar in Du Puy et al. (2002: 631 - 633) distinct ive resin pattern on the unde rsid e or the leatlets .
Ormoc1t1t111111 wn~ pl:i~·c:<J by Rudd (198 1) in 1rlbe Acsdl)'nonieneac, su blfl'lle Hecent field investiga tions have fa iled to locate material of tl1ese rare spec~ e~, and
the disjunct areas in w hich they occur conespond to only a very few surviving
Ormoc:lrpln;1e; J.uvi11 ,q / <1/. (2000; 200 In), b:L~-d o n DNA S<."<IUC n<.'C 'ln• lysc., ,
resolve Ormocmtmm :is sister to Ormocnrpop.<IS in 3 t ra ns111~1rul cl:ide (1111. 40); and distu rbed re mnan t frnest patches
see notes under Piclelia
Used for rimber, the whippy sterns are used for hut constru ction and frn.mes; as
medicine, sh£Jde trees for coffee and as a fish poison Weberbauerella u 1br. 1906
2 spp.; s America (endem ic to coastal Peru)
Named aFter the Peruvian botanist August Weberbauer (1871 - 1948) _ .
Perennial he rbs o r subshrubs; seasonally dry trop ical coasta l forest, 111 vegeratton
on sands and sandy hills (lomos)
Refe rence(s): Ferreyra (1951); Za rncchi 0993) . .. .
Placed by lludd (1981) in tribe Aeschynomeneae, subtribe Poiretunae; Lavin et al
(2001a), however, resolved \'Veberbauerella in the Dalberg1a clade (Ftg- 40)

Ormocarpum bernlerianum Photograph by D. Du Puy Weberbauerel/a brongniartioides Drawing by unknown artist

334 LEGUMES OF THE WORLD TRIBE DALBERGIEAE 335

TRIBE
Hypocalypteae
Tribe Hypocalypteae (Yakovlev) A.L.Schutte 1998
Tribe Liparieae (Benth.) Harv. subtribe Hypocalyptinae Yakovlev (1991)
Polhill Cl98lp: 398) placed Hypocalyptus in tribe
Liparieae, but analyses of numerous morphological,
cytological and chemical characters (summarised in
Schutte & Van Wyk, 1998b), show that Hypocalyptus
does not fit comfortably into any of the existing tribes
so a new monospecific tribe was described to
accommodate the genus. All subsequent studies based
on gene sequence data have supported its isolated
position and uncertain tribal affinities. Hypocalyptus
was misplaced in the genistoid tribes, differing from
them in the chromosome number, seed aril shape,
micropyle type and position, ephemeral antipodal
cells, presence of canavanine, absence of alkaloids
(both quinolizidine and piperidyl alkaloids) and
different flower pigments (absence of esters of
anthocyanins and presence of methylated
anthocyanins) [see Schutte & Van Wyk Cl998b) and
references cited therein].
FIG. 41 Diagram of relationships of tribe Hypocalypteae after Crisp eta/. (2000); Kajita eta/. (2001); Wojciechowski eta/. (2004)
336 LEGUMESOFTHEWORLD
byB.-E. vanwyk
Hypocalypteae now appear to be basally branc.htn
to the clad~ co~prising Indigofereae and the millettioi~
- phaseolo1d tnbes as well as the Hologalegina crfbes
Within this basally branching group , Hypoce1/yptu ·
seems most closely allied to the Mirbelieae a n~
Bossiaeeae (Crisp et al., 2000) and the Baphioid clade
of Sophoreae (Pennington et al., 2001; Kajita et al
2001). Wojciechowski et. al. (2004) find mod e rat~
support for Hypocalypteae as sister to the Nli rbeli~e
and Bossiaeeae. The crown clade age e>f
Hypocalypteae -Mirbelieae (representing the age of the
trans-continental split in this South Africa-Au ·tra lian
diversification) is estimated to be c. 55 Mya (Lavin et
al., in press), remarkably old in legume terms (Schrire
et al., pages 21-54, this volume) and unique in age for
southern Hemisphere trans-continental dlsjun rioos.
The Hypocalypteae includes one genus and tluee
species (Fig. 41).
Hypocolyptus sophoroides Photograph by B. -E. Van Wyk Hypocalyptus sophoroides Photograph by B. -E. Van Wyk
Hypocalyptus Thunb. IBOO
3 spp.; Africa (endemic to the Cape region of South Africa)
From hypo- (Gk: beneath) and calyptra (Gk veil), referring to the flower buds
that are hidden beneath leafy bracts (this no longer applies to the genus as
presently circumscribed)
Shrubs, subshrubs o r small trees; medite rranean shrubland (fynbos and forest
margins), often localised at high altitudes, in rocky and sandy areas and alo ng
Baphioid clade streams
Reference(s): Dahlgren (1972) ; Schutte & Van Wyk (1998b); Schutte in Goldblatt &
(see page 228) Manning (2000: 485)
An anomalous genus with no obvious affinities (see Schutte & Van Wyk, 1998b);
the similarities with the Podalyrieae (e.g., the intrusive calyx base) and other
genistoid legumes (e .g., the fused stamens) a1e superficia l o nly
Hypocalyptus HYPOCALYPTEAE
Mirbelieae (see page 339)
Bossiaeeae (see page 355)
lndigofereae (see page 361)
Miltettioids
Phaseoloids
Hologalegina
11YPoca/yptus oxalidifolius Photograph by 8.-E. Van Wyk
TRIBE HYPOCALYPTEAE 337
TRIBE
Mirbelie ae by M. D. Crisp, J. A. Chappill, R. De Kok and P. Jobson
·be Mirbe licae (Benth.) Polhill & Crisp 1982; Mirbelieae (Be nth.) Polhill 1981, no m. inval.
Tfl Tribe Pod:i lyrieae subtribe Mirbe liinae Be nth. (1837) , as 'Mirbelieae'
Thi treaunem r ecognises 25 gene ra and (6 6)-(688)
- (689) pe ies in the e nd mic Australasian ( ustralia
'fll mania and Papua ew uin a) trlbe Mirbelieae
(Fig. 2 . It comprises tw group , one w ith g ne ra
ha ing -nu l ate e mbryo-sacs and g iant a ntipo da l
'.Ind the other with 21 g ne ra having usually multipl ,
5-nuclea~ embryo sacs and no antipodals. Most p i
ill thi Lril e are ricoid brub with y LI w and .red
('egg and ba o n' flo wers. They are c nspic uo u ,
01 ti me domina nt , unde r t re y m e mbe r o f
50
sclero1 hyU nununitie (heathla nd a nd u caly pt-
dominated woodland and fore. t , m stly n po r o il
of the south-west, south and east coast of Australia.
Polhill's (1994) classification of Mirbelieae was
based on a cladistic analysis using morphology (Crisp
& Weston, 1987), which found the tribe to b e the
monophyl e tic sister group of Bossiaee ae . A
reworking of the morphology (Crisp & Weston, 1995)
with th e addition of embryological data from
Cameron & Prakash (1994), found two groups within
Mirbelieae, one being a large clade in which the
embryo-s a cs are re duced to five nuclei, with
ephemeral antipodals. The second group was
paraphyletic, containing the above group and genera
in which the embryo-sacs are normal Polygonum-
type, but with giant antipodals. This latter embryo-sac
type is shared with Bossiaeeae.
Phylogenetic analysis of DNA seque nces (Crisp et
GIANT ANTIPODALS GROUP
Bossiaeeae
(see page 355)
Gompholobium Sphaerolobium
II
II
Ii
I .I
II
FIG. 42 Diagram of relationships in tribe Mirbelieae after Crisp et al. (2000); Crisp & Cook (2003a)
LEFT Sphaerolobium sp. aff. macranthum Photograph by B. Fuhrer
al., 2000; Crisp & Cook, 2003a) has found the
Bossiaeeae to be monophyletic but part of a polytomy
with the major clades of Mirbeliea e . It is unclear
whether the giant antipodals group (including
Bossiaeeae) is the monophyletic sister group of the 5-
nucleate group, or paraphyletic and containing the
5-nucleate group (Crisp et al., 2000; Crisp & Cook,
2003a). However, the 5-nucleate group is supported by
both morphological and molecular data . Support for
the clade containing both Bossiaeeae and Mirbelieae is
w eak or non-existent, and it would be premature to
recircumscribe these tribes . Analyses of ITS , rbcL and
matK sequence d ata indicate that Mirbelieae and
Bossiaeeae are most closely related to the small South
African tribe Hypocalypteae, and then to the Old World
tropical tribes (including Indigofereae and the
millettioid and phaseoloid tribes) and Hologalegina,
la rgely comprising the temperate herbace ous
papilionoids (Crisp et al., 2000; Doyle et al., 2000;
Kajita et al., 2001; Wojciechowski et al., 2004).
The clade with 5-nucleate embryo-sacs comprises
two strongly supported sister taxa: Isotropis and the
Mirbelia group (Crisp & Cook, 2003a & b).
Circumscription of the c. 19 genera within the Mirbelia
group has never been stable and it has been proposed
to subsume all into an expanded concept of Pultenaea
(Fig. 42; Crisp & Cook, 2003b; Orthia et al., 2005, in
press a).
5-NUCLEATE GROUP
MIRBELIEAE
Daviesia lsotropis Pultenaea sens. lat.
Erichsenia
Viminaria Jacksonia, Leptosema,
Latrobea, Euchilopsis, Phyllota,
Otion, Aotus, Urodon,
Stonesiella, Almaleea, Eutaxia,
Dillwynia, Pultenaea, Mirbelia,
Chorizema, Oxylobium,
Podolobium, Callistachys,
Gastrolobium
I
TRIBE MIRBELIEAE 339
 Ooviesia grossa by M. D. Daviesia euphorbioides Photograph by G. P. Lewis
Gampholobium minus Photograph by M. D. Crisp Photograph Crisp

Gompholobium sm. 1798 Daviesia sm . 1798

135 spp.; Aus1ralia (throughout, but mainly SW and SE)
B1111011 iu R.B1'. ex W:f,Aiton (1811)
Named for Hugh Davies 0739-1821), Welsh bot;inist
44 spp.; S\V (trn1in rnea of clivetsity), C, E & N Australia; two spp. ex te nding to
Shrubs or 1arely trees; sclerophyll forest, woodland, ;md shrublancl including
New Gu inea and tlle Lesser Sunda Islands
From gompho - (Gk.: wedge-sha ped n;iil or bolt) and /obio11 (Gk.: pocl), 1den"ing sandy heathland
Reference(s): Elliot &Jones (1984 , 3: 200-212); Crisp (1995b); C1isp in Hmclen
to the innacecl pods
(2002: 522 - 528); Crisp & Cook (2003c)
Shrubs; seasom11ly d1y tropical ro warm te mpe rate sclerophyll forest, woodland
Monophyletic and e:otsily diagnosed by the rriangularpods, sre1ile bracts on the
and heathland, often in co:_1stal heJths, on mountain slopes and whlelands in
peduncle ~rnd scleromorphic, phyllod c- like leaves; closely l'clated to
sandy or gravelly soils
Sphaero/obium, Vimi1za1 ·ia and h"richsenfa within rhe giant antipodals group
Hcfe rence(s): Stanley & Hoss (1983: 248-250); Elliot &Jones (1986, 1: 376-383);
Some speci es of 'bitter peas' h<ive been used as a substitute for· hops, in mec.licine,
Wheeler in Ma1chant et al. 0987: 262-266); Jeanes in Walsh & Entwisle (1996:
to m::ike a bilte1· tea and as J dye; c1lso u.secl as orna111entals 1 in revegetation, ns
719-752); G rieve (1998: 376-381); Wiecek in Harden (2002: 518 - 52 1l
windb1eaks and as a wildlife refuge
A morphologically divel'se genus ret<iiniog plesiomorphic featl1res such as
compound leaves; diagnosed by a co111bina1ion of leaf structure, elonga 1e and
v;1 lvarc calyx lobes, globosc pods, often numerous ovules and long funidcs;
belongs to rhe giant antipod<1ls group but its precise rel~Hionships are unclea1;
under revision by). Chappi ll
Some species of 'wedge pe;.i' Jre cultivated HS ornamentals

Sphaerolobium Sm. 1so5

22 spp. ; S Australia (especially the SW)
From sphaero- (Gk.: sphere) and /obio11 (Gk.: pod); referring to the sphe1ical pods
Usua lly sm::dl, diffuse shrubs; forest, woodland and he~1thh1nd, often in damp 0 1
wet peaty habit::1ts
Rcfe rence(s): Jeanes in Walsh & Entwisle (1996: 752-754); Grieve (1998:
396 - 399); Wheeler in Marchant el al. (1987: 295 - 297); Butcher & Clrnppill
(2001); Wiecek & Murray in Harden (2002: 521)
A distinctive genus wi1h a usually ru h-like Jeancss h::1bit, o fren paired fl owe fs and
style mostly w ith eithe1 a longirudirnll wing or " ling of hairs around 1he stigina;
falls wirhin the giant antipodals group but its precise relationships <ire unclc;ir;
uncle1 revision by R. Butcher and]. Chappill

Sphaeralobium pulche/lum Photograph by R. Butcher Daviesia cordata Photograph by M. D. Crisp

TRIBE MIRBELIEAE 341

340 LEGUMES OF THE WORLD
 /ocksonia sp. aff. anguloto Photographer unknown jocksonio orgenteo Photograph by J. Chappill
Erichsenio uncinoto Drawing by M. Smith Viminorio junceo Photograph by H. Thompson

Erichsenia Hemsl. 1905 jacksonia R.Ilr. ex Sm. 1811

Piptomeris Turcz. (1853)
1 sp. ; SW Australia (wheatbelt)
74 spp.; SW, E and N Au stralia
Named fo1 F Erichsen, engineer on the \\'lestern Australian Goldfields \X'arer Na med for George Jackson (]780- 1811), gardene r and Scottish botanica l
Scheme illustr<Hor (later ed itor of BotCJnical Reposil01)1), who w<i s in charge of th e
Shrn bs; shrubland including beathLrnd on ltHeritic (yellow) soils
he 1bal'iurn of Ay lmer Bourke L<1mbe1t
Reference(s): Gl'ie ve 0998: 396) Sl1bshrubs, siltl 1bs 0 1• small trees; arid and .semi-add zones (few specie.<.; to tropical
Diffuse sh1ub w ith hooked phyJ/ocles, striped calyx and sma ll ornate pod; related meas) or sclerophyll forest, wood land and he~nhland , o fte n on rocky ridges or
to Daviesia and Viminaria w ithin the giant antipodals group
mountain slo pes
Reference(s): Crisp in Jessop (198 1: lli8- 149): Stanley & Ross (1 983: 251 ); Elliot &
Jo nes 0990, 5, ii63 - 473); Wh eele r in Wh eele r el al. 0992: 414 - 417): Chappill in
Viminaria sm. 1805 Grieve (1998: 381-395); Wiecek in Hard e n (2002: 535 - 537)
1 sp.; S\XI and SE Austr'Jlia Dingnosed by leafless ad ult branchlets, unifol io late seedling leaves, peltate hairs on
From vimiueus (L.: wand-like, having lo ng flexible shoots) stems and calyx, lo ng and more or less equal va lvate cal yx lo bes often recurve d
Ta ll shrubs or small trees; near-coastal, usually in sw~1111ps (these m ~1 y be sc;1sonril) aga insc the pedicel, and hai1 s inside the pod; fo nns a cbcle ·wi th Leptosenw, of
and margins o f .stre11ns unclea r affinity within the P11/tenaeo sens. lat. (5-nude:He embryo-sac) g1u up
Reference(s): Jeanes in Wa ls h & Entwisle 0996: 754-755): Wiecek in l-J;11den °'
Many species (kno wn ~s dogwood) have namenta\ value, p:uticularly].
(2002: 522) scoparia Sm. (winged broom pea) which is also used for foclde1; some species
Distinctive by ics season:illy wet habitat, willowy b1oorn -Jike growth h;lh it and used for traditio nal medicine
semi-succu lt:nt indchisc:ent fruit; !'elated 10 Daviesia ~incl E1"ichseHia " ' it!1in rhe
g iant antipoda ls group
Vi11tinaric1juncea (Schracl .) Hoffm~mn s. (swis h bush, go lden sp1ay, rn sh L1 100111) is
used <1s ;in orna mental , fot reveget;iting watercourses and fm• fibre

Isotropis Benth. 1837

10 spp. ; mos t parts of Au stra lia except tile SE
Fmrn iso- (G I<-• equal) and tropis (Gk : keel), the kee l is equa l in length to the
wings
Herbs and subshn.ibs; 1c111pe1arc heathland 10 tropical (euc~dypt) woodl;1nd , often
in drier communities (in cluding the central clesetts), usually on s~a1cl}' so il s
Reference(s): Maconochie in Jessop (1981 : 146-1 7); Ston ley & Ross (1983: 218):
Elliot &Jones 0990, 5: 453 - 455); Grieve (1998: 374 - 375): Portcners in Ha1<len
(2002: 528-529)
Diagnosed by rhe semi-he1baccous h~bit , unifoliolalc \e;1ves (when present),
conspicuou s veins on tli e rea 1· of the st;rnclarcl, obovoid-oblong pods a nd
reticulate seeds; phyloge netically isolated within th e 5-nucleate embry o-s:1c giour
The foliage of some species ( known as Poison) is toxic to domestic ]i\ estoc k
1

lsotropis cuneifolio Photograph by B. Fuhrer lacksonio scoporio Photograph by H. Thompson

TRIBE MIRBELIEAE 343

342 LEGUMES OF THE WORLD
 Latrobea diosmlfolia var. glabrescens Photograph by M. D. Crisp Phy/Iota luehmannii Drawing by M. Smith Phy/Iota phylicoides Drawing by L. Elkans

Leptosema Benth. 1837 Phyllota (DC.) Benth . 1837

13 spp.; W, C and N Austra lia Pulte11aea sect. Phy/Iota DC. (1825)
From lepta- (Gk.: slender) and serna (Gk. : a sig n 01 banner), referring to the narrow 11 S[1p. ; te mp era te SE and SW Australia
Fmmphy/1011 (Gk.o leaf) and 01os (G lc ear), refcl'ling to the leaf-like br'oc teoles of
standa1d petJl
Shrubs; sclerophyll commun ities (heath, mnllee, spini fex) to semi-al'id some .~pecies
Shrubs; open woodland and open fo res[, mallec wood land and heaths, from
<.:ommunities on sandy and skeletal soils
coastal areas to monwnc (bul not alpine) ;rnd semi-arid regions
Re ference(s)o Elliot &Jones 0993. fr 108-111); Ciisp 0999)
Rcference(s): Jancey (1966); Stones & Curtis 09780 li52); We ber 09860 671-672);
Low-growing Je~fless plants w ith conspicuous bird-pollinated flowc1s; differs from
Jeanes in Walsh & Entwisle (19960 764 -765); Grieve 09980 424-425); Wiecek in
ils sister taxon,jacksonfa, in rhe enlargement and modification of the nowers for
birc.1-pollination; unclear affinity within the Pu lteuaea sens /al , (5 -nuclc:lle Harden (20020 539- 540)
Diagnosed within the Pultenaea sens /al , (5-nucle<ite e mbryo -sac) gmup by a
embryo -sac) group
combination of lea f-like bracteoles, incu rved beaked keel, sta mens partly fused to
Some species are grown as ornmnentals for their showy flowers
base of petals, thickened bcaf<lecl style <rnd hai1s tending to hispid

Latrobea Me isn. 1848

6 spp.; for SW of Western Australia
Otion Crisp & P.H.Weston ined ,
Nair1cd in honour of Charles La Trobe (1801-1875), Lieutenant Gove11101• of c. 8 spp.; SW, C ancl N Austrnli o
From otion (Gk: small car), refe11ing to the presence ofb1acteoles (in conr1~1st ro
Victoria
Shrubs; temperate forest, wooJ!and and hem hlancl in higher rainfall regions on Ao/us, which h1cks them)
sandy soils or in sw~1rnpy areas Shrubs a11c1 st1bshrubs; forest , woocll;:ind ancl heathland
Reference(s} Wheeler in Marc hant et al. (19870 278 -279); Elliot & Jones 0993, 6o Refcrence(s)o Ci isr & Weston (19950 264 - 265)
Diagnosed by a combi nation of a shallow abaxial lobe on the keel, presence of
61 - 62); Grieve 0998: 425-426)
Dbgno.-cd hy unffonn cn ly~ lobe>. P<">kC.'<l btondard, ond ~ompn:..;; •<l lrl:rngulnr- b1actcoles and hairs inside the pod; simihll' to Aotus ttnd Urodon within the
P11//e11aea se ns. lat. (5-nucleate e mbryo-sac) group
~lgJlloid pods; bclon115 within the P11/tc1111M .w11s Im. -nu k"1tc t.•nh!)'O· cl
gro up

Euchilopsis r.Muell . 1882

1 sp.; SW Australia, more or less 1est1•icted ro the coastal plain ~md adj:Kcnt hills
De rived from 'Euchilus-like'; the type species of Euchilus is now plnced in
Pultenaea, so a new generic name was publi shed by Mueller to accommochlTC
E. lineaits (Benth .) l'.Mt1ell.
Sh rub; forest, woocl l;md :incl heithla nd 1 peaty or sanJy soils near sw<Hnps
870
Reference(s} Elliot &Jones 0986, 4o 2·13- 244); Wheeler in Morclrnnt el al. 09
256 - 257); Grieve Cl998o 415)
Dingnosed within the P11/1u11f/ct1 Sl!llS. Jar. (5-nuclf!111c "nihryo-sacl group h)'
sollt"ry axillary nowcrs on slender pcdl ds, and a cnlyx wilh two lo~e ruund
upper lobes and th1 ee minute lower teerh

Euchilopsis linearis Photograph by M. o. Crisp Otion microphyllum (unpublished name) Photograph by M. o. Crisp

344 LEGUMESOFTHEWORLD TRIBE MIRBELIEAE 345

Aotus genistoides Photograph by M. O. Crisp Urodon cop/totus Photograph by B. Fuhrer Almaleea subumbellata Photograph by M. o. Crisp Eutoxia myrtifo/ia Photograph by M. O. Crisp

Aotus Sm 1805 Almaleea Crisr & P.H.Weston 1991

15-18 srp.; tem r ernte S Australia, esrecially SW 5 srr.; SE Austtalia
Prom a-(Gk: without, lacking) ancl otos (Gk: ear); 1ereriing to the abs<.:nce of
N111ned in honour of Alma Lee (1912-1990), Au!'t1•a lian bown ist and legume
brncteolcs resea1cher
Shrul>s: Of>CJ\ woodlund '1nd open fort-st, ti.tile..: woodland, h.::nho :oncl """""P5; Sh1ubs; in freshwa ter bogs and wet hea ths
from CO:tsrnl nrc:ts 10 mon1n11c (hut not alpin :inti .-emf-arid n.>gjons Reference(s); Crisp & Weston (1991); Jeanes in Wa lsh & Enlwisle 0996: 793 -794);
ttefcrcncc'(s): Elliot < Jone• ( 1982.• 2: _ J0 - 212}. Stanley & Ross ( 191!~: 251 -255); Westo n & Crisr in Har·den (2002: 540-511)
Webe r' 0986: 658); Jeanes in W"lsh & Entwisle (1996: 763 - 764); Gric,•e ( 1998: Belongs in the Pullenaea sens lt1t (5 -nuclea te embryo-sac) group; simila1 co
415 - 417); Wiecek in Harden (2002: 537 - 538) Sto 11 esie/fa, Ditltllynia, EHtaxia rind P11/1enaea; diagnosed within tbc grotip by an
D iagnosed by us11 ~1lly whorl ed le1ves, bck of stipul cs and bracreoles (;_1ltl10ugh involucre of pe1sistent bracts often wirll enl <irged 3-lobed stip ubr bases
b 1~1cteol es arc present in some W/ Au snali;.in speci es), and da rk rc;1 1 o f the
S(andarcl; si111iln1· ro Olion and Urodo11 wit hin lhe Pul1e11nea sens. /al . (5- nuck:~rc
Eutaxia R.13r. ex w .T.Aiton 18 11
embryo-sac) g roup Sc/ero1bani11us R.nr. ex \YI T.Ailon (1811)
Used fo1 g1ound cover JO srr.; S Austra lia
From eu- (Gk .: good, or well -developed) and taxis (Gk: 01'Cler or arrangt:menr),
refe11ing to the 1egularly decusS<llC leaves of most speci es
Urodon Turcz. 1849 Sh1ubs; frn esr, woodland, mal\ee <ind heathlrind, often ne01r the coast
Refercnce(s); Elliot & ) on es (1986, 4: 262-264) ; Weber (1986: 667 - 668); Jeanes in
4 spr .; 2 undescribed; SW Austr"11ia Walsh & Entwisle 0996: 795 - 796); Gl'ievc (1998: 426 - 128); Wiecek in Harden
From 11ra- (Gk.: tail) and odon (Gk.: tornh), probably refen'ing to 1lic long
(2002; 5•12)
<1cu mim1te apices on the ca lyx lobes ('teeth')
Di~1gnosed by usu<11ly opposite, decussate le.aves wi th 3 nc 1'\'es on the <Jbaxia l
5111 uhs; fo1est, \Vc>od land and heatl1l~1nd
face, and (in most species) a sho11, thick, hooked style; simih11 to 1Jillu~) 1 11ia,
Hefe re nce(s): Gr ievc 0998: 424)
Pu/tenaea , Almaleea and Stonesiella within the Pultenaea seus~ lat. (5-nucleate
Diagnosed by a combi nation of long, filifol'm bractcoles, basally-lobed sr:1nda1d
cmb1yo -sac) group
;:ind te1minal inllore.o.;cencc; simib1 to 0 11011 and Aot11s \V ithin the P11/feJ1aen sens,
PioneeJ' species, used in reveget;:ition
lat. (5 -nuclea tc embryo-sac) group
Dillwynia srn. 1so5
c. 40 spp.; S <ind E Austi alia
Stonesiella Crisp & P.H.Weston 1999 Na med afle r Lewis Weston Dill wyn (1778 - 1855), a botanist based at Swansea in
1 sp.; E coast of Tas111ani;1 (very 1esttic1cd) \Vales and a close fliend of J. E.Smi rh ~ Dillwyn mainly worked on freshwater
N~1me<l in honorn of E. Ma 1garet Stones 0920 - ), botanical a1tist w ho iUustr<ncd nlgae, but h e also descri bed a numbe1 or species fiom the e~1 s1 coast of lndia in
rhis species in 'Jbe E11demic Flora o/Tas111a11ia Ho11us ,11alabari c11s
Sh1 ubs; in closed he~Hhlrin.cl on mo ist site:; on floodplains ancl nea 1 drainage lines Shrubs; temperate open sclerophyll fo1est and mallee woodl'1.nd, heaths and
in hills swamps on oligolrophic soils; f1om coastril to alpine rireas
Rderence(s): Stones & Cuitis (1978: t, 237, as 'P11//e11etea seletgi1Joides Jiook f'l; Heference(s): lllakely 0939); Lebler (1976); Elliot & Jones 0984, 3: 275-280);
C1Jsr el al Cl 999) Weber (1986: 664 - 667); Jeanes in Walsh & Entwisle 0996: 796 - 802); Grieve
Oclongo In the /'11//Cl/flitl .<en s lt1t. {S·nudc:nc cmbt)'O-s:ic) gmup and i~ pooh;rbl)" 0998: 428 - 429); Jobson & Wes1on 0998, 1999. 2001); Weslo n & Jobson in
closest LO , 1/111f1/fP!tr; tltignOS<.'(i In r:orr II)' keeled k":11·c. wilh minute ~lip11ltJ5 ;incl Ha rden (2002: 542 -549)
c<iducou s, lea f-like biacts Belongs to 1he Prtlteuaea smrs. lat. (5-nucleate cmb1 yo-sric) group; sim il;ir to
Sfo 11 esiel/o, Almaleea, Eutaxia nncl P11/1e11aea; cli:"tgnosccl with in the group by
subte1c tc (o ften t1 iquetrous) Jerwes wirh an 8dax ial groove or ch~mn el , ~1 broacl
standa rd and dilated win g apiccs
Several species of 'pa n·ot peas' are showy sp1ing omament<1 ls, also used as low-
Stonesiel/o selagino/des Painting by E. M. Stones Dillwyn/a floribunda Photograph by M. O. Crisp levcl cove r in wi ndbreaks

346 LEGUMES OF THE WORLD TRIBE MIRBELIEAE 347

 Pultenoeo divoricoto Photograph by M. D. Crisp Pultenoeo reticuloto Photographer unknown Mlrbelio dilototo Photograph by B. Fuhrer Mirbelio plotylobioides Photograph by M. D. Crisp

Pultenaea sm. 1794 Mirbelia s m 1805

E11cb i/11s R.13r. ex W.T.Aito n 0811) 32 spp.; Aust ra lia, mainly in the SE and SW . .
104 spp .; S '1 nd E Australia amed in honour of Charles FJ3. de Mirbel (1776 - 1854), French plant phys1olog1st
Named for Richard Pu lte ney 0 730 - 180 1), Etlglish s urgeo n and botanist, at rhe M useum Narionale d'Histoire N atu1 elle, Paris
biog1ctpher of Linnaeus Shnibs; sclerophytl com nnrnicie~ in eucalypt fo rest, woodland and he:Jth, mostly <.it
ShnJbs; temperate an<l subtropical , coa.stal to montane euca \ypt fo res t, woodland, low altitude on sa ndy and skeletal soils .
hea th and swamps, very ra re in rnin foresr •tnd ab.senr in the arid zone
Re fere nce(s): Stan ley & Ross (1983: 246 - 248); Crisp & Taylor 0987); Elltm &
Refe re ncc(s): St«n ley & Ross 0983: 257 - 263); Weber 0986: 672 - 687); Corrick Jones 0 993, 6: 425-433); Jeanes in Walsh & Entwisle ( 1996: 747 - 749); Gneve
in Walsh & Entwisle 0996: 765- 793); Grieve 0 998: 417-423); Westcm K De Kok (1 998: 345 - 350); Prnteners in Harde n (199 1: 530 - 53 1); Cnsp & Co ok (2003b)
in 1-fa rden (2002: 549 - 565); De Kok & \Vest (2002 , 2003, 2004); Orthia el al. Closely related to Oxylobfwu and Choriz ema w ithin the Pultenct~a sens , .'~"· C?-
(in press b) nucl eate embryo-sac) gi o up; diagnosed by an often late-developmg p a1t 1t1o n m
Proposed fo r expansion to subs ume nll 19 i;cnem and 470 ~pcc:ic' In the 'i- the ova1y and pod
nu lcatc e mbryo-sac group. except Lsol rop l. (On hia el a l., 2005, In pre.s a); Potentia l use as ornamentals
dingnoscd within the group by prominent r<."tl-brown stipuks wllh i,cariou.<
margins and va riably fused on the a<la xial side of the peti ole
Seve ral species of 'bush pea' have showy flowe rs and are cu l riv~tl~d a:->
orna mentals ( 1h 1iving in dry areas)

Pultenoeo brochytropis Photograph by M. D. Crisp Mirbetio specioso Photograph by M. Fagg

348 LEGUMESOFTHEWORLD TRIBE MIRBELIEAE 349

 Chorlzema retrorsum Pho tog rap h by B. Fuhrer Chorizema ilicifo/ium Pho tographer unknown
Chorizema Lobill. 1soo
27 ~pp.; A1c<1r.11J:i (rc61rlCIL'tl 10 1e1111><:r:ue and and ~ · Ausrrnlio. except c.
pwv(/10111111 Ocnrh .• which ocon~ along the ronsi of Queensland :md Ne.\\ South
Wa les)
!'mm t:bwi:n•/11 ( k.: 10 sep.amtc) and SIJJ/ltt (Ck.: ' lgn or IY.lnncr), posslhly
n:forrlng 10 the h1!(hly ronrr:1s1i11g L-.,nrr:il hlo<dt on the ""nclnnl; an ahcrn.11lvc
c.' ])bnurion Is the combination nf ~bo,.lw/11 and 11q11111 Gk •• llbmcnt). rcferrin8 10
the free t:11nin:1I Ola11k>n1.>: no nlcnlion i~ m:.de In 11,., original ti ·S<,-riprlon.
howe••cr, of rhe more colourful lntl'tprernrlon 1ha1 the n1111le l'Omntetnomt.:s (whh
dancing) the finding of fres h drin king w<ilt:r near ro where the plnm w;i s first
cli sco\'erecl by l abill ardiCre (from c/Jor o- 1 k. : d an ce] and ~u11111 I k.: drink])
Shrub.< or :1uh$hru1»1oclc1'0ph~ll rommunillc,; (cu aly1)1 forc«t, woocllnn<I :incl
h<~:11h). 1110~1 l y o n :.:mdy :md skclctnl ~fl:;; extend111ii lnro the :u id wnc:
flefercn :(s): Elll04 & Jones (1984, !I: jl - ;l6); Tn)•lm & Crisp ( 1992); Crisp &
·n1ylor ( 19')3): Wlc ck in I lnrden (2002: 529l; Crisp & Cook 200311)
11·~' •Ir n:lmcd to ,l//r/x.//". nd U'J./00/11111 whhln the l'11fll>Jtt1l!t1 s1111$ 1111. C5·
nucleate emb1yo -s:tc) group; di:1g110.,c;d by kee l sho1te.- than the wings and lack
of the ov<.ny partition .seen in 1\lfl'be/fn
Some species ol 'fhirnc pea' are grown as oina111enl;1ls for the alt/a ctive flm,~1s ,
scrnmbling habi t nod holly-like leoves (e.g •. C. i/icijo/i11111 Llbi ll .)
Oxylobium Andrews 1807
6 spp .; SE Ausnalia
Prom 0.\)1- (Gk.: .sharp) :lnd lobioll (Gk.: pod), refeiring to the ricum ina tc .i pex
th e pod
Shrubs or sma ll rrec~i ma inly in dry scleroph yll (eL1ca lypt) foJ'est and woocll:1ncl,
fro 1n coasra l ro monrnne regions
llt:fcrent'l..'(s). Crisp & \X -.ion 0987. 8?. 1995: .62): Jc:mes In \°U'Jl.'lh & El\lw1•le
J996, 7 l - 7· 3); Wiccck fn Harclen (2002: 532 - 533, plu.• Mirlxd/t1 a\:rlubloidt>$
F.Mu · II. I.e. p. 530): Crisp & Cook (20031>)
Closel)' rel~ned to Cburiz ema and 1lt!i1he/ia within the P11/tenaea sans. !rt! . (5-
11uclc:11c embryo·.s;•c) j!roup: distjngulslK'<I by the wlnJl.S being <-'CJU•I to th • kccl
and the lack of" p:111ilion In the ovnry; formerly fnduded n re than 30 ~f><."<i
now 1mnsfcrrcd to Ga.<1ro/l)l>i11111, l'0tf"1t>/1/11m and Oiiier gencm
Some species o f 'shtlggy 1.,.,,.· :ire grown :i.• omr1111cn1:ib
Oxy/obium arborescens Pho tograph by M. o. Crisp
350 LEGUMESOFTHEWORLD
Podolobium ilicifo/ium Photograph by M. D. Crisp
or
Gastro/obium bilobum Photograph by H. Thompson
Callistachys /anceolata Pho tograph by H. Thompson
Podolobium R.lll'. ex W.T.Airon 1811
6 spp.; SE Aust"ilia
Frompodion (Gk.: Foor. e.g ., of o va.,e) and lobio11 (Gk.: pod), referring to the
stipe of the f 1\1it
Shnibs; dry sclerophy ll (euca lypt) forest, woocl lond and heath
Refe1cnce(s), Crisr & Weston (1995, 262-263 & 279- 281); Wiecek in Ilorclc n
(2002; 533-535): C1·isp & Cook (2003b)
Oi:agnosccl by spre.ading subulate stipules and reticulate leaves and pods; belongs
LO the P11fle11aea selfs /al . (5-nucle me enlbryo-sac) g1oup; 1ecently segrega ted
f10 111 Oxylobi111111 b ut molec ul ar claw suggests thnr tile genus is bi phyl etic, wit h
th1 ee species dosely related lo Ca/lislacbys, whil e the o th ers :ire placed w ith
eithe 1 Gas1rolobi11 !!I (l m L-FJ m C/Jo1·ize111a (ITS) (Crisp & Cook, 2003b)
Callistachys ve nt, 1805
1 sp.; S\V Au stralia, most ly ne~ll' th~ coast, and ir1ainlancl SE Austra lia
from ca/Ii· (Gk, beau tiful) and stacbys (G k., spike), refening to the conspic uous
crncl ';lttractive inOorescences
Low shrubs to sm:ill t1ees; swampy places, oflen seasonal, and along drainage
Jines (\VJ Australia); coasta l to sub-alp ine sclerophyll co1rnmmi l ie.'i (SE Au stralia)
Re fe1encc(s): Wheele r & Crisp in Marchon t et al. (1987: 291 , Fig. 92, os O>.ylobiwn
/a11ceo/(lf11111 (Ve nt.) Drn ce); Crisp & Weston (1937, 83 - 117); Clisp & Cook
(2003b)
Di:1gnosed by conspicuous 1 ~1cemcs of yellow nowers, ptese nce of h racteoles and
retic ul ate, v ill ous pods; within the P11/1e1wea sel7s fell . (5 -nuclea te c111 b1yo-s<.1c)
gro up , close l)' re late d to iVfirbelia, OA:ytobium <.i n cl C/Jorizema
Gastrolobiu1n R.flr. ex W.T.Aiton 1811
Bracbyse!!la R.fl 1·, ex WT.Aiton (]Sll);fcmson ia Kipp ist ex Lindi. (1817); Ne!!lcia
Domin (1923); C11pulc1111b11s Hutch. (1964)
109 spp.; mostly restricted to S\XI Ausu'Jlia; two in C rind N Aust1'Jlia
F1om gas1er(Gk .: belly) and /obion (Gk., pod), 1efe1ting to the swollen f111it
Shrubs; sclerophyll cmnm Lmities in eucalyp1 forest, woodland, he~th, dry
shrobl~nd and hummock g1assland
(continued on next page)
TRIBE MIRBELIEAE 351
Gostro/obium rubrum Photograph by M. o. Crisp Gostro/obium pyromldo/e Photog raph by M. o. Crisp Leptosemo tomentosum Ph otograph by 8 . Fuhrer Leptosemo doviesioides Photograph by A. s. George

Gastrolobium (continued)
Refe re nce(s} Elliot & Jo nes (1 986, 4: 336 - 349); Ci isp & Westo n 0987: 83 -11 7);
Cris p in Eggleto n & Vane-Wright 0994: 28 1- 309); Crisp & Westo n 0995:
262 - 263, 282); Crisp 0995a); Chand ler el al, (2001 , 2002)
Although there is !;trong molecula1· support for this genus ~is a cl ade i nclud ing
Brachysema, ja nsonia and Nemcia , no 11101phologic:.i l syn apomorphy is known;
diagnosed wi thin th e Pultenaea sens. lat (5-nuclea te e mbiyo-sac) group by a
combination of ru gose-lobed arils, opp os ite or who rl ed phyllotaxis, abse nce of
b1acteoles and lo ng sle nde r stipu les
Ma 11y species produce flu oro-acetate, w hich is highly to xic to many vcnehr ~Hes,
including domestic gra zing animals (widely known as po ison o r p oison bushes);
all species formerl y included i n Bracbysema andjunson.ia, ri nd sevcial o f those
from Nemcia , have large fl ow ers modified in shape and colo ur for bi1·tl
pollination; this gro up appea rs to b e rn onophyletic and nes ted deeply w ithin
Gaslrolobium ; on ly a single species (G gra ndiflonwi F.Muell.) is both wxic and
bird-po llin" ted, but it is not included in the bird- pollinated clade
Used as o rnamentals, esp ecia lly bird-pollinated spec ies; those fonneily included
in Bracbysema <mdjanson/a have an :attracti ve lo w-growing grow th hahic :ind
conspicuous flowers; used for ground cover

Gostrolobium brocteolosum Photograph by 8. Fuhrer Dav/esio incrossoto Photograph by 8. Fuhrer Gompholobium copitotum Photograph by A. s. George

352 LEGUMES OF THE WORLD TRIBE MIRBELIEAE 353

IR I BE
i;ossiaeeae by J. H. Ross and M. D. Crisp
'be Bossiaeeae (Benth.) Hutch. 1964.
ffl Tribe Genisteae subtribe Bossiaeinae Benth. (1865), as 'Bossiaeae'
}.S createc.I here Bossiaeea compri e 6 n ra and c.
species (Fjg_ 43 , distributed tbrouP-hout Au tralia
72
i;:lllt mo. t f the diversity is in the outh-west and south-
ea r. Most sp cie are hrubs with yell w and r d 'egg
llncl bacon') flow 1·.Th y are con pi uou , ometime
deit1inanc und r c r y m m er, f s J rophyll
coinmuniti.es (he thland and euca l ypt-dominat d
woodland and foresr), on po rs ils of rh s uth-v est
south and east coast of Australia.
Polhill (1994) placed Bossiaeeae between
Brongniartieae and Mirbelieae, largely because he
considered the Templetonia group a link between
Bossiaeeae (where he placed this group) and
Brongniaitieae (see also Polhill, 1981m & n). Previously,
Crisp & Weston 0987) bad transferred the Templetonia
group to Brongniartieae, based on a cladistic analysis
using morphology. Recent phylogenetic analyses,
especially those using DNA sequences (Crisp et al.,
2000; Doyle et al., 2000; Hu, 2000; Hu et al., 2000;
Pennington et al., 2000a, 2001; Wojciechowski et al.,
2004), support a placement of Brongniartieae either
next to, or within, the Genistoid clade. These same
studies indicate that Bossiaeeae and Mirbelieae are
related , not to the genistoids, but to Hypocalypteae,
lndigofereae, Millettieae and Phaseoleae, and the large
!Iologalegina group of tribes.
FIG. 43 Diagram of relationships in tribe Bossiaeeae after Crisp et al. (2000); Crisp et al. (unpublished)
LEFT Bossiaea dentata Photograph by M. Fogg
According to morphological evidence, the
Bossia eeae is the monophyletic sister group o f
Mirbelieae (Crisp & Weston, 1987). Phylogenetic
analyses based on DNA sequences, however, are more
equivocal. A monophyletic Australian radiati on
including both tribes is indicated by nuclear and
chloroplast DNA, but support for this clade is weak
(Crisp et al., 2000 ; Crisp & Cook, 2003a ;
Wojciechowski et al., 2004). Whilst Bossiaeeae is
clearly monophyletic, its relationship with the genera
of Mirbelieae is unresolved and it would be premature
to redefin e these tribes (Crisp & Cook, 2003a).
Bossiaeeae share embryo-sacs having giant antipodals
with Mirbelieae genera Daviesia, Gompholobium,
Viminaria, Erichsenia and Sphaerolobium (Cameron
& Prakash, 1990), but this embryo-sac group is rarely
monophyletic in DNA analyses (Crisp et al., 2000;
Crisp & Cook, 2003a). Internal relationships of
Bossiaeeae are well resolved by DNA, with Goodia
sister to the rest, and a clade of Muelleranthus,
Aenictophyton and Ptychosema sister to Bossiaea and
Platylobium (Fig. 43; Crisp & Cook, 2003 a) .
Platylobium is nested within Bossiaea and these
genera will probably be combined (Crisp et al. ,
unpublished).
Mirbelieae
(see page 339)
Goodia
Bossiaea
co
0
Platylobium Vl
l/l
)>
rn
Muelleranthus rn
)>
rn
Ptychosema
Aen ictophyton
TRIBE BOSSIAEEAE 355
 Goodlo lot/folio Photograph by 8. Fuhrer Bossloeo sp. Photograph by 8. Fuhrer Platy/obium o/ternifolium Drawing by A. M. Podwyszynski Platy/ob/um triongulore Illustration by 5. Edwards

Goodia Salisb. 1806 Platy l.obium sm. 1793

2 s pp.; Ausrralia, widespread in the Sand E (sou th-west \Y/ Australia to SE
Queensland, b ut with disjuncti ons in Sand \Y/ Australia, and in N T<ismania)
Named for Peter Good(' -1803), Kew gaidener and botanical collectm who
~ cco mpan ied Robert Brown to Australia on board H.A 1/ • lnvestign fOI and died
of dysentery in Sydney in 1803
Shrubs; subt1opical, medire rranea n and temp e1ate forest, woodland, bush land,
th icket and shrub land
Refe rence(s), Stan ley & Ross (1 933 , 272); Elliot & Jones (1986, 4, 410-412); Weber
(1986, 691-692); Ross in Walsh & Entwisle (1996, 819 - 821); Hoss 0997); James
in Harden (2002, 509-510)
Distinguished from other genera in the Bossineeae by the pinnately trifoliola te
leaves. Phylogenetica ll y the s ister group to the other genera in the Bossiaeeae, bu t
diverge nt from them in both morp hology and ONA seq uences
00<1/rt lmlfo/in llsb. Is suspcm:<I of c:iu~lng ck::uh in c:mlc, hydr"l! •n cyanide is:
tJ1e !lctivc compound, opparentl)' in the for111 of un'<l:ihlc , nhydrin (F\ e risl,
1981): rhls >p<:cic> ls. I~> 'uhiv111ed as an ornnmcnml (AQldc11 rip. d<>\'l'f bush or
yellow pc:t) :ind i UM..'<! for ht.xlg"' and , • plon~'Cf!I In rt:V"l.{Cl"1i<:10
4 sp p; E and SE Australia (Wide Bay in Queensland to Kangaroo Island in S
Auslla lia, and in E Tasmania)
From platy- (Gk,, flat) and lobion (G k,, pod), referring to the laterally co m pressed
pods
Shrubs; subtropical, medi terranean and temperate forest, woodlan.d, bushland,
thicket and shrubland
Refe rence(s} Stanley in Stan ley & Ross (1983, 265); Ross 0983); Weber 0986,
694 - 695); Hoss in Walsh & Entwisle 0996' 816 - 819); J ames in Harden (2002,
516)
Closely allied to, and probably nested w ithin Bossiaea (u npublished data), from
which it d iffers in the development of a d ist inct wing beyond the upper sutural
ne rve of the pod, the ext re me enlargement of the two uppe1• calyx-lobes relative
to the lower three (this cond ition found also in Bossiaea spinosa (Turcz.) Domin),
a ha ploid chromosome number of 8 as opposed to 9 in Bossiaea , and pollen with
a well-defined endoaperture
Potent ial as ground cover for erosion control and used as ornamentals (e.g.,
P.formosum Sm., or handsome flat pea, page 359)

Bossiaea vent. 1800

c. 60 spp.; Australia (widesp read, but not in C Austm lia)
Named for I3ossieu d e la Ma1tin iere, medical officer and bol:mist acco mpanying
the expedition of La Perouse on /'A strolabe w hic h was wrcc:kc.:d ne<ir th e isl:rnd of
Vanikoro (Santa Cruz grou p, Melanesia) in 1788
Shrubs; tropical, subtrop ica l, mediterranea n, temperate £ind warm tempew te
fo rest, wood land, b ushl and, thicket a nd sh rub land
Referen ce(s), Ell iot & Jones 0982, 2, 355-362); Stan ley & Ross (1983, 266-268);
Weber 0986, 688 - 691); Wheeler in Marchant el al. (1987, 239-242); Ross in
Walsh & Entwisle (1996, 808 - 815); Ross 0991, 1994a, b, 1998e, 2001 b); Jan1es in
Ha rden (2002, 510-516)
Closely allied to Platylobium; flowers subtended by a se ries of coriaceous or
papery brown bracts, main ly insect-pollinated , but 3 species adapted frn
poll ination by birds
Foliage toxi c ro livestock; used as ornamentals

Bossioeo prlessll Photograph 8. Fuhrer Pfaty/obium obtusongulum (A- E), P. trlongulore (F- J)
Drawing by A. M. Podwyszynski

356 LEGUMES OF THE WORLD

TRIBE BOSSIAEEAE 357
Muelleronthus stipuloris Drawing by P. Halliday Ptychosemo onomolum Drawing by P. Halliday

Muelleranthus Hutch. 1964

3 spp .j Australia: mainly in cbe Eremae;rn or dry zones of \Y/ 1\u stmlia, the N
Territory, nonh-wcstern S Austra lia, Queenshrnd and N New Sou1h \X':il es, but
distiibution disjunct
N;:11ncd for Mueller (O;:u o n F.j.H . Mueller (1 825- 1896), first Government Uurani.st
for Vk to ria ( 1853 - 1896)) and antbos (Gk.: a nowe r); the ge nus is b:1sed on
P01cbosema lrifo/io/ar11111 F.Muell.)
He1bs or sh1ubs; suhtropic~1I , mcdite nane~m and temperate shrubbnd: fi1\0 U11'i
sandy soils, often associ:i tcd with 'Ji·iodin spp. (G ramineac)
llefe l'e nces(s): Lee 0973); Crisp in Jessor (1981: 152) ; Weber (1 986: 09.'--694);
Ell io t & Jo nes (1993. 6: 454 - 455);.J<imes in ][arcle n (2002: 517-518)
Disti nguished f1·0111 other genera of Ilossi::1ee.1c by digitarely tiifoliol;lle k:;n·e.s

Ptychosema Be nih. 1839

2 spp.; Au st1:lli;1 , dist1'ibution \'Cry <llsjunc1 (1 sp. in soutlFwestern \Y/ J\u:-.lw lh1, 1hc
other in the S\XI corner of the N Territory and in the a1·icl interior· of N New ~out h
Wales)
F1om jJlycbo· (Gk: a fold) :ind se11w (Gk .: a sign or banner), app: n e nll~ in
reference to 1he d eepl y norched apex o f the stnnchud peta l
Herbs or shrubs; su lxropica l and medicen ane;rn shn.1b land , fa vou ring 1->: ind y soils
Rel'ere nce(s): Lee (1973); Crisp in Jessop (1981 : 152); Webe r ( 1986: 690 - 696);
/ames in lla1cle n (2002: 517)
.Distinguished from other genera in llossia~eae by imp:iripinnc1tc leaves "ith
opposite lea llcts

Aenictophyton AT.Lee 1973

1 sp ; C Australia (Eremnc;m zone e~1 st and west o f the \V Aust1ali~ and Nouhci n
Terl"ltoiy houncb1y)
from ai11iclo- (Gk~ : hnflli ng) and phy/011 (Gk: a plant), in reference lO rhe
difficult)' experienced by A.T. Lee in classil')'ing the r l;mt
Slifllhs; Slthtropical sbrubl::md, fa\·ouring deep red sand>T soils
Reference(s): Lee (1973); Crisp in.Jessop CJ98l: 151)
Reminiscent of Bossia<?a from which it differs by havinH flowe 1s in l;1 x tcr niin:d
1:lccmes and not su btended by <l series of rxi.pe1y b1·own hl'ac cs

Aenictophyton reconditum Drawing by A. T. Lee & R. M. Polhill P/aty/abium formosum Photograph by M. Fagg

358 LEGUMES OF THE WORLD TRIBE BOSSIAEEAE 359

TRIBE
111digofereae byB. D. Schrire

Jndigofereae Benth. 1859

T~ribe Galegeae subtribe Indigoferinae (Benth .) Benth. (1865), as 'Indigofereae'

p (hill (198lf) recognised 4 genera and c. 710 species
~oindigofereae. This treatment following Polhill (1994),
chrire (1995) , Barker et al. (2000) and Schrire et al.
cio03) recognises 7 genera and c. 768 species in the
tribe (Fig. 44). The Indigofereae are predominantly
}ifrican-Madagascan in distribution, occurring in
easooally dry vegetation types of the tropics and
subuopi s. The genus Indigo/era (third largest in the
i.egumino a ) is pantropical in distribution.
Recent morphological-molecular analyses
(Pennington et al., 2000a ; Crisp et al., 2000;
\'\lojci d 1owski et al., 2000, 2004; Hu, 2000; Kajita et al.,
2001; Hu et al., 2002 and Wojciechowski, 2003) place
Indigof reae at the base of a combined millettioid
group of tribes (including Millettieae, Abreae,
Phaseoleae, Desmodieae and Psoraleeae). This entire
clade is sister to Hologalegina (comprising the
robinioids and the Inverted Repeat Lacking Clade
(IRLC)). Basally branching to these two clades are the
South African Hypocalypteae and Australian tribes
Mirbelieae and Bossiaeeae.
The Indigofereae (Barker et al., 2000; Schrire et al.,
2003) comprises a Cyamopsis, Indigastrum, Microcbaris
and Rhynchotropis (CRIM) clade which is sister to the
Indigo/era - Vaughania clade. The Madagascan
Phylloxylon is putatively the most basally branching
genus in the tribe, although in some analyses in Schrire
et al. (2003), Phylloxylon is sister to the CRIM clade.

Phylloxylon

r= Rhynch0tropis

z
J L Microcharis 0
G)

II ,,0
lndigastrum rn
;:o
rn
)>
rn
; Cyamopsis

Vaughania
lncligofera

Millettieae (see page 367)

Abreae (see page 389)
=- Phaseoleae (see page 393)
Desmodieae (see page 433)
Psoraleeae (see page 447)

Hologalegina

FIG. 44 Diagram of relationships in tribe lndigofereae after Barker et al. (2000); Schrlre et al. (2003)

LEFT lndigofera pendula Photograph by P. Cribb

TRI BEIN DIGOFEREAE 361

 Rhynchotropis poggei Drawing by M. Warwick
Cyamopsls senegalensis Drawing by c. Grey- Wilson Microcharis tritoides Photograph by S. Collenette

Phylloxylon Boi l!. 1861 Microcharis Benrh, 1865

Indigo/era subgen. Microcbaris (Benth.) j ,Il.Gilleu (1958)
Neobaro11ia Baker (1 884) 36 spr .; Afri ca (except largely \VI Africa; c. 16 spp. in Somalia-Masai and c. 18 spp.
7 spp.; Moclagascnr (m ostly N ''"cl \VI, 1 sp widesrread 1'1 om E to SE) in Zambezian to Sudanian regions), Arabian Peninsula and M:ubg<1scar (2 spp.)
from pbyllo- (Gk.: leaO and :..ylou (Gk.: wood), referring to the lcathe1)' or f 1om 111 /cro- (Gk.: s mall) and cbaris (Gk: beau ty, grace), rcfen ing to the graceful
woody phyllodes in the type srecies
and oflen diminutive habit of the genus
Trees ;rnd la rge sh1ubs; seasonally dry to humid tropical forest and\\ oodl;md, Herbs; seasonally d1y tropica l forest n1a1gins, 'vooclland, thi cket, wooded
on sand or rocky (often lime.stone) outc1ops grasslnnd and grnssland, often in <lamp swampy or rive1ine me~s . 01· in shallow
llefcrence(s): Du Puy el al. (1995a); Du Puy & Labat in l)u Puy el ol. (2002:
soil ovc1 rock
451-461) Hefe1ence(s), Gil lett 0 958); Sch1ire 0992; l995); Scl11'ire el al. (2003)
Puta ti ve b:isa lly branch ing genus in the t1i be w itl1 u nce 11<1in ::i ffini ties ou tside the
Sister genus to Rby11cbo1ropls (Ba1·ker el a l ,, 2000)
lndigofere::ie; Du Pu y & Labat in Du Puy el a l. (2002) nme that 3 spec ies <1 1c
critically endange1ed and a fu11hc 1 3 species ;:1 1e endangered
The ext1emely hard wood is used fot fuel, poles. implemc n1s and co11~1 niccio n ; Rhynchotropis Ho1·111s 1901
the bark cont;dn.s fish poisons (rotenones); also L1secl in sorcery
2 spp .; SC Africa in the N of the Za111bezi::1n 1cgional ce ntre of endemism
(White, 1983)
Cyamopsis De. 1825 From rhy11cho- (Gk.: beak) "nd 1ropis- (Gk , keel), referring to the distinctively
Cordaea Spreng. (1831) shared keel petals
4 srp.; Arri ca (mainly in the se mi-arid S\'(f, l sp. disjunc1, also occuning in the Herbs; seasonally dry tmpicnl woodland and wooded g1·ass1and , often in
Sud:anian and Somalia-Masai regiona l cenires of endemism in10 Al":.1bi;1); se~1sonally damp 0 1 open sanely and 1ocky arc~ts
C letragomJ/oba (L ) Taub,, apparen tly unknown in the w ild, but possibly from ll eference(s): Gillett (1958); Sch 1i1e (1992; 1995); Sclll'irc et al. (2003)
India (G illett, 1958) Flowers often before the leaves are produced; sistcJ genus to MioTJcbai.,·s ( l3a1ker
Prom ~Y"'"o- (Gk.: be:in) and -opsis (Gk.: resemblance) el al ., 2000)
Herbs; seasonally dry l ropical 1hom scrub and grassland, oflen in nuoc.l phlins,
stream beds, pans ~ind in open sandy ot rocky a1e<1s
Heference(s): Gillett (1958); Schril'e 0995); Sclll'irc el al (2003) Vaughania s. Moore 1920
C)'c1111opsis is placed bas<i lly i n a c1 ~1 Je uniri ng Jndigastriw1, Rbyncbotropis and
11 spp.; tv1'1dagascar (S, S\V/, \VI and C)
J\ficrocbaris, whic h is sister to an !11digofera chide (B:uker el rJt, 2000) Named after the ll1'itish botanist joh n Vm1ghan Thom pson (1779 -1847)
Cyamopsis lelrttgonoloba (guar, Calcutta or duscer bc<1n) is wide ly cuhiv:t1 ed in Trees and shrubs; seasona lly d1y tmpical fo1est, woodhincl, xeropbytic bushland
tropi cal countries ;1s a source o f seed gum used in rile food, paper and textile and grasshrnd, on san d or rocky (o ften limestone) outc1ops ~incl rive 1ine
industries; also irnpo1t1nt for lrnman food (pocis), fodder, soil improvcmcnr :i nd Refcrence(s): Du Puy el al. ( 1994); Du l'uy & l:lbot in Du Puy et al. (2002:
sr<1bil isation, nnd as green mam11 e
46 1- 474)
Recent moleClllor evidence ( !301 ker el al., 2000; Sch1ire el al, 2003) places
Indigastrum Joub. & spoch 1857 Va11gbania , toge1her with the anomalous Rf-union Island endem ic Indigo/era
/11digofera subgen. Jndlgastmm Qaub. & Srach) J.B.Gilletl (1958) ammoxylum (DC.) Polhill (= Bremouliera ammox_ylum DC.), as elements w il hin
c. 8 spp.; Africa (except larsdy \Y/ Af1ica, mosrl y in Zambezian ro Sudnni:1n 1cgions fndigofera ; both are conside red to be highly modified, fndhrn Ocean I sland, neo·
nnd 3 spp. in the Ka1t)o-Nrnnib l'egion of so uthern Aflic:J); 1 sp. pan tropical endemic li neages of /11digoferc1 , but fonnal ising the llansferof Vcwgbania in to
from -11s1rum (L : resembl ;rnce), referring ro its similmity to !ndiuofera sy nonymy awaits fu1t her s1udy
Herbs; ~nson3 lly chy tropic;1l wood land, bushland , 1hicket or grassland. of1en in The exc remel y hal'd wood is used for ruel , poles, imp lements nnd constructio n;
seasonally damp or open sandy and rocky are[IS other uses are fod d er for livestoc k
Reference(s): Gillett 0958); Schrire 0992; 199)); Scliri re el al. (2003)
A group of c. 4 species, occurring from the drier areas of South Afr ic.1, N:1mibi~
1

and Angola, differ morphologically and may complise a dis1inct subgent1!'>

lndigastrum fastigiatum Pho tograph by B.O. Schrire Used os fodder and for dyes Vaughania depouperata Ph otograph by D. Du Puy

TRIBE INDIGOFEREAE 363

362 LEGUMES OF THE WORLD
lndigofera bosser/ Photograph by D. Du Puy lndlgofera hochstetterl Photograph R.B.G., Kew

Indigofera L. 1753
A11// Miii. (1754); Sph11e1trliopbon1111 Oesv. (1813); Bremontlera DC. (1825);
Acm11ho11otus Benth. 0 849): A11//a Ludw ex Kuntze Cl891)
c. 700 spp.; Africa-Madagascar Cc. 490 spp.); Asia lo Pacific Cc. 115 spp.);
Australasia Cc. 30 - 40 spp.); c. 13 spp. widespread in the Palaeotropics; c. 6 spp.
pantropical; New World c. 45 spp. USA to Argentina Cc. 30 spp. in N < C Americ:i,
c. 1S spp. in S America)
From ind go !Uld -fer (L.: bearing), referring to species of this genus being the
sou rce of the <lye indigo
Herbs, shrubs or small trees; seasonally dry tropicnl to warm temperate forest,
woodland, wooded grassland and grassland, ,;dorophyllous shrubland, forest
margins and disturbed areas
IMcn:n !( ) : Gillett (19511): Sa n~1 pp3 (1995): Oc Ko n & 111ijssc 0 981): rolhill
( 1990). Lievens (l!)<JZ); S.:hri rc ( 1995): Schrirc in Gol<lbbtt & M11nnin!{ (2000:
86 - 492); l)u f'uy & l.nbut In Ou ruy (JI a l. (2002: 476 - 509 : Schrirc t •J a/. (:!O(m
A gCm L• of c. 25 sectio ns In Afrk :i •M:iclagnscnr wh<.'re h ul l/Jflfum s most
d lvcr,;c, incre:1Sing to c. 30 sca ion.s worldwitlc:, nil spcclc belong to one of
four wcll-suppont'<l and bioi:eogrnphfcally dbtlnt1h•c lade~ In th • analyses of
Schrire et al. C2003)
Used as dyes (imponanl species are I. arrecta Hochst. ex A. Rich , I. articulata
Gouan, I. suffrulicosa Mill. and / . tinctorla L.), medicine, fodder, cover crops,
green manure, human food, erosion control and ornamentals; some species are
toxic to livestock, others have insecticidal qualities

lndigofera colutea Drawing by c. Grey-Wilson lnd/gofera heterantha Photograph by B.D. Schrire

364 LEGUMESOFTHEWORLD TRIBE INDIGOFEREAE 365

TRIBE
Millettieae byB. n. schrire
. ·i,e Millettieae Miq. 1855
rri tribe Tephrosieae (Benth.) Hutch. (1964) pro maj. parte
Tribe Galegeae subtribe Tephrosiinae Benth. (1865), as 'Tephrosieae'
Tribe Lonchocarpeae (Benth.) Hutch. (1964) pro maj. parte
Tribe Dalbergieae subtribe Lonchocarpinae Benth. (1860), as 'Lonchocarpeae'
Relation hips among genera of Millettieae have been
notori usly difficult to unravel based on traditional
rphologi ·al evidence and this is exemplified by the
1110
:iJphabeli al arrangement of genera in the tribal
treau11enl of Geesink (1981; 1984) and Polhill (1994) .
Geesink (1981) recognised 44 genera and c . 870
species in tribe Millettieae (as 'Tephrosieae') while 43
genera were accounted for in Geesink (1984) and
Polhill (1994). The genera recognised, however, varied
considerably with only 33 genera in common to both
treatments of Geesink, while the list of Polhill 0994)
combined elements of Geesink (1981, 1984) with new
data accumulated since then. Tephrosia has traditionally
comprised some 400 species but this is re-estimated at
c. 350 species here.
The traditional circumscription of the predominantly
pantropical and subtropical tribe Millettieae is followed
here (Fig. 45), with 45 genera and (904)-909-(914)
species being recognised, (i.e. excluding the two genera
and 11 species transferred to Brongniartieae, see Table
8), although the concept of what comprises Millettieae
sens. strict. is changing rapidly based on evidence from
molecular phylogenies. Sequence data for millettioid
genera comes from the plastid rhcL gene (Doyle et al.,
1997; 2000; Kajita et al., 2001; Hu & Chang, 2003),
phytochrome nucleotide genes (Lavin et al., 1998), the
plastid trnK-matK region (Hu et al., 2000) and the
nuclear ITS region (Hu, 2000; Hu et al., 2002). Molecular
data, together with reinterpreted evidence based on
chemistry (Evans et al., 1985) and wood anatomy
(Gasson et al., 2004), have been the basis for recognising
a number of informal suprageneric groupings and for
transferring Cyclolobium and Poecilanthe to tribe
Brongniartieae (Table 8; Fig. 45).
The most far-reaching result of the above molecular
analyses was that a substantial part of the traditionally
circumscribed tribe Phaseoleae is more closely allied to
the core-Millettieae than to the Phaseoleae sens. lat.
LEFT Wisteria {loribunda Photograph by G. P. Lewis
clade (see page 393). Circumscription of a revised tribe
Millettieae is not possible at present until genera are
more comprehensively sampled; however, a Millettioid
sens. strict. group might be expected to include some
genera in the basal millettioid and phaseoloid group,
Phaseoleae subtribes Diocleinae, Ophrestiinae and in
small part the Erythrininae, tribe Abreae and the core-
Millettieae (Fig. 45). The basal millettioid and phaseoloid
group comprises 17 genera (94 species) that may belong
either in the Millettioids sens. strict. or Phaseoleae sens.
lat., or to a clade sister to both these groups (e.g., Kajita
et al., 2001). The core-Millettieae clade comprises c. 22
genera and c. 777 spp., with some additional generic
segregates being necessary within the 'canavanine
group' (Evans et al., 1985), to accommodate species of
Millettia sens. lat. and Fordia sens. lat., which on the
basis of molecular and chemical evidence are excluded
from Millettia and Fordia sens. strict.
Relationships between the major groups of genera
centred on Lonchocarpus, Derris, Millettia and
Tephrosia remain obscure, and still reflect a
geographical bias in segregating them, i.e. distributions
are limited largely to the New World in the
Lonchocarpus group, and the Old World in the other
groups. The suggestion that the Andean South
American genus Apurimacia might be sister to the
largely Old World Tephrosia rather than to
Lonchocarpus (e.g., Kajita et al., 2001) is possibly
indicative of other Old World-New World sister groups
yet to be found. Further molecular evidence will
probably result in an overall reduction in the number
of genera recognised, particularly in the Tephrosia and
Lonchocarpus groups where various small or
monotypic 'one-organ' genera may be better placed
within larger genera. Ptycholohium, Requienia and
Paratephrosia, for example, are difficult to distinguish
from Tephrosia, but for the emphasis traditionally
placed on their atypical pods.
TRIBE MILLETTIEAE 367
 Pyranthus
TABLE 8 Informal groups currently recognised within the traditional concept of tribe Millettieae based on Evans et at ( Chadsia
Lavin et al. (1998), Hu (2000), Hu et al. (2000), Kajita et al. (2001), Hu et al. (2002), Hu & Chang (2003). • 198 Mundulea
Tephrosia
INFORMAL GROUPS NUMBER OF GENERA NUMBER OF SPECIES DISTRIBUTION
Apurimacia

---
Paratephrosia
A Millettloid groups 23 c.131 Requienia
CL Ptycholobium
1) Transferred to tribe Cyclolobium Tropical S America ::::l
Brongniartieae (page 253) Poecilanthe ,_
0
l9 Millettia sens. strict.
2) IRLC 'millettioid group' c. 6 c. 37 Tropical & temperate Asia; Cal/erya also in Aust~ Q)
Wisteria also in N America a, UJ c Pongamiopsis
<( c
UJ re
3) Basal millettioid and 17 c. 94 Mc;>stly tropical Africa, Madagascar and SE Asia· · I- >
re Derris
phaseoloid group Atistrosteenls(a in Australia; Platycyamus in tro'pical I- c
South America UJ re Paraderris
_J
_J
y
c
0
B Core-Millettieae 22 c. 777
:2: z Dahlstedtia
UJ
~ ?Deguelia
5) Canavanine group c. 1 (plus generic c. 12 + Tropical Africa-Madagascar for Philenoptera 0 Lonchocarpus
segregates ex Millettia u Behaimia
and Fordia)
I I
Bergeronia
6) Non-canavanine group ( Margaritolobium
I Muellera Hesperothamnus
a) Piscidia group 2 c. 12 Mexico, CAmerica and Caribbean with one species to Piscidia Millettioid
I western S America
sens. strict.
b) Lonchocarpus group group
7 c. 144 Tropics and subtropics of the New World Philenoptera
Millettia pro parte
c) Derris group 2 I c. 70 SE Asia with few species to Africa & Australia Fordia pro parte
(? = lmbralyx)
d) Millettia group c.2 s 153 Tropical Africa, Madagascar & SE Asia but some taxa will
be transferred elsewhere, e.g. to the 'canavanine group'
or basal millettioid and phaseoloid group Phaseoleae
I
(subtribes Diocleinae
e) Tephrosia group 8 c. 386 Mostly tropical Africa and Madagascar; Tephrosia to & Ophrestiinae)
Asia, Australia and tropical C & N America; Paratephrosia (see page 394)
in Australia; Apurimacia in Andean S America Abreae (see page 389)
Austrosteenisia
?Leptoderris
Dalbergiella
Agan ope
Ostryocarpus
Xeroderris
Fordia
Dewevrea Basal millettioid &
?Platysepalum phaseoloid group
?Sylvichadsia
Schefflerodendron
Craibia
?Disynstemon
Platycyamus
Kunstleria
Burkilliodendron
Craspedolobium

Callerya
?Antheroporum
Endosamara IRLC
Sarcodum 'millettioid group'
Afgekia
Wisteria
Poecilanthe Brongniartieae
Cyclolobium (see page 253)

? "'placement uncertain in generic group

FIG. 45 Diagram of relationships between informal groups within the traditional circumscription of tribe Mlllettieae after Evans et al. (1985); Lavin et
a/, (1998); Hu (2000); Hu et al. (2000); Kajita et al. (2001); Hu et ol. (2002); Hu & Chang (2003)

TRIBE MILLETTIEAE 369

368 LEGUMES OF THE WORLD
Callerya megasperma Illustration by A. Barnard Antheraporum pie11ei Drawing by P. Halliday Sarcodum scandens Drawing by P. Halliday Afgekia sericeo Photagrapher unknown

CaUerya Endl. 1843 Sarcodum Laur. 1790

c. 3 spp; Asia (S China, Inda-China, Malesia, Papuasia)
Padbntggea Miq. (1855); lf'/Jfl{ortliodt•ndron Elmer 0910); Adinobot1ys Dunn
From sarco- (Gk.: fleshy), fo 1 the fl eshy exocarp o f the pods
091 1); Millellia sect. Eur)'bollytJ<• Dunn (1912)
Lianas; tropical rain forest
c. 20 spp.; Asia (Indian subcon tinent, China, Inda-China, Malesia and Papuasia)
and Austra lia Re fere nce(s): Merrill (1928); Adema 0999)
Allied to a g roup o f millettioid genera (Geesink, 1984) now placed apart f1om the
From ca/Ii- (G k.: beau tiful) and eryo- (Gk.: to gua 1d), the flowe r buds are
core-Mi llettieae in the Inverted Repeat Lacking Clade (IRLC) in the Hologalegina
protected by bracts and bracteoles
alliance (Hu et al, 2000)
Ll anas ro scandent shrubs (2 spp. are trees); tropica l rai n forest (some times
Used as ornamentals
inundated) and seasonally dry forest (often riverine), woodland, wooded
grassland, bushland and th icket
Rcrcren c(s): SdK,Jt (19') ) ; Ph:m Kc t llc & Vidtil (200 1): llu & Chnnt: (2003)
111e i;ruup of genera from Call<'1)vi 10 Wf>101'fd I l'Jg. 45) b now pl.1 ·~od apart from
Afgekia Cra ib 1927
th · rore-M illet1i<.':lc in the Inven~'<i Rc~"t Ln king Cl.1dc (lllLC) in 1hc: 3 spp.; Asia (S China, Inda-China (cultivated in Papuasia))
Ho loga legi na alliance (Hu et al., 2000; Hu & Cha ng, 2003); lf'islerla and Afgekia Named after Dr A.F.G. Kerr who collected the specimen on which the type
are nested within Cal/e1ya in the ana lysis of Hu el al. (2002), rende ring the latter species of the genus is based
paraphyletic Lianas; 1 sp. in tropical 1ain forest, the others in seasonally dry vegetation types
Used as ornamentals Reference(s): Lock & Heald 0994: 85 -86); Phan Ke l..Oc & Vidal (2001)
Resembles Cal/e1ya (Geesink, 1984), which is now placed apart from the co re-
Millettieae in the Inve rted Repeat Lacking Clade (IRLC) in the Hologalegina
AntheroporumGagnep. 1915 alliance (Hu et al., 2000); in the latter analysis the single species sampled of
c. 4 spp.; Asia (SW China and Inda-China) Afgekia is p laced with good support in a clade together with two Australian
From anlhera- (L.: a nth er) and poru111- (L.: pore), referring to the anthers, species of Callerya
originall y thought to deh isce by arical pores Used as ornamentals
Trees 0 1 shrubs; seasonally d1y tropical evergreen forest to shrubland, often on
limestone nnd granite hills or along rivers
Reference(s): Wei 0981); Phan Ke Loe & Vidal (2001)
Geesink 0984) considered this genus to be rather isolated in the Millettieae and
perhaps most similar to Cal/e1ya; rotenones are present in the seeds (Allen &
Allen, 1981)

Endosamara R.Geesink 1984

Millellia sect , Bractealae Dunn (1912)
1-2 spp.; Asia (I ndi an subcontinent , Inda-China and Malesia)
Named for the dehiscent pod which separales into rwo valves comprising the
exocarp only; the endocarp forms an envelope around each seed with a flnr wing.
so that each unit resembles a samaroid fruit
Lianas; tropical rain forest to seasonally dry woodland and th icket
llefc:1 ·n.:c( ) ; Geesink 19 ); Phan Ke L<ic & Vid;1l (2001); Hu & Chan~ (2003)
Conslden:<l close to .flllc•1J"' (Geesink, 1984; Schot, 1994), nnw pi:tt'C<l 111>art froin
!he oore-~ Jillettieae in the lm'ertecl Rep~:u I.: ckln1t Clade (IllLC) 111 tlw
Hologalegina all iance (Hu el al., 2000)
Fruits used as a fish poison
Endosamara racemosa Drawing by M. W01wick Afgekia sericea Drawing by P. Halliday

TRIBE MILLETTIEAE 371

370 LEGUMES OF THE WORLD
 Aganope thyrsiflora Photograph by R. Geesink
Austrosteenlsia blackli Drawing by c. Speighl Dalbergiella nyassae Photograph by 8. 0. Schrire

Wisteria Nu tt 1s1s
Dalbergiella Baker f. 1928
3 spr.; 2 spp. in WC Africa (Gu ineo-Congolian region); 1 sp. in SE Zambezian
Kraubnia Steud. (1840); Rebsonitt Srritch (1984)
reg ion
c. 5- 6 spp.; c. 4 spp. i n China and E Asia (Ko1ea and japan), 1- 2 spr in rrom el/us- CL, diminutive fo rm) of Dalbergla. to which it w;is thought to be
ternpern te N America
closely related
Traditionally named after D1 Casrar Wistar 0760 -18 18), rhil an throrisr and active Trees 0 1 sca ndent shnibs; tropical ra in fo rest and seasona lly dry forest (often
prornotor of science at the Unive1si ty of Pennsylvania; rhe ort hographic v:uiant 1iverine), woodland and bushland, often associated wirh rocky outcrops
\Visteria is conserved over rhe spelli ng \'(lisfaria. Anoth e1 exp l~mati on is that the Refe1ence(s} Polhill in Milne-Redhead & Polhi ll (1971, 93 - 95), Lock (1939,
name commemorates Charles Jones WiSler Sr 0782-1865), a dose f1iend of 350 - 351); Lock in Brummiu el al (submiued)
Thomas Nutrnll, the author of the generic name Placed by Hu el al (2000) among a basa l millett ioicl and phaseoloid grour o f
Li ann.s or eiecr shru bs; wmm temperate forest to shrublancl on 1ocky hillsides, and
genera
along 1i ve rbanks
Used as timber, medici ne, ornamentals
Reference(s), Valder (1995)
T his genus is now placed away frorn the core-Milleuieac in the In veitcd Repeat
Lacking Clade (JRLC) in the Hologa legina alliance (Hu er al., 2000)
Used as ornamentCJls (with 1rnmy cu ltivars), bark fibres for· textiles, essen tial oi ls
Aganope Miq . 1s55
Os/l yodeiris Dunn (19 11)
and fragrances
c. 7 spp.; WC Africa (4-5 spp .); Asia CS China , Indian subcontinent, Indo-China,
Malesia and Papuasia, 2 spr.)
Possibly from agcmo- (Gk , mild, pleas,.nt) and -opis (Gk ' appearance), 1efcn-ing
Austrosteenisia R.Geesink 1984
to the pleasant appearance of the plants
4 spp.; E coasLof Aust1alia (also reco1ded from Papua New Gu inea as Verris sp. C Lianas or scandent shrubs to sma ll trees; tropical ra in forest, mangrove or th icket
by Ve1dcour1 1979, 329) Ileference(s} Polhill (1971); Phan Ke Loe & Vidal (2001)
Named after Prof. C,G G.j. van Steen is who stimulated Geesink's ent h usi~1s111 for Placed among a basal mi lleuioicl and rhaseo loid group of gener•
Ma lesia n botany, and australis (L.: south, southern) from ils distribution
Used as ff1edicine
Lianas; tropical ra in forest, seasonally dry forest and vine lhickets, to subtropical
woodland :rnd scrub, including distu1 be<l areas
Reference(s} Dixon 0997) Ostryocarpus Hook.f 1849
Hu el al (2000) places this among a basa l millettioid and phaseoloid g1oup of
1-2 spp.; WC Africa (Guineo-Congolian region)
genera and Kaji ta et al (2001) place it as basally bianching to the Phaseoleae
From osllyo- (Gk ., shell) and cc11pos (Gk.: fruit), refening to the shell-like pods
sens fat clade
Llanas or scandent shrubs; tropical 1ain forest to seasonally dry forest, often
riparian, and mangrove
Reference(s} Dun n (1911)
Leptoderris Dunn 1910
Placed among a basal millettioid and phaseoloid group o f genera
c. 20 spp.; mostly WC Africa (Guineo-Congolian to Lake Victoria regions); 2 spp.
Used as fish poisons and for fish nets ( fib re) and rope
in th e Zambezian and one in the Zanzibar-lnhambane regions
From lepta- (Gk .. : slender, narrow) and derris, referri ng to a distinguishing
combination of narrow pew ls and ca lyces and a frui1 resembling Den·is
Lianas, sometimes shnibs; tropical rain forest ro season:ill)' di")' fo rest, woocl l::incl
and bushland
Reference(s): Dunn (1910); Geesink (1981); Lock 0989, 351-353)
Evans el al. (1985) p1ov ide chemical evidence that Leploderrls may be :1Jnong ;l
basal millettioid and rhaseoloid grou r of genera, a position supported by the
ana lysis o f Kaji ta el al. (2001)
Used as medicine and fish poisons
Leptoderris nobilis Drawing by P. Holliday Ostryocorpus riparius Drawing by P. Halliday

TRIBE MILLETTIEAE 373

372 LEGUMES OF THE WORLD
 Dewevrea bilablata Drawing by P. Halliday Platysepalum inopinatum Drawing by P. Halliday
Xeroderris stuhlmannii Photograph by P. Van Wyk

Xeroderris Robe ny 1954 Dewevrea Micheli 1898

1-2 spp.; WC Africa (Guineo-Congolian region)
1 sp.; Af1ica (Sudanian and Za mbezhlll 1egions)
Named after Alfred Oewevre (1866-1897), Belgian botanist who collected in
From .xero- (Gk.: dry) and derris-, i.e . related to Derris bu t from drie 1 habitats
Co ngo (Kinshasa) and died there of malaria
Trees; tropi cal seasonally dty woodland and bushland
Lianas; tropical rain forest and seconda1y forest, often riverine
Referen ce(s) : Polhill in Milnc-1\edheacl & Polhill (1971: 91-93); Vcn1 c 1 & Venter
Re ference(s): Hauman 0954a)
0996: 280-281); Lock in 13rummitt et al (submitted)
Geesin k (1984) notes that the distinct hypanthium in Deweurea (wh ich is rare in
Placed in the basal millettioid and phaseoloid group of genera (Hu el nl., 2000)
lribe Millettieae) may indicate an affinity with the basal secrion Podocarpae of
and close to Aga11ope (Hu el al, 2002)
Millellia; placed in a basal mille rtioicl and phaseoloid gioup of genera by Kajita
Used as timber, fibre, medicine, food 01ddit ives (ground seed mea l) :ind for
tanning ~ inc.I fodder
er al (2001)
Used as a fish poison, insecticide and for human food (vegetable)

Fordia HemsL 1886 Platysepalum Welw. ex Baker 1871

Millellia sect Albijlorae Dunn 0912); !mbralyx ll.Geesink (1981 )
7 - 8 spp.; mostly WC Africa (Gu ineo-Congolian region), 1 sp in the Zanziba r-
18 spp.; Asia (S China, Inda-China and Malesia)
Named after Charles Ford (1844-1927) supe rintende nt of the Hong Kong Botanic Inhambane region
From platy- (Gk: broad) and sepalum (L sepal), the upper two calyx teetll are
Garden, who discovered a number of new pl:rnts in H ong Kong and Guangzliou,
conna te into a broad cucullate lip as long as the corolla
China
T1ees, shrubs or lianas; tropica l rain forest to seasonally d1y lowland forest, often
Trees or shrubs; tropi ca l min forest (sometimes inundated), often livednc, from
in distu rbed sites
lowland to upland regions
Re ference(s): Gille tt Cl960b); Lock 0989: 365)
Rcferen ce(s): 13uijsen (1988); Dasuki & Schot Cl99ll; Schot (1991)
A genus in need of revision ; Geesi nk (1984) considered Plalysepalum to be
Apparenrly a polyphylelic genus; a basal millettioid and phaseoloicl group
closely related to Millettia, but tentatively placed he1e near Dewevrea
position for Fordia is indicated in the ana lysis of Kajita et a l (2001) fo1 the type
Used for matelia ls (timber, rope, resin), medic ine and as ornamentals
species F. cautijlora Hems!., and also in the phytochrome phylogeny of Lavin el
al, 0998) for F splendid1ssima (l3lu111e ex Miq ,) Buijsen; the latter species,
however, is placed within the core-Mille ttieae in the trnK-matK and ITS analyses
of Hu el al. (2000; 2002). Ford t1 lopm/.lolrys (Dunn) Schot, Dasuki & Uv•jsen ("
Sylvichadsia Du Puy & Labat 1998
Millellia leplobollya Dunn) wa.s J)lacL-cl in fmbralyx (lluijsen, 1988; Sd10t, 1991) 4 spp.; E and N Madagascar
and in the analyses of Hu (2000) and Hu er al. (2000; 2002) this appeaos among From silva/sylva (L,: forest) and C/Jadsia (q .v.), referring to this forest d well ing
the b<1sal millettioicl and phaseoloicl group of genera genus resembling Chadsia, with its similarly beaked orange flowers
Used fo 1 medicine and cu1ing tobacco Trees, shrubs or Jianas; tropi cal rain forest, often near riv e rs or streams
Reference(s): Labat & Ou Puy (1998); Du Puy & Labat in Ou Puy et al (2002:
374-378)
Sylvichadsia may be basal among che millettioid and phaseoloid group of genera
and similarities with Chadsia are interpreted as being due to convergence to a
bird-pollinated syndrome (Labat & Du Puy, 1998)
Used as fish poisons

5Ylvichadsio grondifo/io (A-G), 5. macrophylla (H-L)

Fordio o/bifloro Drawing by/. Wessendorp Drawing by A. /.Beaumont

TRIBE MILLETTIEAE 375

374 LEGUMES OF THE WORLD
 Croibio zimmermonnii Photograph by P. Van Wyk Platycyomus regnellii Photograph by G. P. Lewis Kunstlerio sarowokensis Drawing by J. H. van Os

Schef.flerodendron Ha rms 1901 Platycyamus Benth. 1862

2 spp.; S America ( Brazil , Peru, Bolivia)
4 spp.; WC Af1ica (Guineo-Congolian region), with I sp extending to the
From platy- (Gk.: broad) and cyama- (Gk , bean), probably referring to the
Zanzibar-lnhambane region
Named after Georg Scheffler (fl. 1899 - 1900), collector of the type speci men of laterally compressed seeds
Trees; tropica l rain forest (Amazonian and E coasta l Bra zil)
the genus in the Usambarn Mou ntains, Tanzan ia, and dendron (Gk. : tree)
Trees; tropical 1ain forest and seasonally dry forest, including distu1bed sites Reference(s): Geesink (1984)
Hu el al. (2000) include this among a group of basal phaseoloid genera; the on ly
Refe1ence(s} Leonard & Latour 0950); Lock 0989: 367)
New World genus in the basal millettioid and phaseoloid group
Placed among the basal millettioid and phaseoloid group of genera allied to
Craibia Used as ornamen tals
Scheflleroclendron usa/llbarense H anns (rnsase) is used for timber (constru ction,
handles, poles) , firewood and charcoa l
Kunstf.eria Pra in 1897
8 spp.; Asia (Malesia and Indian subcontinent [I sp ])
Named after H.H Kunstler ( 0. 1882-1886), a German explorer from Australia who
Craibia Hanns & Dunn 1911
collected in the Malay Peninsula for the Roya l Botanic Gardens, Calcutta
JO spp.; Africa (c. 4 spp. WC [Guineo-Congolian] region; 2 spp Zambezian;
Uanasi tropica l lowland (often dipreroca rp) min forest
2 spp Sudanian; 2 spp, Zanzibar-I nhambanc ro Afromontane or Tongaland-
Reference(s), Ridder-Numan & Kornet (1994)
Pondoland)
Ridder-Numan (1996) used Kw1S//eria as an outgroup in a rev ision of Spalbolobus
Named after William Grnnt Craib (1882 - 1933), Assistan t for India at the Royal
in uibe Phaseole~e
13otanic Gmdens 1 Kew and larer Professor of Botany at Aberdeen
T rees or shrubsi tropical rain fo rest to seasonally dry lowland and uphind forest,
wood land and wooded grnssland, often on rocky slopes
Reference(s): Gillett (1960a); Verdcourt in lliummitt el al (submitted)
Burkilliodendron sastry 1969
Placed among the basal millettioid and phaseoloici group of genera (Hu el al., Burkil/ia Ridley (1925); Alloburkillia Whitmore (1969)
2002) 1 sp ; As ia (Malaya [Perak])
Named after JJ'! . Burk ill (1870-1965), Direc tor of the Botanic Ga 1dens, Singapore
Used as ornamentals, shade t1 ees, for timber (furniture and construction) <tnd
charcoal (1912-1925)
Shrubs; tropical lowland dipterocarp rain fo1est
Reference(s): Adema (2000b)
Geesink (1984) considered Bu.rkilliodendron to be transitiona l between tribes
Disy nstemon R.Vig. 1951
Milleuieae and Phaseoleae; rnre with only two collections known
1 sp.; SW Madagascar
From di- (Gk.: two), syn- (Gk , together), slemon- (Gk , stamen); the genelic naine
derives from the anthers being held in two distinct groups, where the upper
fil aments of the stamens are fu~ until near rhe tip.) wh ile the lower rn.unen ts are
extended well beyond the sheath
Lianas; seasonal! · dry tropical woodland
Reference(s} Pehier 0977); D u Puy & Labat in Du Puy el al. (2002: 391-395)
Placed among tl>e basal millettioid and phaseoloid group of genem

Disynstemon poullinioides Photograph by D. Du Puy Burk/lliodendron album Drawing by P. Halliday

376 LEGUMESOFTHEWORLD TRIBE MILLETTIEAE 377

 r-----

0
~
;~

Piscidia grandifolia Photograph by C. E. Hughes Piscidio carthogenensis Photograph by c. E. Hughes

Craspedalobium schochii Drawing by P. Halliday Phi/enoptera vio/oceo Drawing by P. Halliday

Craspedolobium Harms 1921 Piscidia L. 1759

Can izaresia Britton (1920)
I sp.; Asia (Sa nd SW China, Jndo-China) c. 7 spp.; SE USA (Florida), Mexico, C America, Caiibbean, wit h 1 sp. to western S
From craspedo- (Gk., margin, border) and lobion (GL pod); the pods are
A1nc rica (Ecuitdor, Peru, Coloni.bia, Venezuela)
dehisce nt with the valves splitting along the margins (as disringuished f1orn the Frompisces- (L: fish) and caedo (L kill) , the hark and roo ts , w hen placed in
non-dehisce nt pods of Derris)
wa ter are used to stun and ca tch fish
Lianas; seasona lly dry lropic;:1I and subtropical woodlf!nd Trees, rarely shrubs; seasonally chy tropica l ra rest, woodl a nd , 01- bushlancl, ofte n
Referen ce(s), Ying el al (1993, 345- 347); Wei (1994, 189, 191 , 192); Phan Ke L6c
on rocky hills (some restri cted to limesto ne)
0998)
Re ference(s} Rudel (1<)69)
Hu (2000) places this a1Y1ong a basal millettioid and ph;1seoloid g1oup of gcnern
Sousa & Delg"do (1993) note Piscidia aprcars most closely related to
Hespero1ha111nu s; Hu (2000), Kajita et al. (2001) ancl Hu et al . (2002) pla ce
Piscidia sister to the combined Millettia, Tephrosia, Derris a nd Lonchoca1•pus
Phile11optera Fc nzl ex A.Rich , 1847
groups in the core-Millenieae
Lonchoca1pus sect. Paniculali Denth, (1860); Capassa Klotzsch (1861) Used as fish poisons, fodder, medicine (the bark and esse ntial oil o f P e1~ythrina
12 spp.; Africa (4 spp. Za mhezian, 3 spp. Sucfaninn, 2 spp. Zanzibar-Jnhambane to L., the J.im aica dogwood, are used in commercial herbal re medies) and limber
Tongaland-Poncloland, 1 sp~ Somalia-Mf!sa i, l sp. Guineo-Congoli~rn) ~md (construction)
Madagasc<i r·
Origin un cc nain but pbilenlo- (Gk.: tracrab le) and pteron (Gk: wing) coukl refer
co the pod w ing perhaps nssisring dis pe rsal of the fruit Dahlstedtia Ma lme 1905
T1ees, shrubs or more rarely lianas; seaso nally dry tropical fores! , woodland,
2 spp.; S Ame ri ca (lli<tzil, Argentina)
wooded g 1•asslan cl and bush land Named after Gustav Adolf Hugo Dahlstedt (1856- 1934), Sw edish botanist
Reference(s} Schri1e (2000); Lock in Drum 111itt el al. (submitted) Small trees or (sometimes sca ndent) shrubs; t1'0 pical (mostly Atlantic) rain fores t,
Hu el al. (2000; 2002) place Philenop1era in a grour sister to the rest o f the corc-
mainly rive rine, to seasonally d1y forest on rock y slopes
Millcttieae
Re[erence(s} Va nni & Rod1iguez 0999); Teixeira & Ranga (2004)
Used as a dye (indigo), for medicine , human food (preparacions made from
Hu (2000) rl aces Dahlsledfia close to LonchocatpHs subge nus Punctati
flowers, leaves, young shoots and p ods), as goat Fockler, bee plants, in otc'cticides,
Used as a fish poison (roots) and for insecticides
general purpose timbe r, ground wate r indicators, Jnd has various cultuT<il uses

Hesperothamnus llrnndegee 1919

c. 5 spp. ; l\l~ico 1 mainly in west; species have restri cted distributi o ns
From baSfJU/'IJ- (GL w este rn) and thamnos (Gk., s hru b), probably al luding to the
western distribution of chis genus
Small trees 0 1' shrubs; seasonally dry tropical forest to spiny sh1ubland
Reference(s} Rydberg (1924, 235- 237)
Sousa & Delgado (1993) note this genus is one of th e most ·primitive'
represenwtives of th e tribe in Mexi co, wi th Piscidia as its cl nwst re lative; Kajita et
al, (2001 ) place Hesperolhamnus near the base of th e co re-Mill etri c:ie

Hesperothamnus pentaphyllus Photograph by M. Lavin Dahlstedt/a pinnato Photograph by G. P. Lewis

TRIBE MILLETTIEAE 379

378 LEGUMESOFTHEWORLD
Degue/io densifloro Photograph by G. P. Lewis Lonchocorpus obtusus Photograph by G. P. Lewis Behaimia cubensis Drawing by A. J. Beaumont Bergeronio sericeo Photograph by R. M. Harley

Deguelia.Aubl . 1775 Behaimia Griseb. 1866

l.o n ch oca1p 11s sect. Fc1scic11/afi Bcnth. ( 1860); Lo n cbocmpus suhge n 1 sp.; Cuba
Phace/a111b11s l'iltier (1917) 1am ed after Ma lt in Reh aim (1 459 - 1507) , Genna n na vig<iwr a nd gcogrnphe1
c. 17 sp p.; A m~zo ni a n S Am erica to P:i narna mos1ly Dr;ui l (4 spp. in J·'. and SE
1
known fo1 making the earliest extant globe of the world
Brazil) Small 1rces or shrub.<>; .seasonnlly d1y tropical woodlnnd and rh ickct
Abbrevia te d from assa-ha p,;1-gara u11degucla, the nam e fo1 this gi ve n by the Rel'erence(s} llo1hidi el al. (1 978) ; Le wis (1 988)
Galibi in G uya na App;lre ntl y allied to Loncboca1p11s.; 2 spp ~ire described from Cuba but they ~11-e
Lian;1s; 1ropic~ll Ama zonian and Atlancic rain forest, often se~1 sonall y lloocled or now co nsidered co be conspecific
ri veline Used fo1' timber
Refel'cnce(s) , Azevecio·Tozzi (1989)
Inclucles th e Amcrirnn species p1eviou sly placed in Derris; Evans et ol. ( l985)
provid e evid ence of different chemical pro fil es betw een speci es of s r\ 111 c 1icrn Bergerottia Micheli 1883
Derris and Loncbocwpus ll71 nsfcned to Deguelia by Azeved o-TozL:i ( 1989); the l sp .; S AmeriG1 ( Brazil, Boli via, Pa1ag uay and A1 gemina)
number or species cited nl>ove m;1 y thu s be in questio n Named aftel' B Bergeron (A 1880 - 1885), bownical illusuator in Palis
Used as in.secti cides (roteno ncs) T.-ees or shwbs; seasonall y di) ' tropical woodland outcrops in seasonal\)'
inundated gr;.1s.slancl (Pantanal)
Refel'encc(s): Btrl'kort (1952)
Lonchocarpus Kunth. rnz4 Alli ed to l o11cbocm p 11s (Geesink, 1984) ; possib ly rnre and habitat enclange l'ed
Wil/ard ia Rose ( 189 1) ; Tema St"ncil. & I'.]. Herm. (1949) Used <1!':i ornamenta ls
c. 120 spp.; S America (c. 22 spp. Ama zoni:in <lnd c. 20 spp. no n-Am;1zo ni;10);
C Arnc1 it~ . C.1rihbe:rn and Mexico (c. 80 spp.) wirh one ~1111phi -Albn t ic sp.
widespread also in Africn
F1·om lo ncbe- (G k. : lance) and cmpos- (Gk.: frt1il), refe iring w the hrnce-slwpcd
pods
Mosrly t1 ees, less often shrubs; tropical l'<lin fores t ancl sc~1 so n ~11ly clry fo 1 c~t ,
woodland or rocky sllrubland
Refcrence(s} PittiCI' (1 9 17); McVaugh (1987' 555- 577); Aze vedo-Tozz i (1989);
Sousa (1 992)
The concept of Loncb ocwpus as treated here excludes Degue/ia Aubl. , .l/ue!lera .
l..f. and Pbilenoptera Fenzl ex A. Rich .; generic delirnication from rcl~11cd gcner;J in
1he J..onchocnrpus i:roup (.'l(!c p:ijie 381- 382) L< poorly unc.lcrs1ood
V:trio us sp<'..'d t$ used M thnhcr e.g., black c:tblmgc b:ork, mo hidll.', >in1lj>ple), lo•
fu rnitu re, co nstru ction and noodng; al so used as insectic ides (1 o tc11on es), fi sh
poisons, dyes, 01rnrn1en1uls, focJ der, fi bre:: nnd medicine

Lonchocorpus guil/eminionus Photograph by G. P. Lewis Bergeronio sericeo Drawing by B. Bergeron

380 LEGUMES OF THE WORLD TRIBE MILLETTIEAE 381


Muel/era frutescens Drawing {ram Mart. Fl. Brasil
Margaritolobium Harms 1923
1 sp.; S America (Venezuela)
Del'ived fro m Margarita Island in Venezuela w here the genus was first collected,
a nd lobion (Gk.: pod), fo r th e fmi t whi ch di sti nguished chis species apa rt from its
o riginal placement in Gliricidia
Trees or shrubs; seasonall y dry tropi cal wood land
Refere nce(s} Hanns (1923)
Conside red to be allied to Lonchocmpus and Bergeron /a (Geesink, 1984)
Muellera Lr. 1732
Coublandia Aubl. 0775)
2 spp.; both w idespread, o ne fro m Brazil, Surinam, Guyana, Colombi;1 , C America,
Caribbean ~ nd Mexico; th e other from Brazil, Boli via, Paragu ay and ArgentinJ
Named after Otto Friedrich Mueller (1730-1784), Danish Bot,inist
Trees or (so metimes scanclent) shrubs ; nrnthern species in tropical swamp or
riverine fore.st and mangrove; southern species in seasonally city rropica l
shnibland alo ng rive rs and streams
Heference(s): Burkart (1969)
Considered to be closely allied to Loncboca1pus (Geesi nk, 1984)
Used as fish poisons
Derris Lom 1790
Bracbyptemm (Wight & Arn.) 13enth. (1837); Den1s sect. Brachypten1111 (Wight &
Arn .) Benth. (1860)
c. 55 - 60 spp. (see note below); mainl y Asia (S Ch ln:a, Indian subcontinent, Indo~
Chim1 , Malesia and Papuasia); 1 mangrove sp. extending from E Afrirn to Australia
and \Y/ P<tcific; 2 frnther sp p . in Australia
from derris (Gk.: skin, leather coat), 1eferring to the me mbranous pod
Mainly li ~n<1s, few trees a nd shrnbs; tropical rain fores t to seasonally dry forest,
bushland o r thi cket, ofle n swampy or ri verine
Reference(s): Verdcou1t (1979); Thothathri (1982); Phan Ke Loe & Vid:i l (2001);
Adema (2000a, 2003c)
Derris rob11s1a (Roxb. ex DC.) 13enth. (• Brachypterum robustum (Hoxb. ex QC.)
Dalzell & A.G ibson) is in a basally branching position w ell remov ed from the
Derris clade in. the analysis o f Hu et al. (200 2), but further sampling is needed to
reassess this a nd whether Brachypterum should be uphe ld as a distinct genus;
esrimates o f species number range from about 40 (Gee.si nk, 1984) to nea rl y 70
(Lock & Hea ld, 1994); the current estimate is bosed o n so me species be ing
transferred to Paraderris, Aganope <ind Deguelia
Used as co mmercial insect icides, fish roisons, medicine, green 111am11 e, shade
plants and for timber
Derris trifo/iata Photograph by D. Du Puy
382 LEGUMES OFTHE WORLD
Paraderris el/iptica Drawing by H. E. Ireland Millettia aurea Photograph by D. Du Puy
Paraderris (Miq.) R.Geesin k 1984
Derris Lour, sect Paraderris Miq. (1855)
c.. 13 s pp.; Asia (S China, Indian subcontinem, Indo-China, Malesia and P~tpuasia)
From para (Gk,.: near, comp~u ed with) and Derris (q ,v~), 1eferring to its close
proximity to Den1s
Lianas; tropica l rn in forest to .seasonally chy fores t; often marginal or rive rine
l\e ference(s): Aclema (2003a)
Distinguished from Derris by the hairy anthers and by its chemical p rofile (Evans
et al., 1985)
Petraderris elliptica (\Vall.) Ade ma is widely used for c:o rnme rci<1 l in.secri cide.s;
ro tenones ;:ire ex tracted largely f1'om the 10ots <1nd sold as e.g ., derris powder
Millettia Wight & Arn . 1834
Pongamia Vent. (1803); Lonchocmpus sect. Ca udaria Dunn 0911); Neodunnia
R.Vig. 0950)
c 150 spp. (see taxonomic notes); c. 90- 100 sp p. in Africa and Madagascar (in
Africa c . 65 spp. occu <' in WC regions; 14 spp. Za nzibar·Inhambane; 6 spp.
Za mbezian; c. 4 spp . Somalia-Masa i; c 2 spp. Sudanian; c. 1 sp. Afromo ntane;
8 spp. endemic to Madagasca r); to China , the Incli<m subcontinent, Indo-China
and Malesia
Namecl after Charles Millett ( fl 1825-1834), of Canton, China; in the se<v ice of the
British East lndia Company and a collector o f Chinese pl'1nts
Trees, shrubs o r lianas; tropical rai n forest a nd se<1 sonally dry lowland to upland
Farese and fores r margins, woocll<ind, thi cket and wooded grnssland, and
secondary vegetation types
Refe1ence(s): Dunn 0912); G illett in Milne-Redhead & Po lhill (1971: 122- 146);
Labat & Du Puy (1995); Adema (2000c; 2001); Phan Ke Loe & Vidal (2001);
Du Pu y & La ba t in Du Puy el al. (2002: 379 - 387); Verdcou1t in Hrummitt et al.
(submitted)
This genus is in need of major revision since recent mo lecular evidence (Hu et al .,
2000; Kajita et al., 2001; Hu et al., 2002) has shown Milletlia to be poly phyletic;
estimates of rhe nu mber of species vary from 90-200 and the rotal give n above is
based on a review of current lite1atu re and he rbariurn collections
Used as fish poiso ns, insecticides, timbei (e.g., M laurenlii De Wilcl . [wenge] ; M
sluh/mannii Tmib. [panga pang<i, mpande], fo r flooring, furn iture, cabine t work,
co nstr1.1ction, veneers, joine1y ancJ agricultu1 al implements)i as nitroge n fixe rs in
rigroforestiy (so il rehabilitation), fuelwood , o rn.a1nentals and for medicine
Millettia nathaliae Photograph by D. Du Puv
TRIBE MILLETTIEAE 383
 Pangamiapsis amygdallna Photograph by D. Du Puy Chadslo grevei var. latifolla Photograph by D. Du Puy Chadsia versicalar Photograph by G. P. Lewis

Pongamiopsis R Vig. 1950 Chadsia Bojer 1842

3 spp.; N, W and S Madagascar 9 spp; N, W and S Madagascar
Ponsamia (from the Malabar na me pongam) and -apsis (Gk : resemblance), Named after Sir Henry Chads (1819 - 19o6), later Admira l, Roya l Navy, w ho in
referring to it.s similarity to Ponsamia Vent (now a synonym of MiUellia) 1835 offered the botanist W Bojer a passage to Madagasca r, as a result of wh ich
Trees or shrubs; seasonally dry tropical woodland and xerophytic shnibland; often many new plants became known to sdence
associated with limestone outcrops Shrnbs or small trees; seasonall y dry tropical forest , woodland and xerophytic
Reference(s): Labat & Du Puy (1995); Du Puy & Labat in Du Puy et al (2002: shrubland. often on limestone
387-391) Reference(s): Du Puy & Labat in Du Puy et al. (2002: 431 - 443)
Closely allied to Millettta; some species endangered or vu lnerable Closely related to Mundulea and Tepbrosia (Hu, 2000; Hu et al., 2002)
Used for timber (construction, planks), firewood and as fish poisons Used as fish poisons, timber (const ru ction), fodder and medi cine

Pyranthus Du Puy & Labat 1995 Mundulea (DC.) Benth 1852

6 spp.; W, S and C Madagascar Tepbrosia sect. Mundulea DC. (1825)
From pyr- (Gk : fire) and antbos (Gk,: flower), referring to the species' strong 12 spp.; Madagascar, 1 widespread sp. (M sericea (Willd .) A. Chev.) also in Africa,
resisrance to grassland fires, the stock resprouting after bu rning, and also the lnd ia and Sri Lanka
scarlet or 1ed flower colou r From mundu/us- diminu tive of mundus CL.: pure), the pistil is mostly smooth
Shnibs or small trees; seasonally dry tropical woodland and grassland, on dunes Shrubs or trees; seasonally dry 1rop ical fo rest (sometimes in mo re hu m id
or rocky outcrops evergreen fo rest), wood land, grassland, bushl and, th icket and xerop hyt ic
Reference(s): Du Puy & Labat 0995); Du Puy & Labat in Du Puy et al. (2002: shrubland , in sand and on rocky ou tcrops
374 - 378) Reference(s): Vente r & Vente r (1996: 72- 73); Du Puy & Laba t in Du Pu y et al
Appears to be somewhat intermediate between Mu.ndulet1 and Cbadsia (D u Puy (2002: 416-425); Verdcourt in Brummitt el al. (subm itted)
& Labat, 1995) M 11ndu/ea is veoy closely related to Tephrositl and it is doubtfu l whether it should
Used as fish poisons, medicine, cosmetics and timbel' (construction) be ma inta ined as a d istinct genus (Du Puy & Labat in Du Puy el al, 2002)
Used as fish poisons, dye p lants, medi ci ne and ornamentals

Pyranthus tutlearensls subsp. tul/earensls Photograph by D. Du Puy lttundu/ea stenaphylla Photograph by D. Du Puy

384 LEGUMES OF THE WORLD TRIBE MILLETTIEAE 385

 PG0 1•
Tephrosia phylloxylon Photograph by D. Du Puy Poratephrosia lanata Drawing by P. Halliday Requienia sphaerospermo Drawing by H. Wood

Tephrosia Pers. 1807 P~ratephrosia Domin 1912

Cracca L. (1753); Caulocatpus Baker f. (1926); Lupinopbyllum Hutch. (1967) I sp.; Australia (W Australia, N Territ.ory, S Australia, Queensland)
c. 350 spp.; pantropical, concentrated in C and tropical N America (c 45 spp.), From para- (Gk.: near) and Tepbrosia, indicating the close relationship between
Africa-Madagascar (c. 170 spp.), Asia (c. 40 spp.) and Australia (c. 90 spp.) tl)ese genera .
From tephro- (Gk.: ashen) after the grey-green leaves of many species Shrubs; seasonally dry tropical to warm temperate wooded grassland and open
Shrubs or herbs; seasonally dry tropical wood land, bushland, thicket and sandy or stony scrub
grassland, often in open and disturbed sandy or rocky areas Reference(s): Wheeler et al. 0992; 422-423)

0
Reference(s): Wood (1949); Brummitt (1968a); Gilleu in Milne-Redhead & Polhill Wheeler et al. 0992) note that this monotypic genus is very close to and perhaps
(1971: 157-21 1); Bosman & De Haas (1983); Tellez (1986); Lock (1989: 367-386l; not really distinct from Tepbrosia; Geesink (1984) placed it in synonymy under
Du Puy & Labat in Du Puy et al, (2002: 395-416); Brummitt in Brummitt et al. Tephrosia, but Paratephrosia is still widely maintained in Australia
(submitted) J
A taxonomically complex genus; Pedley (pers comm) notes that c. 12 species

~
from N Australia, characterised by leaves with reticulate veins, could be Requienia De. 1825 ,
segregated as a separate genus Tepbrosia sect. Requienia (DC) Be11th. (1865)
Used as fish poisons, cover crops, livestock fodder, insecticides, ornamentals and 3 spp.; W to NE Africa (Sahelian zone); southern Africa (S Zambezian and
for medicine, e,g T. virginiana (L.) Pers. (goats rue, devil's shoestring, hoary Kalahari.Highveld regions) ,
pea), T. vogelii HookJ. (fish-poison pea), T.pu.purea (L.) Pers. and T. candida Named after Esprit Requien (1788 - 1851), French botanist at Avignon
(Roxb.) DC. (white tephrosia) Herbs; seasonally dry tropical wooded grassland, shrubland and bushland, on
sand
G Reference(s): Brummitt (1989); Brummitt in Brummitt et al. (subm itted)
Apurimacia Ha1·ms 1923 Further evidence may place this as a lineage within Tephrosia
c. 2 spp.; S America (Andean Peru, Bolivia and Argentina)
Named after the Apurimac region in Andean Peru where the type species occurs
Shrubs; seasonally dry tropical montane shrubland on rocky slopes Ptychol.obiumHarms 1915
Reference(s): Macbride (1943); Burkart 0951); Lavin & Sousa (1995) Sylttra E. Mey. (1836)
In the ana lysis of Kajita et al. (2001), the genus (based on the exemplar species 3 spp.; NE Africa and Arabia (Somalia-Masai region); southern Africa (S
A. micbelii (Rusby) Harms [correct name - A. boltviana (B1itton) Lavin]) is sister Zambezian and Kalahari-Highveld regions)
to the Tephrosia group; the other species, A do/icboca.pa (G riseb.) Burka1t may Fromptycbo- (Gk: folds) and lobion (Gk.: pod), referring to the folded pods
have an affinity elsewhere characteristic of the genus
Used as fish poisons and insecticides Shrubs or herbs; seasonally dry tropical woodland, wooded grassland and
shrubland, on sand
Reference(s): Biummitt ( 1980); Brummitt in Brummitt el al (submitted)
Further evidence may place this as a lineage within Tephrosia

Apurimacia michelii Drawing by P. Halliday P'tycha/obium contortum (A- J) Drawing by H. Wood

386 LEGUMESOFTHEWORLD TRIBE MILLETIIEAE 387

TRIBE
j\breae by B. D. schrire
'fribe Abreae (Wight & Arn. ex End!.) Hutch. 1964
Tribe Phaseoleae subtribe Abrinae Wight & Am. ex End!. (1840), as 'Abrineae'
'fhe tri ba l pla m nt of the i lated genu Abru.
(with 17 pe ie ) ha long b en pr bl roa cic (P lb ill
198le: 243- 244). Th most recent mole ular ana ly s
of Doyle et al., 2000; Hu, 2000; Hu et al., 2000; Kajita
eta!., 2001; Wojciechowski, 2003 & Wojciechowski et
al., 2004), however, consistently place Abrus near
the base of the clade comprising the core-Millettieae
and various elements excluded from Phaseoleae sens.
/at. (Figs. 45-47). This basally branching position
together with the unusual combination of paripinnate
le aves, 9 stamens and a chemical profile of
tryptophane-derived alkaloids (abrine and
hypaphorine) and pyridine-based alkaloids
(pre ca torine and trigonellin e), supports the
- Abrus
-
FIG. 46 Diagram ofrelationships oftrlbeAbreae after Doyle
LEFT Abrus precatorius Photograph by G. P. Lewis
maintenance of Abrus as a separate monogeneric
tribe for the present. Improved understanding of
relationships is likely to result from greater sampling
within the millettioid-phaseoloid complex.
The Abreae are predominantly Afro-Madagascan in
distribution with 3 species endemic to the Horn of NE
Africa. The remaining species are Asian and 2 species
are pantropical. New World representatives (from
Amazonian Brazil and Venezuela) of the pantropical A .
melanospermus Hassk. subsp. tenuiflorus (Benth.) D.
Harder [= A. pulchellus Wall. ex Thwaites subsp.
tenuiflorus (Benth.) Verde.]), are possibly distinct from
Old World material of this subspecies (Verdcourt,
1970b).
lndigofereae (see page 361)
Basal millettioid and phaseoloid
group (see page 369)
ABREAE
Phaseoleae subtribe Diocleinae
(see page 394)
Phaseoleae subtribe Erythrininae
(in small part only, e.g. Rhodopis)
Phaseoleae subtribe
LJ Ophrestiinae (see page 394)
core-Millettieae (see page 369)
Phaseoleae sens. lat. (see page 394),
Desmodieae (see page 433),
Psoraleeae (see page 447)
eta/. (2000); Hu (2000); Hu eta/. (2000); Kajita eta/. (2001); Wojciechowski eta/. (2004)
TRI BE ABREAE 389
Abrus conescens DfOw/ng by E. M. Stones Abrus precotorius
i •. , ' ,'I'
'
Photograph
,, , 'I
by G. P. Lewis

Abrus Adans. 1763

c. 17 spp.; Africa (~. 8 spp.), Madagascar (5 spp ), India (1 sp.) and !ndo-China (1
sp.), 2 spp. 'widespread in the 'old'Wol'ld (both also occur__:_ probably introduce~
~
1

- in the New' World)

Fro1i1 habro- (Gk.: delicate, elegant, pretty, soft), referring to the delicate Foliage
and pretty flow~rs '
Cli1l1blng subs~1rubs (sometimes lianas) or herbs; seasonally dr)i tropical forest,
woodland, wooded grassland, grasslind, shrublancl, bushland ancl thicket, often in
Open'disturbed vegetation, al'on·g mafglns and in seasonally wet or in rocky area.S
Reference(s), Breteler (1960; 1994a); Verdcourt (1970b: 235 - 253); Labat (1991);
Thulin 0993: 393 ..! 394); Du Puy & Labat in Du Puy et al (2002: 363 - 370);
Ha'rder'in' Bmmi11itt et al. (sllbtrlltted)
sb1i1e spe~ies are diffi'cult to clas'Sify and species limits have been much discusse~,
1

with Verclcowt (1970b) and Breteler (1960; 1994a) putting opposing opinions; the 1
classification Of Verdcourt is follmVe<l here ·
The type species A pre'catorius'L is 'corilmonly known as false or Indian
liquorite, crab's-eye, jequirity Or prayer bead; 'the attractive shiny bicolou re<l
seeds (red and blac~) are commonly used in necklaces a'nd ~ostume jewellery,
but are highly 'pdiscin'ous and can be lethal if ingested; other uses are as '
lnediCine f6IJ a wide rahge of condition.-! (medical supe rvis io n essential); for fibre;
green 'manure; 01'n'amCrifol:s; a swc~tcncr for diinks :111d 'he root can bC a ''
substitute for liquorice (althoUgh be-Ware of toxic' propeLties); also has various
cultural 'and spiritual uses

Abrus oureus subsp. aureus Pl10109rnpl1 by D. Du Puy Abrus precatorius Photograph by o. Du Puy
I '11 I I;,''

TRll3!0 ABREA!O 391

TRIBE

phaseoleae byB. n. schrire

fribe Phaseoleae (Bronn) DC. 1825

subtribe Phaseolinae Bronn (1822), as 'Phaseoleae'
Tribe Elythrineae (Benth.) Hassk. (1844)
Tribe Glycineae Burnett (1835)

previous accounts of the Phaseoleae by Baudet 0978)
and Lackey 0981) recognised 90 and 84 genera and c.
1540 and 1480 species respectively in the tribe. In an
equivalent, i.e. traditionally held view of Phaseoleae, 89
genera and 0554)-1 567-(1580 species are treated
here (Table 9; Fig. 47). Changes between Baudet 0978)
and this treatment are that eleven genera are now in
synonymy or have subsequently been placed in
Millettieae, two genera have been transferred from
Desmodieae and eight new genera have been added.
Vigna has traditionally been thought to comprise some
150-200 species, but Vigna sens. strict. may contain
fewer than 100.
Recent molecular analyses of the tribe, however,
have emphasised both the polyphyletic and
paraphyletic nature of Phaseoleae as traditionally

TABLE 9 Suprageneric groups, both formal and informal, currently recognised within the traditional concept of tribe Phaseoleae

INFORMAL GROUPS NUMBER OF NUMBER OF DISTRIBUTION

GENERA SPECIES

A Millettioid groups 16 (212)-216-(219) Throughout the tropics and subtropics, to warm temperate (in
(see also page 369) the N Hemisphere)

1) Basal millettioid and phaseoloid See page 369

group

2) Phaseoleae subtribe Diocleinae 13 c. 194 Mostly Neotropics and subtropics [c. 151 spp.], Palaeotropics
[c. 43 spp. - 2 genera endemic to Malesia & Papuasia]

3) Phaseoleae subtribe Ophrestiinae 3 c.22 Africa and Madagascar [17 spp.], Asia [5 spp.]

4) Tribe Abreae See page 389

B Phaseoleae sens. lat. clade 73 (1342)-1352-(1361) Throughout the tropics and subtropics

5) Phaseoleae subtribe Clitoriinae 5 c. 106 Mostly Neotropics and subtropics [c. 91 spp.]; Palaeotropics
[c. 15 spp.]

I) Phaseoleae·Desmodieae group

6) Basal Phaseoleae sens. lat. c. 9 c. 140 Mainly warm temperate to montane and lowland tropical Asia and
(ex Erythrininae) and Kennedinae Australia (c. 14 spp. of Mucuna in the Neotropics)

7) Tribe Desmodieae See page 433

ii) Core·Phaseoleae
(Phaseoleae·Psoraleeae group)

8) Basal core-Phaseoleae 8 c. 175 Palaeotropics and subtropics [c. 105 spp.]; Neotropics and
subtropics [70 spp. of Erythrina]

9) Phaseoleae subtribe Cajaninae 10 c. 495 Mostly Palaeotropics to warm temperate Old World [c. 400 spp.],
Neotropics and subtropics [c. 95 spp. in 2 genera]

10) Phaseoleae subtribe Glycininae 20 c. 122 Mostly Old World [c. 96 spp.]; Neotropics and subtropics
[c. 26 spp. principally in 3 genera]

ll)Tribe Psoraleeae See page 447

12) Phaseoleae subtribe Phaseolinae 21 c. 314 Mostly Palaeotropics and subtropics [c. 196 spp.]; Neotropics
and subtropics [c. 118 spp., principally in 8 genera]

LE FT Clitoria ternatea Photograph by G. D. Carr

TRIBE PHASEOLEAE 393

 circumscribed (Bruneau & Doyle, 1990; Doyle &
Doyle, 1993; Delgado Salinas et al., 1993; Bruneau et
al., 1995; Doyle et al., 1997, 2000; Kajita et al., 2001;
Goel et al., 2001; Lee & Hymowitz, 2001). This has
required a radical realignment of elements of the
phaseoloicls (Table 9; Fig. 47), with at least two major
clades being evident: Phaseoleae subtribes Diocleinae
and Ophrestiinae which together with tribe Abreae
are allied to the core-Millettieae (Fig. 45), and the
remaining groups comprising a Phaseoleae sens. lat.
clade. The rhcL phylogeny of Kajita et al. (2001) and
the iT · analysis o f Hu et al. (2002) are e qu iv cal,
to w h ic h cl a d e s ubtrib e Cli o ri inae b J J on~·ls
Pha eol ae sens. la t. also includes two m1ditiona1t ·
ind pend nr tribe , rh ·modi ae a nd P ora le ~,:
Delimiting a re ir ~um cribed Pba. eolea ·e11 . str;c;
Lhus v ry pr bl rn a ti c . o lu ti n may b l
r - g ni. a b roa d trlb Pba e · lea , com pris ing u ~ 1
ubtrib Ke nne diina Caj·minae, Phas oli oae aild
Glycinina , as orred ba ally bra nching ge nera, and
tribes Des mo dieae and Psora l eae both treated ai:.
subtribal level).

Abreae (see page 389)

I ' VI;' I • I •uo:..
SuBtrioe E>1o·demae

sJbtHbe cii)hrestilha~ c~~ciciasia, ophrestia, f>seuci~eriosema

Core- Millettieae
(see page 369)

B~s,al m\e.ttiqjq ,a~d

phaseeloid group

s'Jbfrib~ ctitoH\~ae C!ltqria,, .13~~bieria; C:e~tr~ se~a. Peria~ dra,

Clitoriopsis

A~,iq~. rosh'tl~~th~s. Sh.~teria, ?Mastersia, Dioc/ea sp. aff. grandifloro Photograph by G. P. Lewis Luzania purpurea Drawing by H. E. Ireland

.'
Desmodieae (see page 433)
?Diphyllarium
Dioclea Ku nth 1824
c. 10 spp .; S America (c. 35 + spp . lrnostly Il1azil with a few extencling to the
Caribbean, C Arnerica and Mexico], c. 4-5 spp . also occuning widely in the
Palaeotropics); Asia (c. 3-4 spp.)
Named for the Greek physician Diocles of Carystos (240-180 BC), who lived
shortly after Hippocrates and was famous for • his knowledge of plants
Lianas or shrubs; tropical lowl<md rain forest, 1iverine or sw<1mp fo1est ;:rncl thicket;
seasonally d1y tropie<1l to subtropical woodland, wooded grassland and scrnb
I 'ii l l'. ,i'l t, i ' «'' i I •
Refe1ence(s): Maxwell (1969); JVbxwell & Taylor (2003); Queiroz el al (2003)
Spatholobus, Butea, Meizotropis Maxwell (in cor.-esponclcncc at the Kew I-Ierba1·iurn) considerecl the genus to
comprise some 50 species. A difficult genus because of proble111.s of species
delimitation within Dioclea, and poorly delimited from neighbou1•ing genera,
Ad,~r~d~licn9"1?~.Pa .~l'alv;$ .. so1usafra, e.g. Cra(ylia. A global ieview of the genus is neces.sa1y; the tribe Diocleinac
sJbtribe C:aiah\hae Qarrl~sea~ Ch ~!iPSG!as, RhY.TJehosia, Erlosema, together with Oph1estiinae and the genus Rhodopis (originally in the
E1ythrininae) are placed in a Millettioicl sens. strict . group, somewhat 1emovecl
Dunbaria, Calanll!!, ,ftemi11gia f1orn the Phaseoleae sens. la/ . (Bruneau el al,, 1995, Doyle el al., 2000), Maxwell
& T;1ylor (2003) ancl Queiroz el cil. (2003) find Dioclea to be polyphyletic,
E~~thrl~a. Psophocar~us, Dys~lobium, Dioc/ea sens strict. is mo1e closely allied, among others, to Cymbosema ancJ the
Bionia clade of Camplosema (Queiroz et al, 2003), while the Pachylobium clade
Otoptera, Decorsea
of Dioc/ea is siste1· to a g1'oup of genera that includes these elements
Common d1i ft seeds; used as ornamentals, medicine and insecticides

Luzonia Elmer 1907

1 sp.; Philippines (Luzon ancl leyte)
Named after tbe largest of the Philippine islands where the type species, L.
pwpurea Elmer, was collected
Subtribe Glycinfnae Lianas or scandent sh1ubs; t1'opical 1<iin fo1est and thicket

, i
I
Reference(s): Elmer (1907); Adema 0998); Maxwell & Taylor (2003)
Elmer 0907) excluded L pwpw"ea f1om Dioc/ea or Pueraria and placed it

I I
I
nearest to Canavalia; Luzania is unknown in fruit but flower clissections (Lewis,
pers. comm ., 2001) suggest that Elmer's conclusion was incorrect and that L.
p1t1purea would be better accommodated within Dioc/e{{; see 8lso note under
I Psoraleeae (see page 447) Nfacropsycha11th1-1s
WaJira .Sp~~postvh. N~sphoslyjls, Alistilus,
~_H~rqpol)cJ1R, r De,ll~hlj>S' Ma,cfotyto.ooa, Macropsychanthus Harms 1900
subtribe Phase~Unae @iJ?,ogori, ablf3b, Spathioqe.mi!, Vatovaea,
c. 2 spp.; Papuasia, Micronesia (possibly Philippines)
Phy~asOgm~ ~ Vigna, ,Q"-yr~Yflt1'.l,\s. From macro- (Gk.: large), p~vche- (Gk.: butter·tly) and arilhos (Gk .: tlower) for the
R~~seolus, R~nif ~ella, Strophostyles, . . ch£1racte1istically large papilionoid flowers
bollehopsis, MacroptiOum .. Mysanthus, oryxis Licinas; tropical rain forest, often along rive1s
? =placement uncertain in generic group Reference(s); Verdcourt (1979: 466-470); Adema (1998); Maxwell & Taylm (2003)
One species has 4 varieties (Adema, 1998); Maxwell & Taylor (2003) fincl Luzania
FIG. 47 Diagram of relationships between phaseoloid groups after Kajita et al. (2001); Hu et al. (2002); Wojciechowski et al. (2004) Nlacropsychanthus lauterbachii Drawing by G. Bartusch and 1Hacropsychan1/Jus fo r m a clade embeclclccl within Dioc/ea

394 LEGUMES OF THE WORLD TRIBE PHASEOLEAE 395

Canava/ia villasa Photograph by G. P. Lewis Canavalia madagascariensis Photograph by D. Du Puy
Canavalia Dc. 1825
Wenderothia Schltdl. (1838)
c, 60 spp.; c. 33 spp. Neotropics (c. 18 spp. endemic in S America; 10 spp. in N &
C America <ind c 5 neotropical-wide); c. 10 spp~ in Pacific Is,; 8 spp. in Asia and 1
sp. in Madagasca1; c. 5 spp. widespread in Palaeotropics or pantropical
Derived from the vernacular name kanavali in Malaba1', lndia for Canaualia rosea
(Sw.) DC ,
Li£1nas to slender vines; tropical flooded or swamp forest, river banks and fo1est
margins to seasonally dry coastal vegetation, thicket, open woodl<md, wooded
grassland and rocky hillsides, often climbing over or trailing through othe r
vegetation
Reference(s), Sauer 0964); St John 0970); Adema 0997); Polhill in Mackinder el
al. (2001: 36-40); Maxwell & Taylor (2003); Queiroz et al (2003)
Many more species than me currently 1ecognised were described from Hawaii by
St. John 0970); Queiroz el al. (2003) find that Canavalia, Cleobu/ia ancl Cmtylia
form a chide
Used for hurnan food, cove1 crops 8nd green manures, c.g~, C. gladiala Qiicq,)
DC. (sword bean) and C. ensi[ormis (L.) DC (jack bean); also as ornamentals,
medicine, hu1rnm food (veget<ible) and forage; common drift seeds
Canava/ia rasea Photograph by J. Murata
396 LEGUMESOFTHEWORLD
Cymbosema roseum Drawing by C. J. Seliger C/eabulio multiflora Drawing by P. Halliday
Cymbosema Benth. 1840
1 sp.; Amazonian Brazil, Colombia, Peru, Venezuela; C America; Mexico
From cymbe (Gk,, boat) and semeia- (Gk.: standard), for the shape of the
standard petal
Vines or lianas; tropical lowland rain forest and dverine forest
Reference(s): Maxwell (1970); Zamora (2000); Maxwell & Taylor (2003); Queiroz
et al. (2003)
See note under Dioclea; Zamora (2000) considers this genus to be synonymous
with Dioclea
Cf.eobulia Mart. ex Benth. 1837
3-5 spp.; Brazil, Mexico
Named after the Greek sage Cleobulus f1om Lindos on Rhodos who died in 560 BC
Vines or lianas, or shrubs; tropical Amazonian rain forest, seasonally dry forest,
oak and pine fo rest
Reference(s): Maxwell 0977; 1982); Maxwell & Taylor (2003); Queiroz et al, (2003)
A 1eview of the generic limits between Dioc/ea and Cleobulia is needed to
establish if the latter is a natural genus; sister to Canava/ia in the analysis of
Queiroz et al (2003); the name Cleobulia Ma1t, ex 13enth. requires conserving
over the earlier homonym Cleobula Veil. (1825)
Camptosema Hook. & Arn . 1833
Bionia Mart, ex Benth. (1837)
10 spp.; two centres in tropical and subtropical S America (EC Brazil; type species
from S Brazil, Argentin<i, Paraguay and Uruguay)
From campto- (Gk.: curved) and semeia (standard), after tl1e reflexed standard of
the type species; all other species have a straight standard
Li8n<is, vines or shntbs; tropical seasonally dry to wet forest or upland woodl<ind,
wooded grassland (cerrado) to rocky shrubland or grassland (caatinga)
Reference(s): Queiroz (1999); Queiroz et al. (2003)
Generic delimitation from Galactia is problematic fo l' the type species,
C. rubicundum Hook. & Arn.; the genus is polyphyletic in the analysis of Queiroz
et al (2003), with Camptosema sens strict. related to Calactia, the Bionia clade to
Cymbosema and Dioclea sens strict ,, and two fu1the r species to the Cratylia clacle
Used as ornamentals
Camptosema coriaceum Photograph by G. P. Lewis
TRIBE PHASEOLEAE 397
 G.a/ac~la jussiaeana Photograph by G. P. Lewis Lackeya multiflora Drawing by E. Catherine Rhadopis planisiliqua Drawing by P. Halliday
I ·

Cratyliq, Ma rt ex Benth 1837 Lackeya Fortunato, LP.Q uei1oz & G l'.Lewis 1996
<". 7 ~pp. : Brazil.' Bolivia, Peru, N Argentina l sp.: SE USA
::~:: cm10: (Gk.! s~rong} anil cau(os- (G~.: ste~n), the plants have firm woody Named afte1•James A Lackey, a specialist on the family Leguminosae who has
published wid ely on the tribe Phaseoleae <tnd added much 10 our know ledge of
tian~ or <!1ct1 to .c1ndcm •hrul : N:~snn.1lly <lo; • 10 wet tropica l forest and forest the group
r1111rg1n,, woodland, thorn thicket, scrub ~n<l woo<it.·d w:is>lnnd Climbing herbs; warm temperate to subtropical ri verine wood land, wood land
llclimm ,( ): Qwiro~ ( 1991); Q11droi <!I al. 2003) ma rgin!'i a nd grassland
:c not • und.,r D/oclen 1111d Cm11fl/<).w11w for relationships Hcference(s): Fortunato el al. (1996); Quei roz el al (2003)
See nme un<ler G'alaclfa for relationships

Galactia P.Browne 1756

55- 60 •m>.; . ~O . pp. In • ,\mcri . ; c Ul - 2S sp1>. in • .~ C 111<:1'1 l ; Alnci (2 Rhodopis Urb. 1900
>pp.), trbplcal >i:t It) Chln.1 :Intl J:tJY.ln (c. (\ •11p.) and Aus1r.1ha {c. 2 ·pp.J: 1- 2 2 sp p.: C""ibbean (Dominican lle pu bl ic, Haiti)
'PP· p:1n1 picr1I From 1bodopv- (G k .: rosy: rosy-faced), re ferring to the colou r of the nowers
F Ill lffllttcl/J- (Ck.: milk), rcforring w the milk)• S<lp in wmc SJ>'-'C!es Perennial climbing herbs; tropical C0<1St a l and riverine shmbland to montane
l>uhshni~ or f>t:n:nnlal hc1bs: =i:son1tlly dry iropit"I nnd s11h1ropirnl fo 1 fo rcsr cleat ings and margins
ma<Wns, 1hlcko!l. woodl:md. wooded wassbnd. llmSSl3MI and ro«k)' shn1bL111d Reference(s): J udd (1984)
ltcl'crcncc:(s>: llurkan ( 1971): \\:rdrou11 In Mil1K~lk'(]Jl('lld & l'olhill 0971 : 5711-581); Comments as for Ne01udolpbia 13ritton
TOITC.' Ill<//. ( 198~): Queh ,, I!/ {(I ('.lOOj)
Clalactln fonns :t 11mup in the Dlotlclnnc 10$.'Cthcr wi1h 11111/l/IJM!lllll, lnd.•:1t1 :u><I
Collm·lt (Quciroz c•1 111., 2003), nnd Mt"~" ·II & 1"nyl r (2003) lndudt: RlxxlvplS in Neorudolphia Britton 1924
th~ir Ga1"ct1:1 chute, thn.-e sections •t'C r<,'t:ognl~'ll tn Gnl11c1ln: Odrmla, Coll<t1.irl11 Rudo/pbla Willd. (1801)
:Ind G11l11clf11.
1 sp ; Caribbean (Puerto Ri co)
Os~d as w ild life forage Cmilkpeas often host butterflies) Named afte 1 Karl Asmund Rud o lphi (1771 - 1832), Swedish-born German botanist
I ··I', : ,"
Perennial climbing herbs; seasonally dry tropical fo rest margins, wet woodland
and thi cket
Collaea oc. 1s25 Refere nce(s): Britton & Wilso n (1 924); Maxwe ll & Taylo1 (2003)
'Pt>.: $ Amc1'1c:t (l'cni, llr:till. llolMn, P:ir.1guay to N :111d C Ari:cntln;i l Lackey (1981) and Bruneau et al. (1995) co nside red this genus close to a nd
:1111ctl for l.ulgl Aloy:>lu> .olln (J766- 18 It! , ltalfan In\\ >er ntl lxx:tni.>l. owner of possibly congeneric w ith Rhodopis, and the latter authors place the rwo genera as
:i 1>01anfrnl garden 111 ltirnll n<>:1r 'rorino sister to the core-Millettieae, with dose affinities to the main group of Diocleinae
Subshrul>s; se:ison:tll)• <.lry tropirnl m1<.I sublropla I lowl:intl 10 11t0ntanc ,h rubla n~ geneoa: Kaj ita et al. (2001) a nd Maxwell & Taylo r (2003) place Rbodopis nea r to
•UJd gmssland ·' : ·' Calaclla, however Maxwell & Taylor (I.e .) suggest a relatio nship between
Refcrcnc""(s): Ourkon ( 1952). l'onunnto (1995): Quelroz ell fll. (.!003) Neonidolpbia and Hetpyza in subtribe Glycininae
n1 . f.'COcric circtllllS<Tl1xlnn of Collmw b thal :1dap1cd mm Gr '<t:LY.1d1 (1874)
who, liku Denthnm 01159). Oru<idcn.'(l thc gtnus <iislin<t rrnm Galactfa P.Browne:
sec notC! u11c.lcr G'nlflctln ~ r rcL1llonsh11~~
Po1un1fnl n~ orru1111entnls
I• '< •I I,

Collaea cipoensis Photograph by G. P. Lewis

; ., l ~' · : . · . , ,..,. · •• l'/;11 /n, ;·n n ,' 1 hi• t, !' ! •" ~eorudolphia volubi/is Drawing by P. Halliday

TRIBE PHASEOLEAE )99

 Clitorio brochystegio Photograph by G. P. Lewis
Ophrestio oblongifolio Photograph by A. E. Van Wyk & S. Malan C/ltorio /oscivo Photograph by D. Du Puy

CruddasiaPrain 1898 Clitoria L. 1753

c. 2 spp.; NE Indian subconti nent; lndo-China (Myanmar, Thailand)
c. 62 spp
4
; s America, c Ameri ca, Caribbean and Mexico (c , 48 spp., cent~ed in
no rtherns America), Afiica (4 spp.), Madagascar (2 spp.), Indian subcontinent (I
Named in honour of Lieucenant Crudda.s who assisted the Director of the
sp ), Indo-China, Ch ina, Malesia (6 spp.), Australia (I sp,) .
Botanical Su rvey of India wirh the investigation of the Kachin fl ora
From c/itoria (Gk.: clitolis); lhe memb1anous fo lds of the co10\la were considered
Climbing herbs; seasonally dry tropical forest and forest margins
to resemble the external genitalia of the human female
Reference(s): Adema & Barham (2002)
Sh rubs, lianas, herbs and trees; seasonally d1y to wet tropi cal lowland and
Cniddas/a is very close ro Opbrestia but both generd need revision to cst<1blish
montane forest, woodland or scrubland and wooded grassland
generic limits See notes under Opbreslia
Reference(s): Fantz 0977); Fantz & Predeep (2000)
Subtiibe Clitoriinae, based on the bulk of mo lecular evidence to date, holds a
basally branching position betwee n the millett ioid and phaseoloid. dades
Ophrestia H.M.L.Forbes 1948 Used as ornamentals, forage, green manure 1 cover crops and mechcme
Paraglycine P.j.Herm . (1962); Pseudoglycine F.J.Henn. 0962)
c. 16 spp ; Africa (9 spp. mostly Za mbezian to Zanzibar-Inhambane and
Tongaland-Po ndoland regio ns) and Madagasca1 (4 spp.) ; Indian subcontinent, S Barbieria DC 1825
China, Indo-China (c. 3 spp.)
1 sp.; S Mexico, C America, Caribbean dnd western. S America .
Anngrnm of Tephrosia, because the foliage is suggestive of that genus
Named aftei J . Baptista Gregorie Barbier (1776-1855), doctor and auth o1 on
H<?rl:>S or shrubs; seasonally dry trop ical forest, wood/and, bushland, thicket and
i:ir:issfan<.I medicine and botany . .
Erecr or scandent shrubs; tiopical rive1ine forest, thicket and fo 1est margms or m
Referen ce(s): Verdcourt Cl970b: 257- 262); Thuan (1979) ; Verdcouit (1997; in
areas of secondary growth and open areas
Mackinde r el al., 2001: 24-28)
Reference(s); Fantz (1996)
Molecular evidence presents conflicting views of relationships between the
Used as ornamenrals ~nd ground cover crops
Ophrestiinae and the res[ of Phaseoleae. A basal position near the Diocleinae~
Millettieae is proposed by I3n111eau el al. (1995), while Hu el al. (2000) indicate
that re latio nsh ips are rat her with the Phaseoleae sens. strict., near to Glycininae;
disparate e lements in this species complex are thus indicated, but to elate the
molc<;ular evidence I. baoed only on .-1:u11pl of two Afri .m • Jx:cies of
Op/Jr11slft1: Asian spc ies as well as reprcs<::m:itives of C111ddr1~"'" sho uld be
sampled co clarify the phylogenetic position of Ophrestiinae. Recenc molecular
evidence of Doyle el al. (2000) and Kajita el al. (2001) supports the Bruneau el
al. 0995) placement although here Ophrestiinae is s ister to core Millettieae and
both are sister to the Diocleinae

PseudoeriosemaHauman 1955, non Hauman 1954 nom. nud.

c. 4 spp.; Africa (mostly Zambc~IJm and Sudanian to Somalia-Masai regions)
From pseudo- (Gk.: false) and wfoscma (q.v.)
Herbs or sh n.1bs; seasonally dry tropical woodland to wooded g1ussland and
seasonally swampy grassland
Reference(s): Hauman (1954b); Verdcourt (1970b: 262-263; in Mackinder el al•
2001: 28 - 32)
Included in subtribe Ophrestiinae; two of the widespread species each have two
subspecies
Used as human food (l'oots)
Pseudoeriosemo ondongense Drawing by E. M. Stones Barbierio pinnota Drawing by P. Halliday

400 LEGUMES OF THE WORLD TRIBE PHASEOLEAE 401

 c.. - .·Jfafo"
~ ~

Centrosema plumier/ Photograph by G. P. Lewis Clitorlopsis mollis Drawing by P. Halliday Apios tuberoso Photograph by G. P. Lewis

Centrosema (DC.) Bemh. 1837 Clitoriopsis R Wi lczek 1951

I sp.; Africa (Congo [Kinshasa] and Sud an)
Bradbuiya Raf. (1817)
From Cliloria (q v,) and - apsis (Gk.: like), 1esemb li ng Cliloria
c. 36 spp.; S America (most spp. in Brazil), C Ame rica, Caribbean, Mexico and SE
Shrubs; se"5onall y dry tropi cal forest ma rg ins, wood land lo wooded grnssl•nd
USA; 3 spp widely int roduced in the O ld World
Reference(s): Wilczek (1954: 269 - 271)
From cenlron- (Gk : sp ur) and semeia- (Gk.: standard), the standard petal has a
short d o rsa l spur . Placed in sub1ribe Clitoriinae
Herbs o r subshrubs; tropical and subtro pical bushland formations derived from
seasonal forest, woodland, wooded grassland and s hrubland, also fo rest margins
Refe rence(s): Fevereiro (1977); Schultz-Kraft & Clemen ts (1990) Ap ios Fabr 1759
Placed in subtribe Clitoriinae (see comments unde r Cliloria) Brad/ea A.clans, 0763)
Important forage and pasture crops with a broad range of tolerance of extreme c. 7 spp; China, japan, Indian subcont ine nt (Hima layas), Inda-China Cc. 5 spp .);
conditio ns; also used as cover crops and green manure Canada and E & C USA (2 spp)
From apios (G k : pear), for the shape of the tuber of Aplos americana Meclik
(Ame1ican potato bean)
Periandra Man. ex Benth. 1837 Perennial climbing herbs or scandent shrubs; su blropical and wa1 m temperate
mixed wood land, thicket, scnib rt nd grassland, often near rivers, la kes or swamps
Glycinopsis Kun tze (1891)
6 spp.; Brazil (all spp.), Bolivia Refe rence(s): Handel-Mazzetti (1933); Woods (1988); Isley (1998)
A genus in the unnatural subt 1ibe Eiytbrin inae, siste1 to subtribe Kennediinae and
From pe1i- (Gk.: aro und) and andros (Gk,: man). referring to the largely
tribe Desmoclieae (Doyle el al , 2000); Brun eau el al. 0995) place Apios as the
mo nadelphous stamens being equal around the style
most basal ly branching genus in the above alliance incl udi ng the Glycinim1e and
Shrubs, s ubshrubs o r twining herbs; seasonally d1y and wet tropical rive• inc
Pllaseolinae; genus in need o f revis ion
forest, wood land or wooded to shrubby grassland, often in rocky areas
Used fo r human food (tubers), fodde1, medicine and as o rname nt:als
Refe re nce(s): Mattos & livcira 0973); Funch & Barroso (1999)
Placed in subtribe Clitoriinae
The sweer-rasring roar is used as a substitute for liqu orice
Cochlianthus Benth 1852
2 spp; W Ch ina (Yunna n, Sichuan) and Himalayas (Nepa l)
Fmrn coc/J/o- (Gk.: spira l shell) and an/hos (Gk.: fiow e1), refe rring to th e twisted
keel pet"ls
Climbing herbs; wa rm tempern te montane scmb and g1assland on dry rocky
slopes
Refe1ence(s): Harms (1921); Woods (1988)
Conside1ed :.i cl ose relative to, or possibly congeneric with, Apios Fabr. (Lackey,
1981); leaves and Rowers turn black on dry ing

Perlandra mediterranea Pho tograph by G. P. Lewis Coch/ianthus montanus Drawing by P. Halliday

402 LEGUMES OF THE WORLD TRIBE PHASEOLEAE 403

 ~~
~I
--. -· . c
~ ~__)
f

Shuteria sinensis Drawing by M. Smith Mastersia baker/ (a-e), M. assamica (f, g) Drawing by C. van Laeren & Mucuna gigantea Photograph by G. K. Linney Mucuna macrocarpa Photograph by/. Murata
/. Wessendorp

Shuteria Wight & Arn 1834
4-5 spp.; Asia (Indian subco ntinent, Inda-China, S China, Males ia, Papuasia)
Named after James Shuter (' -1827), who collected many plants in the vicinity of
Mad1as
Climbing he1 bs or· lianas; seasonally dry tropical and subt1opical forest margins,
seconda1y forest, woodland or scnib, ofceO in open places and on limestone
Reference(s): Thuan (1972); Qian (2003)
Sbuteria, usually placed in the Glycininae (Lackey, 1981) has in recent
phylogenies (B1 uneau et al, 1995; Doyle et al, 2000) grouped with a
Kennediinae-Desmodieae clade togethe1 with ce ttain genera in the Erythrininae
Used as g1een manure and cover crops in plantations
Mastersia Benth. 1865
2 spp.; SE Asia (1 sp. NE Indian subcontine nt, IndC>-China [Myanmar], W China
[Xizang]); 1 sp. Males ia)
Named for Maxwell T. Masters (1833- 1907), British physician and botanist
Lianas or twining herbs; seasonally d1)' t1opical forest often in open pl<1ces, f1orn
wet to dry sites
Refe rence(s): Wel ze n & Hengst (1984)
Currenrl y pf aced in subclibe Glycininae (Bruneau et al., 1995) but see comments
under Diphyllaiium
MucunaAdans. 1763
Stizolobium P.Browne (1756)
c, 105 spp.; pantropical (c . 80 spp. in Asia wd China, 1-2 spp. in Australia; c. 12
spp» in Africa, Madagascar and Mascarenes; c. 12-13 spp. in the Neotropics,
centred in C America & the Calibbean)
Derived from the Brazilian Indicrn name 'Mucuna· given to these plan ts
Mostly lianas; tropical wet lowland co rnontane, often coastal rain forest, co
seasonally dry forest, woodland and thicket
Reference(s): Wilmot-Dear (1984 , 1987, 1990, 1991, 1992); Verdcmut in Mackinder
et al. (2001: 14-24); Du Puy & Labat in Du Puy et al, (2002: 524-529)
Stizo/obium is sometimes maintained as a separate genus but is retained under
Mucuna here pending full revision; molecular evidence (Doyle et al., 2000) places
Mucuna sister to tribe Desmodieae
Mucuna pruriens (L.) DC. (velvet, Bengal or Mauritius bean), is widely gro\.·vn for
forage and as a cover crop <1nd green manui-e; the hairs on many species (often
known as buffalo bean) are an intense irri tant; some species are ornamentals

Diphyllarium Gag nep . 1915

I sp.; Indo-China (Laos & Vietnam)
From di- (Gk,: two), phyllon (Gk.: leaO and -aris (L: prov ided with), referring to
the two expanded bracteoles
Lianas or climbing herbs; seasonally dry tropical rive1 ine forest and secondary
vegetarian
Reference(s): Thuan (1979)
Genus poorly known; Drphyllarium and Mastersia are candidates for molecular
sampling to assess if their <iffinity is with Sbuteria (and hen ce the Kennediinae-
Desrnodieae clade) or with the Glycininae

Diphyllarium mekongense Drawing by P. Halliday Mucuna navo-guineensis Phatagraph by/. Murata

404 LEGUMES OF THE WORLD TRIBE PHASEOLEAE 405

 ~
l------------· ,. _.

Kennedia nigricans Photograph by 8. Fuhrer Vandasina retusa Drawing by R. Natadipoero Spatholobus /atistipulus Drawing by J. Wessendorp

Kennedia vem. 1805 Vandasina Rausche1t 1982

Kennedya DC. (1825) Vandasia Domin 0926), n on Velen (1922)
c LS spp ; Australia (most species in \Y./ Australia) l sp.; SE Asia (Papu.asia) and Australia (Queensland)
Nmn ed after Lewis Kennedy 0 775 -181 8), English nurse1yman at Hammersmith Named for Prof. Karel Va ndas (1861 - 1923), Czech botanist at Ilrno
(London) Herbs to woody climbers; seasonally d1y uopical fores[ margins, woodl<ind,
Cl im bing he rbs or subs hrubs; seaso n<1lly d1y tropiG1I to warm temperate fores t, wooded grassland and grassland
woodland, shrubl<ind, heathland and co::1stal sand dunes to arid plains, gcne1ally Reference(s): Verdcomt (1979: 497-498)
at low altitudes See note under Kennedia
Refe1ence(s): Silsbu1y & Ilrittan (1955); Elliot & Jones 0993, 6: 4-10)
Molecular clam. of the 3 gene1a comprising the Kennediinae (Bruneau el al, 1995;
Doyle el al., 2000) places this subtribe siste1 to tribe Desmo<lieae, along \\1 ith Spatholobus Hassk 1842
various genera in the Erythrininae and Shuteria in the G lyci ninae 29 spp,; SE Asia from \YI India to S China, Philippines and \YI Malesia
Used as ornamentals (often vigorous) and g1o uncl-cover1 e .g., K. pros/r{f/(/ R Ilr. From spathe- (Gk.: blade) and lohion (Gk.: pod), the frnit is nat and knife-like
( running postman) 1.ian::is; season<1lly dry to evergreen tropical fo1•est a ncl thicket, often on rocky
slores and in disturbed meas
Reference(s): Ridder-Numan & Wiiiadinata (1985); Ridder-Numan 0992; 1996)
H ardenbergia nernh_ 1837 Spatbolobus is sister to Butea m1ll 1l1elzolropis (Ridder-Numan, 1996), and all 3
3 spp ; Austra lia genera are part of the unnatural subtfibc Erylhrininae, sister ro the subtribes
Named after Councess Franziska von Hardenberg; her brother Baron K~1rl von Glycin in.ae, Phaseolinae <1nd Kennediinae (Bruneau et al., 1995) rn · Glycinirn1e,
Hligel (1794 - 1870) collected plants in \YI Austrnk1 Phaseolinae and Cajaninac (Doyle et al., 2000); affinities 1nay also be to some
Climbing herbs o r subshrubs; wet to d1y sc\erophyll fore.st, scrublancl, heathland genera placed in the basal milleuiokl -pluiseoloid group, e.g., Kunstfelia (Ridder-
and coastal ph1i ns Numan, 1996) and Plalycyamus (Hu, 2000)
Reference(s): Elliot &Jones (1990, 5: 250 - 253) Used as medicine and cordage (fibre from stems)
See note under Kennedia
Used ris ornamenw ls, human drink (leaves), sc1een ing and grou ncl covei
Butea Roxb. ex Willd . 1802
2 spp .; Asia (subcontinental India and \Y/ Inclo-Ch ina); an Indonesian record from
Jav• is doubtfully native (Sanjappa, l989)
Named after John Stua1t - 3'" Earl of Bute 0713 - 1792) , Il1itish Prime Minister
1762-1763 and effective Director o f Kew befo1e Sir Joseph Banks; "a better
p lantsman than politici<in" (Stearn, 2002)
T1ees or lianas, sometimes growing g1egariously; seasonally chy tropical fore~t.
wood lan d and wooded g1assland, sometimes riverine
Refe1ence(s): Sanjappa (1987, pub! 1989)
Comments as for Spatho/obus
Used <ts om.amen tals (e g, B . mono~perma (bun) T~1ub ., parrot lree or Harne of
the fo1est) , medicine and for religious ceremonies, gums, dyes, cordage and
foclde 1

Hardenbergio vlo/acea Photograph by 8. D. Schrire Butea monasperma Phatagraph by M. Sanjappa

406 LEGUMES OF THE WORLD TRIBE PHASEOLEAE 407

 Adenodolichos rupestris Drawing by M. Grierson Bolusafra b/tuminosa Illustration by S. Edwards Carrissoa angolensis Drawing by P. Halliday

Meizotropis voigi 1845 Bolusafra Ku ntze 1891

2 sp p.; Asia (subcontinemal India and W Jndo-China) Fagelia Neck. ex DC. (1825)
From meizo- (Gk,, greater) and tropis (Gk.: keel), referring to the disti nctive ~ 1 sp.; South Africa (W Cape)
Named after Dr Harry Bolus (1834-1911); botanist in the Ca pe region of South
fl owers ~5 Af1ica and founder o f the Bolus H erbarium, Cape Town
Erect or scandent shn.tbs; seasonally dry tropical woodland and grassland,
some times riverine, often on upland dry slopes
Herbs or' subshrubs; mediterranean mo ntane sclerophyllous shrubland (fynbos),
Reference(s): Sanjappa (1987, pub!. 1989) often a\ong strea msides
Comments as for Spalho/obus Reference(s): Bu llock (1965); Germishuizen (2000: 277 - 278)
Apparently allied to Rbyncbosia (q v.) and Cbrysoscias (q.v.), in subtribe
Cajanin ae
Adenodolichos Harms 1902
c, 15-20 spp.; SC Africa (Zambezian region), 1 sp to Sudanian WC Africa
From adeno- (Gk.: g land) and Dolichos (q.v.), a sticky-glandular plant resembli ng
Carrissoa Baker f. 1933
Dolichos species 1 sp.; SW Africa (Angola)
He rbs or shrubs; seasonally dry tropical woodland, scrub, wooded grassland and Named after Luiz Wittnich Carrisso (1886- 1937), Portuguese botanist in Angola
grassland, often appearing soon after fire
and founder of Conspectus F/orae A ngolensis
Reference(s): Wilczek 0954); Verdcourt in Mi lne-Red head & Polhill (1971: Subshrubs; seasonally dry tropical open bushland and scrub, or along
watercourses 1 w ith woody rootsrocks in fire-prone environments
702-707); Lock (1989: 386-388); Verdcou rt in Mackinder et al. (2001: 242- 249)
The most basally branching gen us in the Cajaninae in the analysis of Bruneau el Refer ence(s): Baker 0933)
al. 0995)
Verdcourt in Mackinder el al. (2001: 169) considers C. angolensis Baker f. to be a
Used for human food (vegetable), medicine and cordage Rhyncbosia (q.v.)

Paracalyx Ali 1968 Chrysoscias E.Mey. 1836

6 spp. ; NE Africa (Ethiopia, Somalia) and Socotra (5 spp.); Indian subcontinent, 3-4 spp.; South Africa (S parts ofW Cape)
Indo-China (1 sp.) From chryso- (Gk. : gold) and skia- (Gk,: shadow), in reference to the flower
From para- (Gk.: near, similar to) and calyx, referring to the distinctive colour
morphology of the calyx in this genus Subshrubs or herbs; medi terranean sclerophyllous shrubland (fynbos) on
Herbs; seasonally dry tropical forest, woodland, thicket, bushland and scrub, often sandstone slopes
along water courses and in rocky areas
Reference(s): Baker (1923); Germishuizen & Schutte in Goldblatt & Ma nning
Reference(s): Ali (1968); Thulin 0993: 446 - 447) (2000: 509 [as Rhynchosia])
This species group and their relationship to Rbynchosla are in need of revision;
All species were originally described in the genus Cylisla Aiton (1789), which is
the genus has been included in Rbyncbosia in South African literature, e .g .,
now a synonym of Rhyncbosia (q.v.) [Ali, 1968)
Germishuizen (2000)

Parocalyx scariosus Drawing by P. Halliday Chrysosc/as porvifloro (1-10) and Chrysocias pauc/flora (11, 12)
DrolVing by P. Holl/day

408 LEGUMES OF THE WORLD TRIBE PHASEOLEAE 409

Rhynchosia malocophyl/o Photograph by S. Collene tte Rhynchosio montoroensis var. cuprinervio Photograph by G. P. Lewis Eriosema psoro/eoides Photograph by D. Du Puy Eriosemo robinsonii Drawing by M. Grierson

Rhynchosia Lou1'. 1790 Eriosema me.) Hchb. 1s2s

c. 150 spp.; Africa and Maclagascrn (c. 100-110 spp.), N and S America (Mexico to
Cy/1s1a Aiton Cl 789); Bc111kea Vatke (1881); l~)'cephy/111111 Piper (1924)
N Aigentin ~1, c. '10 spp .) ancl SE Asia to Austr~ilia (2 spp.); most New \'V'odcl spp-
c. 230 spp. ; pant1 op icn l (c. 140 spp. in Af1i ca - Mad~1gascar; c. 55 endemic spp. in
t1 opical and subtropi c.ii Ame ric:1 [c 28 spp. in N ancl C Am erica, 20 in S AmeiiG1 are S Am erican (c. 30 spp.)
From erion.- (Gk. : wool) and sem ein (Gk: sranda1d), refen·ing to the indumen tllm
<lllcl 6 widespread between both] <rncl c. 30 - 35 spp. in warm tempera te co tropic1I
Asia co N Au stral ia !2 endemic spp.D on the back of che standard
F1om rby11cbo - (Gk..: beak), refening to the shape of the keel in the cype !-'pecies Herbs 01 su bshrubs; seasonall y d1y rropical to subrropical fores! margins,
woodland, 1hicket ::incl wooded gr;:1ssland, or open grassland in rocky OJ' swampy
1-Jerbs, ''i nes o r subshrnhs; seasonally d1y foiest, forest m~irg in s, wooclbnd,
thicket, wo oclccl g1<1ssland, sh 1ubbnd ~ incl g1asshmcl, often in open rocky :1re1s 01 <1reas, or in old cultivations and waste gruu nc1
Refe1-c nce(s): Gre:ir (1970); Ve1dcou1t (1971: 112-1 46; in Mil ne-Redhead & Polhill .
:dong streams and in disnirbecl a1e~1 s; m8ny species a1c pymphytes
Hefcrence(s): Ycrdcou1l (1971: 70-ll2; in Milne-Hcdhead & Po lhill , 1971: 711-761); 1971: 761-805); Stirton (1975); Fortunato 0999); Verd cou1t in Nhickincle1· e1 al.
G rea r (1978); Thulin 0981; 1993: 440 - 446); Germishuizen & Schutte in Colclblall
(2001: 211 - 242); Du Puy & Labat in Du l'uy el ell (2002: 601-605)
& Manning (2000: 509 - 510); Fo1tunato (200 1); Vc1dcou 1t in Ma<..:kindc1 et oJ. Placed in subtribe Cajani n ~1e, allied ro R/Jynchosia
(2001: 168-210) ; Du Pu y & Labat in Ou I'u y el nl. (2002: 590 - 600); van de1• Used ~1 s human fa m ine food (Hux h:1111 et al., 1998), medicine a nd fish poison
Macsen (2003)
Pla ced in s ubt1•ibe 01j~m ina e, allied to Eriosema; Je;wcs of R j"i:!rulifolir1 Bcnlh. ex
Hnrv, from rhc c~1pe region of South Af1ica <lrc unusual in bein g ~1ppa1cntly Dunbariaw;ght & Ai n . 183'1
bipinnate, o r once pinn:ne with 5-9 leaflets co mparccl to the pi11m11ely .J-foliohite 20 spp.; SE Asi<1 (centred in lndo-China to S China, In.c.li ~in subcontinent, Males ia
co ndition in th e test of the genus and Paprn1si ~1 ; 1 sp to F Asia ); 2 .spp. extending to N Aust1a li ~1
Used as r~1sture pl;111ts and scveial species 1 co mmonly Gilled 1osary bean, ha\'C NJmed for Professo1 George Dunbar (178/i -1851) at Edinburgh , ftiencl and
atll acti vc red , hlue 1 black, mottled rn bicolou1ed seeds u sed for necklaces t"Cc .; colleague o f \'(light and A1 noct
seeds a lso used as weigh ts or ::1s narcoti{·s; pbnrs a lso famine foods (Huxharn et Climbing he rbs 0 1· subshrubs; seasona ll y dry t1opica l forest 1 thicket, scrn b and
cl/., 1998) gl'~1ssland, from moist to semi-add areas main!>' in the uopics
llcfcreoce(s): van cier Maesen (1998)
Placed in subtribe Cajani11ae
Used for gree n 111anu1e and in edicinei related to Coja1111s (q ,v .) so may be useful
in breeding progia mmes of pigeon. pea (van dcr Maesen, 1998)

Rhynchosio /eondrli Photograph by D. Du Puy Dunborio cumingiono Drawing by Y. F. Tan

410 LEGUMES OF THE WORLD TRIBE PHASEOLEAE 411

 ~
u
Cajanus cajan Photograph by G: P. Lewis Flemingia strabilifera Photograph by H. Ohashi Erythrina peruviana Photograph by G. P. Lewis

Cajanus oc 1s13 Flemingia Roxb. ex W.T.Aiton 1812

0-ljmt Adans. (1 763): AIJ1o.<in Wight, Am. (I '.:1-1); JI11</0111t1//11s Gngncp. 0915) Mogbcrnia j .St.-HiL (1813); Lepidocoma j ungh. (1845)
3'1 ~pp.: E J\sl~ ( Indian subconuncni, China, E Asi:i, lndo·CJ1inro, MaJc., b.. c. 30 - 35 spp.; SE Asia (Indian subcontinenl, lndo-Ch ina, China, Malesia,
P:ipu:osla, Pn<ific; 16 >pp.>. N Aus1r:1li" (15 spp.), w ( udanl:in) Afrl ,1 (I <p.); 2 Papuasia, c. 26 - 29 sp p .) 10 Ausrrnlia (c. 3 - 4 spp.); Africa 1 sp. and F grabamiana
~ill>· wldcsp1.::1cl in lei \Wlrld. including l : ca}clll (I..) ltu1h Wight & Arn wides pread in Africa and Asia
From calj:oni-: (M!ll:iy: bean or pea), 1he original spell ing of the genus was Caja11 Named afte r John Fleming (1747-1829), Scott ish naturalist arul Physician-Ge neral
Herbs or shrul>s; .sc:1sonally dry cropical o pen fo resc to grassland, o ften in rocky or of the East India Company's Medical Establishme nt in Be ngal
parrly dis1url:x.'<I orcas He1bs or shru bs; seasonally dry tropical fo rest , wood land, wooded grassland and
Itefe rc nce(s): van der Maesen Cl985b; 2003) grnssland , some tim es near swamps and streams; also in disturbed and waste areas
TI•c suhrrlbc njn nl nnc rum-. n l>:r. 1 poli•10111y wilh a l:o rgcly l'hnseolin.'IC. Reference(s): Verclco un in Milne-Redhead & Polhill 0 971 : 805- 807); Wei 0991)
Glydninn" nnd I' · rJ l~-e1e cl!ld~ (Do)•lc Cl nl., 2000. Kafhn C'I al., 2001) orb sis!"' Placed in subtribe Cajaninae; genus in need of revision
10 11 combhK'<I Erythrinin:1<:, Kcnncd llnnc, O.,smcxlie:re, l'ha.-;eolin:ic, Glycinln:1c Used as cover crops, green manure, medicine, human food (uncl ergro Lmd tube(s)
fo nd l'ilQr:1lcc.1") lad• (Onme:111 el ti/., 1995): Verdooun in Mackinder ,1 ul. ( ZOO! : and an orange dye (known as waras)
16;3) .>W IC.\ thm in lmer cdi1ion.'5of1he l111ammto1wl C0</o of8ulr111/c11/
NQ1111111clflt11n.' (e.i,:., Ci r ·mer Cl al., 2000), Cqj111111s is m .."11cd ·~• :ltl vnhogm11hic
' ':itlunt of 11ja11, " nd the conserv<!cl •P<?lllns Is :tttrlbuK'd 10 Adnn on; 1hb i< Erythrina L. 1753
not :it•cptcd hen;.~ Vi:rdmun l/Jlli> co111111.) hns re ·mly com.-ctcd 1he :htlh rity c 120 spp ; c. 70 spp. in the Neolropics (c. 50 spp in Mexico, C America and
of Cfljmms caft111 from the wid ·ly us..-d C. caj<111 (L.) Mlllsp .. 10 c. crtjo 11 (I.) Huth Caribbean; c. 20 spp. in S Ame1 ica, a number of which may complise the basa ll y
Used for human food :IS n ln'1jor pulse Or \'~'l!Clal>lc crop (p gCOn fie"~ (lr gullJ:O branching elements of the genus); 38 spp. in Africa and Madagascar; c. 12 spp. in
1ie:1), Ol hcrwl$c w<cd :is pasture lcj!\1111 !S, fomgo:, gn.~n nmnurc. cover crop~ ;incl Asia to Au sllalia
In rnct,lfdnc; C. cnjr111 is wit.lei)• cultl"ncd In the Old "nd " "' \'furld tropic. From e1ythro- (Gk.: red), re fe rring to the flower colour of some species
Trees and shnibs; seasonally dry tropical and sub1ropical lowland to upland fo rest
(so metimes coastal, in inundated <treas or riveri ne), woodland, wooded grassla nd,
bushland, lhicket ancl grassland
Reference(s): Krukoff & llarneby 0974); Fra nklin Hennessy 0991); Brunea u
0996)
A genus in the unnatural subtribe Erythrin inae, in a basally branching clade sister
to sub11ibes Glycininae and Phaseolinae; Bruneau el al 0995) place Elytbrina
siste1 to subtribe Glycininae
Used as orm1mentals (coral or lucky bean rrees), shade trees, timbeJ' (constru cti on,
implements) , living fences and enclosuies, green manure, livestock foclde1,
medicine and seeds a1e used for neckl aces

Cajanus scarabaeaides Photograph by D. Du Puy

Erythrina madagascariensis Photograph by D. Du Puy

412 LEGUMES OFTHE WORLD TRIBE PHASEOLEAE 413

 0 '

Dysolobium grande Drawing by c. M. Grey-Wilson Oecorseo grondidieri Ph otograph by D. Du Puy Strongylodon modogascariensis Photograph by D. Du Puy

Psophocarpus Neck_ex DC. 1825 Decorsea R.Vig. 1951

Vignopsis De Wild. (1902)
c, 10 spp.; Africa and M;1dagasca1, one wiclespi·eacl cultivated sp . (unkno wn in the
wild), possibly Asia n in oiigin ; a further sp introduced in the Mascarcnes,
Comoro IslanJs, Caribbean and S America
rrompsopho- (Gk o noisy) and ca1pos- (Gk.o fruit), referring to the explosive
dehiscence of the pods
Climbing herbs or s ubshrubs; seasonally dry lropical forest and foresr margins,
secondary vegetation, thicker, swamp, moist wooded grnssi<ln<l and gr<1ssland
Refe re nce(s} Verdcou 1t & Halliday 0978); National Research Council 0981);
Harde1• 0996)
Psophoccupus is bas.:tlly b1anching in subtribe Phaseolinae (Brun ea u et al, 1995),
and placed fu11he1 down the tree in a clacle with E1ylbrina, sister to th e combined
Phaseolinae, Glycinin~1 e and Psrnaleeac (Doyle el aL, 2000)
Used as human food; P. lelmgonolobus (L.) DC (winged bean o r Goa bean) is
vel)' widely cultivated as a prorein-1 ich vegetable, the young shoots, young pods,
nowe rs, seeds and tuberous 1ools are all ed ible; mher species used for green
ma nu re, grou ncl cover 8 nd foclcler
Dysolobium (llenth.J Prain 1897
4 spp.; SE Asia CE Indian subcontinen t, Indo-China, SW China, Malesia)
From dys- (GL hard) and lobiou (Gk o pod), referring to the almost wood)'
texLUrc of the pod.s
Trctiling herbs to wo ody climbers; seasonally dry rropical forest, secondary
vegetation, forest margins <incl 1iverine vegera ti on
Reference(s} Welzen & He ngst 0985); M'11 echal el al. 09780 236-240)
This genus is appare ntly allied to Psophoca>pus (Kajita, peis. co111m)
6 spp ; sou th em Africa (3 spp,) and Madagascar (3 spp)
Named after Dr G.-J. Decorse (1873 - 1907), doctor ancl collecto1· in Madagascar
He J'bs or subsh 1ubs; seasonally chy tro pical and subtropica l woodland and
woodland margins, bushland, thicket or xe1 ophytic sc n.1b
Reference(s} Verdcourt (1970co 442 - 447; in Mackinder el a l., 20010 156-159);
Du Pu y & Labat in Du Pu y el al. (20020 555 -557)
Traditioo;:11ly placed in !iubtribe Phaseolinae, perhc1ps allied co Otoptera
Seeds and tubers said to be edible
Strongylodon Vogel 1836
12 spp.; Madagasca1 (2 spp.), H.Cunion, S1 'i Lanka, Anclanwn Isles, E Malesia,
Papu<tsi~1. Pacifica , N Australia mainly in Philippines
From slrongylo- (Gk,o round) and odons (<ooth), the oute r ma rgins of the calyx
lobes a 1e 1ounded
Li;rnas up ro 40 m high; tropical rain forest, thicket 0 1• seconchuy vegetmion, ofren
in sw<1111 ps or nea r water
Refe re nce(s): Huang (1991); Du Puy & Labat in Du Pu)' et al. (20020 521- 524)
A genus in the Linnatural subrribe E1yth1 ininae, basally branching to subt1ibes
Glycininae and Phaseolinae; Bruneau el al. 0995) rlace Slrongylodon sistet to the
Phaseolinae, w hile Doyle el al . (2003) have it sister co the Glycininae; Huang
(1991) distinguishes four sections in the genus; S. perrieri R.Vig. is now in
Sylvic/Jadsia (Mil lettieae)
Used as 01m1ment.ils, notabl y the jade vine, S. macrobollys A_Giay

Otoptera DC. 1825

2 spp ; southern Africa (I sp.) and SW to \VI Madagascar O sp)
From 010- (Gk o ear) and p1ero- (GL wing), refer ring to the ear at the base of the
wing petals
Herbs o r sh1 ubs; seasorndly dry tmpical open wood land , wood ed grassland,
g rassla nd and scrubland, on sand or limestone
Reference(s)o Verdcou1t in Mackinder el al. (2001' 66-68); Du Puy & Laba< in Du
Puy el al. (20020 558-559)
See notes under Decorset1; traditionally placed in subtribe Phaseolina e, but clllJent
research by Delgado Salinas, Thulin, P<isquet, \'Qeeden and Lavin sugges ts tb~t t
Otoplera is 1ather closer to Psophocmpus
Used as vigorous sand binders and soil stabilisers; stems used for twine

Otoptera burchellii Drawing by P. Halliday Strongylodon macrobotrys Photograph by P. Gosson

414 LEGUMESOFTHEWORLD TRIBE PHASEOLEAE 415

 Herpyzo grondifloro Drawing by A. J. Beaumont Neoroutonenlo ficifo/ius Photograph by A. E. Van Wyk & 5. Molan
Cologonio broussonettii Illustrator unkown

Calopogonium Desv. 1826 Herpyza c .wright 1869

I sp.; Cuba (Allen & Allen, 1981, also note its occurrence in Puerto Rico)
Cyanostremma !3enth. ex Hook. & Arn . (1841); Slenolobium !3enth. (1838), non
From herpo- (GL creeping)+ za (Gk: very), referring to the creeping habit of
D.Don. (1823)
5- 6 spp.; America (Mexico, C America and Caribbean to Paraguay and the plant
Herbs; sandy places near streams, seasonally dty tropica l wooded grassland and
Argentina). 1 sp. widely introduced in the Palaeotropics
pine forest
From calo- (Gk.: beautiful) and pogon (Gk.: beard), for the golden-haired Siems,
Reference(s): Lewis (1988); Ma>.-well & Taylor (2003); !3eyra Matos el al. (2004)
leaves and pods of the cype species, C mucunoides
Kavanagh & Ferguson (1981) noted the unique pollen of He1pyza which is unlike
Herbs or scandent subshrubs; seasonally d1y tropical and subuopical fo 1est,
that o f other Phaseoleae and they excluded this genus, together with
woodbnd or thick et, ofLen on margins, usually near rivers or swamps, or in
Calopogonium and Pachyrhfzus, from subtribe Diocleinae 1 a view supported for
bushland, shrubland and distu 1bed grassy areas
the latte1· 2 genera by molecular evidence (Bruneau et al, 1995; Doyle el al.,
Reference(s): ca,,,alho-Okano & Leitao Filho (1985)
2000); the analysis of Maxwell & Taylo1 (2003) also suggested that Herpyza is
Calopogoniu.m and Pachyrhizus were traditionally placed in the Diocleinae, but
related to genera in subtribe Glycininae (but see note under Neomdolphia)
molecular evidence (!31uneau et al., 1995; Doyle el al , 2000) puts these genera in
the Glycininae; Lee & Hymowitz (2001) place Calopogonium sister to Cologania,
while Doyle et al. (2003) resolved Calopogonium as siste1 to Neonolonia '1nd Neorautanenia schinz 1899
Teyleria, and these three as siste1 to Co/ogania 3 SPP•i Af1ica (Zambezian , Su<lanian and Som<ilia-Masai regions)
U sed as fodder and cove 1• crops; C. mucunoides Desv. is widely used as a g1een Named after Rev. Martti Rautanen (1845-1926), Finnish missionary and botanist
manure; also a weedy esca pe in Namibia
He1 bs or subshrubs; seasonally dry tropical open woodland, bushland, wooded
Cologania Kunth 1824 grassland and grassland, often in rocky places
c. 12 spp~ ; centred in Mexico to SW USA, C America, Colombia, Venezuela, Reference(s): Verdcourt 0970b: 300 - 307; in Mackinder el al., 2001: 74-80)
Ecuador, Peru, Bolivia and Argentina , largely confined to montane habitats Traditionally placed in subtribe Phaseolinae, but cu1Tent research by Delgado Salinas,
Named in honour of the Cologan family, navigators and naturalists from Puerto Thulin, Pasquet, Weeden and L1vin suggests this should be in the Glycininae
Orotava, Tenerife, who rendered i111po1tant services to the sciences and ans The tuberous roots have insecticidal properties and planrs are used as fish poisons
Perennial climbing he rbs; tropical to warm temperate montane forest, woodland (roots also known to kill bilharzia-ca riying freshwater snails)
and thicket to wooded grassland and open bushland on mountain slopes
Reference(s): Fearing (1959); Turner 0992) NeonotoniaJ.A.Lackey 1977
Placed in subtribe Glycininae and sister to Calopogonium by Lee & Hymowitz
Nolonia Wight & Arn. 0834), non DC (1925)
(2001); Doyle el al (2003) have a Calopogonium - Cologania - Neonotonia clade
2 spp..; Africa , Arabian Peninsula, rndian subcontinenc and Malesia; inrroduced
sister to a Dumasia - Glycine clade, with Co/ogania sister to tribe Pso raleeae (but
elsewhere in the Old World
see note under Amphicmpaea)
From neo- (Gk: new) and Nolonia (a~e 1• !3enjamin Noton (1812-1835), an English
Pachyrhizus Rich. ex DC. 1825 botanist who cont1ibuted to the botany of the Neelgherry (Nilgiri) Hills, India)
Scandent or climbing herbs; seasonally dry tropical forest, woodland, thicket,
Cacara Rumph. ex Thouars (1806)
scrub, wooded grnssland and grassland, or in cultivation
S spp ,; Mexico, C America and W tropical S America (Colombb, Venezuela,
Reference(s): Lackey 0977); Mackinder in Mackinder el al. (2001: 54-58)
Ecuador, Peru, Bolivia, \YI Brazil, Guyana), 3 species widely cult ivacecl in tropical
Three subspecies are 1ecognised in N. wighlii (Wight & Arn .) j.AJ.ackey;
and subtropical America , one also extensively in the Palaeotropics
Neonotonia contains species once placed in Glycine; Lackey (1977) notes some
From pachy- (Gk.: thick) and rbiza (Gk.: root), for its tuberous roots
species of Pueran·a, when better known, may be placed here; lee & Hymowit z
Herbs or subshmbs; seasonally dry tropical forest and thicket, often on 1m1rgins,
(2001) place Neonolonia and Teyleria in a clade with Pachyrhizus and some
and in scrub vegetation and open grassy areas
Pueraria species, while Doyle et al. (2003) resolved them as sister to
Reference(s): Sorensen (1988; 1996); Sorensen el at. 0998)
Calopogonium
See under Calopogon tum for taxonomic affinities
Exten sively cultivated as livestock pasture, forage, green manure and ground
Used for human food, e.g., P. erosus (L.) Urb. (yam bean, jicama), pods also
Pochyrhizus erosus Photograph by G. P. Lewis Neonotonlo wlghtii var. longicoudo Photograph bys. Collenette cover; also used as human food (leaves) and for medicine
edible; other pmts used for fibre and medicine

416 LEGUMES OF THE WORLD TRIBE PHASEOLEAE 417


Pueraria labata var. thamsonii
418 LEGUMES OF THE WORLD
Dumosia vi/losa Drawing by D. Erasmus
Teyleria Backer 1839
3 spp.; SE Asia Ondo-China, S China, W Malesia)
Named after Pieter Teyler van der Hulst (1702-1778), silk manufactu1'er and
scientific benefacto 1 at Haa1 lem, The Nethel'\ands
Twin ing herbs; se8sonally d1y tropical forest, often on hillsides
Refe1e nce(s} va n d er Maesen Cl985a); Lock & Heald (1994: 122)
Placed in s ubuibe G lyc ininae (see notes unde r Neonotonia) ; van der Maesen
0985.a) effected two new co mbimltions in Teyleria , a genus previously
cons idered monospecifi c
Dumasia DC. 1825
c. 10 spp.; Asia (Indian ::;ubcontinent, Myanmar [I3urma]. China, Korec1, J~qx111); 1
sp. widespre~d in tropical and subtropica l Asia, Africa and Indicin Ocea n
'anK..:l o(tcr J<!:ln Ba pl Dumn.< (1800-18&1), ProfL..,.<-01 o f Phiitm•mlogy :tn<I
orgnni<; chcntislr)' :11 the 111<.-'<ll al fo ·1iit)1 In l':iri.<; fric nd of A.I'. de ·111(1011 •
Cl imbing herbs; mostly season:::dly clry tropica l to w~11m temperate upland fo1est,
forest margins and riverine secondary growth
Re ference(s): Wei (1995)
Ilasally brnnching in subtribe Gl yclnl noc (Ooy l<: & Doyle, 1993: I.cc & 11 mowlt7.,
2001), while Doyle et al. (2003) pine<: D11111t1slo as bas:•ll)' brond1111s; in " Dumasi•
- Glyci ne clade which is sister to ~ ,Llopogonh1111 -Cologa.nia. 'conotooiu d !1dc
Seeds used as beads
Pueraria DC, is25
Neustanlhus Ilenth. (1852)
c, 18 spp.; Asia (iowl::i ncl ro montane Indian subcontinent, Inda-China, S\'\f China1
Malesia, Papuasia , Pacific); the Kudzu vine (P. monlana (Lour.) M e1T. v:ar. /o!Jata
(Willd.) Maesen & S.M ,Almeida) widespread to tempernte E Asia, Australia and
introduced into Africa and SE USA; tropical Kudzu (P. phaseo/oides (Roxb,)
13enth) is widely introduced in Africa and rhe Neocropics
Na med after M.N. Puera ri (1765 -1845), a Swiss-born Professo.-nt Copenhagen, a
fri e nd of A.P. de Candolle a nd of the botanist Vohl
Lianas, shrubs or climbing herbs, usually with hlrge tuberous roots; seasonally dry
tropica l and subtropicH I forest, rain fo1 est, fo 1es t margins <lfld scrub vegetation,
o ften on open limestone and in rocky are:LS
Re ference(s): van d e r Maescn Cl985a)
Placed in subtribe G lycininae; Lee & Hymowitz (2001) show Pueraria to be
polyphyleric, with species of the non-typical sections Brevirann1/ae and
Schizophyllon being basally branching to th ose of the typic;il section Puerarin ;
Doyle el al. (2003) resolve a Pueraria-Pseudovigna clade sister to an
Amphicarpaea-Glyci ne clade, with Dumas;a being sister to borh these claclcs
Used <1s pasture, fodder and cove1- crops, green manure, human food (tubers),
thi ckening agent (sta1ch from tL1bers), cordage (fib1e); Kudzu vine is often an
Photograph by H. Ohashi invasive weed Sinodolichas /agopus
Nogro fi/icoulis Drawing by V. C. Gordon
Eminia antennulifera Drawing by 0. Erasmus
Nogra Merr. 1935
Grona sensu Benth . (1852), non Lour. (1790)
c. 3 spp .; Asia (Indian subcontinent, S China, Jnclo-Chin a)
Anagram of Grona (Gk.: cave, g1otto), for the deep keel petals
Herbs) seasonally dry tropical forest, o fte n in open grassy aieas
Reference(s): Panigrahi & Tiwa ri (1975); Sanjappa (1992: 220-221)
Lackey (1981) notes that the unifol iolate leaves in Nogra probably unite species
which would otherwise belong in 3 different genera; Nogra grabamii (Wall. ex
Benth .) Merr. is srro ngly supported as sisrer to Pueraria sens. strict. by Lee &
Hymowitz (2001)
Eminia Toub. 1891
c. 4 spp.; Africa (Zambezian region)
Named in honour of Dr Emin Pascha (1840-1892), German doctor and explorer
in Sudan and the Lake Albeit region of Africa
Herbs or shrubs; seasonally d1y tropic<1l woodland, riverine t hicket, wooded
grassland, bushland and shrubland, often on l'ocky slopes at mid to higher oltitucles
lleference(s), Pauwels (1983); Lock 0989' 398); Mackinder in Mackinder et al.
(2001: 42-45)
Placed in su btribe Glycininae (see note under Pseudeminia) and closely re lared to
Pseudeminia Verde~ and Pseudovigna Verde
Used as human food and drink (roots contain amylase used in brewing bee r)
Sinodolichos Verde. 1970
2 spp.; SE Asio (NE Indian subcontinent, SW China, Indo-Chi na)
From sino- (L chinese) and Dolichos (q.v.); both species were previously placed
in that genus
Twining herbs; seosonally dry tropical forest and thicket on sondy soil
Reference(s): Verdcou11 (1970c: 398 - 399); Lock & Heald (1994, 120)
Verdcourt notes a possible affinity w ith Dyso/obium or Pueraria, and a
morphological similarity to Pseudovigna; Doyle el al. (2003) place Sinodolichos
sister to Pseudeminia and Pseudovigna
Drawing by P. Halliday
TRIBE PHASEOLEAE 419
Pseudeminia comosa Drawing by D. Erasmus Pseudovigna argentea Drawing by E. M. Stones Teramnus - various species Drawing by D. Erasmus Glycine max Drawing by D. Erasmus

Pseudeminia Verd e. 1970 Teramnus r.Browne 1756

9 spp .; principally Old World t1'0pics (Africa, SE Asia [Indian su bcontinent, S
4 spp.; Af1ica (Zambezian region)
China, Indo-China, to Malesia]); 1 sp. neotropical and 2 pantropical
From pseudo- (Gk.: false), and Eminiu (q.v.)
F1om teramnon- (Gk.: room or chamber), referring to the seeds being in separate
Herbs or subshrubs; seasonally dry tropical woodland, thicket, bt1shland, wooded
grassland and scrubland or in old cultivated areas compattrnents in the pods
Climbing herbs o r subslm1bs; mainly seasonally d1y t1opical bushland and thicket,
Reference(s): Verdcou11 (1970c: 393 -398)
gra.ssland, wooded g1asslaod and forest clearings, often in open and rocky d1y
Pseudeminia and the closely related Pseudovigna Ve rde. are placed in s ubtl"ibe
G lycininae on molecul ar data (Bruneau el al., 1995; Doyle et al., 2000; Lee & areas
Reference(s): Veiclcou1t Cl970b: 263- 284); Lee & Hymowitz (2001)
Hymowitz, 2001)
Placed in subtribe Glycininae; a natural genus but species delimitation is difficult
Used for ground cover; roots with saponins used for soap
Used for ground cover, has pote ntial as pasture (forage) plants

Pseudovigna (Harms) Vel'dc. 1970

Glycine wilkl. 1so2
2 spp.; E and W Africa (P. argentea (Willcl.) Ve rde. is ± coastal in both E and \YI
Africa) Soja Moench (1794)
19 spp .; principally Australia (16 spp.), 2 of which widesp1ead to Pacific Islands,
From pseudo- (Gk.: false) ancl Vigna (q.v.)
Taiwan, J apan and S China; 2 spp end emic to China, Taiwa n, G. max (L.) Merr
Herbs; seasomilly d1y tropics; the low altitude species occurs in grnssland or low
(soyabean; soybean) from E Ru ssia, Korea, China and japan but widely cultivated
bushland, often in culrivated l<md or near sw::in1ps; the other is from forest, near
From glycys- (Gk.: sweet), refel'ring to the sweet sme ll or taste of the roots and
rivers and in clearings, or in woodland or wooded g1 ;:1ssland
leaves of a .species no longer included in Glycine (now in Apios q.v .)
Reference(s): Verdcourt Cl970c: 390-393); Em 0980)
Twjning herbs; seasonally dry tropical to wam1 temperate open woodland,
See comments on 1elariooships under Pseudemin.ia
thicket, wooded grassland, clearings, riverbanks and cliy hillsides
Reference(s): Hermann (1962); Hymowitz & Newell (1981); Ko llipara el ed. (1997);
Lee & Hymowitz (2001); rfeil & Craven (2002); Doyle el al. (2003); Sakai et al.
Amphicarpaea Elliott ex Nutt 1818
(2003)
Amphica1pa Elliott ex Nutt. (1818) Glycine has had a convoluted taxonomic bislmy; see also under Neonotonia
4 - 5 spp.; C and E USA, SE Canada to Mexico (I sp.); E Asia (Russia, Japan, Soyabean i:s one of man's p1 incipal pulse and food producr crops; also used as
Korea, China) to Himalayas (2- 3 spp.); montane tropical C and E Africa (1 sp.)
fodder
From amphi- (Gk.: of two kincls) and ca1pos (Gk: fruit), refening to the two fruit
types p1oduced, aerial and below ground
Herbs; se;1sonally dry montane tropical fores1 (in Africa) or coasrn l to monca ne
tem perate forest, woodland, lhicket, bushland or wooded grassland
Phylacium Benn . 1840
2 spp. ; Inda-China (both spp.) to Malesia (excluding llomeo), Papuasia and N
Reference(s): Turner & Fearing (1964)
Amphicmpaea has been conserved against rh e original spelling of Ampbicmpa,
Australia
From phylax (G k.: guard), 1eferring to the large brncts which continue to expand
and is strongly supported as sisler to Teram.nus and Glycine by Lee & Hymowitz
(2001); Wojciechowski et al. (2004) resolved an Amphicarpaea-Glycine dade frolll and p rotect th e fniits
Climbing (sometimes woody) he1bs; seasonally dry tropical forest, thicket,
which tribe Psora\eeae is derived (see note un<le1 Cologania )
wooded g1assland and scrub, often in disturbed areas
Used as cover crops, for erosion control and human food
Reference(s): Bresser 0978)
Phylacium and Neocolleftia are rn oved f1om tribe Desrn odieae to the Ph~1seo\eae
based on morphological, palynological and molecular evide nce (Doyle et al., 2000;
Kajita et at., 2001)

Amphicarpaea africana Drawing by E. M. Stones Phytacium majus Drawing by M. Smith

TRIBE PHASEOLEAE 421

420 LEGUMESOFTHEWORLD
 © '
-
,

Neocollettia gracilis Drawing by M. Smith Wajiro albescens Drawing by P. Halliday Nesphostylis holosericea Drawing by o. Erasmus Alistilus jumellei Photograph by o. Ou Puy

Neocoflettia Hemsl. 1890 Nesphostylis Verde. 1970

4 spp.; Africa (Za mbezian and Sudanian reg ions, 2 spfJ.); SE Asia (Indian
1 sp.; lndo-China (Myanmar [Burma]) and Malesia (Java)
subcontinent, Indo-China (M ya nmar], 2 spp.)

--
Named fo1 the plant collector Colonel Sir Herny Collett (1836-1902)
Anagram of Sphenostylis (q.v.), from which the genus was segregated
Herb (creeping); seasonally diy tropi ca l forest, wooded grassland and shru blancl
He1bs; seasonally dry tropical and subtropical woodland, wooded grassland and
Reference(s} Steenis (1960); Allen & Allen (1981: 455)
grassland
See note.') under Phylacium; the fruits develop undergrou nd
Reference(s): Verdcou1t (1970c: 296 -300) ; Tatcishi & Ohashi (1977); Potter &
... ~ . Doyle (1994); Munyenyembe & Bisby (1997)
Placed in subt ribe Phaseolinae, allied to Do/ic/Jos (Thulin el al, 2004)
Wajira Thulin 1982
5 spp.; NE Africa (4 spp. in the Somalia-M;isa i region); l sp. (W gmha111ia11a
(Wight & Am .) Thulin & Lavin) widespread in the Sudano-Zambezian region of Alistilus N.EJ3r. 1921
Africa, Saudi Arabia and India to Sri Lanka
3 spp.; Southern Africa (S Za rn bezian region, 1 s p.), SC and S Madagascar (2 spp.)
Na med afler lhe \Vajir district in E Kenya whe1·e the type collection of che genus
From ala- (L: wing) and stihts (L.: stake, pen), refening to the characteristic style
was made
in the genu s
Climbing herbs or subshru bs; seasonally dry tropical bushland, giasslancl,
Herbs or subshrubs; seasona lly dry tropical open woodland , xerophytic scrubland,
woodland and foiest
bushland or grassland, often associa ted with rocky outcrops
Reference(s): Thulin (1982); Thulin el al. (2004)
Wajira is basally branching and sister to the rest of subtribe Phaseolinae (Thulin ~ ~ l\eference(s): Vcrdcourt in Mackinder et a/ , (2001: 84); Du Puy & Labat in Du l'uy
el al. (2002: 561-565)
~
el al. , 2004)
Placed in subtribc l'haseolinae, closely allied to Dolichos and Lab/ab

Sphenostylis E.Mey 1836

Austrodolichos Verde. 1970
7 spp.; Africa (ma inl y Za rnbezian and Sudani an regio ns)
J sp N Australia
From spheno- (Gk.: wedge) and sly/is (Gk : style), referring to the shape of the
From australis~ (L.: southern), refe rring to 'Australian Dolichos'
distinctive cun ea te styl e
H e1bs; seasonaJly diy t1opi ca l woodi<in<l, scru b, grassland or wasteland, often on
Herbs or subshrubs; seasona lly dry tropi cal and subtropical open forest,
sand or alluvi al terraces
woodland, bushland and thicket, wooded grassland and g1assland
Reference(s): Verdcourt (l 970c: 399-400)
Reference(s): Verdcourt in Milne-Redhead & Po lhill 0971: 670-674); Lock (1989:
Auslrodolichos errabundus (M.13 ,Scott) Verde. is placed in subtribe Ph aseolinae,
438-439); Potter & Doyle (1994); Verdcourt in Mackinder el al (2001: 68-73)
possibly allied to Dolichos or Nesphoslylis (Verdcourt, 1970c)
Placed in subtribe Phaseolinae; basally branching in the Macrotyloma-Dolichos-
Nesphostylis clade (Thulin el al,, 2004)
Sphenoslylis stenocaipa (Hochst. ex A Rich) Harms is used as human food, the
seeds and lubers of the African ya m bean <I re eaten; othe r species are used as
ornamentals and for fib 1e, gu ms, resins and fornine foods ( Huxham et al, 1998)

Sphenastylls angustifolia Photograph by A. f. Van Wyk & s. Malan Austrodalichos errobundus Drawing by P. Halliday

TRIBE PHASEOLEAE 423

422 LEGUMES OF THE WORLD
 Dolichos fongista Photograph by D. Du Puy Mocrotylomo axillare var. glabrum Photograph by D. Du Puy Dipogon lignosus Photograph by 5. Andrews

Dolichos L 1753 Macrotyloma (Wight & Arn .) Ve rde. 1970

Chlo1yllis E.Mey. (1836) Kei>tingiel/a Harms (1908)
24 spp.. ; Africa (mostly Zambezian and Sudanian to Somalia-Masai regions; 23
c. 60 spp.; Africa (mostl y Zambezian ::md Sudanian to Soma lia-Masai and
spp.), Macaronesia, Arabia, Madagascar, Indian Ocean Islands; 1 sp~ endemic to
Afromontane regions; c. 55 spp.) and SE Asia (5 - 6 spp. in the Indian
lndian subcontinent, introduced elsewhere
subcontinent, S China, lndo·China and Malesia)
From macro- (Gk : large), tylo· (Gk,: callus) and lama (Gk,: fringe), refen ing to
F1om dolicho- (Gk.: long), re ferring to the long slender stems
Herbs or subshn..ibs; seasonally dry tropical and subt1opical fores t, wooclland,
the appendages on the standard petal
Herbs; seasonally dry tropical and subtropical woodland, wooded g111ssland,
bushland and thicket, wooded grassland and grassland
bushland, thicket and grassland, from swamps, rocky outcrops to dunes
Reference(s): Yerdcourt 0970c: 380-442; in Milne-Redhead & Polhill, 1971:
Reference(s): Veidcourt (1982a); Mackinder in Mackindel'el al (2001: 104-119)
674-696); Lock 0989: 391 - 397); Mackinder in Mackinder et al. (2001: 84 - 103)
Placed in subrribe Phaseolinae, sister to Do/ichos and Nesp/Jostylis (Thulin el al.,
Placed in subt1ibe Phaseolinae, allied to Macrotyloma and Nesphostylis (Thu lin el
al., 2004) 2004)
Used as pasture legumes (M. axil/are CE.Mey) Verde. (Archer Dolichosl, and M.
Used as ground cover, green manure, forage and shade plants; also as soap
unijlmum (L~un.) Verde. !horsegram]); also for fodder, green manure 1 medicine
substi tu tes (from the roots of some species)
and hum<in food (pulse) crops, including Kersting's groundnut, M. geoccopum
(Ha rms) Marecha l & Baudet

Dipogon Liebm. 1854

Verdcourtia R Wilczek (1966)
J sp.; South Africa (Ca pe), widely int1oduced in Austrnlasia, Asia and rhe New World
From di- (Gk .: two) andpogon- (Gk,: beard), for the 2-line bea rd on rhe uppe1'
half of rhe style
Herbs; warm temperate and mediterranean forest margins, scrub forest (Ind
sclerophyllous shrnbland (fynbos)
Refcrence(s): Verdcou rt (1970c: 406- 409); Marec hal et al. Cl 978: 247-248);
Stirton (198 lc)
Placed in subtribe Phaseolinae, allied to Lab/ab (Thulin et al., 2004)
Dipogon lignosus (L.) Verde. is used as a green manure, covet crop , ornamental
and for human food (pods)

Lablab Adans. 1763

1 sp.; Afri ca (mostly Z<1mbezian and Sudanian to Somalia-Masai regions) 1
Madagascar, Asia (Indian subcor:iti nen t), widely introduced elsewhere
From Arabic fo1 'rendril'. refen ing to the 1endril-like climbing stems; the word is
also supposed to mean 'play', the seeds are used in garnes
Herbs; seasona lly dry tropi cal and subtropical forest margins, bushland and
grassland, drought resistant, does well on poor soils
Refe rence(s): Yerdcourr in Milne-1\edhead & Polhill (1971: 696-699); Marechal et
al. 0978: 244-246); Verdcourt in Mackinder et ul. (2001: 80-83)
Placed in subtribe Phaseolinae, allied to Dipogon (Thulin et al., 2004); 3 subsp .
ancl numerous va1ie ties or cultivars a1 e recognised
Lah/ab pwpureus (L.) Sweet (hyacinth or bonavist bean) is widely cultivated in
the tropics for human food (vegetable or pulse), also for foddei', green manure,
ground cover and medicine
Dolichos filifofiolus Drawing by M. Grierson Lab/ab purpureus Photograph by 5. Col/enette

TRIBE PHASEOLEAE 425

424 LEGUMES OF THE WORLD
Spathionema kilimandscharicum Drawing by D. Erasmus Vatovaea pseudo/ob/ob Drawing by D. Erasmus Vigna caracatla Phatograph by J. Hooper Vigna vexi/lata Photagraph by 8. D. Schrire

Spathionema Tau b. 1895 Vigna Savi 1824

Voandzeia Tho uars (1806); Condy/oslylis Piper (1926); Azukla Owhi (1953);
1 s p.; E Africa (Somalia-Masai region)
From spathe- (G k.: spatula) and nema- (Gk.: thread), refeo·ring to the filamentous Haydonia R.Wilczek 0954)
c 104 sp p. (Vigna sens . slricl. p robably about c. 85 -90 spp.); Palaeotropics and
stamens which are spathulate at the apex
subtropics (c 80 - 85 spp., mostly in Africa le . 55 - 60 spp.; 4 spp endemic to
Climbing subshrubs; seasonally dry tropical mixed bus hl and to semi-dese rt
Madagascar) and SE Asia [c. 21 spp.)), c. 22 spp. in Neotropics and subtropics (but
wooded grassland
Reference(s): Verdcourt in Milne-Redhead & Polhill (1971: 664 -666); Marechal et see taxonomic notes)
Named after Dome nico Vigni ('1577 -1647), Professor of Botany at Pisa and ninth
al. (1978: 235 - 236)
curator of the Pisa Botanic Ga rden (rhe o ldest in Europe) which was founded in
Allied to Vatovaea (Thulin et al , 2004)
1543
He rbs; seasonally dry tropical woodland, wooded grassland and grassland, ofte n
in well drained sites with low fertility
Vatovaea Chiov. 1951 Reference(s): Verdcourt (1970d: 507-569; in Milne-Redhead & Polhill, 1971:
1 sp.; NE Africa (Somalia-Masai region) 617 - 664); Marechal et al. (1978: 160-231); Vaillancouot el al. (1993); Pasquer in
Named for A. Vatova (1897 - ?), Italian bota nist and collector o f the type species Mac kinder el al. (200 1: 121- 156); Tomooka el al. (2002); Maxted el al (unpubl.)
V. pseudo/ab/ab (Harms) J.B.Gillett The New World species of Vigna may be significantly redu ced with the re moval
Climbing shrubs; seasonally dry tropical bushland, grassland and semi-desert of subgenus Sigmoidotropis (c. 17 sp p., includ ing V. caraca //a (L) Verde ,
Refeoence(s): Verdcourt in Milne-Redhead & Polhill (1971: 609 -611); Mareclwl et illustoated above) w hich is polyphyletic with respect to the rest of Vigna (Delgado
al. 0978: 234-235); Thulin (1993: 433-434)
Salinas et al,, 1993)
Allied to Spathionema (Thulin el al., 2004) Many species are major pulse, vegetable, fodder and green manure crops, e g.,
Used for human food; the large tubers are edible (Huxham el al., 1998) V. angularis (Willd .) Ohwi & H.O hashi (azuki or adzuki bea n); V. mungo (L.)
Hepper (urd bean, black gram); V. radiata (L.) R.Wilczek (m un g bean, green
g1am); V. umbellata (Thunb.) Owh i & H Ohashi (rice bean); V. aconilifo/ia ()acq.)
Physostigma Balf. 1861 Marechal (moth bean); V. ung11.fculata (L.) Walp. (cowpea, ya rd long bean) and
c, 4 spp ,; Africa (G uinea-Congolian and Za mbezian regions) V. subterranea (L,) Ve rde . (bambara groundnut, bambara bean)
From physa- (Gk.: bladder) , in reference to the dorsal appendage on the stigma
H erbs or subshrubs; wet tropical swamp and riverine vegetation, to seasonally dry
fores t or open woodland and grassland Oxyrhyncbus Brandegee 1912
Re ference(s): Marechal et al. (1978: 232-233); Lock 0989: 42 1); Burkill (1995: Monoplegma Piper (1920); Peeke/ia Hanns (1920)
422-423); Verdcourt in Mackinder el al. (2001: 119-121) 4 spp.; N and C America (3 spp. from SUSA, Mexico, C America, Caribbean to
Placed in subtribe Phaseolinae, in a clade basa lly branchin g to Vigna (q v) Colombia) and SE Asia (1 sp from Papuasia and E Malesia)
(Thulin et al., 2004) From oxy- (Gk .: sharp) and rby11cho- (Gk. : beak), referring ro the beaked keel of
The seeds of the calabar o r ordea l bean (P. venenosum Balf.) contain alka lo ids
the Aower
and are toxic, but they have various medicinal uses; seeds of some species are Climbing herbs or shrubs; sea,onally d ry to wet tl'Opical and subtropical, coasral
common drift seeds
and 1nontane forest
Reference(s): Rudd 0967); Verdcourt (1978); Marechal et al. 0978: 242 - 243)
Genus geographica lly disjunct; placed in subtribe Pbaseolinae, allied to
Ramireze//a (11iulin el al , 2004)
Used for human food and forage

Physostigma mesoponticum Drawing bys. Ross-Craig Oxyrhynchus volubilis Drawing by E.5. w. (in herb K)

TRIBE PHASEOLEAE 427

426 LEGUMES OFTHE WORLD
 Ramlrezella mlcrantho Drawing by H. Ochoterena·Booth Strophostyles he/vola Drawing by P. Halliday
Phaseo/us vu/garis Photograph by U.S.D.A.

Ramirez ella Rose 1903

Phaseolus L. 1753 7 spp.: N Ame rica (Mexico and C America)
Minke/ersia M.Martens & Galeotti (1843); Alepidocalyx Piper (1926) Named for Dr Jose Ramirez of the lnstituto Medico Nacional de Mexico, expert on
c. 6o-65 spp; tropical, subtropical and warm temperate N, C and S America the natural history of Mexico
Cf1om C USA to Argentina); most concentrated in Mexico and C America and with Climbing herbs; seasonally dry tropical forest, secondary vegetation types and
c. 3 spp. in S America endemic to the N and C Andes and Galapagos (Delgado disturbed areas
Salinas, pers. comm.) Reference(s} Marechal et al (1978' 157 - 160); Ochoterena-Booth & Delgado
From phaselos (Gk , little boat), referring to the shape of the kidney bean Salinas (l 994)
Climbing herbs; seasonally dry to wet lowland and montane forest, thicket, scrnb Placed in subtribe Phaseolinae, allied to Oxyrhynchus (Thulin el al., 2004)
and wooded grassland, from hum id to desert conditions
Reference(s), Marechal et al 0978, 133-151); Delgado Salinas et al (1999);
Debouck (2000); Freytag & Debouck (2002) Strophosty l.es Elliott 1s23
Placed in subtribe Phaseolinae, allied to Ramirezetta (q.v.) and O>.yrhynchus
3 spp.; N America (E and C USA, SE Ca nada, N Mexico)
(q.v.) in Thulin et al. (2004) From stropho- (Gk., twisted) and sty/is (Gk,, column, style), referring to the shape
Used for human food as major pulse and vegetable crops; P. acutifolius A.Grny
of the style
(tepary bean), P. coccineus L (scarlet runner bean), P. flmatus L. (lima bean),
Herbs; subtropica l to temperate open dry woodland, wooded grassland, marshes,
P. vulgaris L (common or kidney bean) and the year bean, P. dumosus Macfad.
open grassland or d isturbed and other ruderal areas
(• P. polyanthus Greenm) are cultivated; also used for fodder, green manure and
Reference(s), Marechal el al. (1978' 241-242); Pelotto & Martinez (1998); Isely
as ornamencals
(1998); Riley-Huiting et al. (2004)
Placed in subtribe Phaseolinae, sister to Dolichopsis (Riley-Huiting el al., 2004)
Used for erosion control and wildlife food

Dolichopsis Hass\. 1907

1 sp ; S America (S Brazil, Paraguay, Argentina)
From opsis- (GL li ke), deno tes resemb lance of this genus to Dolichos
Herbs; seasonal tropical and subtropical low humid g1assland and swamps
Refe rence(s} Man~chal et al. (1978: 240); Burkart in Troncoso de Burka1t &
Bacigalupo (1937, 730-732)
Placed in subtribe Phaseolinae, allied to Strophostyles (Delgado Salinas et al.,
1999), and resolved in a c\ade together with Strophoslyles, sister to Mysanth11s and
Macroptilium (11rnlin et al., 2004; Riley-Huiting et al, 2004)

Dolichopsis paraguariensls Drawing by I. van Rentzell & A. Burkart

Phaseo/us /unatus Drawing by D. Erasmus

TRIB E PHASEOLEAE 429

428 LEGUMES OF THE WORLD
Mocroptilium lothyroides Illustration by M. Hort Mysonthus uleonus Drawing bys. Wickison

Macroptiliunt (Uen1h.) urb. 1928

c J7 .') pp ; N to S Americ1 from SUSA to Argcnlin:i, mainl y C'Once ntratcd in
tropical S Ame rica
From macro- (G k.: brge) andpti/on- (G k .: w ing), refcning to the w ing peta ls
w hich are brgel' th ~in che stan dard
Herbs; seasom1ll y d1y ti oplcal :ind subc ropica l disturbed fo1esl , woodland or
thi cket, :>cr'ubl::tnd ~incl g r..1 ssl::m d, o ften wced>'i 2 sp<::cies (below) widel y
n;Hlllalised in E and S t1 o pical Af1 ic 1
Hefere 11cc(s): Ma 1echal el al (1978' 15 1- 157); fevc rei1 0 (1987); Drewes (l '!97)
Plac~ c.I in subt1 ihe Ph;1seolin~w, in a clad e together Willi 111ysa11lb11s, Dolfchopsis
:ind Slropbos(J•les (Hi ley-Hulling el al., 200·1)

f~
ivfnjor pas ture and fodder- legumes, e.g ., ,1,r. ulrop11rp11re11m (DC.) Uib. (si1:1l10)
:ind ,\.f, lr11hyroides (L.) U1b. (phasy bea n)

-m · M)1santhus G.PJ.ewis &

I sp .; S America ( Bro zil)
A.Delga do 1994

From mys- (Gk .: mo use) and m11bos (G k : llowe1'), rcfoning lo the resemhbnce of
the llowers co the foci;1l c1ricalure of a l ~u ge-earcd mouse
Climb ing herbs; se1son;11ly dq• tropical wooded grnssland, chm n scrub, mcky
shrubbnd o r in was1c a1eas
Hcferencc(s): l.cwb & Delgado Salin:is ( 1994)
Placed in sub! t'i he Phaseolinae, allied to Macrop1ili11m (q ,v.) ( Del g~1do Salin;1.'> el
al., 19991, ::1nd sister eirhcr to~ Dolichopsis-S tm pl1oslyl es cl:1de o r to
J-Jacroj>lili11111 in 1hc "n:i lyscs of Hil ey-Hulling et o/. (2004)

Ory xis A Delg.,clo & G P.LelVis 1997

1 sp.; S Am e1i ca (l3r~ 1 7. il [M in~1 s Gc1ai!"))
From OIJ~\·is- (Gk.: cxcava rion rn digging), 1c fcrring 10 1his spc<.:ies being endemic
10 1hc Br.izlllnn tmc <if Mina" Co:rnl5 ( • •neml Mines) wh ch ha.• long lx.'Cn
r.11nOt1S for lhC CXlnl<,lon o f m:iny 1lli11crnfs
Climbing herbs OJ' subshrubs; sea sona lly d1y rropica l rock y wooded giassland and
shrublancl
Rele rcnce(s): Delgado Salinas & Lew is 0997)
Pbced in subtribe Phascolinae, but no l )'Ct sampled in m o lecub1r analyses;
re blio ns hips ai e suggested lo O.\yrby11 cb11s o r Dolichopsis (Delgado Salin<l~ 6-
Lewis, 1997) ;md lo 1l~pst11rtbt1s a nd Macr optilium (Ri ley -Hulling el al, 200 D

Oryxis montico/o Drawing by E. Catherin e

Strongylodon mocrobotrys Photograph by R.B.G., Kew

430 LEGUMES OF THE WORLD TRIBE PHASEOLEAE 431

tRIBE

p esmodieae byH.ohashi

'fribe Desmodi(!}t (Benth.) Hutch. 196

Tribe Hedysareae subtribe Desmodiinae Benth. (1965), as 'Desmodieae'
Tribe Coronilleae subtribe Desmodiina (Benth.) Schulze-Menz (1964)

'fhe tribe Derm cli~e as treat d by Ohashi et al. 19 1
compri ed 27 genera an I c. 540 pecie in thre
subrribe-, th Biyinae, De modiina and L spedezina .
Molecular analyses by Bailey et al. (1997) and Doyle et
al. (2000) show that Bryinae has affinities elsewhere;
Lavin et al. (2001a) place it within the Pterocatpus
clade of the Dalbergieae sens. lat. (see page 309). The
Btyinae are therefore removed from the Desmoclieae
here , as are two genera formerly placed in ubtribe
Lespeclezinae ; Phylacium Benn. and Neocollettia
Hems!., which are moved to tribe Phaseoleae (see page
393) on morphological, palynological and molecular
evidence (Doyle et al., 2000; Kajita et al., 2001). The
two remaining subtribes of Desmodieae are recognised
in this treatment as three groups, the Lespedeza,
Phyllodium and Desmodium groups, based on results
of an analysis of the chloroplast gene rbcl (Kajita et al.,
2001). The Phyllodium and Desmodium groups
correspond to subtribe Desmodiinae, and the
Lespedeza group to subtribe Lespedezinae (with
Campylotropis now comprising 37 instead of 65 species
as in Ohashi et al., 1981).
Desmodieae as circumscribed here comprises 30
genera and (524)-527-(530) species (Fig. 48). The
tribe occurs in the tropical, subtropical and warm
temperate regions of the world, but extends into the
cool temperate and sub-boreal regions of E Asia and N
America (except W of the Rocky Mountains). At generic
level subtribe Desmodiinae is most diverse in tropical
Sand SE Asia (Dy Phan et al., 1994), while temperate
E Asia (Yang & Huang, 1995) and N America (Isely,
1998) are the centres of diversity of subtribe
Lespedezinae. The tribe occurs widely from coastal to
montane areas, but not at high altitudes. Species are
most commonly shrubs or subshrubs, sometimes herbs,
rarely trees and are usually erect and 3-foliolate:
The Desmodieae have been considered similar to
tribe Phaseoleae (Polhill, 1981a) and were recently
shown to be a monophyletic lineage included within
Phaseoleae sens. lat. (Fig. 47, page 394), closely
related to subtribe Kennediinae (Doyle & Doyle, 1993,
Bruneau et al., 1995; Doyle et al., 1997) and possibly
sister to Mucuna (Bailey et al., 1997; Doyle et al.,
2000; Kajita et al., 2001) .

Phaseoleae sens. lat. (see page 394)

Lespedeza group
Campylotropis, Kummerowla, Lespedeza

Phyllodium group
Dendrolobium, Phyllodium, Ougeinia,
Aphyllodium, Ohwia, ?Hanslia,
Arthroclianthus, Nephrodesmus,
Tadehagi,.Aksch indlium, ?Droogmansia

Desmodium group
Monarthrocarpus, Trifidacanthus,
Desmodium, Codariocalyx, Hylodesmum,
Hegnera, Pseudarthria, Pycnospora,
Mecopus, Uraria, Christia, Alysicarpus,
?Desmodiastrum, Meliniella,
?Leptodesmia, ?Eliotis

? = placement uncertain in generic group since not analysed for rbcl

FIG. 48 Diagram of relationships in tribe Desmodieae

LEFT Lespedezo formosa Photgraph by B. D. Schrire

TRIBE DESMODIEAE 433

 Lespedeza thunbergii Photograph by B. D. Schrire Lespedeza maxlmowiczii Photograph by G. P. Lewis
Campylotropls polyantha Photograph by P. Cribb

Campylotropis Bunge 1835 Lespedeza Michx 1803

c. 35 sp p.; mainly China and E Asia (c 25 spp., a few extending to tropica l India
c. 37 spp ; China, the centre of d istrib ution, India to Inda-China (c. 35 sp p.) and E a nd Malesia); and N America (c. 12 spp., wi th about 30 putative natural hybrids);
Asia (2 spp., Taiwa n and Korea)
subgen. Macrolespedeza (c . 12 spp.) is con[ined in E Asia wit h the centre of
Fro m campy/a- (Gk., curved) and trop is (G k. , keel), 1e ferring to the keel petal
dive rsity in Japan and Korea
w hich is d istinctly incurved in contras t to th at of Lesp edeza
Na med for V. M. de Cespedes ( fi. 1784 -1790), Spanish gove1nor o[ E Florida and
Shrubs, seasonally dry tropical (mon ta ne) to te mperate fo rest, woodland or
sponsor of the French botanist Michaux; the genus name was m isspelled in
bushland
publicati on
Re ference(s} Nemoto & Ohashi 0 988; 1996); Barham 0997); Iokawa & Ohashi
Su bshrubs, sh rubs or he1 bs; temperate to seasonally dry tropical woodl and
(2002a, b, c ; 2003)
margins, bushla nd and grassland, in o pen places
Used as ornamentals, fo r fo 1age, bee-food and medicine
Re fe rence(s), Cle well (1966); Akiyama (1985); Ne moto et al. (1995); Nemoto &
O has hi (1997)
Bush clovers are used as g1 ee n manure and cover crops, highly este emed fornge
Kummerowia SchindL 1912 crops, human food and drink (i.e., as an alternati ve to tea , e g., L bicolor Tu rcz.),
Micr olespedeza (MaximJ Makino 0 914) and as bee-food ; also grown as orname ntals, e.g. L. ihunbergii (DC.) Nakai; L
2 spp.; E Asia, in troduced in N Arnerica and Austra lia j imcea (L.f.) Pers. va r se1icea (Thunb.) Lace & Hems!. (sericea Jespedeza o r
Named in honou1• of the Poli sh Professor, J. Kummero w from Pozna n
Chinese bush clover) is a noxious weed in USA
(Q uatt1 occhi, 2000), or possib ly of Paul Kumme r (1834 - 1912), Germa n clergyman
and lichenologist (S taneu & Cowan, 1979)
Annu al herbs, temperate open grassland or on roadsides
Refere nce(s} Aki yama & Ohba (1 985); Nemoto & Ohashi (1993); lokawa et al
(1996)
Used for forage, green manure and medicine

Kummerow/a striata Photograph by H. Ohashi Lespedeza thunbergii Illustrator unknown (ex Kew col/ecrion)

TRIBE DESMODIEAE 435

434 LEGUMES OF THE WORLD
Dendrolobium umbellotum Drawing by E. M. Stones Phyllodium pulchellum Drawing by P. Halliday Aphy/lodium glossocarpum Photograph by B. Maslin Ohwia caudata Drawing by P. Halliday

Dendrol-Obium (Wight & Arn) Bent h. 1852 Aphyll-Odium (DC.) Gagnep . 1916
Desmodium subgen. Dendrolobium Wight & Arn. (1834) Dicerma DC. (1825)
18 spp.; India (3 spp.) to Japan (I s p.) and <1bunda nt in Indo-China (9 spp ) and 7 spp .; 6 spp. N Australia and Papuasia (New Guinea); 2 s pp. Malesia , S China
Malesia to Australia (6 spp) (Hainan), Indo-China, India and Sri Lanka
From dendrrm (Gk : tree) and /obion (Gk.: pod), with reference to s mall t1ees A compound of a- (GL without), phy/lon (Gk : leaf) and -odium. (L.: or -odio11
with woody pods Gk.: implying both li keness and smallness), 1eferring to the abse nce of leafy b1acts
Trees or shru bs, rarely he rbs; seasonall y d1y trop ica l forest, wood land, bomboo as found in Phyllodium
thicket or grassland Shrubs o r subsh rubs; seasona lly d 1y tropical forest, woodland, wooded g1assland
Reference(s): Ohashi (1973a; 1997a; 1998; 2004<1); Ohashi er al. Cl999b) and grnsslancl, often in sanely and open areas
Used for medicine Reference(s): Ohashi Cl973a; 1997b; 2004a); Ped ley 0996; 1999a)

Phyllodium Desv. 1813 Ohwia H.Ohashi 1999

8 spp.; India, SE & E Asia, a few in N Australia Ca re11aria IJenth (1852), nom. illeg; Desmodium subgen. Calenaria (Ilenth.)
From pbyllon (Gk.: leaf) and -odium (L: 01· -odion Gk.: i111plying both likeness Baker (1876)
and small ness), refen ing to the bra cts o f the in001escences which a1e like 2 spp ; India, lndo-China, Malesia, China (2 spp) lo Japan
small leaves Named in ho no ur of the Japanese plant taxono111ist, jisaburo O hwi 0905- 1977)
Shrubsi seasonally dry uopica l forest, wood land, thicket, open scrub or g1:issland Shrubs; seasonall y d1y tropical forest, wood land and gr:is,land, often o n open
Refere nce(s): Ohash i Cl973a; 2004b) hillsides
Ohashi 0973a) deli111its two subgenera in Pby/lodittm References' Ohashi 0973a; 1999; 2004b)
Used for medicine Used for medicine and reputed to be an inseccicide :.igainsl maggots

Ougeinia Benth (1852) Hanslia schindl. 1924

Desmodium subgen. Ottgeinia (Benth.) H.Ohashi (1973) Desmodlttm subgen Hanslia (Schindl.) H.Ohashi (1973)
1 sp,; lndi<1 and W Nepal 2 spp.; Males ia, Papuasia (centred in New Guinea), Vanuatu <i nd Australia (N
Nmned afte r the ancien t town of Ougeine (now Uj jain) in Madhya Pradesh, India; Q ueensland)
seeds of this species were sent to the Botanic Ga rd en in c,dcutta in 1795 by Dr Named for a close friend, Hansli Hegner
\Xlilliam Hunter, the surgeon to the Resident at Ougeine Sh 1ubs; tropi ca l wet and seasonally chy forest, disturbed areas and thicket, or
Trees; seasonally dry tropical forest, woodland or on o pen rocky slopes along roadsides
Reference(s): Ohashi 0973a) l\eference(s): Ohashi 0973a; 2004b)

Ougeinia aojeinensis Drawing by P. Halliday Hans/la ormocarpoides Drawing by P. Halliday

436 LEGUMES OF THE WORLD TRIBE DESMODIEAE 437

 ~
@ 1

'

Akschindlium godefroyanum Drawing by P. Halliday Draogmansio pteropus Photograph by D. Laing

Arthroc/ionthus deplonchei Drawing by P. Halliday Nephrodesmus sericeus (1-8), and N. al bus (9) Drawing by P. Halliday

Arthroclianthus Baill. 1870 Akschindlium !-! ,Ohashi 2003

1 sp.; Inda-China
c. 30 spp. (Nielsen, pe1> comm., 2003); endemic to New Caledonia (J SJ»
Named in ho no ur of A.K. Schindle r (1879-1964), German botanist
extending to Va nuatu)
Subshrubs or shrubs; seasom1l\y d ry tropical to subtropical forest, scrub and
From a11bron (Gk.: a joint) and Clianrbus (a legume genus derived from cleos
g rassland, in rocky areas or dve1ine
(Gk, glory) an d anthos (Gk, ~ower) alluding to its beautiful ~owers), with
Reference(s} O has hi (2003)
reference to ril e clianthu s-like flowers and large laments of this attra clive genus
Transfe1reel from Tadehagi as A. godefroya num (Kuntze) I·I Ohashi
Shrubs or trees; seasonall y dry r1·opical forest, woodland or scru b, on hillsides and
rocky ridges or riverine
Reference(s} Hochreutiner (1909); Schindler 0926)
Accommoclated in tribe Hedysareae sens. strict. (Hut c hin son, 1964)
Droogmansia De Wild. 1902
c~ 5 spp., or often estimated at over 20 spfJ.; SC to W Zarnbezian to Sudanian
Africa
Nephrodesmus schindl . 191 6 Na med for Hubert Droogmans (1858-1938), Financial Secretary General frwthe

'
13elgian Congo
6 spp.; endemic to New Cnledonia
Shrubs or subsbrubs; season:llly dry trorical plateau to mon t:ine woodland,
F1om neph ro- (GL kidney) and desm os (GL chain); with referen ce to the Joment
wooded g1assland, bushbnd or seasonally wet grassland
Shrnbs or trees; seasonal tropical scrub or wooclbnd, on hillsides and roc ky ridges
Refe1e nce(s} Lock (1989' 248 - 250); Verdcourt (2000, 27- 34)
Closely related to Arlhroclianlhus
Ve rdcowt (2000) notes that species arc very poorly defined and based on spa rse
Reference(s), Allen & Allen (1981: 459)
material; only a very rew true species may be 1ecognisable w hen abund ant
material is available; the lnclo-Chinese species D. godefroyana (Kuntze) Sch indL
was moved to Tadehagi (Ohashi, 1973a), and then to Akschindliwn (Ohashi ,
Tadehagi H.Ohas hi 1973
2003) A genus much in need of revis ion
c. 6 spp.; Jndi;i to Inda -China (5 spp.; centre of distribution); China, E Asia,
Used for fomge
Malesia, SW Pacific and N Ausnalia (I sp.)
Derived from the Japanese name of th e genu s
Subshrubs or shrubs; seasonally dry tropical to subtropical forest, scrub an cl

1
Monarthrocarpus Merr 1910
grnssland , in rocky a1eas or iiverine
Desmofischera Holthuis (1942); Desmodium section Monarlbrocmpus (Me ro°.)
Reference(s), Ohashi (1973a; 2003; 2004b)
H O hashi (1973)
O hashi (2003) delimits 2 subgenera in fodehagi
l sp.; E Malesia (S ulawesi, Philippines, Moluccas) and Papuasia (New Gui nea)
Used as folk lore medicine in Chi n£i
From mono- (Gk ., one), m1hron (GL joint), and cwpos (G k,, fruit); with
reference to th e single articled lament
Shn1bs; seasonally dry tropical fo rest and wood land
Reference(s} O hashi (1973a) as Des111odh1111 (M sewriformis (Benth.) Merr.l;
Ohashi (2004b)

fl
v J.r:.

Tadehagi triquetrum subsp. auriculatum Drawing by F. Crozier Monarthrocarpus securiformis Drawing by P. Halliday

TRIBE DESMODIEAE 439

438 LEGUMESOFTHEWORLD
 Oesmodium vorgosianum Photograph by G. P. Lewis Codariacolyx motorius Drawing by P. Halliday Hylodesmum repondum Photograph by a. Du Puy

Trifidacanthus Merr. 191 7 Codariocalyx Hassk. 1842

I sp; Malesia (Lombok and Flo1es, Philippines (Luzon)), Inda-China (S Vietnam) 2 spp.; Sri Lanka, India, Inda-China, Ma lesia, Ch ina and Ta iwan; C. gyroides
and S China (Hainan) (Roxb. ex Link) Hassk. introduced in tropical C Africa
Fro m M- (LJ Gk,: three), -fidus (L: divided) and acantbos (Gk.: thorn); 1efeJ1ing From coda1ion (G k.: little fleece) and calyx, 1eferring to the early cadu cous bracts
to thorns of th e species w hich, like a fleece removed, reveal what is underneath, in this case the nowe1s
Shru b or smal l tree; seasonally dry tropical forest to open scmb or grassland, on that are still in bud
rocky ridges and riverine Shrubsi seasonally dry lowland co montane tropica l forest, foresc margins, open
Reference(s): Ohashi et al. (1996); Ohashi (2004b) scrub an d grassland, on slopes or riverine
Conside red closely related to Desmodium (Ohas hi et al, 1996) Refe rence(s): Ohashi Cl973a; 2004a)
Used as ornamenta ls (for leaAet movement), medicine and for forage

Desmodium Desv. 1813

Mw1onia Craib 0912) Hywdesmum H.Ohashi & R.R.Mill 2000
c. 275 spp.; most diverse in SE Asia (at infrageneric level) and Mexico to S Desmodium subgen . Podocarpium (Benth .) H Ohashi 0973)
America (a t specific level). Occurring from Africa-Madagascar (c 40 spp , c 15 Podocarpium (Benth ,) Y.C.Yang & P.H.Huang 0979)
endemic), trop ical SE Asia (c. 70 spp.), China to temperate E Asia (c 35 spp , c, '14 spp; India through China (IO s pp., cenri e of distribution) to E Asia , and east
JO endem ic), warm temperate N Ame1ica (c. 35 spp.), Mexico (c. 80 sp p" c 50 N Amelica (3 spp.), 1 sp. extending to Africa (H. repa1Uium (Yah l) H.Ohashi &
e ndemic), C Ame1ica, Caribbean and tropica l to subtropical S America (c. 80 spp., R.R.Mil))
21 spp fou nd in Argentina) and Austra lia (c. 14 spp.). A number of species are From byte- (Gk.: fo rest) and desmos (Gk.: chain, refe rring to an abbreviated fo 1m
w idespread in the Old and New Worlds and are often cultivated of Desmodi11m)
from desmo- (Gk : chain) and -odium (L: or -odion Gk : implying both likeness Herbs or subshubs; seasonally dry, often montane, tropical to temperate forest,
and smallness), referring to the fruits being like a small chain forest margins, wood land, or grassland, also along roadsides
Shrubs or herbs, rarel y s1nall trees; seasonally dry to wet tropical , warm ten1pe.rate Reference(s): Ohashi Cl973a) as Desmodi11111; Yang & Huang 0979); Kajita &
a nd temperate forest, wood land, th icket, wooded grassland, bushland and Ohashi (1994; 2001); Ohashi & Mill (2000); Ohash i (2004b)
grasshmd, usually co mm on in open or seasonally wet and riverine areas, also in Some species used as medicine
disturbed and ruderal vegetation
Reference(s): Ohashi Cl973a; 1982b; 1985; 2004a); McVaugh (1987); Ped ley
Cl999a); Verdcourt (2000: 1-24); Va nni (2001) Hegnera Schindl. 1924
Important livestock forage and cover crops (e g, the rick clovers D int01111m 1 s p.; Inda-China (includ ing Myanmar (Ourma)) and Malesia
(Mill ) Urb and D. uncinalum Qacq.) DC.), also grown as intercropping insect Etymology as for Hanslia, i.e. named after a close friend Hansli Hegne r
repellents; w idely used for medicine; cultiva ted as ornamentals (e ,g., D elegaus Twining subshrub, tropical seasona lly dry to everg ree n forest, in swamps, old
DC.), and used for fibre cul ti vated areas or open places
Re ference(s): Ohashi (1973a; 2004b)
Re lated to Hylodesmum (Ohashi Cl973a)

Oesmodlum ve/utinum Photograph by D. Du Puy Hegnera obcordoto Drawing by P. Halliday

440 LEGUMES OFTHE WORLD TRIBE DESMODIEAE 441

Pseudarthrla conferti(lora Photograph by M. Gilbert & M. Thulin Urarla crinata Photograph by H. Ohashi Urarla picta Drawing by D. Erasmus
Pycnospora /utescens Photograph by J. Murata

Pseudarthria Wight & Arn. 1834 Uraria Desv. 1813

Uraiiopsis Schind l (1916)
c. 3-4 spp.; Africa (2-3 spp ), India and Malesia (I sp .)
c. 20 spp,; India, Indo-China, S China, Taiwan , Malesia, and two extending to
From pseudo- (Gk.: false) and artbron (Gk.: joint), re ferring to the frui ts wh ich are
Africa with the centre of d iversity in India to Indo-China
somewhat impressed but not anicu lated
From ura- (Gk.: ta il), referring to U. p1:c1a (Jacq DC , wh ich has long tail-like
Herbs, subshru bs or shru bs; seasonally dry tropical fo rest margins, grassla nd,
in florescences
open or disturbed areas
Subshmbs or shrubs, seasonally dry tropical woodland 0 1· grassland
Reference(s): Verdcourt in Milne-Redhead & Polhill (1971: 483 - 487); Lock (1989:
References: De Haas el al (1980); Yang & Huang (1981); Dy Phon Cl987a);
250-251)
With furth er research traditional estimates of 4-6 spp. in Pseudarthn·a may be Verdcou11 (2000: 34 -35)
Some sp ecies used for medicine; seeds used in human food
reduced; a number of African species may be better treated as infraspec ific
variants of P. bookeri Wight & Arn.; this genus was placed in cribe Pseudarthrieae
(Hu tchinson 1964) o r subtribe Pseuclarthriinae of tribe Desmoclieae (Praminik &
Thothathri, 1989) Christia Moench 1802
Used as medicine Lourea Neck ex Desv (1813)
c. 10 spp.j Jndia ro China, Malesia and Aus1ralia 1 a nd most diverse ln Indo-China
(6 spp) and Ch ina (5 spp.)
Pycnospora R.13r. ex Wight & Arn. 1834 Na med for the Ge rm an botanist and clergyman, J.L. Christ (1739 -181 3)
Herbs; seasonally dry tro pica l grassland and scrub, also on roadsides
1 sp.; Africa (Somalia-Masai and Lake Victoria regions), India, SE & E Asia and
Refe rences: Dy Phon (1987b); Polhill (1990: 109); Barham 0996)
Australia
From pycno- (Gk.: dense) and sporos (G k,: seeds), re ferring to the many-seeded Used fo r medicine
fruits
Diffuse herb or subshrub; seasonally dry tropical woodland, bushland, scrub and
grassland, either in swampy areas or on rocky hillsides
Reference(s): Verdcourt in Milne-Redhead & Polhill (1971: 481-483); Allen & Allen
0981: 573)
Used as a green manure

Mecopus Benn. 1840

l sp.; India, lndo-China, S China (Hainan) and Malesia (Java)
From meco- (Gk.: length) and podo- (Gk.: foot), referring to the stipe at the base
(foot) of the pod that elongates and which, together with the recurving pedicels,
resul ts in the deve loping pods being pushed back into the dense heads of the
inflorescences
He rb or subshrub; seasonally dry tropical forest or shrubby grassland on hills, also
along roadsides
Reference(s); Hutchinson (1964, 482); Allen & Allen (1981: 423)
Used for medicine

Mecopus nidulans Drawing by P. Halliday Christia vespertillonis Illustration by C. King

TRIBE DESMODIEAE 443

442 LEGUMES OF THE WORLD
Alys/carpus glumaceus Melliniella micrantha Drawing by unknown artist
Leptodesmia - various species Drawing by M. Tebbs
Photograph bys. Collenette Alys/carpus pubescens Photograph by M. Sanjappa

Alysicarpus Desv 1813

Melliniella Harms 1914
1 sp ; WC Africa
25-30 spp.; Africa Cc JO spp , c, 5 endemic), India, Inda-China, Malesia, China, E
Named for Lt. Mellin, who collected the species in Togo
Asia and Japan (c. 20 spp.) and Australia (8 spp., c. 2 endemic); endemic species
Herb (mat-forming); seasonally dry tropical wooded grassland, often on laterite
are mostly in India (15 spp)
plateaux or rocky outc1ops
From hylysis (Gk.: chain) and catpos (Gk,: fruit), refefl'ing to the segment.' of the
Reference(s): Hutchinson 0964: 360); Allen & Allen (1981: 430); Lock (1989: 250)
pod
Potential for livestock fodder
He rbs, seasonally dry tropical woodland, wooded grassland and grassland, often
in seasonally wet or open sandy and rnderal areas
Reference(s): Endo & Ohashi 0990); Pok le (1999); Verdcourt (2000: 36 - 43);
Pokle (2002); Adema (2003b)
Leptodesmia (Benth .l Benth. 1865
Used for livestock fodder, medicine, green manure and as a cove1 crop and 3 spp.; Madagascar, 1 sp. also in India
famine food From lepro- (Gk.: slender) and desmos (Gk : chain), probably with reference to its
more slender habit than Desmodlwn
Herbs or subsh rubs; seasonally dry tropical woodland, grassland and cultivated
Desmodiastrum (Prain) A.Pramanik & Thoth. 1986 lands, oflen in moist open areas and rocky oulcrops
Reference(s): Du Puy & Labat in Du Puy el al. (2002: 626 - 629)
Alyslcmpus subgen. Desmodiaslrum Prain (1897)
Used for ca ttle forage 1 and as a medicinal drink by humans
4 spp.; India and Malesia (E Java)
From desmo- (Gk.: chain) and -astrum (L : incomplete resemblance), with
reference to its similarity to Desmodium
He rbs; seasonally dry tropical (often moist) hillside grassland
Eleiotis oc. 1325
Reference(s): Pramanik & Thothathri (1986); Ohashi (2004a) 2 spp.; India, Inda-C hina (Myanmar) and Sri Lanka
Sanjappa 0992; unpublished) maintains this grou p of species under AlysicatjlllS from eleios (Gk: dormouse) and otio- (Gk ,: ear), referring to the shape of the
leaflets
Prostrate herbs, seasonally dry tropical forest and woodland, in open grassy or
rocky areas
Reference(s) : Hutchinson (1964: 487); Allen & Allen (1981: 261); Sanjappa
0992: 171)

Desmodiastrum belgaumense Drawing by P. Halliday E/eiotis rottleri Drawing by P. Halliday

TRIBE DESMODIEAE 445

444 LEGUMES OFTHE WORLD
TRIBE

psoraleeae by C. H. Stirton

Tribe Psomleeae Lowe 1862, as 'Psornleae'

'fribe G:ilegeae subtribe Psoraleinae A.Gray (1863), as 'Psoralieae'

'fhe tribe Psoraleeae as delimited by Stirton (198la),
comprised 6 genera and c. 135 species, and until very
recently (see below) the tribe has been of uncertain
lineage. Key revisionary studies since 1981 have been
on Otholobium (Stirton, 1989), the New World members
of Psoraleeae (Grimes, 1990), and Cullen (Grimes,
1997). Hallia was subsumed into Psoralea on the basis
of data from inflorescence and flower morphology and
leaf anatomy (Tucker & Stirton, 1991; Crow et al., 1997).
Lectotypifications of infrageneric taxa in Psoraleeae
were made by Grimes (1988). As treated here, the
Psoraleeae comprise a monophyletic group of 9 genera
and 185 species (Fig. 49). The only generic problems
remaining to be resolved are a) the generic position of
Bituminaria acaulis (Steven) C.H. Stirt. (Stirton, 198lb;
Grimes, 1997) currently included in Bituminaria
subgenus Christevenia Barneby ex C.H. Stirt.; and b) the
status of eight species of South American Andean
psoraleas included by Grimes (1990) in Otholobium, an
otherwise southern and eastern African genus.
Prior to 1977 (Stirton, 198la), the Psoraleeae was
considered closely related to Amorpheae. Evidence
from a range of morphological , anatomical, floral
development, phytochemical, nodulation and recently
molecular studies, however, show that Amorpheae are
basally branching in dalbergioid legumes (Lavin et al.,
200la; Wojciechowski et al., 2004), whereas Psoraleeae
are nested within the Phaseoleae sens. lat.
The Psoraleeae are sister to Phaseoleae subtribe
Glycininae (see Figs. 47 & 49) in a well supported clade
based on rbcl sequences of Otholobium and
Bituminaria (Doyle et al., 1997). In addition Cullen
(but cited as Psoralea) is part of a fully supported clade
with other Phaseoleae based on trnK-matK data (Hu,
2000; Hu et al., 2000). Adams et al. (1999) and Doyle
& Doyle (2000) indicate a similar result using data from
the respiratory nucox-II gene (encoding subunit 2 of
cytochrome oxidase). Psoraleeae are placed sister to
Glycine with the basally branching Cullen sister to
Otholobium, Psoralidium and Rupertia in the matK
analysis of Wojciechowski et al. (2004). More sampling
of Psoraleeae is needed in molecular analyses to
ascertain if the basally-branching genera are the
southern African Otholobium and Psoralea (as
suggested by Grimes, 1990) or Cullen (e.g., in
Wojciechowski et al., 2004).

Phaseoleae subtribe
Glycininae (see page 394)

0tholobium

Psoralea

Orbexilum

Hoita ""ti
(J")
0
;;o
Rupertia )>
r
rn
rn
)>
Psoralldium rn

Pediomelum

Biturninaria

Cullen

FIG. 49 Diagram of relationships In tribe Psoraleeae after Grimes (1990)

LEFT Psorolea pinnata x P. aculeata Photograph by c. H. Stirton

TRIBE PSORALEEAE 447


Otho/obium thomii Photograph by B. E. Van Wyk
Orbexilum onobrychis Drawing by P. Halliday
448 LEGUMES OF THE WORLD
Hoita macrostachya Photograph by G. P. Lewis Rupertia physodes Drawing by P. Halliday
Psoralea tenuissima Drawing by A. J. Beaumont
Otholobium C.H. Stin 1981
Hoita Rydb_ 1919
3 spp ; W USA (California) to N\V Mexico (Baja California)
61 spp; 53 spp. Sand E of sout hern Africa (Cape "nd Afromontane regions,
From the Indian name frn the type species, the fib1'e o f wh ich has been usecl
especially common in meditenanean Sand SW Cape, with outliers to Angola ancl
for threads
E mountain areas of Kenya); possibly including 8 spp. from S Ame1ica, in th~
Herbs; meditemrnean lowland to monrane woodland, sh1ub hrnd, grassland, wet
middle to high altitude Andes 1 from Columbia and Venezuela southwards to Chi le
places and open disturbed habitats
and Argentina (Grimes, 1990)
Refe1ence(s): Grimes (1990)
Prom otho~ (Gk., push) and lobio1t (Gk , pod), referring to the distinctive way
Uses include edible leaves, soil stabilisers, fibre and medicine
pods 'push out' from the accrescent calyces <luting development
Shrubs, trees or herbs; seasonally d1y tropical and mediterranean lowli!ncl to
mo nrane shrubland, forest margins, grassland and seepage areas (Andenn species
in montane forest, woocllomd, shrubland and xeric grassland)
Rupertia.J.W.Grimes 1990
Reference(s): Stirton (1986a; 1989; 1991); Giimes 0990); Stirton & Schutte in 3 spp.; w USA (Va ncouver Island to California) and NW Mexico (Baja California)
Named after the distinguished legume raxonornist Rupe1t C. Bameby (1911 - 2000),
Goldblatt & Manning (2000' 500 - 503); Verdcourt (2000, 46-48)
Closely allied to Psora/ea; relationships re1nain unclea1· between the African rnc1nbers see also genus /Jarnebye//a on page 482
~
of the genus and the e ight species from the Andes in S Ame1irn (G rimes, 1990) He rbs; warm temperate to meditenanean wood land, shnibland and g1assland
Used as med icine and for Aavouring food by smok ing Referencc(s), G!imes 0990)
Psoralea L. 1753
~ P1eviously included in Hoila by Rydberg (1919) but differs in its non-accrcscent
calyx and unique secondary inte1rn1l wall of the fruit
Hr;/lia Thunb. Cl 799); Lotodes Kuntze (1891) pro pa11e Used for medicine
c. 50 spp.; S and E of sou thern Africa (Cape and Afromontane regions, especially
common in mediterranean S and SW Cape, wilh fewer species in E parts to
Zimbabwe); outliers in Angola and St. Helena Is land (now extinct) Psoralidium Rydb. 1919
From psoraleo- (Gk,, scabby), referring to the calyx and leaves dotted w ith bbck 3 spp.; widespread in N Amelica in the Great Plains and co rd ille1an regions of S
or trnnslucent glands Canada to N Mexico
Shrubs, trees or herbs; seasonally dry tropical and meclitetTanecm lowland to Frompsora/eo- (GL scabby) and -idion (Gk,, diminutive) , 1efe11ing to its affinity
montane forest margins, shrnbland and grassland, in wetter habitats than Otho/obi11m with the genu.s Psora/ea
Heference(s): Fotbes (1930); Sti1ton (1989); Stirton & Schutte in Goldblatt & Herbs; continenwl and wa1·m temperate woodland, grassland and desert sc rub, on
Manning (2000: 505-507); Ve1dcou1t (2000: 48- 50) rocky slopes or in sandy a1·eas, dunes and alluvi<1l plains
The genus comprises many rare, little-known and undesc ribed species; Reference(s): Gr imes Cl 990)
monograph in preparation by Sti11on Grimes' (1990) concept of Psoralidium is more rest1ictive than that of Rydberg
Used as ornamen tals (1919) and the species limits aie broader
Used as a medicinal snuff
Orbexilum Haf. 1832
Rhytidomene Rydb (1 919)
8 spp., E USA from Virginia to Flrnicla, westwards through Kans<1s, Oklahoma Jnd
Texas; Mexico (south to Chi apas)
Rafinesque names are frequently obscure without obviolls de1ivation, bw in this
case he notes the rounded vexi llum petal as a distinguishing character of the genus.
If, however, he intended a combination based on orbiculan's - (L : circular) and
vexlllum (L' standard petal) then the suffix of the generic name is to be co nside1ecl
a misspelling
Herbs; wann tempe1ate woodland and g1ass land; montane in Mexico
Reference(s): Grimes (1990)
Used fo r medicine Psoralidium /anceolatum Drawing by P. Halliday
TRIBE PSORALEEAE 449
Pedlome/um cyphocalyx Drawing by P. Halliday Bltuminoria bitumlnosa Drawing by M. Y. Saleem

Pediomelum Rydb 1919

Psora/ea subge n Pediome/um (Rydb.) Ocke ndon 0965)
21 spp.; SC Canada, USA (Greac Plains and Great Basin) to C Mexico
From pedio- (Gk.: p lain) and mefo- (Gk : apple), equivalent to ics Fre nch
vernacu lar narne 'pomme de prairie'
H erbs; wa rm and continental temperate woodland and grassland to barren areas
in desert
Reference(s): Grimes (1990); lsely 0998)
Roots of P. escufentum (Pursh) Rydb (piairie turnip or breadroot) are eaten; also
used for medicine

Bituminaria Heist. ex Fabr. 1759

Aspa//hium Meclik. (1787)
2 spp; Maca ro nesia, Medite1ra nean (S Eu1ope, W Asia and N Africa), Tu rkey to
Caucasus
From hituminosus (L: of pitch), refeiring to the ta1-like smell of che planes
Herbs; mediterranean lowland to subalpine woodland, shrubland a nd grassland,
ofren in open dis1urbed areas or on rocky slopes
Re fere nce(s): Stirton (1981b)
Used as ornamen1als, forage and for medicine

Cullen Medik. 1787

Meladenia Turcz. (1848); Bauerel/a Schindl. (1926); Baueropsis Hutch (1964)
c. 34 spp.; mostly N\XI' but also C to E Australia (23 spp.), 1 sp. also to Papuasia
and Philip pines; I sp. Indonesia; 1 sp. Malesia, S China to India, Oma n and
Somalia; 2 spp. Mediterranean; 3 spp. \YI Asia, c. 3 spp soutl1ern Africa (mostly
Ka lahari-H ighveld region to Na ma-Karoo biome)
Possibly named after William Cullen (1710-1790), Professor of Botany, Glasgow
Herbs, shrubs and (rarely) trees; seasonally dry tropical to warm temperate forest
underscory, dry scrub, grassland or desert vegetation, mostly in open sanely or
seasonally wet places
Refe1ence(s): Sti11on (J981b); Grimes 0997); Verdcourt (2000: 43 - 45)
Livestock find some species palatable while others are toxici used for medicine

Cullen abtusifallum Photograph by B. E. Van Wyk Otholabium mexicanum (Andean species) Ph otograph by G. P. Lewis

450 LEGUMES OF THE WORLD TRIBE PSORALEEAE 451

TRIBE
Sesbanieae by M. Lavin and B. D. Schrire
Tribe Sesbanieae (Rydb.) Hutch. 1964
Tribe Galegeae subtribe Sesbaniinae Rydb. 0923)
Polhill & Sousa 0981) recognised 21 genera and c. 145 Robinieae (e.g., Lavin et al., 2003), or to the Loteae
species in tribe Robinieae (including Sesbanieae). This (incl udin g Coronilleae) clade with both being
treatment, following modifications of Lavin (1988, strong ly supported as sister to the Robinieae
1993, 1995, unpublished data) and Lavin & Sousa (Wojciechowski et al., 2004) . This, in addition to the
0995), recognises a separate tribe Sesbanieae (Fig. 50), putative position of Sesbania as the sister clade to the
comprising one genus, Sesbania, with c. 60 species. rest of Robinieae in the morphological analysis of
Recent molecular phylogen ies of robinioid Lavin & Sousa 0995), warrants its segrega tion here
legumes suggest that Sesbania is either sister to as a distinct tribe.
Sesbania macrantha var. macrantha
Loteae incl. Coronilleae
(see page 455)
Sesbania SESBANIEAE
Robinieae
(see page 467)
Inverted-Repeat-Lacking
Clade (IRLC)
FIG . 50 Diagram of relationships of tribe Sesbanieae after Lavin & Sousa (1995); Lavin et al. (2003)
Sesbania tomentosa
452 LEGUMESOFTHEWORLD
Photograph by C.S./.R.O.
Photograph by R.B.G.. Kew
co
CVJj~~
- 9 o~ ..,,.
·)f__.,__.""-
Sesbanio quadrata (1-10) Drawing by L. M. Ripley
Sesbania Adans 1763
Agati Adans. (1763); Glotlidium Desv. (1813); Da.,bentonia DC . (1825);
Daubentoniopsis Rydb. (1923)
c. 6o spp.; Africa-Madagascar (c. 30 spp.); Asia to Pacific (c. 9 sp p .); Australia (c. 7
spp.); 2 spp . widespread in Palaeot1 opics; New World (c. 9 sp p. USA to Argentina,
but mostly N & C America); c. 3 sp p . pantropical
Derived from the Arabic name of the plant, sisaban or sesaban
Herbs, shrubs or trees; seasonally dry tropical, subtropical and wa 1m temperate
areas, in seasonally wet, flooded or swa mpy habita ts; riverine forest, woodland,
wooded grassland and grassland, on lake ma1gins, river banks and in coastal
areas
Reference(s): Gillett (1963b); Bu i bidge (1965); Lewis (1989a); Du Puy & Labat in
Du Puy et al. (2002: 444-449); Lewis in Brummitt et al. (submitted)
four sections are recognised and o nly in warm temperJte to tropical N America
are all four represented
Used for forage, fibre (frnm bark), wood, pulp & paper, dye, gum, cover crops,
gree n manure, hu man food (e.g., flowe1 s of S gra ndiflora (L ) Pers,), medicine,
ornamentals and fish poiso ns (useful species include S grandijlora [agati or
corkwood tree]; S bispinosa (jacq.) W.Wight [dhaincha] and S. exa/tata (Raf.)
Cory [Colo1ado River hernpD; some species are invasive weeds and are toxic to
livestock (e .g., S punicea Benth)
TRIBE SESBANIEAE 453
TRIBE
Loteae byo. o. sokotoffandJ. M. 1.ock
Tribe Loteae DC. 1825
Tribe Coro nilleae (Bronn) Burnett (1835)
Subtribe Coronillinae Bronn (1822), as 'Coronilleae'
Tribe Anthyllideae Buchenau (1894)
The Loteae have been usually considered as the
closest relatives of other temperate tribes, especially of
the astragaloid part of Galegeae (e.g., Polhill, 1981k:
371- 374), and the monospecific genus Podolotus was
either allied with Lotus, or merged with Astragalus.
Recent molecular data, however, have revealed that
Galegeae, Cicereae, Hedysareae, Trifolieae, Fabeae
and Millettieae (in small part) lack the chloroplast-
DNA inverted repeat (IR) which is present in the
majority of Leguminosae including Loteae (Liston,
1995). A study of the chloroplast gene rbcL also placed
Loteae and other temperate tribes in different clades
(Doyle et al., 1997), the Loteae being in a robinioid
clade and the temperate tribes in an Inverted Repeat
Lacking clade (IRLC) . In recent supe rtrees of the
Hologalegina alliance (Wojciechowski et al., 2000,
2004), Loteae sens. lat. are sister to Sesbania , and the
combined Loteae-Sesbanieae clade is itself sister to
the Robinieae.
Loteae differ from Robinieae in a suite of characters
which were listed by Dormer (1945) for his 'epulvinate
series', i.e., often herbaceous habit and leaves mostly
distichous , usually without a pulvinus. These
characters are now of less phylogenetic importance
since the 'epulvinate series' is no longer considered to
be a monophyletic group. An obvious synapomorphy
of Loteae, shared by all extant members of the tribe ,
is stamen filaments dilated upwards. This is an
adaptation for secondary pollen presentation. Another
apomorphy shared by almost all Loteae is the capitate
or umbellate partial inflorescence, while Robinieae
possesses racemes. A most unusual (and possibly
synapomorphic) morphological character of many
Loteae is the presence of a foliage leaf on the peduncle.
This leaf is often described as a bract, but it has neither
a flower nor other strnctures in its axil. Trne bracts in
Loteae usually lack a blade and are membranous or
glandular. Phylogenetic evidence suggests the presence
or abse nce of the foliage leaf on the peduncle is
homoplastic within Loteae.
The circumscription of Loteae has recently been
expanded to include genera formerly placed in
Coronilleae (Polhill, 198lk & l; 1994). These two tribes
LE FT Lotus bertholetii Photograph by P. Gasson
were previously distinguished by the lomentaceous
frnits and branched root nodules in Coronilleae (fruits
non-lomentaceous and root nodules unbranched in
Loteae sens. strict.). The Coronilleae were earlier
placed in tribe Hedysareae (Bentham, 1865) because
of their lomentaceous fruits.
The merging of Loteae sens. strict. and Coronilleae
is supported by both morphological (Polhill, 198lk;
Lassen, 1989; Diez & Ferguson, 1990, 1994, 1996;
Tikhomirov & Sokoloff, 1996a) and molecular data
(Doyle, 1995; Liston, 1995; Doyle et al. , 1997; Allan,
1998; Allan & Porter, 2000; Allan et al., 2003). These data
indicate that lomentaceous fruits have arisen
independently in Coronilleae and Hedysareae, and
perhaps even in different genera of Coronilleae. Allan
& Porter (2000: Fig. 2A) suggested, however, that
lomentaceous fruits were a plesiomorphic condition
for Loteae sens. lat. In our opinion, an ancestor of
Loteae might have had dehiscent fruits divided into
regular compartments by thin transverse septa (as in
Sesbania and Lotus sens. lat.). The genera formerly
placed in Coronilleae do not form a natural taxonomic
unit at any level. Regarding fruit anatomy, lomentaceous
fruits of Securigera and Coronilla are related to
dehiscent fruits in the Lotus group, while lomentaceous
fruits of Ornitbopus and Antopetitia share important
features of pericarp structure with the indehiscent non-
lomentaceous fruits of Anthyllis and Dorycnopsis.
Recent publications have led to the removal of
some groups from Lotus sens. lat. (Lassen, 1986;
Tikhomirov & Sokoloff, 1997; Sokoloff, 1999, 2000),
Coronilla sens. lat. (Lassen, 1989) and Anthyllis sens.
lat. (Lassen, 1986; Tikhomirov & Sokoloff, 1996a, 1997).
Morphological and molecular data relevant to the
relationships between Old World and New World
Loteae have recently been published (Diez & Ferguson,
1990, 1994, 1996; Kramina & Sokoloff, 1997a; Allan,
1998; Allan & Porter, 2000; Allan et al., 2000, 2002,
2003; Degtjareva et al., 2003). The placement of
Podolotus in Loteae is now well supported by
morphological data. Allan & Porter (2000) and Allan et
al. (2003), using nuclear ribosomal ITS, concluded that
Old World and New World Lotus sens. lat. belong to
TRIBELOTEAE 455
 distinct clacles. In the latter analysis, Old World Lotus kl Worl I Ornithopu · (altb ugh with 1 sp. in th ~ Ne
forms a moderately supported monophyletic group if World) and Kebirita am n others. In this rr arme"'
Tetragonolobus and Dorycnium are included, while ·s consK
Loteac 1 · 22 genera and . nt
. Ie recI to comprise
2 2
New World Lotus is paraphyletic, also containing the species Fig. -n.

Robinieae (see page 467)

Sesbanieae (see page 452)

I I Hippocrepis

Scorpiurus
1 I
Securigera

Coronilla

Podolotus

Anthyllis
Hymenocarpos

Pseudo lotus Hippocrepis scobro Drawing by M. Y. Saleem Hippocrepis bo/earico Photograph by G. P. Lewis

Antopetitia Hippocrepis L 1753

I Emems Mill. (1754)
0 c. 34 spp; Europe, mainly Mediterrnnean region (including Saharan N Africa and
Hosackia --l Macaronesia), Arabia and \V Asia 10 Pakistan and Turkmenistan
rn From hippo- (Gk.: horse) and crepis (Gk.: shoe); from the shape of the fruit
)> segments (in most species)

Ornithopus rn Sh1ubs, suffrutices or he1bs; d1y subtropica l, mediterranean, warm temperate ;:ind
temperate shrubland, grassland and desen, and in disturbed places
Reference(s): Hrabetova-Uhrova (1949-50); Dominguez 0976); Schmidt (1979);
Lassen (1989); Talavera & Dominguez in Talavera el al. (2000: 897-935)
* Dorycnopsis Lassen (1989) transfened Coronil/a emerus sens lat in to Hippocrepis; this opinion
has been supported by studies of pollen morphology (Diez & Ferguson, 1996),
fruit anatomy (Sokoloff, 1997), and moleculrn phylogenies (Allan & Prnter, 2000;
Kebirita Allan er al, 2003), and is adopted he re; Hippocrepis emems (L.) Lassen possesses
pu1atively less-derived cha1acters such :1s long fruits with straight (not horsc-shoe-
shapecl) segments, and a shrubby habit
Ottley a Hippocrepis einerus js cultivated as an ornamental; othe1· species (horseshoe
vetches) are used frn medicine
Acmispon
Syrmatium
Scorpiurus L.1753
2 val'iable spp . (sometimes treated ;:is 4 spp .); S Europe, mainly Med iterranean
Lotus (including N Aflica), Macarone~ia and \V/ Asia
From sc01pio- (Gk.: scorpion) and oura (Gk .: tail), for the shape of the pod
Dorycnium Herbs; medite 1ranean shrubland, g1assland and disturbed places
Tetragonolobus Referencc(s): Heyn & Raviv (1966); Talavera & Dominguez in Tabve1a el al.
(2000: 935-942)
The gen us occupies an isolated position in the t1 'ibe; the simple leaves with
Tripod ion (t1sually) three main veins are most unusual in th e family
Sco1piu.rus verm iculatus L. (prickly scoq:>ion's rail) is used for forage; leaves and
Hammatolobium pods used for human food
Cytisopsis

FIG. 51 Diagram of relationships in tribe Loteae based on molecular (Allan et o/., 2003; Degtjareva et o/., 2003) and morphological data (Sokoloff, 2003a)
Nodes marked* are not supported in all analyses Scorpiurus muricotus Drawing by L. M. Rip ley

TRIBE LOTEAE 457

456 LEGUMES OF THE WORLD
Securigera securidaca Drawing by E. M. Stones Coranilla valentina subsp. glauca Photograph by w. F. Downes Anthyl/is montana subsp. jaquinli Photograph by G. P. lewis Anthyl/is vulneraria subsp. maura Drawing by M. Y. Saleem

Securigera DC. 1805 Anthyllis L. 0753)

13 spp.; C and S Europe and Mediterranean region (including N Africa), Somalia Cornicina (DC.) Boiss. (1840) sens. strict.
22 spp.; Europe, mainly Meclite11 anean region (including N Africa) and Madeira
and mainly W to C Asia (W Siberia), naturalised elsewhere
east to Caucasus and Iran, soulh to Ethiopia; natu1 ali.sed in some other countries
From securis CL.: axe) and gero (L.: bear), for the shape of the fruit in the type
From antbo- (Gk: nower) and iou/os (Gk: first soft beard), referring to the
srecies S. secw1daca (L.) Degen & Doi fl.
pubescence of the calyx or the whole inflorescence in A vu.lneraria L. and
Herbs; medirerranean, warm-temperate and continental temperate woodl:and and
grassland to tropical bushland, in disturbed places other species
Shrubs, suffrutices and herbs; mediterranean and tempe1are to montane tropical
Reference(s): Uhrova (1935); Zoz 0970); Jahn (1974); Schm id t 0978); Lassen
grassland and shn1bland, sometimes in disturbed places
(1989); Yakovlev et al. 0996)
Referen ce(s): Cu llen (1968, 1976); Akulova (1985); Miniaev & Akulova 0987);
Securigera has usually been accepted as a monospecific genus, separated f1om
Benedi-Gonzalez (1998); Benedi in Talavera et al (2000: 829 - 863); Sokoloff
Coronilla by the non·l omentaceous fruits; Schmidt (1978) clearly demonstrntccl
that S. securidaca is closely 1elated to annual species of Coronitla; Lassen (1989) (2003a)
Tripodion and Do1ycnopsis are best excluded from Anlbyllis (Tikhomirov &
transferred about half the species of Coronil/a to Securi.gera, and recent studies of
Sokoloff, J996b); morphologically Do1ycnopsis is close to Antby/lis, but also
pollen morphology and fruit anatomy neither suppo11 no r contradicr Lassen 's
resembles Ornithopus in, e.g., fruic anatomy; A. vulneraria is veiy variable and
treatment; Sokoloff (2003b) merged Securigera with Coronilla
20-30 inf1 aspecific taxa have been distinguished (Cullen , 1976)
Secun'gera varia (L.) Lassen (crown vetch) is cultivated as a forage, covet crop,
Anthyllis vulnerm·ia sens. lat. (sand clover, kidney vecch) is cu ltivated as fodder,
weed suppressor and for erosion control; also used as medicine
forage (especially for goats and sheep), g1ound cover and as soil binders; also fo1•
medicine (with a wide 1ange of uses, particularly skin care), as a tea substitute, a

Coronilla L. 1753 dye and as ornamentals

A11ro/obium Desv. (1813), pro pa rte
9 spp; C and S Europe, Mediterranean region (including N Af1 ica) to W Asia
From the diminutive of corona (L.: crown), referring ro the shape or lhe
Hymenocarpos Savi 1798
inflorescence
Cornicina (DC Boiss. pro pmte, excl . type
Shrubs, suffrutices or herbs; mediterranean and warm temperate grassl:ind,
1 variable sp.; Mediterranean, \Y/ Asia
From hymen (Gk.: membrane) and ca1pos (Gk.: fiuit), referring to the wide wing
shrubland and woodland; also in disturbed places
Reference(s): Uhrova 0935); Zoz 0970); Jahn (1974); Schmidt (1979); Lassen of the fruit
Heib; mediterranean grassland, shrubland and frnest, often in disturbed places
0989); Lock 0989: 124 - 125); Garcia Martin & Talavera in Talave1a el al. (2000:
881 - 891); Sokoloff (2003b) Reference(s): Zoha 1y (1987)
TI1e genus has usually been treated as monospecific; Lassen (1986) transferred the
See the notes under Securlgera and Hippocrepls
3 spp of Antbyllis subgen. Comicina to Hymenoca1pos, but Tikhomirov &
Used as 0 1namenta ls
Sokoloff Cl996b) disagree and have returned these to Antbytlis; Allan et al. (2003)
find strong support placing H. circinna/us CU Savi in a clade with Antbytlis
Podolotus Royle 1835 /o/oides L.
The monospecific Hymenocmpos accepted here differs from Anlhyllis in its
Kers/ania Rech_f. 0958) broadly winged fniits, petal shape. and basic chromosome number (x - 8, nor 6
l sp; SW Asia, including Oman, Irnn, Afghanistan, Pakistan and N\V India
or7), however, it is close to Antbyllis; Akulova 0985) and Sokoloff (2003a) placed
From podo- (Gk,: foot) and Lo/us, referring to the long-stipitate pod in a genus
Hymenocc11pos into synonymy under Anlbyllis
orherwise like Lotus
Herb; seasonally dry tropical and subtropical to continental montane warm-
temperate shrubland and woodland
Reference(s): Rechinger 0 984a)
The genus combines the characters of the Lotus and Coronilla groups (Lassen, 1989)
Podalatus hasackioides Drawing by P. Halliday Hymenacarpos circinnatus Drawing by E. M. Stones

458 LEGUMES OF THE WORLD

TRIBE LOTEAE 459

Pseudolotus villosus Drawing by P. Halliday
Hosackia gracllls Photograph by G. D. Carr
460 LEGUMES OFTHE WORLD
Antapetitio abyssinica Drawing by C. Grey- Wilson
Pseudolotus Rech.f. 1958
l s p~; W Asia (Oman, United Arab Emirates, I ran and Pakiscan); records from
Afgh~ni stan are e1mneous
From pseudo- (Gk: false) and Lo/us (q.v.)
H erb; dry tropical and subtropical desert, wadis, sandy places, rocky slopes
Refe1 ence(s): Recl1i ngei- (l984b); Ali & Sokoloff (2001)
Lassen (in Rechingel', 1984b) suggested that Pse11do/0111s is 1elated to Vermiji11x(llere
treated as DOJycnopsis) and Syrmalium, but recent morphologica l analysis does not
supprnt this; Pse11dolo1us resembles Kebi1ila both morphologica lly and ecologic:oll y
Antopetitia A.Rich . 1840
I sp,; Africa
Named afte1 Antoine Petit (?-1843), doctor and zoologist on th e Lefeb1e Expedilion
to Abyssinia (now Ethiopia), 1838 - 1844, taken by a crocodile while swimming
~tcross the Nile
H erb; t1 opica l montane giassland, often on shallow soil over rock
Reference(s): Gillen in M ilne-Redhead & Polhill 0971: 1049 - 1051); Verdcou1t in
Pope et nl. (2003: IO)
A distinct genus combining a basal inflorescence strucru1e and specialised fn1its
with a dehiscence type unique in the tlibe; Anlopelilia has been n-iergcd with
Ornithop11s by several authors, but recent stud ies of pollen mo1phology cmd fn.1it
anatomy do not support this; Antopetilia is the only genus of LoteHe endemic to
tropica l regions
Hosackia Douglas ex Lindi. 1829
11 spp.; SW Canada, W USA, Mexico and Guatemala; mainly in California
Named after David Hosack (1769- 1835), Ameoican physician and botanist
H e1bs; temperate, waJ'm temperate and mediterranean w oodland, shrnbbnd ~ind
giassland; oflen in seasonally or permanently wet sires (unusual in Loteae)
Reference(s): Greene (1890); Ottley (1923); Isely (1981); Ki1 kb1 ide (1999a);
Sokoloff (2003a)
Until recently Hosa ckia was usually treated either as a synonym of Lotus sens. lat '
or as a sepa rate genus comprising all New World Loteae; molecu lar data (A llan &
Po11er, 2000; Allan el al., 2003) give strong support to the separation of the New
\Xforld species from Lotus sens. strict,, however, 1ehnionships between rhe
Am erican raxa and re lated Old Woild genera have still to be cla1ified. As treat ed
lleie, Hosackia sens strict comprises a g1oup o f species chaiacterised by such
presumably plesiomorphic charac1ers as memb1anous or foliaceous stipules,
lriape1tu1are pollen gi ains, dehiscent fruirs, ancJ symmetrica l fl owers, however, the
d"ta o f Arambarl'i (2000) and Allan & Porter (2000) suggest that Hosackia sens
sMcl. may be paraphyletic; Sokoloff (2003") and Degtjareva el al. (in p1ep.) find
sttong suprort for th e rnonophyly of Hosack/a
Hosc1ckia gracilis Oe mh. is occasionally cultivated as an o rnamenral (Isely, 1981)
Dorycnopsis gerardii Drawing by A. Ladete
Ornithopus sotivus Photograph by R.B.G., Kew
Ornithopus L 1753
Arlrolobittm Desv. (1813), pro paite
c. 5 spp.; mainly W Eu rope (no1th to S Scotland) and Mediterranean region (incl .
N Afri ca) to Macaronesia, east to Caucasus and Iran; 1 sp. endemic to N
Argent·ina, Uru gua y and S Br.1Zi1
From omis, o.-nilh o- (Gk : bird) and -pus (Gk : foot), in reference to the pod
clusters l'Csembling a bird's foot
He1bs; rnediterranean to temperate or subtropical grassla nd and shrubland, often
on sandy soils
Reference(s): Griesinger (1939); Hanelt (1962); Talavera & Arista in Talavera el al.
(2000: 873 - 880); Degtjareva et al. (2003)
Other native Loteae are absent from eastern S America; Loteae wi th lomentaceous
fruits are absent f1orn the New \Vorld, so Ornilbopus is unusual in this respect;
this is the only genus of Loteae that incl udes both Old and New World species
Ornilbopus salivus Broe. (serrad ella; common birdsfoot) and other species are
cultivated for forage , soil cover, green manure and for bay and silage
Dorycnopsis Boiss 1839
Ver111iji11x J.13 .Gillett (1966); Helminlhocaipon A.Rich. (1847)
2 spp.; I sp in W Mediterranean, the other in NE Africa and adjacent Arnbian
Peninsulti
From Do1ycnium (q.v.) and -apsis (Gk .: like), for its resemb lance to Do1ycniu111
H erbs; seasona lly dry tropical to mediterranean grassland, shrubland and forest
Reference(s): Tikhomirov & Sokoloff (1997)
Ottley 0944) conside1ed He/mintbocn1pon (- Vemiiji·ux) to be close to the N
Americ•n Lotus subgen . Syrmalittm; Polhill (1981k) listed Vermiji·ux as a synonym
of Lollls; Tikhomirov & Sokoloff (1997) found significa nt differences between
Vermifrux and Lotus in inflorescence stnicture, fruit anatomy and micropylar
orientat ion pattern, and showed thar Vermifrux shou ld be sunk into D01ycnopsis
Kebirita K1amina & D .D .Sokoloff 2001
Acmispon sensu aucl., pro parle non Raf.
1 sp.; N\V Saharan Africa (Mauritania to Tuni sia)
Derived from the type locality, Aln Kebirita, Tunisia
Herb; d1y tropical and subtropical rocky desert, sometimes in dry stream-beds
Reference(s): Lassen (1986); Kramina & Sokoloff 0997b, 2001)
Kebiriln ro11dairei (Bon net) Krnmina & D.D.Sokoloff has trad itionally been included
in Lotus; Lassen (1986) trnnsfe1Ted Lotus roudairei to the New \'(lorld genus
Ac mispon, placing il in the section Simpeteria; morphological studies (Kramina &
Sokoloff, 1997b) have shown that Ac111ispo11 sens sllicl (-Lotus sect Micro/olus),
secti on Simpelen·a sens strict., and Lotus roudairei are quite different taxa, thllS
placement of these into a single genus Acm ~pon would be unsatisfactory; the
sec tion Simpeten·a has been described as a genus 011/eya (q.v) and L roudairei ls
segregated in the monospecific genus Kebi1ita
Keblrlta roudairei Drawing by P. Halliday
TRIBE LOTEAE 461
 Lotus corniculatus Photograph by G. P. Lewis
Lotus maculatus Photograph by G. P. Lewis
Ottleya strigosa Drawing by P. Halliday Acmispon subpinnatus Drawing by P. Halliday adapled from 0. Vilchez & A. Burkart

Lotus L 1753
Ottkya DD.Sokoloff 1999 Helnekenia Webb ex Ch rist (1887); Benediclella Maire (1924)
Lotus sect. Si111pete11a Ottley 0944) c. 125 spp. as treated he1e (c. 180 spp. in the broad sense); N Tempera te; here
13 spp .; SW USA and Mexico treated as exclusively Old World , mainl y Europe, Medi tenanean (including
Named afte1 Alice M. Ottley (1882 -1 971), American botanist who first Africa) and Macaronesia, eastwards to C, SW and E Asia (Sand \VI Siberia,
distinguished this group Mongoli<:1, Himalaya, China, Korea and japan); c. 2 spp. endemic to Australia and
Herbs or suffrutices; wa11n tempe1ate ornd mediterranean desert, serniclesc1t, 1 sp. to New Caledonia and Vanuatu , and c. 20 spp. in mo.s1ly 1n ntane tropict1l co
shrubland, bushland, woodland, and rudernl habitats Somal ia-Masai regions in Africa and Arabia
Reference(s), Ottley (1944); Sokoloff (1999); see also the references under Hosackia Fro1ll Greek Lolos; a name applied to various plants
Allan et al (2003) find no suppoit for the separation of Ottleya from Acmispo11 Herbs, suffrutices nnd shnibs; temperate, mediterranean, subtropical and t1opicnl
and Syr111ati11m; Degtjareva et al. (2003; in prep.) and Sokoloff (2003a) find good mo ntane grassland and shrubland, desert and disturbed places
support for the monophyly of 011/eya as well as Acmispon and Syrm aliu m Reference(s), Ilrand (1898); Chrtkova-Ze1tova 0966; 1973; 1984); K io·kbrid e (1994,
1999a); Krnmina (1999); Valdes in Talavera el al. (2000, 776-812); Allan et al,
(2002); Martins in Pope el al. (2003' 1-9)
Acmispon Raf. 1832 Po lhill (1981 k) adopted a broad concept of the genus, but Lassen 0986) and
Lotus sect Micro/01us Bentlt. (1836) Sokoloff (1999, 2000) accepted a number of segregates; the American species a1e
8 spp.; SW Canada, USA and Mexico; 1 sp. endemic to Chile treated he1e under Hosackia, 011/eya, Ac111ispo11 and Syl'lnatii1111 (Sokoloff, 1999);
From acme ( Gk : point, apex) and rhe rest of the name was unidentified; th e genus is taxonomically difficulr at specific as we ll as at sectional and
Rafinesque's names are often difficu lt to inte1prcr, blH he indicated that the name subgeneric levels; severnl species are extremely va1iable, e.g., L. col"niculalus L.
meant that the pod is hooked at the apex '1nd L auslralis Andr., and hybridisation is f1equent
He1bs; warm temperate and mediterrnnean desert, grassland, shnibland, Lotus co1·niwlaws (birclsfoot trefoil) and L pedu11cu/a1us Cav sens /al (big trefoi l)
woodland and ruderal h'1bit::1rs are major ag1'0nomic species, being valuable forage, pasture, green manure and
ltefeoence(s), Buokart el al. (1972); Sokoloff (2000); see also the references unde1 cover plants; a1her species are used in cosmetic skin care and, e g., L berlbo/etii
Hosackia and note under Ottleya M::1sf., page 454, and L. 111acula111s 13reitf. are cu\tivMed as omamentals

Syrmatium Vogel 1836

14 spp; West N Ameiica; Ilritish Columbia to Mexico
F10111 syrma (Gk., anything uailed or dragged), for the tail ed fruit
He1bs, suffrutices or shrubs; temperate, warm temperace and rnedirerranean
desert, semidesert, shrubland, bushland, woodland and ruderal habitats
Refe1·ence(s), see under /-Josackia and Ot//eya
This genus (like Oitleya) is taxonomically difficu lt at specific level, and
hybiidisation is frequent (e g, Liston et al 1 1990); see comments under
Do1ycnopsis and Ou/eya

Syrmatium eriophorum Drawing by P. Halliday

Lotus aduncus Photograph by}. Hooper

TRI BE LOTEAE 463

462 LEGUMES OFTHE WORLD

Dorycnium pentophyllum Drawing by H. Wood
Dorycnium Mill. 1754
Bonjeanea Rchb (1832)
8 spp.; C, Sand E Europe, mainly Mediterranean, eastwards to Caucasus and \V/
Asia (Iran)
From Greek Do1ycnion (Convolvulus), later transferred in meaning to these plants;
according to Dioscorides, D01ycnion is a sea-shore shrub, while in Pliny the name
refers to the toxic sap of the plant which was used for oiling arrows (DatJ' -
arrow, cnaein. - co lubricate); Dorycnium in the modern sense does not have
toxic sap
Shrubs, suffrurices or herbs; mediterranean woodland and shrubland, d1y slopes;
D. strictum (Fisch. & C.A.Mey.) Lassen often in saline habitats
Reference(s): Rikli 0901); Demiriz in Davis 0970: 512-518); Vural & Tan 0983);
Diaz Lifante in Talavera et al (2000: 863-866)
A generally accepted genus, but limits mu ch disputed; corrections to Rikli's (1901)
monograph sho uld be: 1) transfer Anthyllis/ulgurans Porra and Lotus strictus
Fisch. & C.A Mey. to Do1ycnlum (Lassen, 1979; 1986): 2) transfer Dorycniwn sect.
Canaria Rikli to lotus, as a subgen11s (Gillett, 1959); the basic distinction of
D01ycnium from Lotus is the smooth (not papillose) style; further studies ma y
show that Lotus and Do1ycnium should be merged, as proposed by Polhill
(198lk) and as indicated in the phylogeny of Allan et al. (2003); Sokoloff (2003a)
transferred all species of Dorycnium to Lotus; the D. penlapby//um Scop. complex
is very variable and problematic
Dotycnium b irsulum (1.) Ser. (hairy Canary clover) and other species al'e used as
a forage crop, to connol soil erosion and are cultiv<ited as ornamentals
Tetragonolobus scop. 1772
6 spp.; mainly Mediterranean to E Europe and Caucasus
From tetra (Gk.: four), gonia (Gk.: angle) and lobion (Gk , pod), for the 4-winged
fruit (although some species have 2-winged fruits)
Herbs; mediterranean and warm temperate grassland
Reference(s): Dominguez & Galiano (1979): Valdes in Talavera et al. (2000:
823-828)
Close to Lotus and often considered a section of it; Tetragonolobus can easily be
separated from Lotus by frnil, style and leaf morphology, but the phylogenies of
Allan & Porter (2000) and Allan et al. (2003), and Sokoloff (2003a) indicate that
Tetragonolobus is firmly embedded within Lotus
Tetragonolobus pwpureus Moench (winged pea, asparagus pea) is an important
crop cultivated for its edjb}e pods and tubers 1 and as a culinary herb; seeds arc
roasted as a substitute for coffee; also used as a green manure, winter cover and
hay crop
Tetragonolobus purpureus Photograph by G. P. Lewis
464 LEGUMESOFTHEWORLD
m
~ 4
Hommatolobium lotoides (1-9), H. ludovic/a (10-13) Drawing by P. Halliday
Tripodion tetraphy/lum Drawing by M. Y. Saleem
Tripodion Medik. 1787
Physanthyllis 13oiss. (1840)
1 sp.; Meditenanean
From tri- (Gk.: three) and podion (Gk : little foot), probably because the seedling
usually produces 3 shoots
Herb; mediterranean grassland and ruderal habitars
Reference(s} Tikhomirov & Sokoloff (1996a); 13enedi in Talavera et al (2000:
866-868)
Formerly often included in Antbyllis; Lassen (1986) merged Tripodion with
Hammato/obium, repoiting that both have lomentaceous fruits, but Tikhomirov &
Sokoloff CI996a) have shown that the fruit of Tripodion is de hiscent and not
lomentaceous (see note under Hammato/obium); Allan & Porter (2000) place
Tripodion as sister to a combined DmJtcnium and Old World Lolus dade, well
separated from Anthy!lis vulneraria
Hammatolobium Fenzl 1842
2 spp.; 1 sp in Algeria and Morocco, the other in NE & E Mediterranean (G reece,
Turkey, Syria)
From hammato- (Gk: knot, halter) and /obos (Gk: lobe), referring to the coherent
basal lobes of the wings, a feature common in orher Loteae genera
Suffrutices; rn editerranean woodland and shrubland, often o n limestone
Reference(s): Chamberlain in Davis 0970: 547); Tikhomirov & Sokoloff 0996a)
Lassen (1986) merged Hammato/obium with T1;podion, but morphological study
of the two genera (Tikhomirov & Sokoloff, 1996a) supports their segregation
Cytisopsis Jaub. & Spach 1844
Lyatlleya Maire (1919)
2 spp., although probably I with 2 subsp; Mediterranean; I sp in Morocco, the
other in Turkey and Middle East
From Cytisus (q.v.) and -apsis (Gk.: like), indicating its resemblance to Cylisus
Shrubs; dese1t and semi-desert shrn bland; meditenanean bushland, forest and
thicket
Reference(s): Maire (1919); Chamberlain in Davis 0970: 511-512)
The genus has a number of fealu1es considered basal in Loteae, such as a shrubby
habit and deruscent fruits, but it also possesses seve1al characters regarded as
derived in the tribe, e.g., sessile leaves with a suppressed rhachis and complete
absence of stipules; Degtjareva et al. (2003) suggest a close relationship to
Hammatolobium and Tripodion
Cytisopsis dorycniifolio Drawing by P. Halliday
TRIBE LOTEAE 465
TRIBE

Robinieae by M. Lavin and B. D. Schrire

tribe Robinieae 13 nth.) Hutch. 1964

Tribe a legeae subtribe Robiniinae Benth. (1865)
Tribe Galegeae subtribe Corynellinae Rydb. (1924).

The treatment of Polhill & Sousa 0981) recognised 21
genera and c. 145 species in Robinieae. This treatment,
foll wing modifications of Lavin (1988, 1993, 1995),
Lavin & Sousa 0995), Polhill 0994), Lavin et al. (2003),
reduces the tribe to 11 genera and c. 72 species (Fig. 52).
The phylogenetic positions of genera in Robinieae
(excluding Sesbanieae) have largely remained the
same since Lavin & Sousa (1995), although recent
phylogenetic analysis of mo! cular (ITS/5.8S, /.rnl inrron
and matK locus) and morpho l gical data m .d l fu1the r
modification (Lavin et al., 2003). The Cuban Hebestigma
is well supported as sister to the Mesoamerican Lennea
in the combined analysis of Lavin et al. (2003), and in
most analyses the two genera are sister to the Gliricidia
and Robinia groups. Within the monophyletic Gliricidia
group, Gliricidia is now demonstrated to be
paraphyletic with respect to Hybosema, so the two are
amalgamated. The Gliricidia group still includes the
Antillean Poitea together with Gliricidia.
The monophyletic Robinia group comprising the
'barbistyled' genera , Olneya , Robinia , Coursetia,
Peteria, Genistidium and Sphinctospermum, is sister
to the Gliricidia group in most analyses of Lavin et al.
(2003), although this is not strongly supported. The
only significant modification within the Robinia group
is that four eclectic species, Coursetia hypoleuca (Speg.)
Lavin and C. orbicularis Benth. (unifoliolate species
formerly composing the genus Poissonia) , along with
the Peruvian C. weberbaueri Harms and the Argentine-
Bolivian C. heterantha (Griseb.) Lavin (formerly
composing the monotypic Neocracca) form a well
supported monophyletic group, here referred to as
Poissonia, which in the analyses of Lavin et al. (2003)
is either sister to Robinia or to Sphinctospermum.

Loteae incl. Coronilleae

(see page 455)

Sesbanieae (see page 452)

= Hebestigma
- Lenn ea
I! Gliricidia group
1, Gliricidia
"'- :::0
Poitea
~

0
co
Olneya z
Robihia rn
Robinia group )>
Poissonia rn
~
c;0·ursetia
Peteria
Genistidlum
Sphinctospermum

Inverted-Repeat-Lacking
Clade (IRLC)

FIG. 52 Diagram of relationships in tribe Robinieae after Lavin et al. (2003)

LEFT Gliricidla brenningii Photograph by C. E. Hughes

TRIBE ROBINIEAE 467

 !l
() (\ (1

Pa/tea punicea Illustration by F. W. Horne

Lennea melanocarpa Illustra tion by c. F. Schmidt Poiteo campanilla Photograph by M. Lavin

Poite a vent. 1807

Hebestigma urb 1900 Sabinea DC. (1825); C01ynel/a DC. (1825); Notodon U1b. (1899); Bembicidlum
1 sp.; Cuba Rydb. (1920); Sa11vallel/a Rydb. (1924); Cajalbanta Urb. (1928)
From /Jebe- (Gk., youth, down of puberty) and stigma (Gk,, a st igma), referring to 12 spp.; Caribbean (Cuba, Hispaniola, Puerto Rico, Virgi n Islands and Dominica)
the long-ciliate stigma cha1acr.eriscic of the genus Named after P.A. Poiteau (1766 - 1854), French botanist and botanical artist;
Trees; seasonally dry lropical lowland foiest and scrub, abundant on calca1cous author of Flore Parisienne (1808-1813), who worked in Haiti 1796 - 1801
substrates Trees and shrnbs; seasonall y dry tropical lowland to montane forest, thicket and
Reference(s), Lavin 0987); Lavin & Sousa 0995) thorn scrub, often in secondary vegetation and on steep slopes or roads ides
A taxonomica ll y isolated genus, and apparently the least mo1phologica ll y Reference(s), L.win 0987, 1993); Lavin & Sousa 0995)
modified in the tribe; sister to Lennea in most analyses of Lavin et al. (2003) See under Gliricfdia fo r 1elationships; most of the genera in synonymy above
Used for wood (poles) were recogn ised by Polhill & Sousa (1981)
Used as ornamentals and revegetators

Lennea Klotzsch 1842

3 spp.; S and E Mexico and C America Olneya A.Gray 1855
Named after the Pnissian director of the Gardens of the German Crown Prince, 1 sp.; SW USA and NW Mexico (Sonoran Dese1t)
Peter Joseph Lenne (1789-1866) Named after Stephen Thayer Olney, American botanist (1812 - 1878)
Trees and large shrubs; trop ical riparian (and coastal) areas of lowland forest, Jess Trees or shrubs; subtropica l desert shrubland 1 in gravel washes and open lower
frequent ly in seasonally dry tropical forest hillsides
Refcrence(s), Lavin (1987); Lavin & Sousa 0995) Refe rence(s} Lavin 0987); Lavin & Sousa (1995)
Lennea is sister to Hebestigma in most analyses of Lavi n et a l. (2003) Most closely relaled to Robinia, Poissonfa, Peteria, Genistidfr.ttn and
Used as ornamentals and potentially useful as revegetators and living fences Spbinctosperm1'11l and possibly basally branching to all or some of the above
(Lavin, 1995; Lavin et al., 2003)
An economically important species, 0 , tesota A.Gray (Sonora ironwood, palo
Gliricidia Kunth 1823 Fierro or tesota) is used for forage, as a hardwood (for implemen ts, construction
Hybosema Harms (1923); Yucarato nia Bu rkart (1969) a nd crafts), fuel (fi re wood and charcoal), huma n food and beverage (seeds and
5 spp.; Mexico and C Ame rica, with 1 sp. in Ecuador to Peru; G. sepium Qacq .) pods, also ground into a flour or meal), medicine and wildlife refuge, pote ntially
Steud. is widely introduced in tropical areas throughout the world useful as an ornanlental; the species is threatened by over-exploitation
From glis- (L., dormouse) and caedo- (L_, kill), referring to the powdered baik
and seeds being used as a rodent poison in the tropics
Trees and sbn.1bs; seasonally dry trop ical, often lowland forest, th icket and thorn
scrnb, on hil lsides and in exposed or d isturbed a reas; 1 sp. in montane wood land
Reference(s), Lavin (1987); Lavin & Sousa (1995); Lavin el al. (2003)
Closely related to Pol/ea; Lavin (1987; 1995) treated Hybosema sepaiately from
Gliricidia
The economically important G. sepium (madre de cacao, madricacao) is widely
used for living fence systems, shade trees for crops, weed and erosion comrol,
livestock fodder, medicine, insecticides, fuelwood, poles 1 green ma11u 1e,
ornamentals and bee forage

Gliricidia sepium Photograph by G. P. Lewis 0/neya tesota Drawing by P. Halliday

TRIBE ROBINIEAE 469

468 LEGUMES OFTHE WORLD
 Coursetia rostrata Pho tograph by G. P. Lewis Coursetla grand/flora Photograph by G. P. Lewis
Poissonio hypo/euca Photograph by M. Lavin

Robinia L. 1753 Coursetia oc. 1825.

Cracca 13e nth. (1854). non L. (1753); Balboa Liebm. (1854); Humboldlie/la
4 spp ; USA and N\V Mexico; some species wiclely cu lt ivated
Ha m1s (1923)
Named afte r the F1e11ch botanist Jean Robin (1550 -1 629), a nd his son Vespasian
c.: 35 spp.; S\Y/ USA, Mexico, C Am e1ica, Ca1ibbean and S America (Guyana,
(1579 - 1662), 1oyal garclene1s to Herny JV of France
Venezuela, adjacent Brazil ::md the Ande:ln co rdillen.t co N Argentina; also with
Tree~ and large shrubs; cool to wa1m temperate montane regio ns in Appalachia
extensions to Paraguay and EC Brazil)
to subtropical S\Y/ USA - NW Mexico in seasonally dry woodland, mostl y as
Named after the French agronomist George Lou is Marie Dumont de Coursct
1hickets at forest edges, often on f'ocky hillsides, road curs, and along su c;rn1
(1746 -1824)
banks
Trees, shrubs rind herbs; seasonnlly dry tropical rin d subtiopical forest and
Hefe rence(s): Jsely & Peabody 0984) ; Lavin & Sousa 0995)
monrnne w ood land
Re lated (as th e Robinia gro up of Lavi n, 1995) to the S Am e1'ican Polsso nia,
Refe ren ce(s): Lavi n 0988)
a lthough with low bootstrnp supp mt (Lavin at al, 2003)
Altho ug h Lavi n (1988) and Lavin & Sousa 0995) recognised 5 sections, 1ece nt
Robinia psa11doacacia L (blac k locust) an d R. hispida L. (b1'istly loct1st) are
phylogenetic a nalysis of molecular data (Lavin el al., 2003) revea ls that two of
widely cu lti vated as oiname nlals, both are root suckeiing and 1he for me r,
these, Po ;ssonia and Neocracca, are besc relega ted to a dislinct g1oup, genus
especi all y, e!-ltablishes ilself aggressively in many areas; a number of hybrids :111d
Poisso nia (along with Courselia weberbauen); of the re 1m 1ining th1ee sections,
1rn111y cultivars exist Other uses include soil ern ichment and sta bili.~mtio n , wood
Cracco ides anc.l Course/la should be amalgamated. Only the Mesoamerican
(constru ction, implements and crn fts), fu elwood, hun1an food and beverage
sectio n Madre11ses forms a monophyletic subclade w ith in Coursetia
(seeds, pods and flowe1s; ~1 tea is infused from the bark) bu t ca1e is necess;uy ;1s
Used for gums and wood
plants h:1ve toxic p1ope1ri es, medici ne, bee food (honey), ba1k fi bre (text iles and
paper), essent i::ll oi ls and dyes

Poissonia 13aill 1870

Chiovandaea Speg. 0917); Naocmcca Kllntze (1898)
4 - 5 spp.; S America (Pe111, Argentina, Bolivia)
Named afte r th e fre nch botanist Ju les Poisson (1833-1919)
Ti ees, shru bs and herbs; tropical and subt1•opic<1l seasonall y d1y forest and
shrublan d, along riveJ' and stream banks, :l lso in open arid vegeta ri a n
Referencc(s): Lavin er a l. (2003)
T 1ea te<l as seclio ns Poissvnia and Neocracca of Cou rselia by Lavin (1988)
Used for edible roots and as a soil stabiliser

Poissonio heterontha Photograph by M. Lovin Coursetia glandulosa Photograph by G. P. Lewis

TRI BE RO BIN IEAE 471

470 LEGUMES OF TH E WORLD
Peteria thompsonae Drawing by B. Angel Genistidium dumosum Drawing by P. Halliday

Peteria A.Gray 1852

4 sp p.; mid-West to SW USA and N Mexico
Named after Dr Robert Peter (1805 -1894), American botanist
Herbs or subsh rubs; cool to wa rm temperate and subtropica l desert thorn scrub,
dry grassland and montane pine forest, on gravel washes and slopes of dry hills,
mesas or m oun tain valleys
Re ference(s} Porter 0956); Lavin 0987); Lavin & Sousa (1995)
Closely related to Genislidium and Sphinctospermum (Lavin el al., 2003)
The tuberous roots of P. glandu/osa (A.Gray ex S.Watson) Rydb. (ca mote del
monte or wild swee t potato) are used for human food

Genistidium I,M.Johnst 194 1

1 sp.; USA (S Texas) and NE Mexico (Ch ihuahuan Desert)
From idion- (Gk.: diminutive su ffix), referring to a likeness to the genus Genis/a
Shrubs; subtropical to warm temperate desert scrnb on high cliffs and rocky or
clayey soils
Refe re nce(s), Lavin (1987); Lavin & Sousa (1995)
Sister to either Peteria or Sphinctospermwn in the analyses of Lavin et al. (2003)

Sphinctospermum Rose 1906

I sp ; Mexico along the Pacific slope and S USA (Arizona)
From sphincto- (Gk : const ri ct) and -sperma (G k., seeded), referring to the seeds
of Sphinctospermum which have a distinct ive vertical consrriction making them
almost dumb-bell shaped
Herbs; seasonall y dry tropical to subtropical Pacific coastal rorest, oak woodland ,
grassland and desert thorn scrub, often in disturbed vegetation
Reference(s), Lavin 0990); Lavin & Sousa 0995)
Lavin 0987) transferred Sphinctospermum to Millettieae, but later returned it to
Robinieae based on molec ular evidence (Lavin & Doyle, 1991)

Sphinctospermum constrlctum Drawing by P. Halliday Robinia pseudoacacia Photograph bys. Andrews

TRIBE ROBINIEAE 473

472 LEGUMES OF THE WORLD
TRIBE
Galegeae hyJ. M. 1..ockandB. n. schrire
'fribe Galegeae (Bronn) Dumort 1827
Subtribe Galeginae Bronn (1822), as 'Galegeae'
Tribe Astragaleae (DC.) Dumort (1827)
Subtribe Astragalinae DC. (1825), as 'Asu-agaleae'
Tribe Carmichaelieae Hutch. (1964)
Polhill (l 981a, h & i) recognised Galegeae and
carmichaelieae as distinct tribes. Within the Galegeae
he distinguished four subtribes: Coluteinae,
Astragalinae, Galeginae and Glycyrrhizinae. Polhill
(1994) added a fifth subtribe Alhagiinae . More recent
studies suggest that the Carmichaelieae are best treated
as another subtribe within Galegeae, and this is
followed here. However, these studies also point
strongly to the polyphyly of Galegeae (Wagstaff et al.,
1999; Wojciechowski et al. , 2000; 2004). Polhill's
concept of Galegeae appears to be in the process of
disintegration into a number of smaller tribes, but
decisions on this would be premature until constituent
taxa are sampled more comprehensively, and putative
monophyletic groups can be substantiated by
morphological and other systematic data.
Glycyrrhizinae Rydb. was recognised by Polhill
(1981h) as vety distinct, but he left it as 'an outlying
subttibe of Galegeae'. Molecular analyses have confinned
the outlying position of Glycyrrhiza (Fig. 53). Although
it is a member of the Inve1ted Repeat Lacking Clade
(IRLC), it forms a basal grade or sister group, along with
such woody millettioid genera as Callerya and Wisteria,
to the whole of the rest of the IRLC (Sanderson & Liston,
1995; Wojciechowski et al., 2000; 2004).
Galeginae, containing the single genus Galega (Fig.
53), is also isolated from the rest of 'Galegeae'. In
Wojciechowski et al. (2000), it falls near the base of
their Vicioid clade (Fig. 53) and is sister to the Cicereae.
In Wojciechowski et al. (2004), Galega is sister to a
combined Cicereae-Trifolieae-Fabeae clade, although
with relatively poor support. Such placements have not
been suggested before and careful morphological
studies are needed to see if they support the
relationships suggested by molecular analyses . One
of the prominent features of Cicereae is the
craspedodromous venation of the leaflets and while the
leaflet venation in Galega is not truly craspedodromous,
Polhill (1981h) does give 'leaflet-nerves to margin or
nearly so' as one of the characteristics of Galeginae.
Astragalinae contains, inter alia, what is probably
the largest genus of flowering plants - Astragalus -
with 2300-2500 species. Wojciechowski et al. (1999;
2000) have shown that their Astragalean clade (Fig.
LEFT Swoinsono formoso Photograph by R.B.G., Kew
53) is itself made up of several distinct clades. The first
of these includes the vast majority of the species of
Astragalus. Wojciechowski et al. (1999) and Kazempour
Osaloo et al. (2003) find good support for Astragalus
sens. strict. as a monophyletic group, i.e., excluding
only a very few outlying species (mentioned below).
Wojciechowski et al. (1999) also showed that genera
such as Oxytropis, Sutherlandia and Swainsona, which
are morphologically very similar to Astragalus
(although they have never been formally combined
within it), are in fact distinct and not nested within
Astragalus. Sanderson & Liston (1995) and
Wojciechowski et al. (1999, and references therein)
have clearly shown that the vast majority of New World
Astragalus form a single clade in which most of the
species have chromosome numbers in an aneuploid
series: n = 11, 12, 13, 14 or 15 (Spellenberg, 1976). The
Old World taxa, on the other hand, have chromosome
numbers that are euploid: n = 8, 16, 32 etc., and
polyploids are common (Spellenberg, 1976). All the
studies of Liston and his group (e.g., Liston & Wheeler,
1994) demonstrate that Astragalus, far from being a
taxonomic ragbag, does in fact form a monophyletic
group in which speciation has been particularly active;
possible mechanisms have been discussed by Barneby
(1964) and Polhill (1981h).
A second small clade, sister to Astragalus, includes
Biserrula pelecinus L. and Astragalus epiglottis L. These
are both annuals of the Mediterranean Basin and N
Africa. Both have only five fertil e anthers and
dorsiventrally compressed pods . The position of
Biserrula has long been disputed but it has often been
regarded as no more distinct from Astragalus than
some of the other monotypic genera that have from
time to time been split off, usually on account of their
distinctive pods (Barneby 1964: 26). The position of
Biserrula at the base of the Astragalus clade (e.g., in
Wojciechowski et al., 1999, 2001; Kazempour Osaloo et
al., 2003), however, supports its current treatment as a
separate genus. Astragalus epiglottis, however, does
not seem to have been treated anywhere above
sectional rank.
The third clade (Fig. 53) within the Astragalean
clade of Wojciechowski et al. (1999; 2001), i.e., the
TRIBE GALEGEAE 475
 Coluteoid clade, contains much of Polhill's 0981h)
Coluteinae, plus Astragalus sinicus L. and A.
complanatus Bunge and the Southern Hemisphere
carmichaelioid group. The group has a very scattered
distribution, with the two Astragalus species occurring
in E Asia, Lessertia and Sutherlandia in southern
Africa, the carmichaelioids (including Swainsona) in
Australia and New Zealand, Colutea widespread in
continental Eurasia and NE Africa, and various smaller
genera such as Smirnowia and Ere mosparton
restricted to C Asia. Wojciechowski et al. 0999) point
sam pling within these groups has r ultecl in different
a nalyses sug esring differenc dlvisjons and plac 111 nts
, om of the genera are pc>orly known and of r lricted
di tribution (e.g., Oi·eophy a and Tibetia. , b'ut Olher
are more wide pr· ad, in luding Che neya an<1
G11eldenstaedtit1. Glycyrrhiza
Erophaca baetica, together with Chesneya anct
Gueldenstaedtia (note that G. himalaica Baker, the Wisteria, Callerya, etc (see page 369)
exemplar of the genus in the supertree, is now placed
in Tib etia) are part of the sister group to the
Astragalean clade (Wojciechowski et al., 2001). Hedysareae (see page 489)

out that Astragalus sinicus and A. complanatus Caragana , Calophaca and Halimodendron, on the
resemble o ther Coluteinae in possessing non- other hand, form a monophyletic group sister to the Chesneya, Spongiocarpella,
interlocking wing and keel petals and a ciliate style Hedysaroid clade, and are here placed in Hedysareae. Gueldenstaedtia, Tibetia
(Lavin & Delgado, 1990). However, they also note that Sanderson & Wojciechowski 0996) and Wojciechowski
Wenniger (1991) has found stylar hairs to be quite et al. (2000; 2004) show that Alhagi appears to be Erophaca
widely sca ttered among sections of Old World best pl ace d in Hedysareae, in agre ement with (= Astragalus lusitanicus Lam.)
Astragalus, and suggest that the character may have Hutchinson 0964).
arisen several times. Kazempour Osaloo et al. (2003) In a tribe that is so clearly polyphyletic it is difficult Oxytropis
found that Astragatus vogelii (Webb) Bornm., from N to know how to arrange the constituent genera. For the
Africa, Arabia, W Asia, Pakistan and India, grouped purposes of this book, genera recognised in the tribe Biserrula, Astragalus epiglottis )>
with Colutea in their analyses, and they erected a new largely follow Polhill 0981h; 1994), although w ith Vl
-I
genus Podlechiella Maassoumi & Kazempour Osaloo modifications resulting from more recent research. The Astragalus, Ophiocarpus, ;:;o
(p. 22) to account for this species. This decision two major realignments since Polhill 0994) are the Barnebyella )>
G')
appears premature before detailed molecular analyses transfer o f the ge nera me ntioned a bove to the )>
of the Coluteinae are available, and it is not accepted Hedysareae and the inclusion of the Carmichaelieae Colutea, Oreophysa, Smirnowia, I
rn
here. The genera separated as Carmichaelieae by (Polhill, 1981i) within Galegeae. The arrangement of Eremosparton, Sphaerophysa )>
Polhill 0981i) also appear best placed in this clade genera in this treatment is that suggested by the z
(Heenan, 1998a; Wagstaff et al., 1999). They have been supertree of Wojciechowski et al. (2001). Not all genera ("")
Lessertia, Sutherlandia r
shown to form a monophyletic group, confined to have b een sampled; genera not represented in the )>
Australia and New Zealand. supertree have been intercalated in positions that 0
Swainsona rn
The fourth clade (Fig. 53), is made up of a appear to be appropriate from other data in , e.g.,
monophyletic and distinct Oxytropis. The isolation of Polhill (1981h). Likewise , with the excep tion of Montigena, Clianthus,
Oxytropis is reassuring but somewhat surprising as Astragalus lusitanicus, here treated as Erophaca, the Carmichaelia, Streblorrhiza
me mbers of the genus are often morphologically isolated species of Astragalus revealed by the supertree
extremely similar to species of Astragalus and can only have not been treated separately. Podlech's 0994)
Parochetus (see page 500)
be distinguished by the pointed keel petals and by placement of three isolated Astragalus species as
the pod septum that arises from the adaxial suture, genera (Biserrula, Ophiocarpus and Barnehyella) has
Galega
not the abaxial as in Astragalus. Species of Oxytropis been followed here. <
are widespread in the north temperate regions, often Although we have suggested that it is premature to ("")

on mountains. re-circumscribe tribal limits in this group, the bulk of Cicereae (see page 496) 0
Three further groups of genera, exemplified by a) taxa (i.e., thos e comprising Polhill 's subtribes 0
Erophaca haetica Boiss. (=Astragalus lusitanicus Lam.); Astragalinae [for the most part] and Coluteinae, and Trifolieae: Trifolium (see page 500) ("")
b) Chesneya plus Gueldenstaedtia and c) Caragana, tribe Carmichaelieae), are likely to become recognised I
)>
Calophaca and Halimodendron, are sister either to as a more narrowly defined tribe Astragaleae. As treated 0
Fabeae (see page 505)
the Astragalean clade, or to the Astragalean plus Vicioid here the Galegeae sens. lat. comprises 24 genera and rn
clade (Wojciechowski et al., 2000). Relatively poor
'I (2880)- 3030 - (3180) species.
Trifolieae: Ononis, Melilotus,
Trigonella, Medicago (see page 500)

FIG. 53 Diagram of relationships in tribe Galegeae sens. lot. after Wojciechowski et al. (1999; 2000)

TRIBE GALEGEAE 477

476 LEGUMES OF THE WORLD
 ~
w

fl
.

Gueldenstaedtia multiflara Tlbetia himalaica Drawing by unknown artist

Chesneya rytidosperma Drawing by L. M. Ripley Drawing by unknown artist

Gueldenstaedtia Fisch . 1823

Glycyrrhiza L 1753
c. 10 spp.; Sino-Himalayan region to Mongolia and Siberia
Mel'istotropis Fisch. & C A.Mey (1843) Named after J.A. von Gueldenstaedt (1745 - 1781), Latvian botan ist and explorer
c. 20 spp.; mainly Eurasia; Europe (and Medi terranean inclt1Cling Africa) to w ith S.G. Gmelin in SE Russia
\Y/, C and E Asia, also N America Cl sp.), tempe1ate S America (1 sp.) and Herbs; continental temperate grassland and alp ine zones to 3000 m
Australia (l sp.)
Refe1e nce(s): Ali (1962, including Tibelia)
From glycys (Gk: sweet) and rhiza (Gk.: root) The taxon representing the genus in the supertree of Wojciechowski et al (2001)
Herbs; mainly mediterranean, warm temperate and continental tempe rate
is now placed in Tibelia
grassland, shrnbland, bushland and woodland, often in sandy, marshy and
Used in medicine
disturbed areas
Reference(s): Chamberlain in Davis 0970: 260 - 263); Yakovlev et al 0996)
Molecu lar data show Glycyrrhiza, with Wisteria and Cc//le1yc1, to be sister and
basall y branching to the whole of the IRLC (Wojciechowski et al., 2000)
Tibetia (Ali) H.P.Tsui 1979
4 spp.; mainly Sino-Himalayan regio n
Glycyrrht'za glabra L. {liquo1 ice, licorice) roots and its extra cts are used in
Named after the countiy, Tibel, whe re most of the species occur
medicine, navouring of confectionery and tobacco, human food (roots) and dri nk
Herbs; conri nental, monrane and altimontane grassland
(tea), soil bincle1s and as foam ing agents (in indust1ial products)
Reference(s): Ali 0 962); Tsui 0979)
Ali (1962) split Gue/denstaedtia into two well-defined sections; Tsui's suggestion
of generi rnnk is accepted here; G bimalaica 1 used to place Gzteldenstaedtia
Chesneya Lindi. ex Endl. 1840 close co C/J('SJl<!)~I in the molecular supertree of Wojciechowski et al. (2001), is
Kostyczewa Korsh. (1896) now treated as a species of Tibelia: T hima/aica (Baker) H.P.Tsu i
c, 30 spp.; temperate S\V Asia to Sino-Himalayan region, most diverse in C Asia co
Mongolia
Named after Ge neral f.R. Chesney 0789-1872), British soldier and plant collecmr
Shrubs or herbs; continental temperate grassland and shrubland, ofren rnonrane or
altimontane
Reference(s): see listing in Yakovlev et al. 0996)
Placed in the basally branching gro up of genera to the Ast1aga lea n clade
(Wojciechowski et al., 2000)
Some species have ornamental v~1 lue

Spongiocarpella Yakovlev & N.Ulziyk h. 1987

c 7 spp.; SC Asia to \Y/ China, mosc diverse in C Asia
From 'li ttle spo nge frnit', alluding to the spongy pericarp
Herbs; con tinental temperate semi~desert grassland , some monrane
Reference(s)o Yakovlev & Sviazeva 0987)
Small cushion-forming herbs with woody rootstocks; most were formedy placed
in Cbesneya but there are no molecular dara to confirm th at th ey are sister taxa

Spongiocarpella purpurea (1-7), 5. nublgena (8-14), S. spinasa (15-2 1).

Drawing by unknown artist Tibet/a hima/a/ca Photograph by H. Ohashi

TRIBE GALEGEAE 479

478 LEGUMES OF THE WORLD

Erophoco boetlco Drawing by R. Tavera Oxytropis holler/ Photograph by C. Grey-Wilson
Erophaca Boiss. 1840
As/raga/us lilsilanicus Lam. (1783)
1 sp. (with two subspp.); Mediterranean region (Spain, NW Africa ; Turkey, Lebanon)
From eros (Gk.: dawn, spring) and phaca (a genus of legumes in synonymy under
As/raga/us), referring to its early flowering
He1bs; mediterranean shrubland and bush land
Reference(s): Podlech 0993a); Podlech in Talavera el al. (1999)
Originally placed in Astragalus but sepa rated by various authors; Podlech (1993a)
suggested that it belongs in Sophoreae but in Flora Jberica (Podlech in Talavem el
al., 1999: 347-350) , he treated it as a subtribe within Ga legeae; it cannot be a
member of Sophoreae since il lacks the Inverted Repeat (Liston, 1995);
Wojciechowski el al. (2000) place it as sister to their Astragalean clade
Oxytropis De. lso2
A ragal/11s Neck. ex Greene (1897)
300 - 400 spp.j Eurasia, most numerous and diverse in subarctic and mounrainous
regions of C Asia, a long a be lt of the Tianshan, Pamirs-Altai, Baerh asita i and Altai
mountains to N & NW Ch ina, south to \YI Asia and Himalaya; N America (Canada
and W USA, c, 22 spp, [Barneby, 1952], reduced to 10 e ndem ic spp. and 3
circumboreal spp. by lsely [1998])
From oxy- (Gk: shallJ) and tropis (Gk,: kee l), for the characteristically beaked keel
Herbs or cushion-like subshrubs; mainly cold and continental temperate
grassland; many montane or alcimontane
Reference(s): Bunge (1874); listing in Yakovlev et al. (1996); Zhu & Ohashi (2000)
for N Eurasia and China; Barneby (1952); lsely 0998) for N Ame rica
Usually regarded as very close to As/raga/us and differing from it only in teclrnical
characters such as the pointed kee l with vascular bundles reaching the top of the
keel petals; rnolecular data, however, show that it is monophyletic and also not
even the sister group of Astragalus, although it is a me mbe1 of the Astragalean
clade (Wojciechowski et al.. 2000); Zhu & Ohashi (2000) recogn ise 3 subgenera
and 20 sections (c. 125 spp.) in China
Used in medicine, as a food ingredient and as o rnamen tals; some species are used
fo r forage, others r1re poisonous to Jivestock (causing locoism)
Biserrula L 1753
I sp, (with 2 subspp.); Mediterra nean region and Canary Islands, E & NE Africa
From bi- (L.: twice) and semtla (L.: a little saw); referring to the toothed crests on
the pods
Herbi mediterranean and tropical montane grassland, shrubland and disturbed areas
Reference(s): Mathews in Davis (1970: 48-49); Thulin (1983: 190-192)
Often included in As/raga/us but there are only 5 stamens and the winged pods
are distinctive; its position in the supertree of Wojciechowski et al. (2001) supports
it being treated separately
Biserrulo pe/ecinus Photograph bys. Collenette Usefu l pasture legume for acidic soHs and for hay
480 LEGUMES OF THE WORLD
Astrogo/us coccineus Photograph by M. Wojciechowski
Astrogolus monspessulonus
Astrogolus splnosus Photograph bys. Collenette
Astragalus L. 1753
Phaca L. sens strict (1753); Tragacantha Mill. (1754); Neodie/sia Harms (1905);
Astracantha Podlech (1983)
At least c. 2300 (largely based on curre ntly accepted names in Podlech, 1999) and
perhaps up to 2500 spp. in some 2 5 roxonomic sections; mainly N 1cmpcr:11c,
from S and C Europe (chiefly Medil •m1ne<t11 region, including N Afrl .1 l, Mltldle
East, SW Asia and Sino-Himalayan region to W China (c 1200 spp, most diverse
in Turkey, Iran and Afghanistan); also E Europe to C Asia, Mongolia, Siberia, NE
China and Japan (c. 620 spp., mostly in former USSR); western N America (c. 380
spp.); S America (c, 100 spp.); Af1ica (1 sp .)
From aslragalos (Gk . wo1d fo r a species of this ge nus)
Shrubs and herbsi mainly medi terranean , warm to cool temperate, and semi-arid
to arid continental temperate grassland and shrubland, a few montane to dry
tropica l and subtropical
Re fe rence(s): valious sectional revisions by Pocllech and co-workers (Demi, 1972;
Podlech, 1975a & b, 1983, 1984, 1988, 1990, 1993b, 1994; Agerer-Kirchhoff, 1976;
Agerer-Kirchhoff & Agerer, 1977; Ali, 1977; Ott, 1978; Podlech & Kozik, 1983;
Tietz, 1988, 1994; Wenniger, 1991; Gazer, 1993; Maasso umi, 1993; Tietz & Zarre,
1994; Zarre & Podlech, 1994; 1995). Regional accou nts for fo rme r U.S.S.R.
(Gontscharow in Komarov, 1946); Turkey (Chamberlain & Matthews in Davis,
1970: 49-254), Flora lberica region (Podlech in Talavera el al., 1999: 279-338); N
Ame rica (Barneby, 1964; Isely, 1998), and S America (Johnston, 1947). Recent
phylogenetic analyses by Sanderson & Listo n (1995); Sanderson & Wojciechowski
(1996); Wojciechowski el al 0999); Kazempour Osaloo et al. (2003)
Astracantha has been returned to Astragalus on both molecu la r and
morphological grounds (Liston & Wheeler, 1994; Sanderson & Liston, 1995; Zarre
& Podlech, 1995); single taxa have been split off as Eropbaca, Biserrula,
Barnebyella and Ophiocarpus (q.v.) Kazempour Osaloo el al. (2003) find the
latter two genera to be well supponed within Astragalus; molecular data support
th e separation of the first two genera and several other isolated species, bu t wide r
sa mpling is still needed
A few species are significant economicallyi A . gummifer Labill. (gum tragacCJ.nlh), is
used for its gum in a wide range o f indust ries (ar least 20 other species are noted
to produce gums [Duke, 1981]); A mongboliws Bunge var. daburicus (DC.)
Podlech (=A membra11aceus Bunge) is a w idely used traditional medicine; other
species (m ilk vetches) are used for human food (edible pods, but some species
toxic), forage, erosion co nt1 ol, fuelwood and as selenium and u ranium indica tors
(Allen & Allen, 1981); some species are poisonous to livestock (locoweeds)
Photograph by c. Grey-Wilson
TRIBE GALEGEAE 481
 _ _ _ _ _.-fl

Smirnowio turkestana Drawing by P. Ha lliday

Oreophyso microphyllo Drawing by P. Halliday
Bornebyello co/ye/no Drawing by P. Halliday
Oreopbysa (Ilu nge ex Boiss.) Borom. 1905
Opbiocarpus (Bunge) Ikonn 1977 Colutea sect, Oreophysa Bunge ex Boiss. (1872)
Aslragalus L. sect. Ophiocarp11s Bunge
""~
1 sp.; N Iran (Elburz Mts.)
1 sp.; fraq to Afghan istan and NW India From oro-(Gk.: mou ntain) andpbysa (Gk.: bladder) , re fening to its habitat and
From oph io- (Gk.: snake) and carpos (Gk.: fruit) , alluding to the long curved pod ~ inOated fruit
Herb; continental temperate and medite1ranean grassland and desert H erb; continental temperate montane sh1ubland, open hillsides
Reference(s): Podlech (1994) Refe rence(s): Browicz (1963b)
Kept distinct fro m As/raga/us by Podlech pending molecular ev idence, mainly A poorly-known taxon
because of the long narrow subcylindrica l unilocular pods, often somewhal
constricted between the seeds; Kaze mpour Osaloo el al, (2003), however, find
0 aitcbisonii (Baker) Podlech to be well supported within Astraga/11s Smirnowia Bunge 1876

CJ)
1 sp ; Afghanista n, I ran, Tu rkmenistan , Uzbekistan
' Named for M.N. Smir•nov (1849 - 1889), Russian bota nist
Barnebyella r odlech 1994 J
J Shrub; dry sandy rlaces in semi-desert
\ Reference(s): Vassilczenko in Komarov (194 1: 315; Engl . trans l. : 235 - 236); Allen &
As/raga/us sect. Mirae Sirj & Rech.f.
1 sp.; fran, Afghanistan , Pakistan ~
' Alle n (1981); Yakovlev et al. (1996)
Named after R.C. Barneby (1911-2000), British-born legume specialist at New Smirnowia turkestana B unge is used as a 1raditional m edicine, the isoflavans have
York and monographer of Astragalus in N America, as well <is several other anti-protozoa n prope1ties, e g. aga inst malaria
legume genera
Herb; arid and semi-arid mediterra nean and subtropical grassland and shrubland
Reference(s): Podlech (1994) Eremosparton Fisch . & C.A.Mey_ 1841
Originally desc ribed as a Do1ycnium, but acco rding to Podlech (1994) it is neither 3 s r p.; SE Russia and Caucasus to C Asia and NW China
th is nor As/raga/us; Kazempour Osaloo el al (2003), however, find B . calyc ina From eremia (Gk : desert) and spa11on (Gk.: broom), resembling Stipa
(Stocks) Podlech to be well supported within As/raga/us; the pod is unilocula r tenacissima (esparto grass)
and 1-seeded, the peta ls are haily and the calyx e longates d uring anthesis Shrubs; continental temperate semi-dese1t shrubland on plains
Reference(s): Vassilczenko in Komarov 094 1: 31 0 -311; Engl. tra nsl.: 232-234);
Yakovlev et al. 0996)
Colutea L 1753 Used for forage, re habilitation of arid lands and medici ne; weedy
c. 28 spp.; C and S Europe a nd Mediterranean region (including N Africa) east to
\YI and C Asia (where the majority of spp. occur), N China and \YI Himalaya and in
NE tropica l Africa
From colulea (the Gk. name for this p lant)
Shrubsi medi terranean, warm or continental temperate w oodland, shrubland,
bushland a nd montane grassland
Reference(s): Browicz (1963a); Yakovlev el al. (1 996)
Used as ga rden orname ntals (notably the bladder senna, C. arborescens L. and
hybrids); pasture p lants; for soil stabilisation, erosion control and hedges; also for
wood (fuel and timber); a multi-purpose legumino us shrub for arid climates

Eremosporton ophy/lum Drawing by P. Halliday

Coluteo orborescens Photograph by G. P. Lewis

TRIBE GALEGEAE 483

482 LEGUMES OF THE WORLD
 Lessertia minlata Photograph by 8. -E. Van Wyk
Sphaerophysa DC. 1825
2 spp.; Tu rkey, SE Russia and Caucasus, C As ia ro Siberia, Mo ngolia and N Chi na
From sphaero-(Gk.: sphere) andphysa (Gk: bladder), referring ro the inFlared
f1u ir
Herbs or subshmbs; halophytic; continen tal temperate g1assland and shrublancl to
semi-d ese1t
Reference(s): Clrnmberlain in Davis (1970: 44-46); Go rschkova in Ko1m11ov
(1941: 312; Engl. tiansl.: 235)
Used fo r forage and medici ne; S salsula (Pa ll ) DC. (A11strian peaweed), is a
noxious weed
Lessertia De. 1so2
c . 50 spp.; southern Afri ca (mainly \YI and NW Cape), 4 spp extend ing no rth to
the Kalahari-Highveld reg i on~[ transi ti on zone of BotswanLJ, Nam ibia and \V/
Zambia, 1 sp. extend ing to Angola and Ta nza nia; othe r spp largely in
Afromontan e regions to E & N of southern Afrie<1 , wit h l. perennans DC. to
Zimbabwe
Na med afte r Baron J.P.B. Delesser1 0773-1847), French industrialist, banker,
philanthrop ist and amateur botan ist; his herbariu rn fo 1ms the nucl eus of the
present herba rium a t Geneva
H e 1bs or subshrubs; rnedite 11'anean shrubland (fynbos), sem i-dese1t and
su btropical to tropical shrubland, th icket, wooded grassland and montane
g 1assland, orten in roc ky or sa ndy areas (K~lahari Sands in the tropi cs) , and in
seasonally w et habitats
Refere nce(s): Bolus (191 5); Lock (1 989) ; Ba lkwi ll & Bal kwill 0999); Schutte in
Goldblatt & Manning (2000: 494-495); D ini z in Pope et al (2003: 15- 19)
The genus is in need of revision
Several species possibly poisonous ro livestock
Lessertia lanata Ph otograph by 8 .-E. Van Wyk
484 LEGUMES OF THE WORLD
Sutherlandia montana Photograph by 5. Andrews Swainsona formosa Photograph by A. 5. George
Sutherlandia R.Br. ex \Yl,T.Aiton 1812
Trad itio nall y 5- 6 spp .; recent stu d ies, however, suggest only 2 spp shou ld be
recogn ised (Moshe et al., 1998); South Africa, Na mi bia, [Jotswa na
Named afte1James Sutherland (c 1639-1719), superintendent of the Botan ic
Gardens and Professor of Botany, Edi nburgh
H e 1bs or shrubs; mediterranean, se mi-d ese rt to subtrop ica l shmbland and
grassland , and coastal stmds
Refe rence(s): Phill ips & Dyer (1934); Sch rire & Andrews 0 992); Diniz in Pope et
al (2003: 12-15)
Th ree su bspecies recognised in S fmtescens (L.) R. Br. (Moshe et al., 1998) ;
Gold blatt & Man nin g (2000: 708) have placed Sutherla nd ia into synonymy unde r
lesserlia
Sutberlandiafn~tesce ns (cance1 bush, balloon 11ea, duck pea) is grow n as an
ornamental ; species widely li sed as a profound trad itional med icine in southern
Africa; 111ay have ca ncer-inhibiting propert ies
Swainsona salisb. 1so6
Cyc/ogyne [Jenth ex Lindi. (1839); Diplo/obiwn F.Muell (1863); \Vi/ldampia
A.S.George (1999)
84 spp.; Au stralia
Na med a fter Isaac Swainson (1746 - 181 2), London phys ician who deve loped a
private botan ic garden in Twickenham; he was co usin to th e eccentric and more
famous Willit1m Swainson w ho was thought , mistakenly, to have had the genus
named after hi m
H e1bs or subshrubs ; trop ical, medite1ranean and warm temperate regio ns, chie tly
arid and semi-arid grassland and desert , but also ex tend ing into shrubl<1nd, open
woodland and fo1est of coastal and montane 1egions and <il pi ne grass land
Reference(s): Lee (1948); Tho mrson 0993); Heenan 0 998a); Wagstaff el al
(1999)
The ge nus \Vi /lda mpia was erected by Geo rge 0 999) for the surerficia ll y
disti nctive Sturt's dese rt pea (S ,fo rmosa CG.Don) Joy Thomps .. p reviously know n
as Clian tbztsformosus (G Don) Ford & Vickery), bu t the analysis of Heen~n
(1998a) places it in the middle of Swainsona
Used as o rnamentals ( the fl owe1s have a wide 1ange of co lours) and fo r medicine;
some spec ies (poison pe as) are toxic to stock and affect the nervou s system; other
species a1 e used Lis forage
Swainsona sp. Photograph by G. P. Lewis
TRIBE GALEGEAE 485
 --
=--=-1
____-.:.,
;;:-

Streblarrhiza speciosa Illustration by Drake Ga/ego officinalis Photograph by T. Cope

Montigena novae-zelandlae Drawing by P. Halliday Clianthus punlceus Photograph by R.B.G .. Kew

Montigena Heenan 1998 Streblorrhiza End!. 1833

1 sp., now extinct; Phillip Island, near Norfolk Island (between Australia and New
1 sp.; New Zealand
Zealand)
From mons (L: mountain) and ·-gena' (L born), refen'ing to the habitat From strehlo- (Gk.: crooked) and rhiza (root), referl'ing to the shape of the root
preferences and disrribution in the mountains of E South Island
Scandent shrub or liana; probably subtropical thicket
Subshrub; warm temperate, subalpjne and alpine screes
Reference(s): Allen & Allen (1981); Heenan (2001)
Reference(s): Heenan 0998c); Wagstaff et al (1999) Streb/orrhiza speciosa Endl. (the Phillip Island glory pea) is extinct in the wild; it
Removal of M. novae-zelandiae (Hook.f.) Heenan from Swainsona makes the was formerly cultivated but is now almost ce1tainly lost (Lucas & Synge, 1978)
latter monophyletic and exclusively Australian (Heenan, 1998a), however, the ITS
Used as an ornamental
phylogeny of Wagstaff et al. 0999) places Montigena within Swainsona; the
validity of recognising Monligena thus remains equivocal

Gal.ega L. 1753
6 spp.; C and S Europe and Mediterranean region (including N Africa), east to
Clianthus Sol. ex Lindi. 1835 Turkey, Caucasu s and Pakistan; monrane E Africa from Ethiopia to Kenya
2 spp.; New Zealand (N Island) From gala (Gk .: milk) and agetn (Gk.: to act); a reputed galactogogue (stimulating
From cleos (Gk.: glory) and anthos (Gk,: flower) both the production and flow of milk in nursing mothers)
Shrubs; subtropical thicket and low forest, often on cliffs and in alluvial or Herbs or subshrubs; warm temperate and t1opical montane grassland, bushland,
colluvial sites
woodland and forest
Reference(s): Wagstaff et al. 0999); Heenan (2000) Reference(s): Gillett 0963a); Gillett in Milne-Redhead & Polhill 0971 : 1051 - 1054);
Clianthus punfcetis (G.Don) Sol. ex Lindi. is critically endangered with only one Garcia Murillo & Talavera in Talavera et al (1999: 267-274)
wild population (Wardle, 1991; Heenan, 2000); it is, however, widely cultivated as Molecular studies (Wojciechowski et al., 2000) suggest that its closest relative
the kakabeak, lobster claw or glory pea; C. maximus Colenso is more widespread may be Cicer in the Vicioid clade, rather than the other taxa treated as part of
in east N Island but still vulnerable (Heenan, 2000); C.formosus has been
Galegeae
transferred to Swainsona (Thompson, 1990) Ga/ega officinalts L. (goats me) is a traditional medicine; species also used as
Used as ornamenrals
ornamentals, as herbs in cheese making 1 for soil improvement and as bee plants;
but apparently poisonous to sheep and goats
Carmichaelia R.Br. 1825
Notospa1tium Hook.f. (1857); Corallospanium J.B.Armstr. (1881); Chordospaitium
Cheeseman 0911)
23 spp.; New Zealand (mainly S [but also NJ Island)
Named after Captain Dugald Carmichael (1722-1827), Scottish army officer and
botanist
Small trees, shmbs or subshrubs; temperate and subtropical montane and
submontane screes, moraines and disturbed ground
Reference(s): Simpson 0945); Heenan 0995, 1996, 1998a & b); Wagstaff et al.
0999)
Heenan 0998a & b) has shown convincingly that Chordospartium, Notospaitium
and Cora/lospa11ium should all be sunk into Carmichaelia; in a morphological
cladistic analysis the three genera are nested within Carmichae/ia, and all three
hybridise between themselves and with Carmichaelia
Used as ornamentals

Carmichaelia glabrata Photograph by G. P. Lewis Galega lindblomii (1-9), G. battiscombel (10-12) Drawing by H. Wood

TRIBE GALEGEAE 487

486 LEGUMES OFTHE WORLD
TRIBE
Hedysareae byJ.M.Lock
Tribe Hedysareae DC. 1825
Tribe Alhageae (DC.) Burnett (1835)
The history of Hedysareae is summarised by Polhill
(198lj). Originally the tribe, as defined by De Candolle
(1825) and Bentham (1865), incorporated most of the
leguminous taxa with jointed fruits, in six sub-tribes -
Coronilleae, Eu-Hedysareae, Aeschynomeneae,
Adesmieae, Stylosantheae and Desmodieae. Hutchinson
(1964) was the first to treat the tribe in its present,
restricted, sense. As well as the genera here placed in
Hedysareae, he included Corethrodendron and the
enigmatic Baueropsis (Bauerella) which is a synonym of
Cullen (Polhill & Raven, 1981: 56), possibly based on C.
badocanum (Blanco) Verde. (A.S. George,pers. comm.)
(= Baueropsis tomentosa (Schindl.) Hutch.). Polhill
regarded the Hedysareae in this restricted sense as a
single complex centred on Hedysarwn. and Onob1ychis,
with a few minor segregate . Choi ' hashi (1 96), on
the basis of pollen morphology, held that there is little
basis for maintaining Taverniera and Stracheya as
distinct from Hedysarum, and they supported Polhill's
(1981j) decision to sink Corethrodendron into
Hedysarum . Polhill (1981j) himself had pointed out that
it is difficult to separate Taverniera and some sections
of Hedysarum unambiguously, and that Ebenus and
Onobrychis are also very similar. Choi & Ohashi (2003)
placed Stracheya into synonymy under a more nan-owly
circumscribed Hedysarum, and recognised Sulla as a
distinct generic segregate; they also resurrected
Corethrodendron and maintained Taverniera as a
distinct genus. Choi & Ohashi (1998) proposed to
conserve the name Hedysarum with a new type species,
since the designated type species, H. coronarium L.,
should now be placed in Sulla. Some 180 species
currently in Hedysarum (estimates are increased here
based on Yakovlev et al., 1996), would otherwise need
new combinations under Stracheya (the next available
name with Corethrodendron and Sulla resurrected as
genera). Recent molecular sequence data
(Wojciechowski et al., 2000) suggest that four genera
usually placed in a polyphyletic Galegeae, i.e. Alhagi,
Caragana, Calophaca and Halimodendron, form a
single clade sister to Hedysareae and are best placed
within it - a course that is followed here.
The tribe as a whole usually occurs in dry open
places with a continental temperate or mediterranean
climate, and is restricted to Eurasia, N America, and the
Horn of Africa with Socotra. The tribe, as treated here,
is dominated by three large genera, Hedysarum,
LEFT Ebenus cretica Photograph by P. Cribb
Onobrychis and Caragana. Hedysarum occurs in the
mediterranean, warm temp erate and continental
temperate zones of Eurasia and N America (where there
are four species with several subspecies and varieties).
Polhill (1981j) listed the number of species as 'about 100'
but Yakovlev et al. (1996) feel that this is much too low
and suggest 200 (reduced to c. 160 species here,
excluding the recently resurrected generic segregates
Corethrodendron and Sulla (Choi & Ohashi, 2003)).
Onobrychis with c. 130 species occurs only in Eurasia
and its centre of diversity is in the continental temperate
and warm-temperate zones of the Irano-Turanian region.
The four genera sometimes regarded as part of Galegeae
but forming the Hedysaroid clade (Wojciechowski et
al., 2000) , are mainly Asian in distribution.
Halimodendron includes a single species, a tree of
usually saline arid or semi-arid C Asia. The species of
Alhagi are all spiny desert or semi-desert shrubs,
sometimes becoming weeds because of their ability to
sprout from the rootstock; specific limits are
controversial and it is possible that there may be just one
variable species. Calophaca is a small genus (perhaps 8
species) of shrubs and small trees, occurring from SE
Europe to C Asia, usually in semi-arid sites. The largest
genus in this group is Caragana, which may contain
upward of 70 species, all shrubs, occurring from SE
Europe to China.
Hedysareae is part of Polhill's 'Temperate Herbaceous
Group' and has been traditionally placed close to
Galegeae. Sanderson & Wojciechowski (1996), using
nrDNA ITS data, found a strongly supported Hedysaroid
clade within the Inverted Repeat Lacking Clade (IRLC).
This also included Alhagi, a genus whose position has
long been controversial. This clade was sister to a group
including Caragana, Calophaca and Halimodendron.
Wojciechowski et al. (2000; 2004) obtained similar results
using the chloroplast matK gene; this analysis placed the
'Hedysaroid clade' as sister to the 'Astragalean clade'.
However, their supertree (Wojciechowski et al., 2001)
placed the 'Hedysaroid clade ' outside both their
Astragalean and Vicioid clades. The Hedysareae remain
undersampled, and a full analysis including all the
genera is overdue, both to improve our understanding
of the relationships of the tribe as a whole, and also to
help in the definition of generic limits within it. In this
treatment, the Hedysareae comprises 12 genera and
( 400)- 427- (453) species (Fig. 54).
TRIBE HEDYSAREAE 489
 Calophaca

Caragana

Halimodendron

I
Alhagi rn
0
-<
(/')
?Eversmannia )>
Al
rn
)>
Hedysarum rn
Corethrodendron
Sulla
Taverniera

Onobrychis Caragana pygmaea Photograph by G. P. Lewis

Calophaca wo/garica Illustration by J. Harr
Sartoria
?Eben us Calophaca Fisch . ex DC . 1825
c. 5-8 spp .; 111ainly rnontane C Asia ; from E Ewope, Caucasus to Mongolia and
China
From calo- (Gk: bea utiful) and phacos (Gk.: le ntil)
Chesneya et al. Shrubs; continental ternpe1ate mon lane g1assland, shrubland find d1y forest
(see page 477) Refeie nce(s): listing in Yakovlev et al. 0996)
Used as ornamencals, fibre and coa rse cordage ( f101n the bark); seeds used as feed

Astragalean clade
(see page 477)
Caragana Fa br. 1763
c. 70- 80 spp.; E Europe, mainly Caucasus, \YI an d C Asia to Mongolia, Chi na and
Himalaya, and into the Russian Far East
D erived fro rn the Latinised version o f 1he Mongolian name Khargana used for
Vicioid clade these pla nts
(see page 477) Shrubs or t1 ees; mainly continental tempe rate desert, grassland, (xerophyti c)
s hrubland, w oodlan d and forest, often montane
Refere nce(s): Fu (1993: 13 - 67); Ya kovlev et al, 0996); Sanchir (1999; 2000)
Some species are morphologically extremely similar to some species of
Astragalus, sharing the cu sh ion-forming habit wi th persistent spinescent lea f
rac hises
Used as orname nt als (C arhorescens Lam. or Siberian pea tree), for fo1age,
erosion contro l, shelter-belts and wi11dbreaks 1 fuelwood, dyes, medicine, human
? =uncertain placement in generic group food (pods) and as bee plants

Halimodendron Fisc h, ex DC. 1825

1 sp ; E Eu rope, Tmkey, Caucas us, \YI and C Asia to Mongo lia, Ch ina and N to
Siberia
F1'0m ha/imo- (Gk.: ma ritime, salty) and dendron (Gk.: tree)
Trees or shrnbsi war m and con rin ental temperate shrublan d and bushland
lleference(s): Go isc hkova in Komarov (1941: 323-324; Engl transl,: 242 & 244);
Cha mberlain in Davis (1970: 547); Yakovlev et al, (1996)
Used for forage, e1osion control, as a bee plant, dye, fue hvood, and ornamental
(H /Jalodendro11 (Pa ll.) Voss, or Siberian sa ndthorn)

FIG. 54 Diagram of relationships in tribe Hedysareae after Wojciechowski et al. (2000) Halimadendron argenteum Illustration by 5. Edwards

TRIBE HEDYSAREAE 491

490 LEGUMES OFTHEWORLD
 Hedysarum varium Corethrodendron multijugum Photograph by G. P. Lewis
Alhaglgraecorum (1-10),A. maurorum (11) DrawingbyM. Grierson Alhagi graecorum Photograph bys. Collenette Drawing by H. Wood

Alhagi Gagnebin 1755 Hedysarum L. 1753

Stracbeya Benth (1853)
c. 3 spp.; E Meditenanean (including N Africa) 10 Wand C Asia, Himalaya, India, c. 140-180 spp.; Europe, Mediterranean (i ncluding N Africa), mainly in Caucasus
Mongolia and China; naturali sed in Australia 10 W and C Asia, Himalaya, Mongolia, China, Siberia and E Asia; N America ( 4
Derived from the Arnbic Al-Haggi/Al-Hajji - the pilgrim; said to be the Arabic spp_, 2 of which are circumboreai) from Canada to S USA
name for the plant
From bedy- (Gk.: sweet) and saran (GL broom)
Shrubs or herbs; medit·errn nean and continental temperate desert, semi-dese1t, Herbs or shrubsi mediterranean, temperate, wa rm and contine ntal temperate
shrubland and bushland
gra:.slJnd, shnibland and forest
Reference(s), Chamberlain in Davis (1970, 596-597); Yakovlev Cl979b} Lock & Kcrcn.·n~ •(s), Fedtschenko (1902); Hedge in Davis (1970, 549-560); Yakovlev el al.
Simpson 0991' 26-27); Awmack & Lock (2002) ' (1996) list more than 130 species from N Asia; Isely 0998); Choi & Ohashi (2003)
The tribal placement has been much disputed between Hedysareae and Galegeae, A large and complex genus with cent1 es of diversity in the M editerranean region
but molecular data suggest rhat its affin ities may lie with th e former and in C Asia; species delimitation is difficult, and the gem1s as a whole is in
(Wojciechowski et al, 2000); the numbe1 of species is disputed and there may serious need of revision; a genus in an 'active process of spedation' (Yakovlev et
only be a single variable species The generic name is often attributed 10 al., 1996); Choi & Ohashi (2003) recognise four sections in Hedysamm, sect.
Adanson, but Gagnebin published ii first Hedysarum with c. 30-35 spp., sect. M11/tlcau/ia with c. 100-140 spp. (the latter
Used for edible gums (e.g, manna which is a sweet exudate from twigs of /1 figure extrapolated from Yakovlev et al., 1996), and sect. ftttembranacea and sect
graecorum Boiss,), famine food (roots), medicine forage, dye and fuelwood ;
1
Stracheya which are monospecific
useful in erosion control and as bee plants, but weedy in some areas whe re Many of the species are actually or potentially va luable forage plants; some are
introduced Ce g , N America)
used as medicine and for soil improvemen t

Eversmannia Bunge 1838 Corethrodendron Fisch ex Basiner 1845

4 spp .; SE Russia and Caucasus, monrane N Iran, AFghaniscan, C Asia to China
4 spp.; C Asia to E Siberia
Named afte1 E.F. van Eversmann (1794-1860), Geiman botanist and zoologist at From core/bro- (Gk., broom) and dendron (Gk ., tree), for the shrubby habit of the
Kazan
:species
Shrubs; continental temperate desert, shrubland and grassland Shrubs; continenta l temperate, inontane grassland, forest and desert
Reference(s} Yakovlev et at. 0996)
Reference(s} Choi & Ohashi (2003)
Unlil rece ntly this genus was regarded as monospeci fic, but three local endemics Includes species placed p1eviously in Hedyst1rum sect Fruticosa B.Fedtsch.
have been described from C Asia (Yakovlev et al., 1996). Mironov & Sokoloff
(2000) have pointed out that the pods do not break into segments in the manne1
typical of the tribe; further work is needed to see if this really indicates that the
genus is misplaced in Hedysareae
Sulla Medik. 1787
7 sp p ; W Mediterra nean (including N Africa)
Used as an ornamen tal
Probably derived from the Spanish vernacular name for the p lant
Annual he1 bs, J sp. perennial (S coronaria (L) Medik ); dry mediterranean
shrubland, bushland, thicket and grassland to semi-desert
Reference(s} Valdes in Talavem et al (2000' 949-955, as Hedysantm); Neila &
Mohamed (2001); Choi & Ohashi (2003)
Includes species placed previously in Hedysanim secl Spl11osissi111a B.Fedtsch
and sect Eleurherotion Basin er
Sulla coronm1a (== Hedysarum coronariunz L.: sulla cl over, Spanish or Ita lian
sainfoin) is a va luable forage plant, also used as fodder, hay, medicine, humCJn
food (roots), g1een manure and cultivated as an ornamental

Eversmann/a subsplnosa Drawing by M. Warwick Sulla caronarla Illustration by unknown artist

TRIBE HEDYSAREAE 493

492 LEGUMES OF THE WORLD
 ~-- -.

Onobrychls corduchorum (1-9) Drawing by M. Grierson Sartoria hedysaroides Drawing by P. Halliday Ebenus cretlca Photograph by J. Hooper

Taverniera Dc. 1825 Sartoria Boiss. & Heldr. 1849

15 spp.; NE Afri ca, Arab ian Peninsula , Iraq and Iran east to Pakistan and India I sp.; S Turkey
Named after Joseph Sartori (1809 -1 880), Bava ri an physicia n, botanist and
Named after j ean Baptiste Tavernier (1605 -1 689), French traveller in the Levant
and India explorer of the Greek flora
H erb; mediterranean shrubland, 1nontane
Shn1bs or herbs; seasonall y dry tropical and subtropical desert, shru bla nd and
bushland Refere nce(s): Hedge in Davis (1970: 589 - 590)
Known only from the type; treated as a good genus by Hedge in Davis (1970); he
Re fe rence(s): Thulin (1985)
stated : 'diffe rs from Onobrycbis in the 3-celled ovary and 2- 3-seeded large ovate -
Ve ry close to Hedysamm and Corelhro<lendron, which diffe r only in the ir
oblong un armed fru it, and from Hedysan.tm by the non-lomentoid fruit, the
m ul tifo liolate, not tri- or unifoliolate, leaves (Thu lin, 1985); Choi & Ohashi (2003),
however, have maintained the genus
flowers with small corollas and the dwarf habit'
Used as a traditional medicine; grazed by ca mel s

Ebenus L. 1753
Onobrychis Mill 1754 c. 20 s pp.; S Europe , mainly E Medite rranean (i ncluding N Africa), but cent red in
Dendrobrychis (DC.) Galushko 0980); Xan1bob1ychis Ga lushko (1 980) Turkey, Arabia, SW Asia and W Himalaya
From the Greek ebenos (a wo rd probably of Egyp tia n origin fo r another pla nt ,
c. 130 spp.; Europe and mai nly Med iterranean region (including N Africa south
to Ethiopia), Causasus, \Y/ and C Asia east in to Siberia, M ongo\i a1 China and \V/ not this)
Suffrurices; medite rranea n sh rubla nd and forest, o n rocky (often limestone) slopes
Hima laya
Reference(s): Huber-Mora th 0 965; in Dav is, 1970: 590 - 596); Aytac (2000)
Apparently fro m ono- (G k.: ass) a nd eithe r bnicbo (Gk.: to eat greedily) or brycbo
Used as an ornamental, e.g., E_ crelica L. (C retan sainfoin)
(G k,: to bray), beca use the smell of the plant was supposed to excite do nkeys to
the point of braying
H e.rbs or shrubs, some forming thorny cushions; mediterranean, tempera te, wa rm
and continental temperate grassland and s hrubland (often xerophytic)
Refe re nce(s): Sirjaev (1925-1 926); Hedge in Davis 0 970: 560 - 589); numero us N
Asian taxa listed by Yakovlev el al. 0996); Menitskii (2001)
The form of the fruit has been much used in taxonomy, but this seems variable
and authorities differ considerably about the nu mber of taxa that me rit
recognition, and the ra nk at which they sho uld be recognised
Many species are valuable fo rage plants (e.g., 0 viciifolia Scop., o r sainfoin) ,
also used for erosion control and as bee plants; a few species are cultivated as
o rn a m e nt~ls

Onobrychis - several species Drawing by l. M. Ripley Ebenus plnnata (A-H), E. armltagei (I) Drawing by M. Y. Saleem

494 LEGUMES OF THE WORLD

TRIBE HEDYSAREAE 495
 TRIBE
Cicereae by J. M. Lock and N. Maxted
I
1 I
I
Tribe Cicereae AJef. 1859
Cicer, the only genus in Cicereae (Fig. 55), occurs
exclusively in the O ld World, with centres of
distribution in W and C Asia. One of the 43 species
occurs in NE tropical Africa. Cicer arietinum L., the
chickpea, is widely cultivated in warm te mperate and
subtropical regions. The genus was formerly placed in
Fabeae (as Vicieae), however, Kupicha 0977; 1981b)
showed clearly that the genus is distinct and reinstated
tribe Cicereae. Steele & Wojciechowski (2003) and
Wojciechowski (2003) find Cicer is best supported as
sister to the Trifoli eae -Fabeae clade.
Polhill Cl981a) had placed Cicereae as part of his
'temperate epulvinate series'. Doyle (1995) and Liston
0995) grouped the Cicereae, Trifolieae, Galegeae,
Hedysareae, Fabeae (as Vicieae) and some genera
currently in Millettieae sens. lat., on the basis that they
lack the inverted repea t (IR). Wojciechowski et al.
FIG . 55 Diagram of relationships to tribe Cicereae after Wojciechowski (2003); Steele & Wojciechowski (2003); Wojciechowski et al. (2004)
496 LEGUMES OF THE WORLD
(2000 pla ed ice rea within th ir Inv it d R Jpeat
la king Clade (JRL and within Lhi , in th ir idoid
dade. Galega and Ci ar si t r roup in thi
'tnaly, i to th r ~ t f th Vicioicl ciad whi h in lud
Fabeae and Trifolieae ( . cepl Parochetus which is
basally branching to the Vicic.id clade). The postulated
relationship with Galega is un xpected and, p erhu ps
because of this , the morpholog-i al similaritie and/ r
diffe rences remain unexplored . The analyses of
Wojciechowski (2003), Steele & Wojciechowski (2003)
and Wojciechowski et al. (2004), suggest that Cicereae
remains a fa irly is lated group w ith respect to Galega
and the Trifo lieae-Fab a clade. In future, tribe
Galegeae sens. strict. is likely to comprise the single
genus Galega, although relationships with Parochetus
and Cicereae need to be studied further to ascertain if
these should also be included.
Cicer arietinum Drawing by E. M. Stones Cicer cuneatum Photograph by S. Collenette
CicerL. c1153).
43 spp.; Med iterr-.i nean reg ion 10 \VI and C Asia, Ca na1y Islands and NE tropical
Africa
The origin of Cicer derives from rhe classical Latin name of th~ chickpea
Herbs, sometimes sh rnbby; medilerranean and warm and conrmenta1 tempera te
gr.1 land and shrubland; perennial species often montane ~r upland ,
Rcfcrc nce(s), va n der Maesen (1972); Santos-Guerra & Lewis (1986); Sefe1ova
Parochetus (see page 477)
(1995); Coles et al 0998) . .
Cicer arielinum L. is a major pulse c1o p (chick pea, pois ch1che, ga rbanzo, Benga l
g ram, chana), loin ing peas and lentils as one of the three main foodpu lses; rhe .
Galega (see page 477) seeds are also giound into flour, used in soup, dha l and bread or are mashed with
oils and spices into hummus; ot her uses include starch, medicine and fodder
Cicer CICEREAE
Fabeae (see page 505)
Trifolium (see page 500)
Trifolieae pro parte
(see page 500)
Cicer canarlense Drawing by G. P. Lewis
TRIBE CICEREAE 497
TRIBE
lrifolieae byJ. M. Lock
Tribe Trifolieae (Bronn) End!. 1830
Subtribe Trifoliinae Bronn (1822), as 'Trifolieae'
Subtribe Parochetinae (1998)
Tribe Trigonelleae O.E. Schulz (1901)
Tribe Ononideae Hutch. (1964)
Trifolieae forms a morphologically distinctive tribe,
although the position of both Ononis and Parochetus
has been questioned (see below). In total there are 6
genera and c. 485 species, of which more than half
belong to Trifolium (Fig. 56). The distribution of the
tribe is centred in the N temperate regions of the Old
World, particularly in areas of winter rainfall. Trifolium
itself has spread into the tropics on mountains, where
there has been considerable diversification, particularly
in Ethiopia. It is also the only genus of the tribe to
occur naturally in the New World. Parochetus occurs
only on palaeotropical mountains. The importance of
some genera as fodder legumes, particularly Trifolium
and Medicago, has led to their introduction to many
parts of the world.
Ononis was placed in a tribe of its own, Ononideae,
by Hutchinson (1964) and this has been followed by
some (e.g., Yakovlev et al., 1996). The distinctness of
Parochetus (and of Ononis) was emphasised by Small &
Jomphe (1989), and Chauclhary & Sanjappa (1998a) have
placed Parochetus in its own subtribe Parochetinae.
Within the core of Trifolieae, there are some
problems in generic delimitation, particularly between
Trigonella, Medicago and Melilotus, with some (e.g.,
Yakovlev et al., 1996) recognising the intermediate
genus Melilotoides. Distinctive species here placed in
Medicago have been variously segregated as Radiata
(Pseudomelissitus), Rhodusia, Crimea, Kamiella and
Factorovskya. This treatment follows Small (1987) and
Small et al. (1987) in recognising an expanded
Medicago including all those species with explosively
tripping flowers. In Trifolium, on the other hand, the
generic boundaries are reasonably clear, but the unit
can be treated either as a large genus with several
well-defined sections (the course followed here), or as
the separate genera Amoria, Cbrysaspis, Lupinaster
and Trifolium sens. strict. (see below).
Trifolieae forms part of the 'temperate epulvinate
series' of Polhill (198la). In the same volume Heyn
0981) was unable to suggest a clear relationship to any
LEFT Trifolium repens Photograph by G. D. Carr
other tribe. The morphological cladistic analysis of the
whole family by Chappill (1995) placed Trifolieae next
to Cicer. Kupicha (1977) had earlier suggested that
Cicer is closest to Trifolieae, with the adnation of the
stipules to the petiole in Trifolieae being the only
differential character; the tribes Cicereae and Trifolieae
also share the characters of long-stalked glandular
hairs and serrate leaflets with craspedodromous
venation. Doyle (1995) placed Trifolieae, along with
Carmichaelieae, Cicereae, Galegeae, Hedysareae,
Fabeae and some Millettieae in a group characterised
by the loss of the inve1ted repeat (IR) (Liston, 1995).
Endo & Ohashi (1997) placed Trifolieae as sister to the
Cicereae and Fabeae (as Vicieae) in a cladistic analysis
based on a range of non-molecular characters.
Wojciechowski et al. (2000) distinguish a Vicioid clade
that includes Trifolieae, Cicereae and Fabeae (as
Vicieae), as well as Galega. Within this clade,
Parochetus is basally branching to the rest of the taxa,
and Galega plus Cicereae form a sister group to a
paraphyletic Trifolieae, with Fabeae emerging as sister
to Trifolium. In a clade sister to Trifolium and Fabeae,
Wojciechowski et al. (2000) and Steele &
Wojciechowski (2003) place Ononis basally branching
to the sister monophyletic clades Medicago, and
Melilotus-Tn'gonella (Fig. 56). The latter three genera
comprise tribe Trigonelleae of Schulz (1901).
Given that molecular phylogenies do not support a
monophyletic Trifolieae in its current form, further
study may reinforce the pattern of relationships
suggested so far by these analyses. A tribe Trigonelleae
could be recognised including the genus Ononis, and
tribe Trifolieae would then only include the genus
Trifolium, sister to tribe Fabeae. The Trifolieae in its
broader paraphyletic sense is maintained here pending
further study. The 'supertree' of Wojciechowski et al.
(2001) is not supportive of the segregate genera of
Trifolium; more thorough sampling of Trifolium and
other large genera is desirable before any final
conclusions can be drawn.
TRIBETRIFOLIEAE 499
 Hedysareae (see page 489)

Galegeae (see page 477)

Parochet us

Galega (see page 477)

Cicereae (see page 496)

Parochetus africanus Drawing by c. Grey-Wilson Trifo/ium rubens Photograph by G. P. Lewis

Trifolium
P a roch etus Buch.-Ham ex D-Don 1825
2 spp.; 1 sp each o n mountains of Tropical Asia (Indian Subcontinent, Indo-
Ch ina. China, Malesia) and E Aflica
From para (Gk : near) and ocbetos (Gk.: brook); from the streamside ha bitat of
these species
Fabeae (see page 505) Herbs; tropical montane grassland and forest edges in damp places
Reference(s): Beckett & Polhill 0991); Beckett (1995); Chaudhary & Sanjappa
CI998b); Vidigal in Pope et al (2003: 35-37)
Paroc/Jetus afi•iccmus Polhill has been ti eared by Vidigal in Pope et al. (2003) at
species level and by Chaudhary & Sanjappa (1998b) as P. commu nis Buch .-Ham.
ex D Don subsp afitca nus (Polhill) L.B.Chaudhary & Sanjappa
Ononis Moderately tender ornamentals (blue clover, shamrock pea) in horticu lture

Trifoliu m L 1753
Amoria C.Presl (1831); Cb1ysaspis Desv. (1827); Lupinaster Fabr. (1759); Bobrovia
Meli lotus A P.Khokhr (1998)
Trigonella c 250 spp .; principally temperate Eurasian (c. 150 spp ., wit h c. 130 spp. in the
Medi terranean region including Turkey), New World (mostly temperate N
America, c 60 sp p ); also tropical and subtropical montane areas (c. 36 spp in
Africa, some in Andes of S Ameri ca)
From tri- (L : three) and/o/i1m1 (L : leaO, for the trifoliolate leaves
He1bs; mainly meditenanean, temperate and tropical monrane grassland
Medicago Reference(s): Zoha1y in Davis 0970: 384 - 448); Zohary & Heller 0984 - gene ri c
acco unt); Steiner et al. (1997); Jsely (1998); Munoz Rodriguez et al in Talavera et
al. (2000: 647 - 719); Wa tson et al. (2000); Vidiga l in Pope et al. (2003: 45 - 52)
Roskov (1990a & b) recogn ises Tlifolittm in~ restricted sense and t1eats Amon:a,
Cb 1ysaspis and Lupinaster as good genera The molecular studies of Watson et al.
(2000), while they support Trifo/ium sens. lat as monophyletic, support ne ither
Amoria nor Trifolium sens. strict. as mo nophyletic; C/11ysaspis is so suppo1ted but
its recog nition would make T!?folium paraphyletic; no members of Lupinaster
were included in the analysis; the nine montane Africa n taxa sampled form a
monophyletic g roup within Trifoli11m
Introduced world wide for fodd er, soil and pasture improvement, as honey plants,
hay and silage, and in hrnticullu re; T repens L. (w hite clover) is probably the most
widely grown species with the greatest impact on agricullure of any cultivated
forage plant; T pratense L. (red clover) is also used fof medicine; some species are
eaten as human food; crefoil dermatitis associated with photosensitisation is a
problem caused by animal ingescion of some TrifoUum species

FIG. 56 Diagram of relationships in tribe Trifolieae sens. lat. after Wojciechowski (2003); Steele & Wojciechowski ( 2003 )

Trlfolium repens Photograph by T. Cope

TR IBE TRIFOLIEAE 501

500 LEGUMESO FTHE WORLD

Ononls natrix Drawing by S. Makar
Ononis natrix Photograph by C. Grey-Wilson
Ononis L. 1753
c. 75 spp.; Europe, principally in the M~dltcrr.tncan reg ion (including N Africa
south to the highlands of Ed1iopia and Keny:1), Mataronesia, \YI Asia east to S
Siberia, China and \YI Himalaya
Derived from the classical Greek name for th e p lane
Herbs or shrubs; temperate, warm temperate and mediterranean grassland and
shrubland
Reference(s): Sirjaev (1932); Losa Espana 0958); Devesa in Talavera et al. (2000:
590 - 646)
Sometimes placed in the separate tribe Ononideae Hutch, because of its
monadelphous stamens and dimorphic anthe '" (by, e.g., Yakovlev et al, (1996)
and Devesa in Talavera el al. (2000))
Used as ornamentals, forage, human food, bee plants, dyes and medicine (e.g., O
spinosa l, or spiny restharrow and 0 repens L. , o r common restharrow)
Melilotus Mill. 1754
20 spp.; temperate Europe, Mediterranean and subtropi cal Asia and N Africa
Derived from the classical Greek name; from meli (Gk : honey) and Lo/Os (Gk.:
lotus, a name given to several fodder plants)
Herbs; mediterrane~n and warm temperate grassland and shrubland
Reference(s): Schul z (1901); Stevenson (1969)
See under Trigonella for notes on generic delimitation
Used fo r forage, cover crops, green manure, medicine, human food, as a bee
plant, hay and silage; widely introduced and oficn norura lised in waste places and
along roadsides; the most widespread species :ire M. a/bus Med ik. (white
swectdovcr) and M. officinalis (L,) Pallas (yellow sweetclover); improperly cured
hay c:in lend to cattle poisoning
Melilotus a/bus Photograph by S. Collenette
502 LEGUMES OF THE WORLD
~
~
Medicago polymorpha Drawing by S. Makar Medlcago arborea Photograph by G. P. Lewis
Trtgonella L. 1753
c . 55 spp.; Mediterrnnean region to C Europe, from the Canary Islands eastwards
to C Asia and Ind ia with an outlying species in Australia
From the diminutive of lrigonus (L: 3-cornered, triangular), re ferring to the
appeara nce of the corolla
H erbsi mediterranean and warm temperate grassland and shrubland
Reference(s): Vassikzenko (1953) (in Russian); Huber-Morath in Davis (1970:
452-482; some species placed here in Medicago); Bena (2001)
The boundaries between Meli/o!Us, Trigone//a and Medicago have long been
disputed; Small et al (1987) have proposed a fairly un ambiguo us definition of
Medicago which includes two sections fo rmerly placed in 7'/igonella, and this
is fo llowed here; it is supported, for the most part, by later work (Small el al,
1989) using seed surface characters; the ge nus Melilotoides He ist. ex Fabr.
(1763) included some iaxa regarded as inte rmediate between Melilotus and
Trigo,.ella; it is accepted by Yakovlev et al. (1996) who regard it as ancestral to
these two genera
Used for fodder, soil improvement, medicine, dyes and an insect repellen t; T.
foenum-graecum L. ( fenu greek, methi) is a pulse crop , and an important
condiment for flavouring food (both seeds and leaves are used)
Medicago L 1753
Radiala Medik, (1789); Faclorovskya Eig (1927); Rhodusia Vassilcz. (1972);
Psertdome/issitus Ovcz, Rassulova & Kinzikaeva 0978); Crimea Vassilcz. (1979);
Kc1111ie//a Vassilcz. (1979)
83 spp . (Small & Jomphe, 1989); Mediterranea n Region and W to C Asia
Derived from the classical name medice, for a crop plant (Gk .: implying 'from
Media' ( Persia) with refe 1ence lo its introduction from there to Europe), with the
female suffix -ago
Hel'bs or shrubs; mediterranean and warm-temperate grassland and shnib1and
Reference(s): Heyn 0963); Lesins & Lesins 0979); Small & jomphe (1989); Bena
(2001); Mehregan el al. (2002)
See notes under Trigonel/a for relatio nships; Small & j omphe (1989) include some
species of Melilotoides in Medicago; Factorovskya appears to be a geocarpic
Medicago (Small & Brookes, 1984); other distinctive species have been given
generic status (see above) but these genera are not recognised here, following
S1m1 II et al. (1987)
Medicago saliva L. (alfalfa; lucerne; purple medic) is a widely cultivated forage
legume for livestock and is a drought-resistant fodder a nd hay plant; species are
also used for medicine, h11man food (e .g., sprouts) , honey, green manu re,
indt1strial enzymes in biotechnology, pulp for fibre and as a source of fu el
Trigonella stellata Drawing by M. Y. Saleem
TRIBE TRIFOLIEAE 503
TRIBE
Fabeae by J. M. Lock and N. Ma~ted
Tribe Fabcae Rchb. 1832
Tribe Vicieae (Bronn) DC. (1825)
Widely known as the Vicieae, the correct name for this
tribe is Fabeae (see Greuter et al., 2000, A.tticles 19.4 and
18.5), since it must be based on the name of the type
genus of the family, Faba Mill. ( = Vicia L.). n1is does not
reflect on the names Leguminosae and Papilionoideae
(see introduction) whose use as alternative names for
Fabaceae and Faboideae respectively is sanctioned in
the International Code of Botanical Nomenclature
(Greuter et al., 2000; Article 18.5).
Fabeae is a well-clefinecl tribe, forming part of the
'temperate epulvinate series' (Polhill, 1981a). It contains
five genera, of which two (Lathyrus and Vicia) are
large . The tribe as a whole is centred in the Irano-
Turanian Region of the E Mediteffanean. Lathyrus and
Vicia, each with about 160 species, have very similar
distributions centred on the Mediterranean but
extending throughout Europe, N Asia and N and
tropical E Africa , with secondary centres in N America
and S America. One large group of species, some in
Vicia and some in Latbyrus, are superficially extremely
sim il ar and can only be distinguished by technica l
characters of the style. This group was in the past
recognised as the genus Orobus L. (Kupicha, 1981a).
Lens has 4 - 6 species and Pisum 2 or 3. Both include
important crop plants and, perhaps because of this,
their taxonomy is controversial. Both are E
Mediterranean genera with outlying species. The
monospecific genus Vavilovia, sometimes included in
Pisu.m, is confined to montane habitats in W Asia.
Kupicha (1981a) was unable to suggest a closest
FIG. 57 Diagram of relationships in tribe Fabeae after Steele & Wojciechowski (2003); Wojciechowski
LEFT Vicio foba Photograph by G. P. Lewis
relative of the tribe; she had previously (Kupicha, 1977)
excluded Abrus (Abreae) and Cicer (Cicereae) from it.
The morphological analysis of Chapp ill (1995) placed
Fabeae (as Vicieae) in a group with Astragalinae,
Galeginae, Loteae, Coronilleae, Cicereae and Trifolieae.
Doyle 0995) included these subtribes and tribes
(except Loteae and Coronilleae) in a clacle
characterised by the loss of the inverted repeat (the
IRLC), with Carmichaelieae (here included in Galegeae
sens. lat.), Cicereae, Ga legeae, Hedysareae, some
Millettieae , a nd Trifolieae. More recent work
(Wojciechowski et al., 2000) places Fabeae at the heart
of a Vicioid clade that includes Trifolieae (q.v.) and
Cicereae as well as Galega - a fragment of a
paraphyletic Galegeae. Fabeae (as Vicieae) appears
embedded within Trifolieae as sister to Tr!folium.
In the analyses of Steele & Wojciechowski (2003)
and Wojciechowski et al. (2004), Fabeae (as Vicieae)
forms a clearly rnonophyletic group in which Pisum is
sister to Lathyrus, and these two emerge as a well
supported clacle within a paraphyletic Vicia. A subclacle
of Vic ia species is sister to Lens. Within Latbyrus, the
cpDNA restriction site phylogeny of Asmussen & Liston
0998) agrees in general with dividing the genus into
sections previously recognised using classical taxonomic
methodology (e.g., Kupicha, 1983).
The publications of the Vicieae Database Project (e.g.,
Allkin et al., 1983 a & b) provide basic information for the
whole tribe. In this treatment the Fabeae is considered
to comprise 5 genera and c. 329 species (Fig. 57).
Trifolium (see page 500)
Vicia pro parte
Vici a pro parte
,,
Lens )>
OJ
rn
Lathyrus )>
rn
Pisum
Vavilovia
et al. (2004)
TRIBE FABEAE 505
 I

Lathyrus splendens Illustration by 5. Ross-Craig

Vicia sylvatlca Photograph by T. Cope Lathyrus japonicus Photograph by T. Cope

Vicia L. 1753
Faba Mill. (1754); Anatropostylia (Plitmann) Kupicha 0973)
c. 160 spp.; mostly N temperate: Europe and Asia (c. 110 spp., principally from
the Mediterranean and lrano-Turanian regions, some spp~ to China, Korea and
Japan); N to E Africa (c. 15 spp.); additional centres in N America (c. 17 spp.,
including 1 sp. in Hawaii) and temperate S America (c. 18 spp.)
Derived from the Larin name for a vetch
H erbs (most ly climbing); temperate, mediterranean and tropical mon cane
grassland, sh rubland and woodland
Reference(s): Davis & Plitmann in Davis 0970: 274-325); Kupicha (1976); Allkin
et al (1983a, b & c, 1986); Maxted (1993); Romero Zarco in Talavera et al. 0999:
361 - 417); Wouw et al. (2001; 2003)
Many species are widely introduced and naturalised (sorne are weeds)i vetches
are used extensively as cove r crops, for forage , hay, silage, erosion control and
green manure; V.faba L. (broad bean, fava bean) is a major pulse crop, also
eaten green, with many cultivars in the trade; V. narbonensis L. is a minor pulse
crop; several other species (e.g., V. saliva L., V. ervilia (1.) Willd and V. viltosa
Roth) are cultivated as fodder; V. saliva (common vetch) is also used for
medicine; in various species occasional toxicity (causing favism) is found from
amino glucosides in seeds
Lathyrus L 1753
c. 160 spp.; mostly N temperate regions: Europe and Asia (c. JOO spp., principally
Mediterranean and Irano-Turanian, some spp. to China, Korea and japan) and N
to E Africa (c. 5 spp.), with additional centres in N America (c. 30 spp.) and
temperate S America (c, 23 spp.)
From /athyros (Gk.: the name of a pea or pulse)
Herbs (often climbing); temperate , mediterranean and tropica l montane grassland,
shrubland and woodland
Refe rence(s): Bassler (1973, 1981); Kup icha (1983); Allkin et al. (1985, 1986);
Asmussen & Liston (1998); Neubert & Miotto (2001); Kenicer (2003)
Mony sped.,,; nrc widely introducc:d :md nnturallscd' used cx1c11sivcly os <:Md'
crops, fur fodder (e.g .. /•. clc~m L, /.. bfls1/111s I.. :ind L. sa//1111s L); as orn:tmc111als
(e.g., l odom111s 1- l>weet pc<•), '" lnt/folf11. I. I "'Crl:isting pc.11 :ind /_ $.J~VC.</1-is L.)
a nd n lnno:on rood (e.g., l . sn1iu11.< (gr.i:;s I"""'• !ndl:on pea, d1Jckllng vet hi. L
ochms (L.) DC. and L. montanus 13ernh. (with edible root tubers) and also for
erosion control, as green manure and for medicine; toxins are present in some
species, causing lathyrism

Lens Mill. 1754

4 - 6 spp.; Mediterranean region to W Asia
Derived from the classical name for the plant; a lens (magnifying glass) is so
named for its resembl ance to a lentil seed
Herbs (sometimes climbing); woodland, mediterranean shrubland and tropical
altimontane grassland; weedy
Reference(s): Davis & Plitmann in Davis 0970: 325 -328); Ladizinsky & Abba
(1996); Ferguson et al. (2000); Mayer & Bagga (2002)
Maye r & Bagga (2002) find strong support for a monophyletic Lens, with L.
nigricans (M.Bieb.) Godr. being basally branching, and L odemensis Ladiz. most
likely sister to L, culinaris Medik.
111e importance of L cttlinaris as a cu ltivated crop - and the search for wild
relatives - has led to numerous taxonornk studies and often to conflicting
conclusions
Lens culinaris (lentil) is a major food (pulse) crop and is the only cultivated
species in the genus; also used for starch extracted from seeds 1 flour and dhal·
species are used as fodder, green manure and for medicine '

Lens cullnaris Drawing bys. Makar Lathyrus boissierl Illustration by M. Grierson

TRIBE FABEAE 507

506 LEGUMESOFTHEWORLD
Pisum sativum (line JI 2822) Photograph by A. Davis Plsum sativum var. arvense Photograph bys. Collenette

Pisum i. 1753
2- 3 spp.; Mediterranean and W Asia; cultivated worldwide in temperate regions
Derived from the Latin name for the pea planr
Herbs (often climbing); mediterranean grassland and shrubland; weedy
Reference(s} Davis in Davis (1970: 370-372); Ben-Ze'ev & Zo haty 0973);
Lamprecht 0974); Maxted (2000); Maxted & Ambrose (2001)
The taxo nomy is complicated with both inter- and infraspecific classification
disputed
Piswn salivum L (common or garden pea) is a major pulse and green vegetable
crop, with many cultivars in the trade; also grown as fodd er, ground cover, green
manure, hay and silage

Vavilovia Al.Fed. 1939

1 sp.; W Asia (Caucasus)
Named for NJ , Vavilov, Russian geneticist, economic botanist, and plant
geogrnpher 0887 - 1943)
Herb; mediterranean montane vegeration (scree slopes)
Reference(s): Davis in Davis 0970: 372-373)
Sometimes treated as pa1t of Pisum but now generally accepted as a separate
genus

Vavilavla formasa Drawing by unknown artist Plsum sativum Illustration by E. M. Stones

TRIBE FABEAE 509

508 LEGUMES OF THE WORLD
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Index to illustrations
Abaremafilamentosa (Benth.)Pittier 205
- lehmannii (Britton & Killip)Barneby & J.W.Grimes 205
Abrn.s aureus R.Vig. subsp. aureus 390
- canescens Baker 390
- precatorius L. 388, 390, 391
Acacia anthochaera Maslin 186
- aphylla Maslin 190
- brandegeana I.M.Johnst. 189
- daemon Ekman & Urb. 191
- hindsii Benth. 189
- hopperiana Maslin 188
- lanuginophylla RS.Cowan & Maslin 189
- nilotica (L.)Delile subsp. kraussiana (Benth.)Brenan
190
- pervillei Benth. 188
- piptadenioides G.P.Lewis 188
- visco Lorentz ex Griseb. 190
- willardiana Rose 190
Acmispon subpinntilus (Lag.)D .D. okol ff 462
Acosmium bracbysracbywn (B nth . y, kovlev 236
- lentiscifolium Schott 236
Acrocarpus fraxinifolius Wight & Arn. 133
Adenanthera pavonina L. 167
Adenocarpus complicatus (L.)J.Gay ex Gren. & Godr. 289
Adenodolichos rupestris Verde. 408
Adenolobus garipensis (E.Mey.)Torre & Hille. 59
Adenopodia paten.s (Hook. & Arn.) J.R.Dixon ex Brenan 182
Adesmia gracilis Meyen ex Vogel 312
- guttulifera Sandwith 312
-Sp. 312
Aenictophyton reconditum A.T.Lee 358
Aeschynomene indica L. 329
- martii Benth. 329
- mediocris Verde. 329
Afgekia sericea Craib 371
Ajzelia quanzen.sis Welw. 91
Aganope thyrsijlora (Benth.)Polhill 373
Ai1yantha schwelnfurthli (Taub.)13nunmitt 245
Alt.scbf11dllum godefroycm11111 (Kuntze)H.Ohashi 439
Alantsilodendron ramosum Villiers 177
Albizia mainaea Villiers 211
- multiflora (Kunth)Barneby & J.W.Grimes var. multiflora
211
- viridis E.Fourn. var. zygoides (Baill.)Villiers 211
Aldina latifolia Benth. 221
Alexa canaracunensis Pittier 229
Alhagi graecorum Boiss. 492
' - maurorum Medik. 492
Alistilus jumellii (R.Vig.)Verdc. 423
Almaleea subttmbellata (Hook.)Crisp & P.H.Weston 347
Alysicarpus glumaceus (Vahl)DC. 444
- pubescens J. .L'lw 444
Amblygonocaipus andongensis (Oliv.)Exell & Torre 167
Amburana cearensis (Allemao)A.C.Sm. 220
Amherstia nobilis Wall. 95
Amicia zygomeris DC. 312
Am.mopiptanthus mongolicus (Maxim.)S.H.Cheng 264
Ammothamnus gibbosus (DC.)Boiss. 242
A morpha fi,.,,;ticosa L. 302
- nana Nutt. 302
Amphicaipaea africana (Hook.f.)Harms 420
Amphi111tt · pteroct11poides Hann · 2 7
Amphithalea ericifolia (L.)E ·kl. ~ Z yh. 269
Anade11a11thera colti/Jri11a lU13renan var. colubrina 179
Anagyrisfoetida L. 264
Anarthropbyllum desideratum (DC.)Benth. 289
Andirafi·axinifolia Benth. 311
- inermis (W.Wright)DC. 311
- legalis (Vell.)Toledo 311
Androcalymma glabrifolia Dwyer 118
Angylocalyx braunii Harms 230
Antheroporum pierrei Gagnep. 370
Anthonotha conchyliophora (Pellegr.)J.Leonard 102
Anthyllis montana L. subsp. jaquinii (A.Kern.)Hayek 459
- vulneraria L. subsp. maura (Beck)Maire 459
Antopetitia abyssinica A.Rich. 460
Aotus genistoides Turcz. 346
Aphanocalyx obscun,;s Wieringa 108
- richardsiae Q.Leonard) Wieringa 108
Aphyllodium glossocaipum Pedley 437
Apios tuberosa Moench 403
Apoplanesia paniculata C.Presl 301
Apuleia leiocarpa (Vogel)J.F.Macbr. 115
Apurimacia michelii Harms 386
Arachis hypogaea L. 326
Arapatiella psilophylla (Harms)R.S.Cowan 151
Archidendron lucidum (Benth.)I.C.Nielsen 202
Arcbidendropsis fulgens (Labill.)I.C.Nielsen 203
- paivana (E.Foum.)I.C.Nielsen subsp. tenuispica
(Harms)I.C.Nielsen 203
Arcoa gonavensis Urb. 132
Argyrocytisus balfandierl (Mair ) IG.1ynaud 290
Argyrolobium i/remoense M.P lei r x Du Puy & Labat 287
Arthroclianthus deplanchei Hochr. 438
A :palatbus linearis (Burm.f.)R.Dahlgren 277
- pe1foliata (Lam.)R.Dahlgren subsp. phillipsii R.Dahlgren
277
- teres Eckl. & Zeyh. subsp. teres 277
Astragalus coccineus Brandegee 481
- monspessulanus L. 481
- spinosus (Forssk.)Muschl. 481
Ateleia berbe11-smithii Pittier 220
Aubrevillea kerstingii (Harms)Pellegr. 166
Augouardia le1es111i P ·llegr. 79
Au.strodolich ermb1111dus (M.B.Scott)Verdc. 423
Austrosteenisia blackii (F.Muell.)R.Geesink 372
Baikiaea plurijuga Harms 82
Balsamocarpon brevifolium Clos 20, 147
Baphia m.adagascariensis C.H.Stirt. & Du Puy 246
Baphiastrum bracbycarpum Harms 247
INDEX TO ILLUSTRATIONS 545
 Baphiopsis parvijlora Baker 224
Baptisia australis (L.)R.Br. 265
Barbieria pinnata (Pers.)Baill. 401
Bark/ya syringifolia F.Muell. 64
Barnebydendron riedelii (Tul.)].H.Kirkbr. 68, 73
Barnebyella calycina (Stocks)Pocllech 482
Batesiafloribunda Spruce ex Benth. 148
Baudouiniajluggeiformis Bail!. 114, 125
- louvelii R.Vig. 114
- rouxevillei H.Perrier 114
Bauhinia grandidieri Bail!. 61
- hildebrandtii Vatke 61
- madagascariensis Desv. 56
- pwpurea L. 61
- weberbaueri Harms 67
Behaimia cubensis Griseb. 381
Bergeronia sericea Micheli 381
Berlinia brnneelii (De Wild.)Torre & Hille. 103
Bikinia evrardii (Bamps)Wieringa 107
- pellegrinii (A.Chev.)Wieringa 107
Biserrnla pelecinus L. 480
Bituminai·ia bituminosa (L.)C.H.Stirt. 450
Blancbetiodendron blanchetii (Benth.)Barneby &
].W.Grimes 205
Bobgunnia madagascariensis (Desv.)].H.Kirkbr. &
Wiersema 217
Bocoa prouacensis Aubl. 217
Bolusafra bituminosa (L.)Kuntze 409
Bolusanthus speciosus (Bolus)Harms 239
Bolusia resupinata Milne-Reclh. 278
Bossiaea dentata (R.Br.)Benth. 354
- priessii Meisn. 356
-Sp. 356
Bowdicbia vi1-gilioides Kunth 237
Bowringia discolor J.B.Hall 246
Bracbycy lix vageleri (Harms)R.S.Cowan 97
Bracbystegia torrei Hoyle 106
Brandzeia filicifolia Ba ill. 74
Brenierea insignis Humbert 60
Brodriguesia santosit: R.S.Cowan 92
Brongniai·tia sp. 256
Brownea gr-andiceps ]acq. 98, 109
- macrophylla Linden 55, 98
Browneopsis disepala (Little)Klitg. 98
Brya ebenus (L.)DC. 316
Bryaspis humularioides Gledhill 332
Burkea africana Hook, 158
Burkilliodendron album (Ridl.)Sastry 377
Bussea massaiensis (Taub.)Harms 152
- occidentalis Hutch. 152
Butea monospenna (Lam.)Taub. 407
Cadia pwpurea (G.Piccioli)Aiton 241
- sp. cf. pubescens Bojer ex Baker 241
Caesalpinia bahamensis Lam. 140
- cassioides Willd. 140
- madagascariensis (R.Vig.)Senesse 126
- putcbem:ma (L.)Sw. 140
Cajanus cajan (L.)Huth 412
- scarabaeoides (L.)Thouars 412
Calia secundijlora (Ortega)Yakovlev 233
Calicotom.e spinosa (L.)Link 293
Callerya megasperma (F.Muell.)Schot 370
Calliandra babicma Renvoize 197
- coccinea Renvoize 198
546 LEGUMESOFTHEWORLD
- houstoniana (Mill.)Standl. var. acapulcensis (Britton &
Rose)Barneby 198 Coch lianthu.s montanus Harms 403 - monetaria L.f. 327
- lanata Benth. 198 Codariocalyx motorius (Houtt.)H.Ohashi 441 Dalbergiella nyassae Baker f. 373
- leptopoda Benth. 198 Cojoba arborea (L.)Britton & Rose 199 Dalea botterii (Rydb.)Barneby 304
- tumbeziana ] .F.Macbr. 197 Collaea cipoensis Fortunato 398 - carthagenensis (Jacq.)].F.Macbr. var. brevis (J.F.Macbr.)
Cologania broussonettii DC. 416 Barneby 304
Calliandropsis nervosus (Britton & Rose)H.M.Hern. &
P.Guinet 176 Colophospermum mopane CJ.Kirk ex Benth.)].Kirk ex - cliffortiana Willd. 304
Callistachys lanceolata Vent. 351 Leonard 76 Dalhousiea bracteata (Roxb.)Benth. 245
Calophaca wolgarica (L.f.)DC. 491 Colutea arborescens L. 482 Daniellia alsteeniana Duvign. 77
Calvi/lea racemosa Bojer ex Hook. 155 - oliveri (Rolfe)Hutch. & Dalziel 77
Calopogonium. caeruleum (Benth.) Sauv. 416
- mucunoides Desv. 416 Conzattia multijlora Stand!. 154 Daviesia cordata Sm. 341
Calpocalyx dinklagei Harms 168 Copaifera palustris (Symington)de Wit 84 - euphorbioides Benth. 341
Calpurm:a a.urea (Aiton)Benth. 271 -sp. 84 - grossa Crisp 341
Camoensia scandens (Welw.)].B.Gillett 245 Cordeauxia edulis Hems!. 135 - incrassata Sm. 353
Campsiandra sp. 159 Cordy/a africana Lour. 221 Decorsea grandidieri (Baill.)R.Vig. 415
- madagascariensis R.Vig. 214 Deguelia densijlora (Benth.)A.M.G.Azevedo, ined. 380
Camptosema coriaceum (Nees & Mart.)Benth. 397
Coretbrodendron multijugum (Maxim.) B.H.Choi & Delonixfloribunda (Baill.)Capuron 154
Campylotropis polyantha (Franch.)Schindl. 434
H.Ohashi 493 - pumila Du Puy, Phillipson & R.Rabev. 510
Canavalia madagascariensis JD.Sauer 396
- rosea (Sw.)DC. 396 Coronilla valentina L. subsp. glauca (L.)Batt. 458 - regia (Bojer ex Hook.)Raf. 154
- villosa Benth. 396 Coulteria (Brasilettia) velutina (unpublished combination) Dendrolobium umbellatum (L.)Benth. 436
139 Derris trifoliata Lour. 382
Candolleodendron bracbystachyum (DC.)R.S.Cowan 219
Caragana pygmaea (L.)DC. 491 -sp. 139 Desmanthus bicornutus S.Watson 175
Carmichaelia glabrata G .Simpson 486 Coursetia glandulosa A.Gray 471 - oligospermus Brandegee 175
Carrissoa angolensis Baker f. 409 - grandijlora Benth. ex Oerst. 471 Desmodiastrum belgaumense (Wight)A.Pramanik & Thoth.
- rostrata Benth. 471 444
Cascamnia astragalina Griseb. 323
Cassia hippophallus Capuron 124 Craibia zimmermannii (Harms)Dunn 376 Desmodium vargasianum B.G.Schub. 440
- javanica L. 124 Cranocarpus mai1ii Benth. 316 - velutinum (Willd.)DC. 440
- moschata Benth. 124 - mezii Taub. 316 Detarium senegalense ].F.Gmel. 84
Craspedolobium schochii Harms 378 Dewevrea bilabiata Micheli 375
Castanospennum australe A.Cunn. & C.Fraser 229
Cratylia hypargyrea Mart. ex Benth. 398 Dialium guianense (Aubl.)Sandwith 120
Cathormion umbellatum (Vahl)Kosterm. 207
Cedrelinga cateniformis (Ducke)Ducke 201 Cristonia biloba (Benth.)J.H.Ross 258 - madagascariense Baill. 120
Cenostigma gardnerianum Tul. 142 Crotalaria calycina Schrank 280 - orientate Baker f. 120
- capensis ]acq. 279 - unifoliolatum Capuron 120
Centrolobium ochroxylum Rose ex Rudd 321
- cornetii Taub. & Dewevre 280 Dichilus reflexus (N.E.Br.)A.L.Schutte 285
Centrosema arenarium Benth. 402
- plumieri (Turpin ex Pers.)Benth. 402 - flavicarinata Baker f. 280 Dichrostachys akataensis Villiers 177
Ceratonia siliqua L. 133 - grevei Drake 279 - cinerea (L.)Wight & Arn. 177
Cercis siliquastntm L. 59 - micans Link 272 Dico1ynia guianensis Amshoff 121
- ocbroleuca G.Don 279 Dicraeopetalum capuronianum (M.Peltier)Yakovlev 240
Chadsia grevei Drake var. latifolia (R.Vig.)Du Puy & Labat
385 - variegata Baker 280 - mahafaliense (M.Peltier)Yakovlev 240
- versicolor Bojer 385 Cruddasia insignis Prain 400 Dicymbe alstonii Sandwith 101
Chaetocalyx longiloba Rudd 55, 314 Crndia choussyana Stand!. 87 - bymenaea Barneby 101
Chamaec1·ista desvauxii (Collad.)Killip 122 - gabonensis Harms 87 Didelotia unifoliolata ].Leonard 103
- nigricans (Vahl)Greene 122 - klainei De Wild. 87 Dillwyniafloribunda Sm. 347
Cryptosepalum pseudotaxus Baker f. 100 Dimorpbandra gardneriana Tul. 157
- ocbnacea (Vogel)H.S.I1win & Barneby 122
-sp. 100 Dinizia excelsa Ducke 166
Chapmannia prismatica (Sesse & Mos,:.)Thulin 324
- reghidensis Thulin & McKean 324 Cullen obtusifolium (DC.)C.I-I.Stirt. 450 Dioclea sp. aff. grandijlora Mart. ex Benth. 395
Cyamopsis senegalensis Guill. & Perr. 362 Diphyllarium mekongense Gagnep. 404
Chesneya 1ytidosperma]aub. & Spach 478
Chidlowia sanguinea Hoyle 160 Cyatbostegia mathewsii (Benth.)Schery 219 Diphysa americana (Mill.)M.Sousa 333
Chloroleucon mangense (Jacq.)Britton & Rose 206 Cyclocarpa stellaris Urb. 330 - sp. 333
Cyclolobium brasiliense Benth. 254 Diplotropis racemosa (Hoehne)Amshoff 237
Chorizema retrorsum J.M. Taylor & Crisp 350
- ilicifolium Labill. 350 Cyclopia intermedia E.Mey. 268 Dipogon lignosus (L.)Verdc. 425
Christia vespertilionis (L.f.)Bakh.f. 443 - maculata (Andrews)Kies 266 Dipteryx alata Vogel 251
Chrysoscias paruiflora (Bolus)C.A.Sm. 409 Cylicodiscus gabunensis Harms 172 Diptychandra aurantiaca Tu!. 160
- pauciflora (Bolus)C.A.Sm. 409 Cymbosema roseum Benth. 397 Discolobium psoraleifolium Benth. 315
Cicer arietinum L. 497 Cynometra abrahamii Du Puy & R.Rabev. 89 Distemonanthus benthamianus Baill. 115
- crassifolia Benth. 89 Disynstemon paullinioides (Baker)M.Peltier 376
- canariense A.Santos & G.P.Lewis 497
- cuneatum A.Rich. 497 - spp. 89 Dolicbopsis paraguariensis Hass!. 429
Cladrastis sinen.sis Hems!. 233 Cytisophyllum sessilifolium (L.)O.Lang 290 Dolichosfangista R.Vig. 424
Clathrotropis macroca1pa Ducke 238 Cytisopsis dorycniifolia Jaub. & Spach 465 - filifoliolus Verde. 424
Cleobulia multijlora Mart. ex Benth. 397 Cytisus proliferns L.f. 292 - trilobus L. 424
- scoparius (L.)Link 292 Dorycnium. eriophthalmum Webb & Be1thel. 464
Clianthus puniceus (G.Don)Sol. ex Lindi. 486
Cliton:a bracbystegia Benth. 401 - pentaphyllum Scop. 464
- lasciva Bojer ex Benth. 401 Dahlstedtia pinnata (Benth.)Malme 379 Dorycnopsis gerardii (L.)Boiss. 461
- ternatea L. 392 Dalbergia cearensis Ducke 327 Droogmansia pteropus (Baker)De Wild. 439
Clitoriopsis mollis R.Wilczek 403 - ecastaphyllum (L.)Taub. 327 Dumasia villosa DC. 418
- lemurica Bosser & R.Rabev. 327 Dunbaria cumingiana Benth. 411
INDEX TO ILLUSTRATIONS 547
Duparquetia orchidacea Bail!. 113
Dussia tessmannii Harms 231
- sp. 231
Dysolobium grande (Benth.)Prain 414
Ebenopsis confinis (Standl.)Barneby & J.W.Grimes 208
Ebenus armitagei Schweinf. & Taub. 495
- cretica L. 488, 495
- pinnata Aiton 495
Echinospartum algibicum Talavera & Aparicio 293
Ecuadendron acosta-solisianum D.A.Neill 95
Eleiotis rottleri Wight & Arn. 445
Elephantorrhiza elephantina (Burch.)Skeels 170
Eligmocarpus cynometroides Capuron 114
Elizabetha coccinea Schomb. ex Benth. var. oxyphylla
(Harms)R.S.Cowan 97
Erninia antennulifera (Baker)Taub. 419
Endertia spectabilis van Steenis & de Wit 85
Endosamara racemosa (Roxb.)R.Geesink 370
Englerodendmn usambarensis Harms 102
Entada polystachya (L.)DC. 169
Enterolobium cyclocaipwn Qacq.)Griseb. 212
- schomburgkii (Benth.)Benth. 212
Eperuafalcata Aubl. 78
- grandiflora (Ducke)R.S .Cowan 78
- jenmanii Oliv. subsp. sandwithii RS.Cowan 78
Eremosparton aphyllum (Pall.)Fisch. & C.A.Mey. 483
Erichsenia uncinata Hems!. 342
Erinacea anthyllis Link 294
Eriosema psoraleoides (Lam.)G .Don 411
- robinsonii Verde. 411
Erophaca baetica Boiss. 480
Errazurizia megaectrpa l.M.Johnst. 302
E1ythrina madagascai'iensis Du Puy & Labat 413
- peruviana Krukoff 413
Erythrophleum couminga Bail!. 157
- lasianthum Corbishley 157
Erythrostemon (Caesalpinia) calycina (unpublished
combination) 141
- gilliesii Klotzsch 141
Etaballia dubia (Kunth)Rudd 306, 321
Euchilopsis lineci1·is (Benth.)F. Muell. 344
Eu ch resta f 01·mosana (Ha yata)O hwi 261
- horsfieldii (Lesch.)Benn. 261
- japonica Benth. 261
Eurypetalum batesii Baker f. 77
- tessmannii Harms 77
- unijugum Harms 77
Eutaxia myrtifolia (Sm.)R.Br. 347
Eversmannia subspinosa (Fisch. ex DC.)B.Fedtsch. 492
Exostyles amazonica Yakovlev 224
- venusta Schott ex Spreng. 224
Eysenhardtia orthocarpa S.Watson 303
Faidherbia albida (Delile)A.Chev. 196
Falcatai'ia moluccana (Miq.)Barneby & J.W.Grimes 202
Fiebrigiella gracilis Harms 324
Fillaeopsis discophom Hanns 171
Fissicalyxfendle1'i Benth. 324
Flemingia strobilifera (L.)W.T.Aiton 413
Fordia albiflora (Prain)Dasuki & Schot 374
Gagnebina pterocarpa (Larn.)Baill. 176
Galactia jussiaeana Kunth 398
Galega battiscombei (Baker f.)J.B.Gillett 487
548 LEGUMESOFTHEWORLD
- lindblomii (Harms)j.B .Gillett 487
- o.fficinalis L. 487
Gastrolobium. bilo~um R.Br. 351
- bracteoloswn (F.Muell .)G.Chandler & Crisp 352
- pyramidale T.Moore 352
- rubrum (Crisp)G .Chandler & Crisp 352
Geissaspis c1'istata Wight & Arn. 332
Genista microphylla DC. 295
- tinctoria L. 295
- sp. (Rivasgodaya nervosa Esteve) 295
Genistidium dumosum I.M.]ohnst. 472
Geoffroea deco1ticans (Gillies ex Hook. & Arn.)Burkart 323
- spinosa ]acq. 323
Gigasiphon humblotianum (Baill.)Drake 62
- macrosiphon (Harms)Brenan 62
- schlechte1'i (Harms)de Wit 62
Gilbeniodendron bilineatum (Hutch. & Dalziei)j.Leonard
104
Gilletiodendron kisantuense (De Wild.)j.Leonard 82
- mildbraedii (Harms)Vermoesen 82
- pierreanum (Harms)].Leonard 82
Gleditsia caspica Desf. 130
- japonica Miq. 130
Gliricidia brenningii (Harms)Lavin 466
- sepium Qacq.)Walp. 468
Glycine max (L.)Merr. 421
Glycyrrhiza lepidota Pursh 478
Gompholobium capitatum A.Cunn. 353
- minus Sm. 340
- tomentosurn Labill. 340
Goniorrhachis marginata Taub. 73
Gonocytisus pterocladus (Boiss.)Spach 294
Goodia lotifolia Salisb. 356
Gossweilerodendron balsamiferum (Vermoesen)Harms 75
Grazielodendron rio-docense H.C.Lima 326
Griffonia physocaipa Bail!. 60
Gueldenstaedtia multiflora Bunge 479
Guibourtia coleosperma (Benth.)J. Leonard 81
- schleibenii (Harms)].Leonard 81
Guilandina bonduc L. 143
- ciliata Bergius ex Wikstr. 143
Guinetia tehuantepecensis L.Rico & M.Sousa 197
Gymnocladus dioiect (L.)K.Koch 130
Haematoxylum brasiletto H.Karst. 135
- dinte1'i (Harms)Harms 135
Halirnodendron ai-genteum (Lam.)DC. 491
Hammatolobium lotoides Fenzl 465
- ludovicia Batt. 465
Hanslia ormocaipoides (DC.)H.Ohashi 437
Haplormosia monophylla (Harms)Harms 235
Hardenbergia violacea (Schneev.)Stearn 406
Hardwickia binata Roxb. 76
Harleyodendron unifoliolatum RS.Cowan 223
Harpalyce arborescens A.Gray 255
- brasiliana Benth. 255
- sp. 255
Havardia campylacantha CL.Rico & M.Sousa) Barneby &
J.W.Grimes 208
Hebestigma cubense (Kunth)Urb . 468
Hedysantm varium Willd. 493
Hegnera obcordata (Miq.)Schindl. 441
Herpyza grandiflora (Griseb.)C.Wright 417
Hesperalbizia occidentalis (Brandegee)Barneby &
J.W.Grimes 210
Hesperolaburnum platycarpum (Maire)Maire 292
Hesperothamnus pentaphyllus Harms 378
Heterostemon mimosoides Desf. 97
Hippocrepis baleai'ica ]acq. 457
- scabra DC. 457
Hoffmannseggia glauca (Ortega)Eifert 145
Hoita rnacrostachya (DC.)Rydb. 449
Holocalyx balansae Micheli 222
Hosackia gracilis Benth. 460
Hovea chorizemifolia DC. 257
- longifolia R.Br. var. montana (Hook.f.)].H .Willis 257
- pu.ngens Benth. 252
- trispenna Benth. 257
Humboldtia unijuga Bedd. var. trijuga].]oseph &
V.Chandras. 94
Humularia rosea (De Wild.)P.A.Duvign. var. reptans
(Verdc.)Verdc. 332
Hydrochorea corymbosa (Rich.)Barneby & ].W.Grimes 204
Hylodendron gabunense Taub. 82
Hylodesmum repandurn (Vahl)H.Ohashi & RR.Mill 441
Hymenaea courbaril L. 80
- mai1iana Hayne 80
- verrucosa Gaertn. 80
Hymenocarpos circinnatus (L.)Savi 459
Hymenolobium alagoanum Ducke var. a!agoanum 310
Hymenostegia afzelii (Oliv.)Harms 94
- floribunda (Benth.)Harms 94
- normandii Pellegr. 94
Hypocalyptus oxalidifolius (Sims)Baill. 337
- sophoroides (P.].Bergius)Baill. 337
Jcuria dunensis Wieringa 107
Indigastrum fastigiatum (E.Mey.)Schrire 362
Indigo/era bosse1'i Du Puy & Labat 364
- colutea (Burm.f.)Merr. 364
- heterantha Wall. ex Brandis 365
- hochstetteri Baker 364
- pendula Franch. 360
Indopiptadenia oudhensis (Brandis)Brenan 170
Inga feuillei DC. 200
- flagelliformis (Vell.)Mart. 200
- sapindoides Willd. 200
- sp. aff. densiflora Benth. 213
Inocarpus edulis JR.Forst. & G.Forst. 318
Intsia bijuga (Colebr. )Kuntze 90, 91
Isoberlinia angolensis (Welw. ex Benth.) Hoyle & Brenan
var. lasiocalyx Hoyle & Brenan 104
Isotropis cuneifolia (Sm.)Benth ex Heynh. 342
jacksonia argentea C.A.Gardner 343
- scoparia Smith 343
- sp. aff. angulata Benth. 343
jacqueshuberia brevipes Barneby 151
julbernardia bi'ieyi (De Wild.)Troupin 105
Kalappia celebica Kosterm. 119
Kana/oa kahoolawensis Lorence & K.R.Wood 176
Kebirita roudairei (Bonnet)Kramina & D.D.Sokoloff 461
Kennedia beckxiana F. Muell. 406
- nig1'icans Lindi. 406
Kingiodendron alternifolium (Elmer)Merr. & Rolfe 75
Koompassia malaccensis Benth. 117
Kotschya africana Encll. var. bequaertii (De Wild.)Verdc.
331
- pe1·rieri (R.Vig.)Verdc. 331
- strigosa (Benth.)Dewit & P.A.Duvign. 331
Kummerowia striata (Thunb.)Schindl. 434
Kunstleria sarawakensis Ridd.-Num. & Kornet 377
Labichea lanceolata Benth. subsp . lanceolata 117
Lab/ab purpureus (L.)Sweet 425
Laburnum anagyroides Medilc 291, 297
- x watereri (Kirchner)Dippel 291
Lackeya multiflora (Torr. & A.Gray)Fortunato, L.P.Queiroz
& G .P.Lewis 399
Lamprolobiumfruticosum Benth. 258
Lasiobema cbampionii (Benth.)de Wit 66
Lathynts boissieri Sitj. 507
- japonicus Willd. 507
- splendens Kellogg 507
Latrobea diosmifolia (Benth.)Benth. var. glabrescens Benth.
344
Lebeckia cytisoides Thunb. 275
- macrantha Harv. 275
Lebruniodendron leptanthum (Harms)].Leonard 87
Lecointea amazonica Ducke 223
Lem.botropis nigricans (L.)Griseb. 293
Lemurodendron capuronii Villers & P.Guinet 171
Lemuropisum edule H.Perrier 156
Lennea melanocaipa Vatke 468
Lens culinaris Medik. 506
Leonardoxa afi'icana (Baill.)Aubrev. 95
Leptoden'is nobilis (Welw. ex Baker)Dunn 372
Leptodesmia spp. 445
Leptosema aphyllum (Hook.)Crisp 344
- daviesioides (Turcz.)Crisp 353
- tomentosum (Benth.)Crisp 353
Lespedeza formosa (Vogel) Koehne 432
- maximowiczii C.K.Schneid. 435
- thunbergii (DC.)Nakai 435
Lessertia lanata Harv. 484
- m.iniata TM.Salter 484
Leucaena trichodes Qacq.)Benth. 174
Leucochloron limae Barneby &J.W.Grimes 206
Leucomphalos cappa1'ideus Planch. 246
Leucostegane grandis Prain 86
Libidibia (Caesalpinia) glabrata (unpublished
combination) 144
- coriaria Qacq.)Schltdl. 144
- scleroca1pa (Standl.)Britton & Rose 144
Librevillea klainei (Harms)Hoyle 103
Liparia hirsuta Thunb. 270
- splendens (Burm.f.)Bos & De Wit 270
Loesenera gabonensis Pelle gr. 92
- walkeri (A.Chev.)].Leonard 92
Lonchocarpus gu.illeminianus (Tul.)Malme 380
- obtusus Benth. 380
Lophocaipinia aculeatifolia (Burkart)Burkart 146
Lotononis corymbosa Benth. 278
- m.agnifica B.-E.van Wyk 278
Lotus aduncus (Griseb.)Nyman 463
- benholetii Masf. 454
- corniculatus L. 463
- maculatus Breitf. 463
Luetzelburgia auriculata (Allemao)Ducke 234
Lupinus conicus C.P.Sm. 288
- mutabilis Sweet 288
- pilosus Murray 287
- p1'incei Harms 288
- tauris Benth. 287
INDEX TO ILLUSTRATIONS 549
- weberbaueri Ulbr. 282
- sp. 'paniculatus alliance' 288
Luzania purpurea Elmer 395
Lysidice rhodostegia Hance 85
Lysiloma tergemina Benth. 212
Lysiphyllum binatum (Blanco)de Wit 64
- cunninghamii (Benth.)de Wit 64
Maackia bupehensis Takeda 242
Machaerium hirtum (Vell.)Stellfeld 328
- millei Stand!. 328
Macro/obium acaciifolium (Benth.)Benth. 99
- latifolium Vogel 99
- unijugum (Poepp. & Endl.)R.S.Cowan 99
Macropsychanthus lauterbachii Harms 395
Macroptilium lathyroides (L.)Urb. 430
Macrosamanea macrocalyx (Ducke) Barneby &
J.W.Grimes 199
Macrotyloma axillare (E.Mey.)Verdc. var. glabrum
(E.Mey.)Verdc. 425
Maniltoa megalocephala Harms 88
Maraniona lavinii C.E.Hughes, G.P.Lewis, Daza & Reyne!
319
Margaritolobium luteum Harms 382
Marina parryi (Torr. & A. Gray ex A. Gray)Barneby 303,
305
Marmaroxylon basijugum (Ducke) L.Rico 201
Martiodendron excelsum Gleason 118
- parvijlorum (AmshofOR.C.Koeppen 118
Mastersia assamica Benth. 404
- bakeri (Koord.)Backer 404
Mecopus nidulans Benn. 442
Medicago arborea L. 503
- polymorpha L. 503
Meizotropis buteifonnis Voigt 408
Melanoxylon brauna Schott 148
Melilotus a/bus Medik. 502
Melliniella micrantha Harms 445
Melolobium macrocalyx Dummer 285
- wilmsii Harms 285
Mendoravia dumaziana Capuron 115
Mezoneuron hildebrandtii Vatke 136
- scortechinii F.Muell. 136
Michelsonia microphylla (Troupin)Hauman 108
Micklethwaitia carvalhoi (Harms)G.P.Lewis & Schrire 88
Microberlinia bisulcata A.Chev. 105
- bmzzavillensis A.Chev. 105
Microcharis tritoides (Baker)Schrire 363
Microlobiusfoetidus (Jacq.) M.Sousa & G.Andrade 181
Mildbraediodendron excelsum Harms 220
Millettia aurea (R.Vig.)Du Puy & Labat 383
- nathaliae Du Puy & Labat 383
Mimosa debilis Humb. & Bonpl. var. vestita
(Benth.)Barneby 162
- loxensis Bameby 183
- tenuijlora (Willd.)Poir. 183
- townsendii Barneby 183
Mimozygantbus carinatus (G riseb.)Burkart 185
Mirbelia dilatata Dryand. 349
- platylobioides (DC.)Joy Thomps. 349
- speciosa Sieber ex DC. 349
Moldenbawera blanchetiana Tu!. var. blanchetiana l 49
- nutans L.P.Queiroz, G.P.Lewis & Allkin 149
- papillanthera L.P.Queiroz, G.P.Lewis & Allkin 149
Monarthrocarpus securiformis (Benth.)Merr. 439
550 LEGUMES OF THE WORLD
Monopte1yx inpae W.A.Rodrigues 232
Montigena novae-ze/andiae (Hook. f.)Heenan 486
Mora exce/sa Benth. 158
Moullava spicata (Dalzell)Nicolson 147
Mucuna gigantea (Willd.)DC. 405
- macrocarpa Wall. 405
- novo-guineensis Scheff. 405
Muellera frutescens (Aubl.)Standl. 382
Muelleranthus stipularis (J.M.Black)A.T.Lee 358
Mundulea stenophylla R.Vig. 385
Myrocaipusfrondosus Allemao 231
Myrospermum frutescens Jacq. 232
Myroxylon ba/samum (L.)Harms 232
Mysanthus uleanus (Harms) G.P.Lewis & A.Delgado 430
Neoapaloxylon madagascariense (Drake) Rauschert 72
- mandrarense Du Puy & R.Rabev. 72
- tuberosum (R.Vig .)Rausche1t 72
Neochevalierodendron stephanii (A.Chev.)].Leonard 93
Neocolletia gracilis Hems!. 422
Neobarmsia baronii (Drake)R.Vig. 239
Neonotonia wightii (Arn.)].A.Lackey var. longicauda
(Schweinf.) ].A.Lackey 417
Neorautaneniaficifolius (Benth.)C.A.Sm. 417
Neorudolphia volubilis (Willd.)Britton 399
Nephrodesmus a/bus Schindl. 438
- se1"iceus Schindl. 438
Neptunia oleracea Lour. 174
Nespbostylis holosericea (Baker)Verdc. 423
Newtonia buchananii (Baker)G.C.C.Gilbert & Boutique 171
Nissolia fruticosa Jacq. 314
- hintonii Sandwith 314
Nografilicaulis (Kurz)Merr. 419
Normandiodendron romii (De Wild.)].Leonard 93
Oddoniodendron micranthum (Harms)Baker f. 105
Ohwia caudata (Thunb.)H.Ohashi 437
Olneya tesota A.Gray 469
Onobrychis carduchorum C.C.Towns. 494
-spp. 494
Ononis natrix L. 502
Ophiocarpus aitchisonii (Baker)Podlech 482
Ophrestia oblongifolia CE.Mey.) H.M.L.Forbes 400
Orbexilum onob1ychis (Nutt.)Rydb. 448
Oreophysa microphylla (Jaub. & Spach)Browicz 483
Ormocarpopsis parvifolia Dumaz-le-Grand 334
Ormocaipum bernierianum (Baill.)Du Puy & Labat 334
- sennoides (Willd.)DC. 334
Ormosia amazonica Ducke 235
- smithii Rudd 235
Ornithopus sativus Brat. 461
Orphanodendron bernalii Barneby &J.W.Grimes 161
Oryxis monticola (Mart. ex Benth.)A.Delgado & G.P.Lewis
430
Ostryocarpus 1"iparius Hook.f. 373
Otholobium mexicanum (L.f.)].W.Grimes 451
- thomii (Ha1v.)C.H.Stirt. 448
Otion microphyllum (unpublished name) 345
Otoptera burchellii DC. 414
Ottleya strigosa (Nutt.)D.D.Sokoloff 462
Ougeinia oojeinensis (Roxb .)Hochr. 436
Oxylobium arborescens R.Br. 350
Oxyrhynchus volubilis Brandegee 427
Oxystigma msoo Harms 74
Oxytropis halleri Bunge ex W.D.].Koch 480
Pachyelasma tessmannii (Harms)Harms 156
Pacbyrhizus erosus (L.)Urb. 416
Painteria leptophylla Britton & Rose 208
Paloue guianensis Aubl. 96
- riparia Pulle 96
Paloveopsis emarginata RS.Cowan 96
Panurea longifolia Spruce ex Benth. 248
Paracalyx scariosus (Roxb.)Ali 408
Paraderris elliptica (Wall.)Adema 383
Paramachae1"ium onnosioides Ducke 321
Paramacrolobium coeruleum (Taub.)].Leonard 100
Parapiptadenia ilheusana G.P.Lewis 181
- pterosperma (Benth. )Brenan 181
Pararchidendron pruinosum (Benth .)I.C.Nielsen 204
Paraserianthes lophantha (Willd.)I.C.Nielsen 203
Paratephrosia lanata (Benth.)Domin 387
Parkia bicolor A.Chev. 178
- gigantocarpa Ducke 178
- igneijlora Ducke 178
Pai·kinsonia aculeata L. 153
- scioana (Chiov.)Brenan 153
Parochetus afi"icanus Polhill 501
Parryellafilifolia Torr. & A.Gray 301
Pearsonia sessilijlora (Harv.)Dummer 281
Pediomelum cyphocalyx (A.Gray)Rydb. 450
Pelleg1-iniodendron diphyllum (Harms)].Leonard 104
Peltiera nitida Du Puy & Labat 335
Peltogyne paucijlora Benth. 79
Peltophorum dubium (Spreng.)Taub. 152
Pentaclethra macroloba (Willd.)Kuntze 166
Pe1"iandra mediterranea (Vell.)Taub. 402
Pericopsis angolensis (Baker)Meeuwen 236
Petaladenium urceoliferum Ducke 238
Petalostylis labicheoides R.Br. 117
Pete1"ia thompsonae S.Watson 472
Petteria ramentacea (Sieber)C.Presl 290
Pbanera bidentata (Jack)Benth. var.fraseri de Wit 65
- williamsii (F.Muell.)de Wit 65
- sp. (Bauhiniajlexuosa Morie.) 65
Phaseolus coccineus L. 428
- lunatus L. 428
- vulgaris L. 428
Philenoptera violacea (Klotzsch)Schrire 378
Phylacium majus Collett & Hems!. 421
Phyllodium pulchellum (L.)Desv. 436
Phyllota luehmannii F.Muell. 345
- phylicoides (Sieber ex DC.)Benth. 345
Phyl/oxylon xylophylloides (Baker)Du Puy, Labat & Schrire
362
Pbysostigma mesoponticum Taub. 426
Pickeringia montana Nutt. 264
Pictetia marginata C.Wright 333
Piliostigma thonningii (Schum.)Milne-Redh. 66
Piptadeniajlava (Spreng. ex DC.) Benth. 180
- viridiflora (Kunth)Benth. 180
Piptadeniastrum africanum (Hook.f.)Brenan 169
Piptadeniopsis lomentifera Burkart 173
Piptanthus nepalensis (Hook.)Sweet 265
Piscidia carthagenensis ]acq. 379
- grandifolia I.M.]ohnst. 379
Pisum sativum L. 508, 509
- sativum L. var. aruense (L.)Poir. 508
Pithece/lobium dulce (Roxb.)Benth. 209
- excelsum (Kunth)Benth. 55, 192, 209
- histrix (A.Rich.)Benth. 209
Plagiocarpus axillar'is Benth. 256
Plagiosipbon longitubus (Harms)].Leonard 88
- multijugus (Harms)].Leonard 88
Plathymenia reticulata Benth. 170
Platycelyphium voense (Engl.)Wild 240
Platycyamus regnellii Benth. 377
Platylobium alternifolium F.Muell. 357
- formosum Sm. 359
- obtusangulum Hook. 357
- triangulare R.Br. 357
Platymiscium lasiocarpum Sandwith 317
- pubescens Micheli subsp. pubescens 317
Platypodium elegans Vogel 318
Platysepalum inopinatum Harms 375
Podaly1"ia calyptrata (Retz.)Willd. 269
Podocytisus caramanicus Boiss. & Heldr. 291
Podolobium ilicifolium (Andrews) Crisp & P.H. Weston 351
Podolotus hosackioides Benth. 458
Poecilanthe falcata (Vell.)Heringer 254
Poeppigia procera C.Presl 113
Poincianella exostemma (DC.)Britton & Rose 142
- pannosa (Brandegee)Britton & Rose 142
Poiretia punctata (Willd.)Desv. 313
Poissonia heterantha (Griseb.)Lavin 470
- hypoleuca (Speg.)Lillo 470
Poitea campanilla DC. 469
- punicea (Urb.)Lavin 469
Polbillia canescens C.H.Stirt. 286
- obsoleta (Harv.)B.-E.van Wyk 286
- pallens C.H.Stirt. 286
Polystemonanthus dinklagei Harms 101
Pomaria stipularis (Vogel)B.B.Simpson & G.P.Lewis 141
Pongamiopsis amygdalina (Baill.)R.Vig. 384
Prioria copaifera Griseb. 75
Prosopidastrum globosum (Gillies ex Hook. & Arn.) Burkart
173
Prosopis julijlora (Sw.)DC. 172
- palmeri S.Watson 172
Pseudarthria confe11ijlora Baker 442
Pseudeminia comosa (Baker)Verdc. 420
Pseudoeriosema andongense (Baker)Hauman 400
Pseudolotus villosus (Blatt. & Hallb.) Ali & D.D.Sokoloff 460
Pseudomacrolobium mengei (De Wild.)Hauman 102
Pseudopiptadenia brenanii G.P.Lewis & M.P.Lima 179
Pseudoprosopisfischeri (Taub .)Harms 168
Pseudosamanea guacha.pele (Kunth)Harms 210
Pseudovigna argentea (Willd .)Verdc. 420
Psophocarpus tetragonolobus (L.)DC. 414
Psoralea pinnata L. x P. acu/eata Andr. 446
- tenuissima E.Mey. 448
Psoralidium lanceolatum (Pursh)Rydb. 449
Psorothamnus schottii (Torr.)Barneby 303
- spinosus (A.Gray)Barneby 298
Pterocarpus angolensis DC. 322
- rohrii Yahl 322
- rohrii Yahl sens. lat. 322
Pterodon abruptus (Moric.)Benth. 251
Pte1·ogyne nitens Tul. 134
Pterolobium stellatum (Forssk.)Brenan 137
Ptycholobium contortum (N.E.Br.)Brummitt 387
Ptychosema anomalum F.Muell. 358
Pueraria lobata (Willd.)Ohwi var. thomsonii
(Benth.)Maesen 418
Pultenaea brachytropis Benth. ex Lindi. 348
- divaricata H.B.Will. 348
INDEX TO ILLUSTRATIONS 551
- reticulata (Sm.)Benth. 348
Pycnospora lutescens (Poir.)Schindl. 442
Pyranthus tullearensis (Baill.)Du Puy & Labat subsp.
tullearensis 384
Rafnia meyeri Schinz 276
- perfoliata E.Mey. 276
Ramirezella micrantha A.Delgado & Ochot.-Booth 429
Ramorinoa girolae Speg. 320
Recordoxylon speciosum (Benoist)Normand & Mariaux 148
Requienia sphaerosperma DC. 387
Retama raetam (Forssk.)Webb 294
Rhodopis planisiliqua Urb. 399
Rhynchosia leand1·ii Du Puy & Labat 410
- malacophylla (Spreng.)Bojer 410
- mantaroensis J.F.Macbr. var. cuprinervia Grear 410
Rhynchotropis poggei (Taub .)Harms 363
Riedeliella magalhaesii (Rizzini)H.C.Lima & Vaz 315
Robinia hispida L. var. kelseyi (Hutch.)Isley 470
- pseudoacacia L. 473
Robynsiophyton vanderystii R.Wilczek 281
Rothia hirsuta (Guill. & Perr.)Baker 281
Rupertia physodes (Douglas ex Hook.)]. W.Grimes 449
Sakoanala madagascariensis R.Vig. 238
- villosa R.Vig. subsp. menaheensis Du Puy & Labat 238
- villosa R.Vig. subsp. villosa 238
Salweenia wardii Baker f. 244
Samanea saman (Jacq.)Merr. 210
Saraca sp. 85
Sarcodum scandens Lour. 371
Sartoria hedysaroides Boiss. & Heldr. 495
Sche.f/lerodendron usambarense Harms 376
Schizolobiu.m parahyba (Vell.)S.F.Blake var. amazonicum
(Huber ex Ducke)Bameby 151
Schleinitzia insularum (Guill.)P.Guinet 175
Schotia brachypetala Sonder 73
Scorodophloeus fischeri (Taub.)J.Leonard 86
Scorpiurus muricatus L. 457
Securigera securidaca (L.)Degen & Dorfl. 458
Sellocharis paradoxa Taub. 289
Senna australis (Vell.)H.S.Irwin & Bameby 123
- leandrii (Ghesq.)Du Puy 110
- pistaciifolia (Kunth)H.S.Itwin & Barneby var. picta
(G.Don) H.S.I1win & Barneby 123
- form taxon 'filifolia' 123
Serianthes grandiflora Benth. 202
Sesbania macrantha E.Phillips & Hutch. var. macrantha
453
- quadrata J.B. Gillett 453
- tomentosa Hook. & Arn. 453
Shuteria sinensis Hems!. 404
Sindora cochinchinensis Baill. 83
Sindoropsis letestui (Pellegr.)J.Leonard 83
Sinodolichos lagopus (Dunn)Verdc. 419
Smirnowia turkestana Bunge 483
Smithia elliotii Baker f. 330
Soemme1-ingia semperflorens Mart. 330
Sophora alopecuroides L. 243
- microphylla Aiton 226, 243
- tomentosa L. subsp. tomentosa 249
- velutina Lind!. subsp. zimbabweensis J.B. Gillett &
Brummitt 243
Spartidium saharae (Coss. & Durieu)Pomel 275
Spartium junceum L. 296
552 LEGUMESOFTHEWORLD
Spathionema kilimandscharicum Taub. 426
Spatholobus latistipulus Merr. 407
Sphaero/obium pulchellum Meisn. 340
- sp. aff. macranthum Meisn. 338
Sphaerophysa salsula (Pall.)DC. 484
Sphenostylis angustifolia Sond. 422
Sphinctospermum constrictum (S.Watson)Rose 472
Sphinga acatlensis (Benth.)Barneby &J.W.Grimes 207
Spirotropis longifolia (DC.)Baill. 248
Spongiocarpella nubigena (D.Don)Yakovlev 478
- pwpurea (P.C.Li)Yakovlev 478
- spinosa (P.C.Li)Yakovlev 478
Stachyothyrsus staudtii Harms 159
Stahlia monosperma (Tul.)Urb. 145
Stauracanthus genistoides (Brot.)Samp. 296
Stemonocoleus micranthus Harms 79
Stenodrepanum bergii Harms 146
Stirtonanthus taylorianus CL.Bolus) B.-E.van Wyk &
A.L.Schutte 269
Stonesiella selaginoides (Hook. f.)Crisp & P.H.Weston 346
Storckiella australiensis ].H.Ross & B.Hyland 116
- pancheri Bail!. 116
Streblon-hiza speciosa Lind!. 487
Strongylodon macrobotrys A.Gray 415, 431
- madagascariensis Baker 415
Strophostyles helvola (L.)Elliott 429
Stryphnodendron pulcherrimum (Willd.)Hochr. 182
- rotundifolium Ma1t. 182
Stuhlmannia moavi Taub. 136
Stylosanthes capitata Vogel 325
- guianensis (Aubl.)Sw. 325
-spp. 325
Styplmolobium japonicum (L.)Schott 233
Sulla coronal"ia (L.) Medik. 493
Sutherlandia montana E.Phillips & R.A.Dyer 485
Swainsonaformosa (G.Don)Joy Thomps. 474, 485
-sp. 485
Swartzia cf. flaemingii Raddi 225
- myrtifolia Sm. var. elegans (Schott)R.S.Cowan 218
- panacoco (Aubl.)R.S.Cowan 218
- simplex (Sw.)Spreng. 218
Sweetiajhtticosa Spreng. 234
Sylvichadsia grandifolia (R.Vig.)Du Puy & Labat 375
- macrophylla (R.Vig.)Du Puy & Labat 375
Sympetalandra borneensis Stapf 159
Syrmatium e1"iophorum A.Heller 462
Tachigali paniculata Aubl. 150
- versicolor Stand!. & LO .Williams 150
- sp. 150
Tadehagi triquetrum (L.)H.Ohashi subsp . auriculatum
(DC.)H.Ohashi 438
Talbotiella batesii Baker f. 86
- eketensis Baker f. 86
Tamarindus indica L. 90
Tara (Caesalpinia) cacalaco (unpublished combination)
138
- spinosa (Molina)Britton & Rose 138
Tara/ea steyermarkii Schery 251
Taverniera aegyptiaca Boiss. 494
Templetonia aculeata (F.Muell.)Benth . 256
- retusa (Vent.)R.Br. 259
Tephrosia phylloxylon (R.Vig.)Du Puy & Labat 386
- vogelii Hook.f. 386
Teramnus spp. 421
Tessmannia macrantha, ined. 83
Tetraberlinia longiracemosa (A.Chev.)Wieringa 106
- polyphylla (Harms)].Leonard 106
Tetragonolobus purpureus Moench 464
Tetrapleura tetraptera (Schumach & Thonn.)Taub. 167
Tetrapterocaipon geayi Humbert 132
-spp. 132
Teyleria tetragona (Merr.)Maesen 418
Thailentadopsis vietnamensis (I .C.Nielsen)G.P.Lewis &
Schrire 207
Thermopsis barbata Benth. 262
- rhombifolia (Nutt. ex Pursh)Richardson var. montana
(Nutt. ex Torr. & A.Gray)Isley 265
Thinicola incana (J.H.Ross)J.H.Ross 258
Tibetia himalaica (Baker)H.P.Tsui 479
Tipuana tipu (Benth.)Kuntze 320
Trifidacantbus unifoliolatus Merr. 440
Trifolium repens L. 498, 501
- rubens L. 501
Trigonella stellata Forssk. 503
Ti"ipodion tetraphyllum (L.)Fourr. 465
Trischidium molle (Benth.)H.E.Ireland (combination in
press) 219
1)1/osema esculentum (Burch.)A.Schreib. 63
- f assoglensis (Schweinf.)Torre & Hille. 63
Uittienia modesta Steenis 119
Uleanthus eryth1"inoides Harms 248
Ulex europaeus L. 296
Umtiza listerana Sim 131
Uraria crinata DC. 443
- picta DC. 443
Uribea tamarindoides Dugand & Romero 234
Urodon capitatus Turcz . 346
Vandasina retusa (Benth.)Rauschert 407
Vatairea macrocarpa Ducke 310
Vataireopsis araroba (Aguiar)Ducke 310
Vatouaea pseudolablab (Harms)J.B.Gillett 426
Vaughania depauperata (Drake)Du Puy, Labat & Schrire
363
Vaviloviaformosa (Steven)Al.Fed. 508
Vici a f aha L. 504
- sativa L. 506
- sylvatica L. 506
Vigna caracalla (L.)Verdc. 427
- vexillata (L.)A.Rich. 427
Viguieranthus ambongensis (R.Vig.)Villiers 199
Viminaria juncea (Schrad.)Hoffmanns. 342
Virgilia oroboides (P.J.Bergius)T.M.Salter 271
Vouacapoua americana Aubl. 161
Wajira albescens Thulin 422
Wallaceodendron celebicum Koord. 204
Weberbauerella brongniai1ioides Ulbr. 335
Wiborgia cf. mucronata (L.f.)Druce 276
Wisteriafloribunda (Willd.)DC. 366
- sinensis (Sims)Sweet 372
Xanthoce1·cis madagascai"iensis Bail!. 230
Xerocladia viridiramis (Burch.)Taub. 173
Xeroden"is stuhlmannii (Taub .)Mendon<;a & E.P.Sousa 374
Xiphotheca canescens (Thunb.) A.L.Schutte & B.-E.van Wyk
268
- phylicoides A.L.Schutte & B.-E.van Wyk 268
Xyliafraterna (Vatke) Drake 168
Zapoteca caracasana (Jacq.)H.M. Hern. 55, 196
Zenia insignis Chun 119
Zenkerella citrina Taub. 93
Zollernia glabra (Spreng.)Yakovlev 222
- modesta A.M.Carvalho & Barneby 222
Zornia spp. 313
Zuccagnia punctata Cav. 146
Zygia macropbylla (Benth.)L.Rico 201
Zygocarpum somalense (J.B.Gillett)Thulin & Lavin 334
INDEX TO ILLUSTRATIONS 553
Index to vernacular names

Abem 103 Apple-ring acacia 196 Bean, guar 362

Abey 212 Arapati 151 Bean, hyacinth 425
Abogo 104 Arariba 321 Bean, ice cream 200
Acapu 161 Arbolsoga 318 Bean, jack 396
Adoum 172 Arbre a semelle 1Tl Bean, jumbie 174
African greenheart 169 Archer dolichos 425 Bean, kidney 428
African locust bean 178 Ardilla 199 Bean, Lima 428
African teak 322 Aridan 167 Bean, locust 10, 133
African walnut 73 Ariella 106 Bean, lupini 287
African zebrawood 105 Aromata 238 Bean, mahogany 91
Afronnosia 236 Ashoka 85 Bean, marama 63
Afzelia 91 Asoka 85 Bean, matchbox 169
Agati 453 Asokarista 85 Bean, Mauritius 405
Agba 75 Assa-ha pa-gara undeguela 380 Bean, mescal 233
Ahianana 79 Auraremo-temo 205 Bean, moth 427
Aila 318 Austrian peaweed 484 Bean, mung 427
Akpa 167 Awoura 105 Bean, nickar 143
Akume 81 Ayan 115 Bean, ordeal 426
Albizia, white 202 Ayanran 115 Bean, phasy 430
Albizia, plume 203 Azobi 172 Bean, rat 147
Alfalfa 10, 503 Bean, rice 427
Algarroba 172 Babai 85 Bean, rosary 410
Algarrobo 80 Bahia powder 311 Bean, salmon 203
Alma negra 238 Balsam of Peru 232 Bean, sataw 178
Almendro 251 Balsam, Jesuit's 84 Bean, scarlet runner 428
Amarante 79 Balsam, tolu 232 Bean, screw 172
Amaranth 79 Bambara groundnut 427 Bean, snake 217
Amargo 310 Bannia 218 Bean, soy 10, 421
Amarillo guayaquil 321 Banuyo 204 Bean, soya 421
Ambatch wood 329 Baraka 89 Bean, spider 175
Amboyna 322 Barbatimao 182 Bean, St. Thomas 169
Amburana 220 Basralocus 121 Bean, sword 396
Amendoim 134 Bastard indigo 302 Bean, tepary 428
American potato bean 403 Batai 202 Bean, tonka 251
American yellow-wood 233 Batibatra 212 Bean, urd 427
Amherstia 95 Bean, adzuki 427 Bean, velvet 405
Amur maackia 242 Bean, African yam 422 Bean, winged 414
Anchico colorado 181 Bean, African locust 178 Bean, yam 416
Andoung 107, 108 Bean, American potato 403 Bean, yard-long 427
Angelim amarelo 310 Bean, azuki 427 Bean, year 428
Angelim amargosa 310 Bean, Bambara 427 Bee brush 303
Angelim da campina 221 Bean, Bengal 405 Beli 105
Angelim de mata 310 Bean, black 229 Berlinia 103
Angelim pedra 166 Bean, boer- 73 Billy Web 236
Angelim rajado 201 Bean, bonavist 425 Bird of paradise, desert 141
Angelim rosa 310 Bean, broad 10, 506 Bird of paradise, red 140
Angelim vermelho 166 Bean, buffalo 405 Bird of paradise, yellow 141
Angelique 121 Bean, Calabar 426 Birdsfoot, common 461
Angico 181 Bean, Calcutta 362 Bitter angelim 310
Angico de bezerro 180 Bean, cluster 362 Black cabbage bark 380
Angico preto 179 Bean, common 428 Black gum 235
Angok 103 Bean, eland's 170 Black locust 470
Anjan 76 Bean, fava 506 Black plum 311
Anzilim 77 Bean, gemsbuck 63 Black wattle 189
Apa 78, 91 Bean, Goa 414 Blackwood 327

INDEX TO VERNACULAR NAMES 555

Bladder senna 482 Cativo 75 Denya 172 Golden rain tree 291 Ironwood 204 Liquorice, false 390
Bloodwood 322 Caviuna 328 Derham mahogany 204 Golden shower 124 Ironwood, Sonora 469 Lobster claw 486
Blue clover 501 Cebil 179 Derris powder 383 Golden spray 342 Irul 168 Locoweeds 481
Blue wild indigo 265 Cedrorana 201 Desert bird of paradise 141 Golden tip 356 Ishpingo 220 Locust bean 10, 133
Boer-bean 73 Cerejeira 220 Detar 84 Gombe 103 Italian sainfoin 493 Logwood 135
Boire 84 Chana 497 Devil's shoestring 386 Gorse 296 Lokundu 171
Bois boco217 Chanfuta 91 Dhaincha 453 Gram, Bengal 497 Jade vine 415 Lucerne 503
Bois rose 98 Charcoal tree 86 Ditah 84 Gram, black 427 Jamaica dogwood 379 Lucerne, Brazilian 325
Bokanga 247 Chari 181 Divi-divi 144 Gram, horse 425 Jamaican ebony 316 Lucerne, tree 292
Bolengu 91 Cheny wood 220 Djedoe 150 Gram, green 427 Japanese pagoda tree 233 Lucerne, wild 325
Bomanga 106 Chica 320 Dogwood 343 Granadillo 317 Jatoba 80 Lucky bean tree 413
Bondu 103 ChickpealO, 496, 497 Doka 104 Grapia 115 Jequirity bead 390 Luknieng 202
Botonallare 237 Chinese bush clover 435 Doussie 91 Groundnut 326 Jering 202 Lupins 12
Brasil 10 Chinese yellow-wood 242 Groundnut, Bambara 427 Jerusalem thorn 153 Lupinus Russell hybrids 287
Dragon's blood 322
Brasil wood 12 Chou dou 178 Drinking vine 169 Groundnut, Kersting's 425 Jesuit's balsam 84
Brauna 148 Chulul 301 Dyer's Greenweed 10, 295 Guacamayo 73 Jeungjing 202 Macacauba 317
Brazilette 135 Clover, blue 501 Guachapele 210 Jicama 416 Macawood 317
Brazilian lucerne 325 Clover, bush 356 Ear pod tree 212 Guaje 174 Joint vetch 329 Machiche 380
Breadroot 450 Clover, Chinese bush 435 East Indian walnut 211 Guanacaste 212 Jolote beard 199 Macucu de paca 221
Bristly locust 470 Clover, haity canary 464 Ebena 97 Guapino 180 Judas tree 59 Madras thorn 209
Broom 292 Clover, purple prairie 304 Ebiara 103 Guapinol negro 89 Jumbie bean 174 Madre de cacao 468
Broom Dalea 303 Clover, red 501 Ebony, Jamaican 316 Guapuruvu 151 Jumby bead 235 Madricacao 468
Broom, Albanian 290 Clover, sand 459 Ebony, mountain 61 Guar 362 Jurema preta 180 Malacca teak 90
Broom, black 293 Clover, sulla 493 Ebony, Texas 208 Guaribeiro 73 Jutahy 120 Manangona 239
Broom, dune 302 Clover, sweet 10, 502 Ebony, West Indian 316 Guava machete 200 Mangium 189
Broom, dyer's 295 Clover, tick 440 Ebony wood 327 Guayamochil 209 Kakabeak 486 Mani 234
Broom, hedgehog 294 Clover, white 501 Ekaba 106 Gum arabic, true 189 Kalappi 119 Manila tamarind 209
Broom, Moroccan 290 C6bana negra 145 Ekhimi 169 Gum copal 77 Kanavali 396 Maniltou 88
Broom, pineapple 290 Cockies tongue 256 Ekop 106 Gum tragacanth 481 Kekatong 89 Manna 492
Broom, round leaf 302 Coco tamarind 210 Ekop rouge 107 Kempas 117 Marblewood 201
Broom, rush 342 Cocobolo 327 Ekpogoi 103 Haiari 229 Kentucky coffee tree 130 Mayo 107
Broom, Scotch 292 Cocuswood 316 Emonga 75 Haiariballi 229 Kentucky yellow-wood 233 Mbanga 236
Broom, Spanish 296 Cohoba 179 Enzieze 167 Hawaiian silverswords 42 Keranji 120 Mbarika 104
Broom, weaver's 296 Colorado river hemp 453 Eperu 78 Hintsi 90 Kersting's groundnut 425 Meblo 106
Broom, white 294 Colville's glory tree 155 Essabem 103 Hog potato 145 Keurblom 269 Medic, purple 503
Brown salwood 189 Common birdsfoot 461 Etabally 321 Hoita 449 Keurboom 271 Mendoravy 115
Brush wattle 203 Common restharrow 502 Evergreen laburnum 265 Honey locust 131 Khargana 491 Mengris 117
Bubinga 81 Copaiba 84 Honeybush tea 268 Kiaat 322 Merbau 90
Bumba 86 Copaiva 84 False indigo 265, 302 Horsegram 425 Kingwood 327 Mesquite 172
Burmese ironwood 168 Copa! 84 False liquorice 390 Horseshoe vetch 457 Kina gum 322 Methi 503
Bush pea 348 Copa!, South American 80 False lupin 265 Huajillo 208 Koa haole 174 Mgodoma 86
Bush sweetpea 269 Copa!, West African 77 False thorn 147 Huayracaspi 201 Kokriki 235 Milk vetch 481
Copa!, Zanzibar 80 Faro 77 Huayruru 235 Kowhai 243 Mimosa, florists' 10, 189
Cabbage tree 311 Cora<;:ao de negro 218 Faveira amargosa 310 Humble plant 183 Kowhai, small-leaved 243 Mirabow 90
Cabreuva 231 Coral tree 413 Feather tree 212 Hyedua 81 Ku shen 243 Mlembela 102
Calliandra 197 Corkwood tree 453 Fenugreek 503 Kudzu vine 418 Monkey bread 66
Cam xe 168 Corotu 212 Firetree 202 Ibirapepe 222 Kudzu, tropical 418 Monkeypod 210
Camel's foot 61 Costilla 318 Flamboyant 154 Icarape 180 Kulavu ena 75 Manta de! rosa 98
Camibar 75 Courbaril 80 Flame of the forest 407 Ice cream bean 200 Kumpas 117 Mopane 76
Camote del monte 472 Cowpea 10, 427 Flame tree 154 Icurri 107 Kurupa'y ra 181 Mora 158
Campeche 135 Cow tamarind 210 Florists' mimosa 10, 189 Illinois bundleflower 175 Kwilau 90 Morabukea 158
Camwood 246 Coyote 317 Frijolillo 210 Imburana de cheiro 220 Morado 328
Canaleto 318 Crab's eye 390 Furze 296 Indian laburnum 124 Lady bug tree 235 Moreton Bay chestnut 229
Canalua 318 Crested wattle 203 Indian liquorice 390 Lala 318 Mountain dahlia 270
Canary wood 321 Cumaru 251 Garapa 115 Indian rush pea 145 Lancewood, red 203 Mountain ebony 61
Cancer bush 485 Cumaru-da-praia 251 Garauna 148 Indigo 10, 364, 378 Lati 247 Movingui 115
Cape Leeuwin wattle 203 Curupay 179 Garbanzo 497 Indigo, bastard 302 Lead plant 302 Mpande 383
Carboncillo 236 Gavilan 166 Indigo, blue wild 265 Lentil 10, 506 Msasa 106
Carbonero 179 Dabema 169 Gemsbuck bean 63 Indigo, false 265, 302 Lespedeza, sericea 435 Msase 376
Carcuera 318 Dahoma 169 Gheombi 83 Inga 200 Leucaena 174 Mtondo 104
Carob tree 10, 133 Dakkar 90 Goa powder 311 Inga cip6 200 Libi-dibi 144 Mtundu 106
Carolina 167 Danto 310 Goats rue 386, 487 Intolwane 170 Licorice 478 Muande 86
Cascara amarga 236 Datach 84 Golden banner 265 Ipil 90 Limbali 104 Mucuna 405
Cassia (of commerce) 124 Dead finish 203 Golden chain tree 291 Ipil-ipil 174 Liquorice 10, 478 Muhimbe 89

556 LEGUMES OF THE WORLD INDEX TO VERNACULAR NAMES 557

Mukarati 158 Pea, flame 350 Red lancewood 203 Sleeping grass 183 Tesota 469 White lupine 287
Mullu 147 Pea, garden 508 Red sandalwood 167 Small-leaved kowhai 243 Texas ebony 208 White sweetclover 502
Muninga 322 Pea, glory 486 Red syringa 158 Smoke tree 303 Texas kidneywood 303 White tephrosia 386
Muvange 236 Pea, golden 265 Redbud 59 Snakewood 201 Texas mountain laurel 233 Wild lucerne 325
Muwa 105 Pea , grass 507 Restharrow, common 502 Snow wood 204 Timba 103 Wild sweet potato 472
Muyati 220 Pea, gungo 412 Restharrow, spiny 502 Soethoutbossie 276 Tindalo 91 Wild syringa 158
Pea, handsome flat 357 Rhodesian teak 82 Sokram 168 Tipa 134 Winter thorn 196
Naga 106 Pea, hoary 386 Rooibos tea 277 Sola 329 Tipa colorado 134 Wisteria 372
Nambar 327 Pea, Indian 507 Rosemary-of-campinas 222 Sola rosario 329 Tipu 320 Woman's tongue 211
Nandu 236 Pea, Indian rush 145 Rosewood 320, 327 Solar topi 329 Tola branca 75 Wood rosemary 222
Narra 322 Pea, parrot 347 Rosewood, bastard 218 Solomon's padauk 322 Tolu balsam 232 Worm bark 311
Nato 158 Pea, Philip Island glory 487 Rosewood, Papua New Guinea 322 Sonora ironwood 469 Tonka butter 251
Necklace tree 235 Pea, pigeon 10, 412 Rosewood, Santos 328 Soroca 180 Tornillo 201 Yawaredan 150
Nedun 236 Pea, poison 485 Round leaf broom 302 Sorrowless tree 85 Trebol 317 Yaya 247
Nere 178 Pea, shaggy 350 Royal poinciana 154 Sotocaballo 201 Tree fuchsia 73 Yeheb nut 135
Nganga 89 Pea, shamrock 501 Running postman 406 South American copal 80 Tree hovea 257 Yellow bird of paradise 141
Nickar bean 143 Pea, Sturt's Desert 485 Soybean 10,421 Tree lucerne 292 Yellow lupine 287
Nickar nut 143 Pea, sweet 12, 507 Sabicu 212 Soyabean 421 Tree wisteria 239 Yellow satinwood 115
Nickels 143 Pea, wedge 340 Saboneteira 219 Spanish sainfoin 493 Trefoil 10 Yellow sweetclover 502
Nigerian satinwood 115 Pea, winged 464 Saga 167 Spiny holdback 138 Trefoil, big 463 Yellow tail 290
Niopo 179 Pea , winged broom 343 Sa-gbembe 100 Spiny restharrow 502 Trefoil, birdsfoot 463 Yopo 179
Nkobakoba 82 Pea , yellow 356 Sainfoin 494 St. Thomas bean 169 Trefoil, stinking bean 264 Yvyra pere 115
Nyala tree 230 Peacock's plume 202 Sainfoin, Cretan 495 St. John's bread 10, 133 Tropical kudzu 418
Peanut 10, 326 Sainfoin, Italian 493 Stinking bean trefoil 264 Tualang 117 Zamang 210
Obang 236 Pearl lupine 287 Sainfoin, Spanish 493 Stinkwood 204 Tulip siris 204 Zambesi redwood 82
Ogea 77 Peaweed, Austrian 484 Sakoanala 238 Sua'alat 85 Tulipwood 327 Zanzibar copal 80
Oil bean tree 166 Petai 178 Salmon bean 203 Sucupira 237 T'zalam 212 Zebrawood 321
Okan 172 Petir 83 Saman 210 Suicide tree 150 Zebrali 105
Okwen 106 Pink cedar tree 133 Saman samando 240 Sulla 493 Ugba 166 Zebrano 105
Omutoyo 171 Plume albizia 203 Sana 123 Sulla clover 493 Uhiuhi 136 Zingana 105
Ovangkol 81 Pod mahogany 91 Sand clover 459 Sunn hemp 279 Umgusi 82
Owala 166 Pois chiche 497 Sandalwood, red 167 Supa 83 Umthiza 131
Poison 342, 352 Sandthom, Siberian 491 Surucucu 180
Padauk 322 Poison bush 352 Sangre de drago 322 Sweet blood 236 Vanillio 182
Padauk, Solomon's 322 Polynesian chestnut 318 Santos rosewood 328 Sweetclover 10, 502 Vanwykshout 239
Palisandro 327 Pomme de prairie 450 Sapo 103 Sweetclover, white 502 Vara dulce 303
Palo de cruz 98 Porcupine tree 321 Sara 85 Sweetclover, yellow 502 Vesi 90
Palo fierro 469 Porcupine wood 321 Sarrapia 251 Sweet potato, wild 472 Vetch 10
Palo mo1tero 320 Powder puff 197 Sataw bean 178 Swish bush 342 Vetch, chickling 507
Palo rosa 98 Pracuuba 158 Satinwood 236 Syringa, wild 158 Vetch, common 506
Paloverde 153 Prairie bundleflower 175 Satinwood, Nigerian 115 Vetch, crown 458
Panga panga 383 Prairie mimosa 175 Satinwood, yellow 115 Tachi 150 Vetch, joint 329
Papua New Guinea rosewood 322 Prairie turnip 450 Screw bean 172 Tachigali 150 Vetch, horseshoe 457
Para-angelim 310 Prayer bead 390 Sea hearts 169 Tad 150 Vetch, kidney 459
Parakwa 166 Prickly scorpion's tail 457 Senna 123 Tahitian chestnut 318 Vetch, milk 481
Paroa-caxi 166 Pride of Barbados 140 Senna pod 10 Tallow tree 84 Vetches 10
Parrot tree 407 Pride of Bolivia 320 Sensitive plant 183 Tamarind 90 Vine, Jade 415
Parrot-bush 256 Pride of Burma 95 Sepetir 83 Tamarind, coco 210 Vine, Kudzu 418
Partridge wood 144, 161, 311 Pula 75 Sericea lespedeza 435 Tamarind, cow 210 Vinhatico 170
Pau brasil 12, 142 Purging cassia 124 Serpentwood 201 Tamarind, Manila 209 Violetwood 79
Pau ferro 328 Purple medic 503 Serradella 461 Tamboril 212 Viraro 134
Pau jacare 180 Purple prairie clover 304 Shan dou gen 243 Tapang 117 Vouacapou 161
Pau roxo 79 Purpleheart 79 Shan dou geng 261 Tara 138
Pea, asparagus 464 Pyinkado 168 Shoestring 302 Tarahuilca 169 Wacap 161
Pea, balloon 485 Pynkado 168 Shola 329 Tarala 251 Wakapu 161
Pea, bitter 341 Siberian pea tree 491 Tarara colorado 317 Wallaba 78
Pea, bush 348 Quamwood 151 Siberian sandthorn 491 Tasaa 113 Wamara 78, 218
Pea, chaparral 264 Sickle bush 177 Tatabu 237 Waras 413
Pea, chick 10, 496, 497 Racehorse tree 320 Sindjaple 380 Tchitola 74 Wenge 383
Pea, common 10, 12, 508 Rain tree 210 Sindor 83 Tea, honeybush 268 West African copal 77
Pea, cow 10, 427 Reach-for-the-sky 151 Sirari 235 Tea, rooibos 277 West Indian ebony 316
Pea , duck 485 Red angelim 166 Siratro 430 Teak, Malacca 90 Whin 296
Pea, everlasting 507 Red bead tree 167 Siris tree 211 Tenaza 208 Whip tree 155
Pea, fish poison 386 Red bird of paradise 140 Sisam 327 Tento 235 White albizia 202

558 LEGUMES OF THE WORLD INDEX TO VERNACULAR NAMES 559

Index to scientific names
(including phylogenetic and biogeographic terms)

Abarema Pittier 15, 48, 193, 204, 205 - elaphroxylon (Guill. & Perr.) Taub. 329
Abarema alliance 195 - subgen. Aeschynomene 329
Abreae (Wight & Am. ex End!.) Hutch. 5, 9, 122, 361, 367, - subgen. Ochopodium (Vogel) ].Leonard 327, 328, 329
389, 393, 394, 505 - subgen. Rueppellia (A.Rich.) ].Leonard 329
Abrus Adans. 17, 51, 390, 505 Aeschynomeneae (Benth.) Hutch. 4, 8, 307, 308
- melanospermus Hassk. subsp. tenuiflorus (Benth.) D. Ajfonsea A.St.-Hil. 193, 200
Harder 389 Afgekia Craib 17, 371
- precatorius L. 122, 390 African 'arid corridor' 24, 43
- pulchellus Wall. ex Thwaites subsp. tenuiflorus (Bentb.) Afrormosia Harms 236
Verde. 389 Afzelia Sm. 13, 34, 69, 90, 91, 92
Acacia coulteri Group 187 Aganope Miq. 17, 373, 382
Acacia Mill. 10, 12, 15, 163, 187, 188, 212 Agati Adans. 453
- albida Del. 196 Airyantha Brummitt 15, 245, 246
- dealbata Link 189 Akschindlium H.Ohashi 9, 18, 439
- coulteri Benth. 187 - godefroyanum (Kuntze) H.Ohashi 439
- farnesiana (L.) Willd. 189 Alantsilodendron Villiers 6, 14, 163, 177
- mangium Willd. 189 Albizia Durazz. 3, 12, 15, 48, 193, 201, 205, 207, 211
- mearnsii De Wild. 189 - julibrissin Durazz. 211
- nilotica (L.) Willd. 188 - lebbeck (L.) Benth. 211
- senegal (L.) Willd. 189 Albizzia Benth. 211
- sens. lat. 3, 187, 193 Aldina End!. 15, 221
- sens. strict. 187 - heterophylla Spruce ex Benth. 221
- subgen. Acacia 6, 31, 187 Aldinoid clade 5, 7, 216, 228
- subgen. Aculeiferum 187 Alexa Moq. 5, 7, 15, 229
- subgen. - sect. Aculeiferum sens. strict. 187 - imperatricis (R.H.Schomb.) Baill. 229
- subgen. - sect. Filicinae (Benth.) Taub. 187 A lexandra R.H.Schomb. 229
- subgen. Phyllodineae 5, 6, 48, 187, 193 Alhageae (DC.) Burnett 489
Acacieae Dumort. 5, 6, 15, 22, 48, 163, 187, 193, 196 Alhagi Gagnebin 9, 19, 476, 489, 492
Acaciella Britton & Rose 187, 188 - graecomm Boiss. 492
Acanthonotus Benth. 364 Alhagiinae DC. 9
Acmispon Raf. 9, 19, 53, 462, 463 Alistilus N.E.Br. 18, 423
Acmispon sensu auct., pro parte non Raf 461 Alloburkillia Whitmore 377
Acosmium Schott 5, 15, 31, 36, 49, 236, 240 Almaleea Crisp & P.H.Weston 17, 346, 347
- panamense (Benth.) Yakovlev 236 Alvesia Welw. 61
Acrocarpus group 127, 128 Alysicarpus Desv. 19, 444
Acrocarpus Wight & Arn. 14, 24, 47, 111, 127, 131, 132, - subgen. Desmodiastrum Schindl. 444
133 Amaria Mutis 61
- fraxinifolius Wight ex Arn. 133 Amblygonocarpus Harms 14, 167
Adenanthera group 6, 163, 164, 165, 168 - andongensis (Oliv.) Exel! & Torre 167
Adenanthera L. 14, 47, 48, 167 Amburana Schwacke & Taub. 15, 215, 220
- pavonina L. 167 - cearensis (Allemao) A.C.Sm. 220
Adenanthereae (Benth.) Benth. & Hook.f 163 Amerimnon P.Browne 327
Adenocarpinae Rouy 284 Amherstia group 69
Adenocarpus DC. 16, 279, 283, 284, 289, 290 Amherstia Wall. 13, 69, 95
Adenodolichos Harms 18, 52, 408 - nobilis Wall. 95
Adenolobus (Harv. ex Benth. & Hook.f.) Torre & Hille. 13, Amherstieae Benth. 69
24, 31, 45, 57, 59, 60 Amherstieae clade 71
Adenopodia C.Presl 14, 182 Amicia Kunth 16, 312, 313
Adesmia clade 5, 8, 50, 308, 309 - zygomeris DC. 312
Adesmia DC. 3, 16, 36, 50, 307, 312 Ammodendron Fisch. ex DC. 15, 242, 279
- loto ides Hook.f. 312 - argenteum Kuntze 242
Adesmieae (Bentb.) Hutch. 4, 8, 36, 307 Ammopiptanthus S.H.Cheng 16, 263, 264
Adinobotrys Dunn 370 - mongolicus (Maxim. ex Korn.) Cheng 264
Aenictophyton A.T.Lee 17, 355, 358 Ammothamnus Bunge 7, 15, 242
Aeschynomene L. 3, 10, 17, 50, 329, 330 Amoria C.Presl 499, 501
- aspera L. 329 Amorpha L. 16, 50, 299, 301, 302

INDEX TO SCIENTIFIC NAMES 561

- canescens Nutt. ex Pursh 302 Argyrolobium Eckl. & Zeyh. 16, 36, 50, 267, 273, 283, 284, - riedelii (Tu!.) ].H.Kirkbr. 73 Bergeronia Micheli 17, 381 , 382
- fruticosa L. 302 285, 286, 287, 289 Barnebyella Podlech 10, 19, 476, 481, 482 Berlinia group 69
Amorpheae Boriss. 5, 7, 8, 16, 31, 36, 50, 299, 307, 447 - zanonii (Turra) P.W.Ball 283 - calycina (S tocks) Podlech 482 Berlinia Sol. ex Hook.f. 13, 103
Amorphoid clade 8, 299 A1illaria Kurz 235 Basal millettioid and phaseo loicl grou p 367, 368, 369, 393, Biancaea Tod. 140
amphi-Atlantic disjunction 21, 23, 37, 40, 42, 43, 44, 46, 50 Arthrocaipum Balff 8, 308, 324, 325 394 Bikinia J.].Wieringa 6, 13, 106, 107, 108
Amphicarpaea Elliott ex Nutt. 18, 52, 416, 420 Arthroclianthus Bail!. 18, 438 Basal Papilionoideae 7, 22, 34, 49, 54, 263 Bionia Mart. ex Benth. 397
Amphicarpa Elliott ex Nutt. 420 Arthrosamanea Biitton & Rose ex Britton & Killip 211 Batesia group 127, 129 Biserrula L. 9, 19, 475, 476, 480, 481
Amphimas Pierre ex Harms 15, 247 Artrolobium Desv., pro parte 458, 461 Batesia Spruce ex Benth. & Hook.f. 14, 47, 127, 128, 148 - pelecinus L. 475
- pterocarpoides Harms 247 Aspalathus L. 3, 16, 50, 276, 277 - floribunda Spruce ex Benth. 148 Bituminaria Heist. ex Fabr. 19, 36, 450
Amphithalea Eckl. & Zeyh. 8, 16, 267, 268, 269 - linearis (Burm. f.) R.Dahlgren 277 Bathiaea Drake 6, 74 - acaulis (Steven) C.H. Stirt. 447
Anadenanthera Speg. 12, 14, 179, 183 Aspalthium Medik. 450 Baudouinia Baill. 13, 31, 46, 111 , 113, 114, 115 - subgen. Christevenia Bameby ex C.H . Stirt. 447
- colubrina (Veil.) Brenan 179 Astracantha Podlech 9, 481 - flu ggeiformis Bail!. 114 Blanchetiodendron Barneby & ].W.Grimes 6, 15, 193, 205
- peregrina (L.) Speg. 179 Astragaleae (DC.) Dumo11 5, 10, 475 - rouxevillei H.Perrier 114 - blanchetii (Benth.) Barneby & ].W.Grimes 205
Anagyris L. 16, 263, 264, 283 Astragalean clade 475, 476, 477, 489 Bauerella Schindl. 450, 489 Bobgunnia ].H.Kirkbr. & Wiersema 7, 15, 34, 42, 215, 217,
- foetida L. 264 Astragalinae DC. 475, 476, 499, 505 Baueropsis Hutch 450, 489 218
Anarthrophyllum Benth. 16, 50, 273, 283, 284, 287, 289 Astragalus L. 1, 3, 9, 10, 19, 54, 455, 475, 476, 480, 481, - tomentosa (Schindl.) Hutch. 489 - fistuloides (Harms) ].H.Kirkbr. & Wiersema 217
Anatropostylia (Plitmann) Kupicha 506 482, 491 Bauhinia L. 4, 13, 57, 58, 59, 61 , 62, 63, 65 - madagascariensis (Desv.) ].H.Kirkbr. & Wiersema 217
Andira group 7, 8 - alpinus L. 43 - acuminata L. 61 Bobrovia A.P.Kokhr. 501
Andira Lam. 16, 50, 161, 307, 308, 310, 311 - aboriginum Richardson ex Spreng. 43 - gossweileri Baker f. 64 Bocoa Aubl. 7, 15, 215, 217, 218, 219
- inermis CW.Wright) DC. 43, 311 - americanus (Hook.) Jones 43 - humblotiana Baill. 61, 62 - prouacensis Aubl. 217
- - subsp. inermis 33 - complanatus Bunge 476 - petersiana Balle 61 Bolusafra Kuntze 18, 409
Androcalymma Dwyer 14, 34, 111, 118 - epiglottis L. 475 - strychnifolia Craib 64 Bolusanthus Harms 15, 239, 240
Angylocalyx group 227, 228 - gummifer Labill. 481 - sens. lat. 57, 58, 59, 60 - speciosus (Bolus) Harms 239
Angylocalyx Taub. 5, 7, 15, 220, 230, 248 - lusitanicus Lam. 476, 480 - sens. strict. 3, 31, 45, 60, 63 Bolusia Benth. 16, 50, 278
Anita Ludw. ex Kuntze 364 - membranaceus Bunge 481 - subgen. Barklya (FMuell.) Wunderlin, K.Larsen & Bonduc Mill. 143
Anil Mill. 364 - mongholicus Bunge var. dahuricus (DC.) Podlech 481 S.S.Larsen 64 Bonjeanea Rchb. 464
Anneslia Salisb. 197 - sinicus L. 476 - subgen. Bauhinia 61 Borbonia L. 277
Antheroporum Gagnep. 17, 370 - vogelii (Webb) Bomm. 476 - subgen. Elayuna (Raf.) Wunderlin, K.Larsen & S.S.Larsen Borbonieae Hutch. 273
Anthonotha P.Beauv. 13, 99, 102 - sect. Mirae Sirj. & Rechf 482 66 Boreotropics hypothesis 34, 40, 43
- noldeae (Rossberg) Exell & Hille. 102 - sect. Ophioca1pus Bunge 482 - subgen. Lasiobema Korth. 66 Bossiaea Vent. 17, 355, 356, 357, 358
Anthyllideae Buchenau 455 Ateleia (Moc;:. & Sesse ex DC.) Benth. 7, 15, 31, 49, 215. - subgen. Phanera (Lour.) Wunderlin, K.Larsen & - Spinosa (Turcz.) 357
Anthyllis L. 19, 455, 459, 465 220 S.S.Larsen 63, 65 Bossiaeeae (Benth.) Hutch. 5, 7, 8, 17, 36, 51, 253, 336,
- fulgurans Porta 464 Atylosia Wight & Arn. 412 - sect. Adenolobus Harv. ex Benth. & Hookf 59 339, 355, 361
- lotoides L. 459 Aubrevillea group 164, 165 - sect. Afrobauhinia Wunderlin, K.Larsen & S.S.Larsen 61 Bowclichia Kunth 15, 237, 248
- vulneraria L. 459 Aubrevillea Pellegr. 14, 34, 166 - sect. Alvesia (Welw.) Wunderlin, K.Larsen & S.S.Larsen 61 - nitida Benth. 237
Antopetitia A.Rich. 19, 455, 460 Augouardia Pellegr. 13, 79 - sect. Amada (S .Mutis) End!. 61 Bowringia Champ. ex Benth. 7, 15, 227, 246
Aotus Sm. 17, 345, 346 Austrodolichos Verde. 18, 423 - sect. Austrocercis (de Wit) Wunder·/in, K.La1-sen & Brachycylix (Harms) R.S.Cowan 13, 97
Apaloxy lon Drake 72 - errabundus (M .B.Scott) Verde. 423 S.S.Larsen 65 Brachypterum (Wight & Arn.) Benth. 382
Aphanocalyx Oliver 13, 108 Austrostee nisia R.Geesink 17, 52, 372 - sect. Bauhinia 61 - robustum (Roxb. ex DC.) Dalzell & A.Gibs . 382
Aphyllodium (DC.) Gagnep . 9, 18, 437 Azukia Owhi 427 - sect. Caulotretus DC. 65 Brachysema R.Br. ex WTAiton 8, 35 1, 352
Apios Fabr. 18, 52, 403 - sect. Gigasiphon (Drake) Harms 62 Brachystegia group 69
- americana Medik. 403 babjit clade 70, 71 - sect. Lasiobema (Korth.) Benth. 66 Brachystegia Benth. 13, 106
Apoplanesia C.Presl 16, 299, 301 Baikiaea Benth. 13, 82 - sect. Lysiphyllum Benth . 64 - cynometroides 103
- paniculata C.Presl 301 - insignis Benth. 82 - sect. Micralvesia Wunderlin, K.Larsen & S.S.Larsen 61 Brachystegioideae Hutch . 69
Aprevalia Bail!. 154 - plurijuga Harms 82 - sect. Palmatifolia (de Wit) Wunderlin, K.Larsen & Bracteolanthus de Wit 57, 58, 64
Apuleia Mart. 13, 111 , 115, 118, 119, 120 Bakerophyton (!.Leonard) Hutch . 329 S.S.Larsen 65 Bradbwya Raf 402
- leiocarpa (Vogel) ].F.Macbr. 115 Balboa Liebm. 471 - sect. Pauletia (Cav.) DC. 61 Brad/ea Adans. 403
Apurimacia Harms 17, 367, 386 Balisaea Taub. 329 - sect. Pseudobauhinia Wunderlin, K.Larsen & S.S.Larsen Brandzeia Bail!. 6, 13, 46, 72, 74
- dolichocarpa (Griseb.) Burkart 386 Balizia Barneby &]. WGrimes 193, 205 65 Brasilettia sensu Britton & Rose 139
- michelii (Rusby) Harms 386 Balsamocarpon Clos 14, 147 - sect. Pseudophanera Wunderlin, K.Larsen & S.S.Larsen Brasilettia (DC.) Kuntze 139, 140
- boliviana (Britton) Lavin 386 Bandeiraea Welw. ex Benth. 60 61 Bremontiera DC. 364
Arachis L. 10, 16, 307, 324, 325, 326 Baphia group 227, 228 - sect. Schnella (Raddi) Benth. 65 - ammoxylum DC. 363
- hypogaea L. 10, 326 Baphia Afzel. ex Lodd. 15, 224, 245, 246 - sect. Semla Wunderlin, K.Larsen & S.S.Larsen 65 Brenandendron H.Rob. 6, 88
- pintoi Krapov. & W.C.Greg. 326 - nitida Lodd. 246 - sect. Telestria (Raf.) Wunderlin, K.Larsen & S.S.Larsen 61 Brenaniodendronj.Leonard 6, 88
Aragallus Neck. ex Greene 480 Baphiastrum Harms 7, 15, 227, 246, 247 - sect. Tubicalyx Wunderlin, K.Larsen & S.S.Larsen 65 Brenierea Humbert 13, 45, 57, 59, 60, 63
Arapatiella Rizzini & A.Mattos 14, 127, 128, 150, 151 Baphioid clade 5, 7, 8, 22, 51, 216, 228, 336 - sect. Tylosema Schwein/ 63, 65 Brodriguesia RS.Cowan 13, 34, 92
- psilophylla (Harms) RS.Cowan 151 Baphiopsis Benth. ex Baker 7, 15, 215, 224, 245, 246 - ser. Canseria (Raf.) Wunderlin, K.Larsen & S.S.Larsen 61 - santosii RS.Cowan 92
Archidendron F.Muell. 15, 34, 193, 202 Baptisia Vent. 16, 263, 265, 283 Bauhinieae Benth. 57 Brongnia1tia Kunth 7, 16, 45, 49, 253, 256
- jiringa Qack) LC.Nielsen 202 - australis (L.) R.Br. 265 Bauhiniinae (Benth.) Wunderlin 4, 57, 60 - ulbrichiana Harms 256
Archidendropsis LC.Nielsen 15, 193, 203 - tinctoria (L.) R.Br. 265 Baukea Vatke 9, 410 Brongniartieae (Benth.) Hutch. 5, 7, 8, 9, 16, 36, 45, 49,
Arcoa Urb. 14, 24, 42, 47, 111, 128, 132 Barbieria DC. 9, 18, 401 Behaimia Griseb. 17, 381 238, 253, 273, 299, 355, 367
Arcto-Tertia1y Geoflora hypothesis 40, 43 Barklya F.Muell. 13, 58, 62, 63, 64 Belairia A.Rich. 8, 333 Broussonetia Ortega 233
Argyrocytisus (Maire) Raynaud 16, 284, 289, 290, 292 - syringifolia F.Mue ll. 64 Be!iceodendron Lundell 223 Brownea clade 71
- battandieri (Maire) Raynaud 290 - subgen. Barklya Wunderlin 64 Bembicidium Rydb. 469 Brownea group 69
Argyrolobium clade 283 Barnebydendron ].H.Kirkbr. 6, 13, 31, 46, 72 , 73 Benedictella Maire 463 Brownea Jacq. 13, 98

562 LEGUMES OFTHEWORLD INDEX TO SCIENTIFIC NAMES 563

 - houstoniana (Mill.) Stand!. var. calothyrsus (Meisn.) Centrosema (DC.) Benth. 10, 18, 402 Cologania Kuntb 18, 52, 416, 418
Browneopsis Huber 13, 98
B1ya P.Browne 16, 307, 308, 316 Barneby 197 Ceratonia L. 6, 14, 24, 47, 111 , 112, 127, 131, 132, 133 Colophospermum ].Kirk ex ].Leonard 13, 46, 76
- ser. Laetevirentes 196 - oreothauma Hille., G.P.Lewis & Verde. 133 - mopane Q.Kirk ex Benth.) ].Kirk ex ].Leonard 76
- ebenus DC. 316
Calliandropsis H.M.Hern. & P.Guinet 14, 163, 176, 177 - siliqua L. 10, 133 Colutea L. 19, 482
Bryaspis P.A.Duvign. 17, 50, 329, 332
Callistachys Vent. 17, 351 Ceratonieae Rchb. 127 - arborescens L. 482
B1yinae 4, 9
Buchenroedera Eckl. & Zeyh. 278 Calophaca Fisch. ex DC. 9, 19, 476, 489, 491 Cercideae Bronn 4, 5, 13, 22, 24, 40, 45, 46, 57, 111 - sect. Oreopbysa Bunge ex Boiss. 483
Calopogonium Desv. 18, 52, 416, 417, 418 Cercidinae 4, 57, 59, 60 Coluteinae Benth. 476
Burkea Benth. 14, 158
- mucunoides Desv. 416 Cercidiopsis Britton & Rose 153 Colvillea Boj. ex Hook. 14, 127, 146, 153, 154, 155, 156
- africana Hook. 158
Calpocalyx Harms 14, 127, 156, 157, 158, 163, 168 Cercidium Tu/. 127, 153 - racemosa Boj. ex Hook. 155
Burkillia Rid!. 377
Calpurnia E.Mey. 16, 49, 271 Cercis clade 24 Condylostylis Piper 427
Burkilliodendron Sastry 17, 377
Bitrtonia R.Br. ex WTAiton 340 Camoensia Welw. ex Benth. 15, 245 Cercis L. 13, 40, 45, 46, 59 Conzattia Rose 14, 127, 128, 153, 154
- scandens (Welw.) ].B.Gillett 245 - canadensis L. 59 - multiflora (B.L.Rob.) Stancil. 154
Bussea Hanns 14, 127, 152
Campecia Adans. 140 - occidentalis Torrey ex A.Gray 59 Copaifera L. 6, 10, 13, 34, 69, 83, 84
- occidentalis Hutch. 152
Campsiandra Benth. 14, 128, 159 - siliquastrum L. 59 - palustris (Symington) de Wit 84
Butea Roxb. ex Willd. 12, 18, 407
- monosperma (Lam.) Taub. 407 Camptosema Hook. & Arn. 18, 395, 397, 398 Chadsia Bojer 17, 375, 384, 385 Corallospartiumj.B.Armstr. 9, 486
- rubicundum Hook. & Arn. 397 Chaetocalyx DC. 16, 312, 314 Cordaea Spreng. 362
Cacara Rumph. ex Du Petit-Ibouars 416 Campylotropis Bunge 18, 433, 434 Chamaecrista Moench 3, 5, 6, 14, 31, 111, 112, 122, 124, Cordeauxia Hems!. 6, 14, 127, 134, 135
Canavalia DC. 18, 395, 396, 397 127, 134, 150 - eclulis Hems!. 135
Cadellia pentastylis F.Muell. 45
- ensiformis (L.) DC. 396 - absus (L.) H.S.I1win & Barneby 122 Corclyla Lour. 15, 215, 221, 247
Cadia Forssk. 15, 49, 241, 267
- gladiata (Jacq.) DC. 396 - mimosoides (L.) Greene 122 core genistoids 260, 263, 267, 273, 283
- pedicellata Baker 241
- purpurea (G.Piccioli) Aiton 241 - rosea (Sw.) DC. 396 Cbamaecytisus Link 292, 293 core Millettieae 52, 367, 368, 369, 389, 394
Candolleodendron RS.Cowan 7, 15, 215, 218, 219 Chamaespartium Adans. 295 core Peltophorum group 127, 129, 148, 149, 151, 153, 154,
Caesalpinia group 127, 129
Caesalpinia L. 6, 14, 47, 139, 140, 146 - brachystachyum (DC.) R.S.Cowan 219 Chapmannia Torr. & A.Gray 8, 16, 31, 42, 50, 308, 324, 325 155
- angolensis (Oliv.) Herend. & Zarucchi 136 Canizaresia Britton 379 Chasmone E.Mey. 287 core Phaseoleae 9, 52, 393, 394
Cantuffa j.F. Gmel. 13 7 Chesneya Lindi. ex End!. 19, 476, 478 Corethrodendron Fisch. ex Basiner 10, 19, 489, 493, 494
- brasiliensis L. 139
Capassa Klotzsch 378 Chidlowia Hoyle 14, 128, 160 Cornicina (DC.) Boiss. 459
- cassioides Willd. 140
Caragana Fabr. 9, 19, 476, 489, 491 - sanguinea Hoyle 160 Coronilla L. 19, 53, 455, 458
- delphinensis Du Puy & Rabev. 143
- arborescens Lam. 491 Chiovendaea Speg. 470 - emerus L. 457
- echinata Lam. 12, 142
Carmichaelia R.Br. 9, 19, 486 Chloroleucon alliance 195 Coronilleae (Bronn) Burnett 452, 455, 505
- eriostachys Benth. 142
Carmichaelieae Hutch. 9, 475, 476, 499, 505 Chloroleucon (Benth.) Britton & Rose 15, 193, 206, 207 - subtribe Aeschynomeninae (Bentb.) Scbulze-Menz 307
- glabrata Kunth 144
- insolita (Harms) Brenan &j.B.Gillett 136 Carrissoa Baker f. 18, 409 Cblo1yllis E.Mey. 424 - subtribe Desmodiinae (Bentb.) Schulze-Menz 433
- madagascariensis CR.Vig.) Senesse 140 - angolensis Baker f. 409 Cbordospartium Cbeeseman 486 - subtribe Patagoniinae (Taub.) Scbulze-Menz 307
- paraguariensis CD.Parodi) Burkart 144 Cascaronia Griseb. 16, 323 Chorizema Labill. 17, 349, 350, 351 Coronillinae Bronn 455
Cashalia Stand/. 231 - ilicifolium Labill. 350 Co1"otbamnus C.Presl 292
- pluviosa DC. 142
Caspareopsis Britton & Rose 61 - parviflorum Benth. 350 Corynella DC. 469
- pulcherrima (L.) Sw. 140
Casparia Kunth 61 Christia Moench 19, 443 Coublandia Aubl. 382
- pumilio Griseb. 145
Cassia L. 5, 6, 12, 14, 111, 112, 122, 124 Chronanthus K.Koch 292 Coulteria Kunth 6, 14, 128, 139
- rosei Urb. 140
- arereh Del. 124 - biflonts (Desf) Frodin & Heywood 295 - mollis Kunth 138, 139
- sessiliflora S.Watson 140
- fistula L. 124 Clnysaspis Desv. 499, 501 Coumarouna Aubl. 251
- spinosa (Molina) Kuntze 139
- grandis L. 124 Ch1ysoscias E.Mey. 18, 409 Coursetia DC. 9, 19, 467, 470, 471
- sens. lat. 135, 137, 139, 141, 143, 144, 147
- javanica L. 124 Cicer L. 19, 496, 497, 505 - beterantha (Griseb.) Lavin 467
- sens. strict. 23, 31, 136
- subgen. Guilandina (L ,) Gillis & Procto1· l43 - roxburghii DC. 124 - arietinum L. 10, 496, 497 - hypoleuca (Speg.) Lavin 467
- subgen. Mezoneuron (Desf) Vidal ex Herend. & - senna L. 111, 123 Cicereae Alef. 5, 10, 19, 36, 53, 54, 455, 475, 496, 499, 505 - orbicularis Bentb. 467
- sieberiana DC. 124 Cincliclocarpus Zoll. 140 - weberbaueri Hanns 467, 471
Zantcchi 136, 137
- sect. Cinclidocarpus (Zoll.) Benth. 140 - subgen. Absus (DC. ex Co/lad.) Symon 122 Cinnamomum aromaticum Nees 124 Cracca Benth. 471
- subgen. Lasiorhegma Vogel ex Benth. 122 Cladotrichium Vogel 141 Cracca L. 386
- sect. Libidibia DC. 144
- sect. Nugaria (DC.) Benth. 140 - subgen. Senna (Mill.) Benth. 123 Claclrastis-Styphnolobium clade 7, 8, 23 Craibia Harms & Dunn 17, 376
- sect. Peltophorum Vogel 152 Cassieae Bronn 4, 6, 13, 22, 24, 46, 111, 112, 127, 128 Claclrastis Raf. 5, 15, 24, 31, 40, 49, 233, 264 Cranocarpus Benth. 16, 307, 308, 316
- subtribe Cassiinae 112 - kentuckea (Dum.-Cours.) Rudd 233 Craspedolobium Harms 17, 378
- sect. Sappania Benth. 140
Caesalpinieae Rchb. 5, 6, 14, 24, 34, 40, 46, 47, 54, 111, - subtribe Ceratoniinae H.S.I1win & Barneby 111, 127, 133 Clathrotropis (Benth.) Harms 15 , 49, 237, 238 Cratylia Mart. ex Benth. 18, 395, 396, 398
- subtribe Dialiinae H.S.Irwin & Barneby 6 - bracbypetala (Tul.) Kleinhoonte 238 Crimea Vassilcz. 499, 503
112, 127
- subtribe Duparquetiinae H.S.I1win & Barneby 112 Cleobulia Mart. ex Benth. 18, 396, 397 Cristonia ].H.Ross 7, 16, 253, 256, 258
- sens. lat. 5, 22, 47, 48
Caesalpinioideae CR.Br.) DC. 1, 2, 3, 4, 5, 6, 13, 21, 22, 49, - subtribe Labicbeinae H.S.Irwin & Barneby 6, 112 Clianthus Sol. ex Linell. 19, 486 Crotalaria L. 3, 16, 36, 50, 273, 278, 279
- sens. lat. 5, 6 -formosus (G.Don) Ford & Vicke1y 485, 486 - cornu-ammonis R.Vig. 278
54, 184, 215
- sens. strict. 5, 6 - maximus Colenso 486 - juncea L. 279
Cajalbania Urb. 469
Castanospermum A.Cunn . ex Hook. 7, 12, 15, 220, 229 - puniceus CG.Don) Sol. ex Lindi. 486 Crotalarieae (Bentb.) Hutch. 5, 8, 16, 36, 50, 253, 263, 267,
Cajaninae 52
- australe A.Cunn. & C.Fraser ex Hook. 229 Clitoria L. 9, 18, 401, 402 273, 283, 285, 287
Cajanus DC. 18, 411, 412
Catenaria Bentb. 437 Clitoriinae Benth. 9, 52, 394 crown clacle 45
- cajan (L.) Huth 10, 412
Cathormion Hassk. 15, 193, 207 Clitoriopsis R.Wilczek 18, 403 crown clade ages 35
Cajan Adans. 412
Caulocmpus Baker f 386 Cochlianthus Benth. 18, 403 Cruddasia Prain 18, 400
Calia Teran & Berland. 5, 15, 49, 227, 233, 243, 260
- secundiflora (Ortega) Yakovlev 233 Caulotretus Rich. ex Schott 65 Codariocalyx Hassk. 18, 441 Crudia group 69
Cedrelinga Ducke 15, 193, 201 Coelidium Vogel ex Walp. 8, 267, 269 Crudia Schreb. 13, 69, 87
Calicotome Link 16, 284, 292, 293
- cateniformis (Ducke) Ducke 201 Cojoba Britton & Rose 7, 15, 34, 193, 199, 200, 202 C1yptosepalum Benth. 13, 100
Calispepla Vved. 287
Cenostigma Tul. 14, 142, 146 - arborea (L.) Britton & Rose 199 - exfoliatum De Wild. 100
Callerya End!. 5, 9, 17, 370, 371, 475, 478
Centrolobium Mart. ex Benth. 16, 321 Collaea DC. 18, 398 - maraviense Oliv. 100
Calliandra Benth. 3, 12, 15, 193, 197, 199

564 LEGUMES OF THE WORLD INDEX TO SCIENTIFIC NAMES 565

Cullen Medik. 19, 36, 52, 447, 489, 450 Dalhousiea Wall. ex Benth.15, 245, 246 Dicorynia Benth. 14, 111 , 117, 118, 119, 120, 121 Ecastaphyllum PBrowne 327
- badocanum (Blanco) Verde. 489 Damapana Adans. 330 - guianensis Amshoff 12, 111, 121 Echinosophora Nakai 243
Cupulanthus Hutch . 351 Daniellia Benn. 13, 74, 77 - paraensis Benth. 121 Echinospaitum (Spach) Rothm. 16, 284, 293, 295
Cyamopsis DC. 17, 361, 362 - oliveri (Rolfe) Hutch. & Dalziel 77 Dicraeopetalum Harms 15, 31, 36, 239, 240 ecological drift 41, 42, 44
- tetragonoloba (L.) Taub. 362 Dansera Steenis 120 - stipulare Harms 240 Ecuadendron D.A.Neill 6, 13, 95, 97
Cyanostremma Benth. ex Hook. & Arn. 416 Daubentonia DC. 453 Dicymbe Spruce ex Benth. 13, 34, 101 Edwardsia Salish. 243
Cyathostegia (Benth.) Schery 7, 15, 49, 215, 218, 219 Daubentoniopsis Rydb. 453 Dicymbopsis Ducke 101 Eleiotis DC. 19, 445
Cyclocarpa Afzel. ex Urb. 17, 329, 330 Daviesia Sm. 3, 17, 341, 342, 355 Didelotia Bail!. 13, 103 Elephantorrhiza Benth. 14, 169, 170
Cyclogyne Benth. ex Lindi. 485 Decorsea R.Vig. 18, 415 - minutiflora (A.Chev.) ].Leonard 103 Eligmocarpus Capuron 13, 46, 111, 114
Cyclolobium Benth. 7, 9, 16, 49, 253, 254, 367 Deguelia Aubl. 17, 380, 382 Dillwynia Sm. 17, 347 Elizabetha Schomb. ex Benth.13, 95, 97
- brasiliense Bentb. 254 Delonix Raf. 12, 14, 127, 142, 146, 153, 154, 155, 156 Dimorphandra group 24, 111, 127, 128, 129 - princeps Schomb. ex Benth. 97
Cyclopia Vent. 16, 49, 263, 268 - regia (Boj. ex Hook.) Raf. 154 Dimorphandra Schott 14, 47, 48, 127, 156, 157, 158 Emerus Mill. 457
- intermedia E.Mey. 268 Dendrobrychis (DC.) Galushko 494 - exaltata Schott 157 Eminia Taub. 18, 419
- subternata Vogel 268 Dendrolobium (Wight & Arn.) Bentb. 18, 436 - gardneriana Tu!. 157 Endertia Steenis & de Wit 13, 85
Cylicodiscus group 6, 164, 165 Dermatophyllum Scheele 233 - mollis Benth. 157 - spectabilis Steenis & de Wit 85
Cylicodiscus Harms 14, 163, 171, 172 Derris group 368 Dim01phandreae Benth. 127 Endomallus Gagnep. 412
- gabunensis Harms 172 Derris Lour. 17, 52, 367, 372, 379, 380, 382 Oinizia group 164, 165 Endosamara R.Geesink 17, 370
Cylindrokelupha Kosterm. 202 - robusta (Roxb. ex DC.) Benth. 382 Oinizia Ducke 5, 6, 14, 34, 157, 166, 184 EngJerodendron Harms 13, 102
Cylista Aiton 410 - sect. Brachypterum (Wight & Arn) Benth. 382 - excelsa Ducke 166 - usambarensis Harms 102
Cymbosema Benth. 18, 395, 397 - sect. Paraderris Miq. 383 Oioclea Kunth 18, 397, 395, 398 Entada group 164, 165
Cymbosepalum Baker 135 Desmanthus Willd. 14, 163, 174, 175, 176 Oiocleinae Benth. 9, 51, 394 Entada Adans. 1, 14, 47, 48, 169, 170
Cynometra group 24, 69 - illinoensis (Micbx.) MacMill. 175 Diphaca Lour. 334 - gigas (L.) Fawcett & Rendle 169
Cynometra L. 13, 34, 42, 69, 88, 89, 131 - pernambucanus (L.) Thell. 175 Diphyllarium Gagnep. 18, 404 - phaseoloides (L.) Merr. 169
- carvalhoi Harms 6 - virgatus (L.) Willd. 175 Diphysajacq. 17, 42, 50, 307, 308, 333 - rheedii Spreng. 169
Cynometreae Benth. 69 Desmodiastrum (Prain) A.Pramanik & Thoth. 9, 19, 444 Diplolobium FMuell. 485 - subgen. Acanthentada Brenan 182
Cytiseae Horan. 283 Desmodieae (Benth.) Hutch. 4, 5, 8, 9, 18, 52, 308, 361, Diplotropis Benth. 5, 15, 49, 237, 238 Entadopsis Britton 169
Cytisinae (Horan.) Benth. 284 393, 394, 433 - purpurea (Rich.) Amshoff 237 Enterolobium Mart. 15, 48, 193, 201 , 212
Cytisophyllum O .Lang 16, 284, 290, 292 - subtribe B1yinae B.G.Schub. 307, 433 - sect. Clathrotropis Benth. 238 Eperua Aubl. 13, 34, 77, 78
- sessilifolium (L.) O.Lang 290 - subtribe Desmodiinae Benth. 433 DipJotropoideae Yakovlev 237 - falcata Aubl. 78
Cytisopsis ]aub. & Spach 19, 465 - subtribe Lespedezinae (Hutch.) B.G.Schub. 433 Dipogon Liebm. 18, 425 - oleifera Ducke 78
Cytisus Desf. 12, 16, 283, 284, 291, 292, 293, 294, 295 Desmodium group 9, 433 - lignosus (L.) Verde. 425 Ephedra L. 24
- battandieri Maire 290 Desmodium Desv. 3, 10, 18, 34, 52, 439, 440, 445 Dipterygeae Polhill 5, 7, 15, 215, 220, 227, 228, 250 epulvinate series 455
- fontanesii Spach ex Ball 295 - elegans DC. 440 Dipteryx Schreb. 15, 250, 251 Eremosparton Fisch. & C.A.Mey. 19, 476, 483
- proliferus L.f. 292 - intortum (Mill.) Urb. 440 - odorata (Aubl.) Willd. 251 Erichsenia Hemsl. 17, 341, 342, 355
- purpureus Scop. 291 - uncinatum (Jacq.) DC. 440 Diptychandra Tu!. 14, 128, 150, 160 Erinacea Adans. 16, 284, 294, 296
- scoparius (L.) Link 292 - subgen. Catenaria (Benth.) Baker 437 - aurantiaca Tu!. 160 - anthyllis Link 294
- sect. Lembotmpis (Griseb.) Benth. 293 - subgen. Dendrolobium Wight & Arn. 436 Discolobium Benth. 16, 50, 315 Erinaceinae Talavera 284
- sect. Petteria (C.Presl) Polhill 290 - subgen. Hanslia (Schindl.) H.Ohashi 437 dispersal assembly 41, 43, 44 Eriosema (DC.) Rchb. 3, 18, 52, 410, 411
- sect. Tubocytisus DC. 293 - subgen. Ougeinia (Benth.) H.Ohashi 436 Distemonanthus Benth. 13, 115 Erophaca Boiss. 9, 19, 480, 481
- subgen. Podocarpium (Benth.) H.Ohashi 441 - benthamianus Baill. 115 - baetica Boiss. 476
Dablstedtia Malme 17, 379 - sect. Monarthrocarpus (Merr.) H.Ohashi 439 Disynstemon R.Vig. 17, 376 Errazurizia Phil. 16, 299, 302
Dalbergia clade 5, 8, 50, 308, 309 Desmofischera Holthuis 439 Dolichonemia Nees 149 - rotundata (Wooton) Barneby 301, 302
Dalbergia L.f. 12, 17, 307, 308, 327, 328 Detarieae DC. 6, 13, 22, 24, 34, 46, 47, 49, 54, 69, 70, 127 Dolichopsis Hass!. 18, 430, 429 Erythrina L. 3, 12, 18, 31, 52, 413
- candenatensis (Dennst.) Prain 327 - sens. lat. 5, 46, 69, 71 Dolichos L. 18, 423, 424, 425 Erythrineae (Benth.) Hassk. 393
- densa Benth. 327 - sens. strict. 71 Do1ycnium Mill. 19, 456, 465, 464 Erythrininae Benth. 367
- ecastaphyllum (L.) Taub. 327 Detarium clade 71 - hirsutum (L.) Ser. 464 E1ythrophleum Afzel. ex R.Br. 14, 127, 156, 157
- latifolia Roxb. 327 Detarium group 69 - pentaphyllum Scop. 464 Erythrophloeum 157
- melanoxylon Guill. & Perr. 327 Detarium Juss. 13, 74, 83, 84 - strictum (Fisch. & C.A.Mey.) Lassen 464 Erythrostemon Klotzsch 6, 14, 128, 141, 142
- nigra (Yell.) Allemao ex Benth. 12, 327 - senegalense J.F.Gmel. 84 Do1ycnopsis Boiss. 9, 19, 53, 455, 459, 460, 461, 462 - gilliesii (Wall. ex Hook.) Klotzsch 141
- sissoo Roxb. 327 Dewevrea Micheli 17, 375 Drepanocaipus G.Mey. 328 Etaballia Benth. 7, 8, 16, 227, 247, 318, 321
Dalbergieae Bronn ex DC. 5, 7, 8, 16, 49, 215, 227, 307, Dewindtia De Wild. 100 Droogmansia De Wild. 18, 439 Euchilopsis F.Muell. 17, 344
433 Dialiinae H.S.Irwin & Barneby 5, 24, 34, 46, 47, 48, 54, 111, - godefmyana (Kuntze) Schindl. 439 - linearis (Benth.) F.Muell. 344
- subtribe Euchrestinae Nakai 260 112 Dumasia DC. 18, 52, 416, 418 Euchilus R.Br. ex WTAiton 348
- subtribe Geoffroeeae Benth. 260 Dialium L. 6, 14, 34, 111, 115, 117, 118, 119, 120, 121 Dunbaria Wight & Arn. 18, 411 Euchresta Benn. 16, 49, 260, 261
- subtribe Lonchocarpinae Benth. 367 - cochinchinense Pierre 120 Duparquetia Bail!. 5, 6, 13, 46, 113 - horsfieldii (Lesch.) Benn. 261
- sens. lat. 4, 9 - dinklagei 121 - orchidacea Bail!. 113 - japonica Hook.f. ex Regel 261
Dalbergiella Baker f. 17, 373 - guianense (Aubl.) Sandwith 121 Duparquetiinae H.S.I1win & Barneby 46, 111 Euchresteae (Nakai) I-I.Ohashi 5, 7, 8, 16, 49, 253, 260, 263,
Dalbergioid clade 5, 7, 8, 299, 309 - pachyphyllum Harms 120 Dussia group 227, 228 267, 273
- sens. lat. clade 5, 7, 8, 50 - subgen . Uittienia (Steenis) Steyaert 119 Dussia Krug & Urb. ex Taub. 7, 15, 220, 231 Eurypetalum Harms 13, 34, 77
- sens. strict. clade 50 Dibrachion Tul. 237 Dysolobium (Benth.) Prain 18, 419, 414 - batesii Baker f. 77
Dalea Lucanus 3, 16, 50, 299, 304 Dicerma DC. 9, 437 Eutaxia R.Br. ex W.T.Aiton 17, 347
- filiciformis B.L.Rob. & Greenm. 304 Dichilus DC. 16, 50, 267, 273, 283, 284, 285, 286, 287 Ebenopsis Britton & Rose 6, 15, 193, 208 Eversmannia Bunge 19, 492
- purpurea Vent. 304 Dichrostachys group 163, 164, 165 - ebano (Berland.) Barneby & ].W.Grimes 208 Exostyles Schott 15, 215, 223, 224
Daleeae Hutch. 299 Dichrostachys (DC.) Wight & Arn. 12, 14, 47, 163, 176, 177 Ebenus L. 19, 489, 495 Eysenhardtia Kunth 16, 299, 303
Daleoid clade 8, 299 - cinerea (L.) Wight & Arn. 177 - cretica L. 495 - texana Scheele 303

566 LEGUMES OF THE WORLD INDEX TO SCIENTIFIC NAMES 567

Faba Mill. 1, 2, 505, 506 Gen1:stella Ortega 295 Halimodendron Fisch. ex DC. 9, 19, 476, 489, 491 Hymenaea group 69
Fabaceae 1, 2, 505 Genisticlium I.M.Johnst. 19, 467, 469, 472 - halodendron (Pall.) Voss 491 Hymenaea L. 10, 13, 34, 42, 69, 79, 80, 81
Fabales 1, 4, 5 Genistinae Bronn 283, 284 Hallia 7bunb. 447, 448 - protera Poinar 80
Fabeae Rchb. 5, 10, 19, 36, 53, 54, 455, 475, 496, 499, 505 Genistoid clade 5, 7, 22, 49, 227, 228, 299, 336 Hammatolobium Fenzl 19, 465 - verrucosa Gaertn. 80
Faboideae 2, 505 Genus A of Nielsen 0981) 193 Hanslia Schindl. 9, 18, 437 Hymenocarpos Savi 19, 459
Factorovskya Big 499, 503 Genus B of Nielsen (1981) 193 Haplormosia Harms 15, 235 - circinnatus (L.) Savi 459
Fagelitt 1 eek. ex DC. 409 Genus C of Nielsen (1981) 193 - monophylla (Harms) Harms 235 Hymenolobium Benth. 5, 8, 16, 50, 307, 311, 310
Faidhcrl i<t A.Chev. 6, 12, 15, 31, 187, 193, 196 Genus D of Nielsen (1981) 193 Hardenbergia Benth. 18, 406 - excelsum Ducke 310
- albida (Del.) A.Chev. 196 Geoffroea Jacq. 16, 321, 323 Hardwickia Roxb. 13, 46, 76 Hymenostegia group 69
Falcataria (LC.Nielsen) Barneby & J.W.Grimes 7, 15, 193, Gigasiphon Drake 13, 45, 57, 58, 61, 62, 64 - binata Roxb. 76 Hymenostegia (Benth.) Harms 13, 92, 93, 94, 95
202 , 203 - gossweileri (Bak.f.) Torre & Hille. 62 Harleyodendron RS .Cowan 15, 215, 223, 224 - afzelii (Oliv.) Harms 94
- moluccana (Miq.) Barneby & J.W.Grimes 202, 203 - hu mblotianum (Baill.) Drake 62 Harpalyce Mot;:. & Sesse ex DC. 7, 16, 253, 255 Hypocalypteae (Yakovlev) A.L.Schutte 5, 8, 17, 22, 36, 51,
Fedorovia Yakovlev 235 - macrosiphon (Harms) Brenan 62 Havardia Small 15, 207, 208 267, 336, 339, 355, 361
Ferreiria Allemiio 234 Gilbertiodendron ].Leonard 13, 99, 104 - pallens (Benth.) Britton & Rose 208 Hypocalyptus Thunb. 8, 17, 36, 267, 336, 337
Feuilleea Kuntze 200 Gilletiodenclron Vermoesen 13, 82 Haydonia R. Wilczek 427
Fiebrigiella Harms 16, 324 Gleditsia group 127, 128 Hebestigma Urb. 19, 467, 468 Icuria J.J.Wieringa 6, 13, 107
Fillaeopsis group 164, 165 Gleditsia L. 12, 14, 23, 24, 40, 42, 47, 111, 127, 130, 131, Hecastophyllum Kuntb 327 Jmbralyx R.Geesink 374
Fillaeopsis Harms 14, 171 132, 133 Hedysareae DC. 5, 9, 10, 19, 36, 53, 54, 455, 476, 489, 496, immigration rates 41, 42, 43, 44
- discophora Harms 171 - amorphoides Taub. 24, 42 499, 505 IndigastrumJaub. & Spach 17, 361, 362
Fissica lyx Benth. 16, 324 - triacanthos L. 131 - subtribe Adesmiinae Benth., as Adesmieae 307, 489 Indigofera L. 3, 8, 17, 31, 51, 361, 363, 364
Flemingia Roxb. ex W.T.Aiton 18, 413 Gliricidia Kunth 9, 12, 19, 42, 382, 467, 468, 469 - subtribe Aeschynomeninae Bentb., as Aeschynomeneae - ammoxylum (DC.) Polhill 363
- grahamiana Wight & Arn . 413 - sepium Oacq.) Steud. 468 307, 489 - arrecta Hochst. ex A. Rich. 364
Fordia Hems!. 17, 374 Glottidium Desv. 453 - subtribe Coronillinae Bronn, as Coronilleae 489 - articulata Gouan 364
- cauliflora Hems!. 374 Glycine Willd. 18, 52, 416, 418, 420, 421 , 447 - subtribe Desmodiinae Ben.th., as Desmodieae 433, 489 - suffruticosa Mill. 364
- leptobot1ys (Dunn) Schot, Dasuki & Buijsen 374 - max (L.) Merr. 10, 12, 421 - subtribe Patagon.iinae Taub. 307 - tinctoria L. 364
- splendidissima (Blume ex Miq.) Buijsen 374 Glycineae Burnett 393 Hedysaroid clade 489 - subgen. lndigastrum (Jaub. & Spacb) j.B.Gillett 362
- sens. lat. 367 Glycininae (Burnett) Benth. 9, 52, 53 Hedysarum L. 3, 10, 19, 54, 489, 494, 493, 495 - subgen. Microcharis (Ben.th.) j.B.Gillett 363
- sens. strict. 367 Glycinopsis Kuntze 402 - coronarium L. 489, 493 Incligofereae Benth. 5, 8, 17, 22, 31, 51, 299, 336, 355, 361
fossil evidence 21, 24, 34, 36, 40, 43, 44, 49 Glycyrrbi za L. 5, 9, 10, 19, 53, 478 Hegnera Schindl. 9, 18, 441 Inclopiptadenia Brenan 14, 170, 171
- glabra L. 10, 478 Heinekenia Webb ex H. Christ 463 - oudhensis (Brandis) Brenan 170
Gagnebina Neck. ex DC. 14, 177, 176 Goldmania Rose ex Micheli 181 Helmintbocmpon A.Rieb . 461 Inga alliance 195
Galactia P.Browne 18, 51, 397, 398, 399 Gompho lobium Sm. 17, 355, 340 Herminiera Guill. & Perr. 329 Inga Mill. 3, 12, 15, 33, 193, 200
- sect. Collaearia (Benth.) Burka1t 398 Gondwana hypothesis 21, 41 , 44 Herpyza C.Wright 18, 52, 399, 417 - edulis Mart. 200
- sect. Ga lactia 398 Goniogyna DC. 279 Hesperalbizia Barneby & J.W.Grimes 7, 15, 193, 210, 212 Ingeae Benth. 1, 5, 6, 15, 22, 48, 54, 163, 187, 193
- sect. Odonia (Berto!.) Burkart 398 Goniorrhachis Taub. 13, 46, 72, 73 Hesperolaburnum Maire 16, 284, 291, 292 Inocarpus JR.Forst. & G.Forst. 7, 8, 16, 34, 42, 50, 227, 247,
Galega L. 5, 10, 19, 54, 475, 487, 496, 499, 505 Gonocytisus Spach 16, 284, 294 Hesperothamnus Brandegee 17, 52, 378, 379 308, 318, 321
- officinalis L. 487 Good ia Salisb. 17, 355, 356 Heteroflorum (ined.) 128, 154 - fagifer (Parkinson) Fosberg 318
Galegeae (Bronn) Dumort 5, 9, 10, 19, 36, 54, 299, 455, - lotifolia Salisb. 356 Heterostemon Desf. 13, 95, 97 intergeneric disjunction 34, 37, 46, 47, 48, 49, 50, 51
475 , 496, 499, 505 Gossweilerodendron Harms 13, 74, 75 , 76 - subgen. Bracbycylix Hanns 97 intrageneric disjunction 34, 37, 45, 46, 47, 48, 49, 50, 51
- sens. lat. 9, 10, 476, 505 - balsamiferum (Vermoesen) Harms 75 Heylandia DC. 279 Intsia Thouars 13, 69, 90, 92
- subtribe Alhagiinae DC. 475 Gourliea Gillies ex Hook. & Arn . 323 Hippocrepis L. 19, 53, 457, 458 - bijuga (Colebr.) Kuntze 90
- subtribe Astragalinae DC. 475 Grass biome 31, 44 - emerus (L.) Lassen 457 Inve1ted Repeat Lacking Clade (IRLC) 5, 8, 9, 36, 53, 299,
- subtribe Brongniartiinae Benth . 253 Grazielodendron H.C.Lima 17, 308, 326 Hoffmannseggia Cav. 14, 128, 141, 145, 146, 147 452, 455 , 475, 496, 505
- subtribe Coluteinae Benth. 475 Griffonia Baill. 13, 45, 57, 59, 60 - glauca (Ortega) Eifert 145 IRLC millettioid group 5, 368, 369
- subtribe Cmynellinae Rydb. 467 Grimaldia Schrank 122 - intricata Brandegee 145 Isoberlinia Craib & Stapf. 13, 104
- subtribe Ga leginae Bronn. 475, 499, 505 Grona sensu Benth. 419 Hoita Rydb. 19, 449 lsomacrolobium Aubrev. & Pellegr. 102
- subtribe Glycyrrhizinae Rydb. 475 Guaymasia Britton & Rose 139 Holocalyx Micheli 15, 49, 215, 222 , 234 Isotropis Benth. 17, 339, 342, 348
- subtribe Jn digoferinae (Benth.) Benth. 361 Gueldenstaedtia Fisch. 19, 476, 479 - balansae Micheli 222 ltaobimia Rizzini 315
- subtribe Psoraleinae A. Gray 299, 447 - himalaica Baker 479 Hologalegina 5, 7, 8, 9, 53, 54, 336, 339, 355, 361, 455
- subtribe Robiniinae Benth. 467 Guibou rtia Benn. 13, 34, 42, 69, 79, 80, 81 Horn of Africa 24, 42, 43, 44, 53 Jacksonia R.Br. ex Sm. 17, 343, 344
- subtribe Sesbaniinae Rydb. 452 - hymenaeifolia (Moricand) ].Leonard 81 Hosackia Douglas ex Lindi. 9, 19, 53, 460, 462, 463 - scoparia Sm. 343
- subtribe Tephrosiinae Benth. 367 Guilandina L. 6, 14, 128, 143 - gracilis Benth. 460 Jacqueshuberia Ducke 14, 127, 128, 150, 151
Gamwellia Baker/ 281 - bonduc L. 143 Hovea R.Br. ex W.T.Aiton 16, 45 , 49, 253, 257 jansonia Kippist ex Lindi. 8, 351, 352
Gas trolobium R.Br. ex W.T.Aiton 3, 8, 17, 350, 351, 352 Guilfoy lia monostylis F.Muell 45 - elliptica (Sm.) DC. 257 Julbernarclia Pellegr. 6, 13, 105, 106, 107, 108
- grancl iflorum F.Muell. 352 Guinetia L.Rico & M.Sousa 7, 15, 193, 197 - longipes Benth. 257 - paniculata (Benth.) Troupin 105
Geissaspis Wight & Arn. 17, 329, 330, 332 Gymnocladus Lam. 14, 23, 24 , 40, 42, 47, 111, 127, 130, Humboldtia Vahl. 13, 69, 93, 94 - pellegriniana Troupin 105
Genista group 283 131, 133 - africana Baill. 94 jupunba Britton. & Rose 205
Genista L. 12, 16, 284, 293, 294, 295 , 296 - assamicus Kanj . ex P.C. Kanj. 130 Humboldtiella Hanns 471
- tinctoria L. 295 - chinensis Baill. 130 Humularia P.A.Duvign. 17, 329, 332 Kalappia Kosterm. 14, 111, 119
Genisteae (Bronn) Dumort. 5, 8, 16, 36, 50, 253, 263, 267, - dioica (L.) K.Koch 130 Hybosema Hanns 9, 467, 468 - celebica Kosterm. 12, 119
273, 275, 279, 283 Hydrochorea Barneby &].W.Gri.mes 7, 15, 193, 204, 205, 207 Kamiella Vassilcz. 499, 503
- subtribe Bossiaeinae Benth. , as Bossiaeae 355 Haematoxylon. L. 135 - (?) acreana O.F.Macbr.) Barneby & J.W.Grimes 204 Kanaloa Lorence & KR.Wood 6, 14, 163, 174, 175, 176
- subtribe Crotalariineae Benth. 273 Haematoxylum L. 14, 127, 134, 135, 137 Hylodendron Taub. 13, 82 - kahoolawensis Lorence & KR.Wood 176
- subtribe Ge nistinae Bronn 283 - brasiletto Karst. 135 Hylodesmum H.Ohashi & RR.Mill 9, 18, 441 Kaoue Pel!egr. 159
- subtribe Lupininae Rouy 283 - campechianum L. 135 - repandum (Vahl) H.Ohashi & RR.Mill 441 Kebirita Kramina & D .D.Sokoloff 9, 19, 53, 456, 460, 461

568 LEGUMES OF THE WORLD INDEX TO SCIENTIFIC NAMES 569

- roudairei (Bonnet) Kramina & D.D.Sokoloff 461 Leonardoxa Aubrev. 13, 86, 93, 94, 95 Luetzelburgia Harms 5, 7, 15, 234 Melilotoides Heist. ex Fabr. 499, 503
- africana (Baill.) Aubrev. 95 Lumbricidia Veil. 311 Melilotus Mill. 10, 19, 499, 502, 503
Kennedia Vent. 18, 406, 407
Lepidocomajungh . 413 Lupinaster Fabr. 499, 501 - albus Medik. 502
- prostrata R.Br. 406
Leptoderris Dunn 17, 372 Lupineae (Rouy) Hutch . 283 - officinalis (L.) Pallas 502
Kennediinae Benth. 52, 433
Leptodesmia (Benth.) Benth. 19, 445 Lupininae Rouy 284 Melliniella Harms 19, 445
Kennedya DC. 406
Leptolobium Vogel 236 Lupinophyllum Hutch. 386 Melolobium Eckl. & Zeyh. 16, 50, 267, 273, 283, 284, 285,
Kerstania Rechf 458
Kerstingiella Harms 425 Leptosema Benth. 17, 343, 344 Lupinus L. 3, 12, 16, 36, 50, 283, 284, 287, 289 286, 287
Lespedeza group 9, 433 - albus L. 287 Mendoravia Capuron 13, 46, 111, 115
Keyserlingia Bunge 243
Lespedeza Michx. 18, 52, 435 - luteus L. 287 Meristotropis Fisch. & CA.Mey. 478
Kingiodendron Harms 13, 34, 42, 74, 75, 76
- bicolor Turcz. 435 - mutabilis Sweet 287 metacommunity 41 , 42, 43, 44
- pinnatum (Roxb.) Harms 75
- juncea (L.f.) Pers. var. sericea (Thunb.) Lace & Hems!. - polyphyllus 287 Mezoneuron Desf. 6, 14, 128, 136
Klugiodendron Britton & Killip 193, 205
Koompassia Maingay ex Benth. 14, 34, 117, 118 435 Luzonia Elmer 18, 395 - angolense Welw. ex Oliv. 136
- thunbergii (DC.) Nakai 435 - purpurea Elmer 395 - kauaiense (Mann) Hillebr. 136
- excelsa (Becc.) Taub. 117
- subgen. Macrolespedeza 435 Lyauteya Maire 465 Mezoneurum DC. 136
Kostyczewa Karsh. 478
Lespedezinae (Hutch.) B.G.Schub. 9 Lysidice Hance 13, 85, 86 Mezonevron Desf. 136
Kotschya End!. 17, 329, 330, 331
Lessertia DC. 19, 484, 485 - rhodostegia Hance 85 Michelsonia Hauman 13, 108
Krauhnia Steud. 372
- perennans DC. 484 Lysiloma Benth. 15, 193, 212 Mickle thwaitia G.P.Lewis & Schrire 6, 13, 88
Kuhnistera Lam. 299, 304
Leucaena group 163, 164, 165 - sabicu Benth. 212 Microberlinia A.Chev. 13, 105
Kummerowia Schindl. 18, 434
Leucaena Benth. 12, 14, 34, 47, 163, 174, 175 Lysiphyllum (Benth.) de Wit 13, 45, 57, 58, 62, 63, 64 - brazzavillensis A.Chev. 105
Kunstleria Prain 17, 377, 407
- leucocephala (Lam.) de Wit 174 - winitii (Craib) de Wit 64 Microcharis Benth. 17, 361, 362, 363
Leucochloron Barneby & J.W.Grimes 7, 15, 193, 206 Microlespedeza (Maxim) Makino 434
Labichea Gaud. ex DC. 14, 34, 111, 116, 117
Leucomphalos Benth. ex Planch. 7, 15, 227, 245, 246, 247 Maackia Rupr. & Maxim. 15, 242 Microlobius C.Presl 14, 180, 181, 182, 183
Labicheinae H.S.Irwin & Barneby 111
Leucostegane Prain 13, 86 - amurensis Rupr. & Maxim. 242, 244 Mildbraediodendron Harms 15, 215, 220, 47
Lablab Adans. 18, 423, 425
- purpureus (L.) Sweet 425 Leycephyllum Piper 410 Machaerium Pers. 3, 17, 50, 307, 308, 321, 327, 328, 329 - excelsum Harms 220
Libidibia (DC.) Schltdl. 6, 14, 128, 144 - isadelphum CE.Mey.) Amshoff 328 Millettia group 368
Laburneae (Rouy) Hutch. 283
- coriaria (Jacq.) Schltdl. 144 - lunatum (L.f.) Ducke 43 Millettia Wight & Arn. 3, 9, 12, 17, 367, 375, 379, 383, 384
Laburninae Rouy 28
Laburnocytisus (Poit.) C.K.Schneid. 291 Librevillea Hoyle 13, 103 - sect. Tipuana Benth. 320 - laurentii De Wild 383
Liparia L. 8, 16, 270 Macrolobieae clade 71 - leptobot1ya Dunn 374
- adamii (Poit.) C.K.Schneid. 291
- splendens (Burm.f.) Bos & de Wit 270 Macrolobieae Breteler 46, 69, 70, 71 - stuhlmannii Taub. 383
Laburnum Fabr. 12, 16, 279, 284, 291
- alpinum (Mill.) Bercht. & ].Pres! 291 Liparieae (Benth.) Harv. 8, 267, 336 Macrolobium group 69 - sens. lat. 52, 367
- anagyroides Medik. 291 - subtribe Hypocalyptinae Yakovlev 336 Macrolobium Schreb. 13, 69, 99, 100, 102, 104 - sens. strict. 52
- x watereri (Kirchner) Dippel 291 Listia E.Mey. 278 Macrolobium auct. non Schreb. 102 - sect. Albiflorae Dunn 374
Lackeya Fottunato, L.P.Queiroz & G.P.Lewis 9, 18, 51, 398, Loesenera Harms 13, 92, 93, 95 Macropsychanthus Harms 18, 395 - sect. Bracteatae Dunn 370
- kalantha Harms 92 Macroptilium (Benth.) Urb. 10, 18, 429, 430 - sect. Eurybotryae Dunn 370
399
Lonchocarpeae (Benth.) Hutch. 367 - atropurpureum (DC.) Urb. 430 Millettieae Miq. 5, 7, 8, 9, 17, 299, 355, 361, 367, 393, 455,
Lamprolobium Benth. 16, 253, 256, 258
Lonchocarpinae Benth. 260 - lathyroides (L.) Urb. 430 472, 499, 505
Larrea Ortega 145
Lasiobema (Ko1th.) Miq. 13, 58, 62, 63, 65, 66 Lonchocarpus group 368 Macrosamanea Britton & Rose ex Britton & Killip 15, 193, - canavanine group 368, 369
Lonchocarpus Kunth. 3, 9, 17, 33, 52, 367, 379, 380, 381, 199 - non-canavanine group 368, 369
Lathriogyne Eckl. & Zeyh. 269
382 Macrotyloma (Wight & Arn) Verde. 18, 424, 425 - sens . lat. 367, 496
Lathyrus L. 3, 12, 19, 54, 505, 507
- sericeus 43 - axillare (E.Mey.) Verde. 425 - sens. strict. 8, 9
- cicera L. 507
- subgen. Phacelanthus Pittier 380 - geocarpum (Harms) Marechal & Baudet 425 Millettioid groups 22, 54, 368, 393, 394
- hirsutus L. 507
- sect. Caudai'ia Dunn 383 - uniflorum (Lam.) Verde. 425 Millettioid sens. lat. clade 5, 7, 8, 51
- latifolius L. 507
- sect. Fasciculati Benth. 380 Madrean-Tethyan hypothesis 40 Mimosa L. 3, 12, 14, 31, 47, 48, 163, 183, 184
- montanus Bernh. 507
- sect. Paniculati Benth. 378 Maniltoa Scheff. 13, 88, 89 - pudica L. 183
- ochrus (L.) DC. 507
Lophocarpinia Burkart 14, 146 - grandiflora Scheff. 88 Mimoseae Bronn 6, 14, 22, 34, 47, 48, 127, 163, 184, 187
- odoratus L. 507
Loteae DC. 5, 9, 19, 36, 53, 452, 455, 499, 505 Maraniona C.E.Hughes, G.P.Lewis, Daza & Reyne! 8, 16, - sens. lat. 5, 163
- sativus L. 507
- subtribe Lipariinae Benth. 267 308, 319 - sens. strict. 5, 163
Latrobea Meisn. 17, 344
Lotodes Kuntze 448 - lavinii C.E.Hughes, G.P.Lewis, Daza & Reyne! 319 Mimosoicleae (R.Br.) DC. 1, 2, 3, 5, 6, 14, 21, 22, 46, 47, 49,
Laurasia 21, 40, 44
Lebeckia Thunb. 16, 50, 273, 275, 283 Lotononideae Hutch. 273 Margaritolobium Harms 17, 382 54, 127, 128, 163, 184, 193
Lotononis (DC.) Eckl. & Zeyh. 3, 16, 36, 50, 273, 275, 278, Marina Liebm. 16, 299, 304 Mimozygantheae Burkart 5, 6, 15, 163, 184
Lebruniodendron ].Leonard 13, 87, 88
279, 281 Marmarm,'Ylon Killip 15, 193, 201 Mimozyganthus Burkart 15, 173, 184, 185
Lecointea clade 215
- bainesii Baker 278 - racemosum (Ducke) Killip 201 - carinatus (Griseb.) Burka1t 184
Lecointea Ducke 15, 222, 223, 234
Lotus L. 3, 9, 19, 299, 455, 460, 463, 464, 465 Martia Benth. 118 Minkelersia Mart. & Galeotti 428
Lecointeoid clade 5, 7, 49, 216, 228
Leguminosae crown clade 45 - australis Andr. 463 Martiodendron Gleason 14, 111, 117, 118 Mirbelia Sm. 17, 339, 349, 350, 351
- bertholetii Masf. 463 Maitiusia Benth. 118 - oxylobioicles F.Muell. 350
Lembotropis Griseb. 16, 283, 284, 292, 293
- nigricans (L.) Griseb. 293 - corniculatus L. 10, 463 Mastersia Benth. 18, 404 Mirbelieae (Benth.) Polhill & Crisp 5, 8, 17, 36, 51, 263,
Lemurodendron Villiers 14, 163, 171 - japonicus (Regel) K.Larsen 12 Mecopus Benn. 18, 442 336, 339, 355, 361
Lemuropisum H.Perrier 14, 127, 146, 153, 154, 155, 156 - maculatus Breitf. 463 Medicago L. 19, 499, 503 - giant antipodals group 339
- pedunculatus Cav. sens lat. 463 - sativa L. 10, 503 - 5-nucleate group 339
- edule H.Perrier 156
- roudairei Bonnet 461 - truncatula Gae1tn. 12 Mirbelioicl sens. lat. clade 5, 7, 8 , 22, 36, 51
Lennea Klotzsch 19, 467, 468
-strictus Fisch. & C.A.Mey. 464 Meizotropis Voigt 18, 407, 408 Moghania].St.-Hil. 413
Lens Mill. 19, 505, 506
- sect. Microlotus Benth. 462 Meladenia Turcz. 450 Moldenhawera group 129
- culinaris Medik. 10
- sect. Simpeteria Ottley 462 Melanosticta DC. 141 Molclenhawera Schrad. 14, 127, 128, 149
- nigricans (M.Bieb.) Godr. 506
Lourea Neck. ex Desv. 443 Melanoxylon Schott 14, 127, 128, 148, 149 Monarthrocarpus Merr. 9, 18, 439
- odemensis Ladiz. 506
Lovanafia M.Peltier 240 - brauna Schott 148 - securiformis (Benth.) Merr. 439
Leonardendron Aubrev. 102

INDEX TO SCIENTIFIC NAMES 571

570 LEGUMES OF THE WORLD
Monopetalanthus Harms 106, 108
- auct. non Harms, pro pai1e 107
Monoplegma Piper 427
Monopteryx Spruce ex Benth. 7, 15, 232, 250
Monoschisma Brenan l 79
Montigena Heenan 10, 19, 486
- novae-zelandiae (Hook.f.) Heenan 486
Moparia Britton & Rose 145
Mora Schomb. ex Benth. 14, 127, 156, 157, 158
- excelsa Benth. 158
- gonggrijpii (Kleinhoonte) Sandw. 158
- megistosperma (Pittier) Britton & Rose 158
Moreae Britton & Rose 127
Morolobium Kosterm. 202
Moullava Adans. 14, 147
- spicata (Dalzell) Nicolson 147
Mucuna Adans. 3, 9, 12, 18, 52, 405, 433
- pruriens (L.) DC. 405
Muellera L.f. 17, 380, 382
Muelleranthus Hutch. 17, 355, 358
Mundulea CDC.) Benth. 17, 384, 385
Murtonia Craib 440
Myrocarpus Allemao 7, 15, 232, 231, 247
- frondosus Allemao 231
Myrospermum]acq. 5, 7, 15, 17, 220, 232
Myroxylon group 8, 227, 228
Myroxylon L.f. 7, 15, 220, 232
- balsamum (L.) Harms 232
- peruiferum L.f. 232
Mysanthus G.P.Lewis & A.Delgado 9, 18, 429, 430
Nemcia Domin 8, 351, 352
Neoapaloxylon Rauschert 13, 46, 72, 73, 74
Neobaronia Bakei- 362
Neochevalierodendron ].Leonard 13, 93
Neocollettia Hems!. 9, 18, 421, 422, 433
Neocracca Kuntze 467, 470, 471
Neodielsia Harms 9, 481
Neodunnia R. Vig. 9, 383
Neoharmsia R.Vig. 15, 239
- baronii (Drake) R.Vig. ex M.Peltier 239
Neonotonia ].A.Lackey 18, 416, 417, 418, 421
- wightii (Wight & Arn.) ].A.Lackey 417
Neorautanenia Schinz 18, 417
Neorndolphia Britton 18, 51, 399, 417
Nephrodesmus Schindl. 18, 438
Neptunia Lour. 14, 47, 163, 173, 174
- oleracea Lour. 174
Nesphostylis Verde. 18, 423, 424, 425
Neustanthus Benth. 418
Newtonia group 163, 164, 165
Newtonia Bail!. 14, 47, 48, 163, 171
- buchananii (Baker) G.C.C.Gilbert & Boutique 171
- sect. Neonewtonia Burkart l 79
Nicarago Britton & Rose 138
Nissolia ]acq. 16, 312, 314
- fruticosa Jacq. 314
Nogra Merr. 18, 419
- grahamii (Wall. ex Benth.) Merr. 419
Normandiodendron ].Leonard 4, 13, 93
Notodon Urb. 469
Notonia Wight&Arn. 417
Notospartium Hookf 9, 486
Obolinga Barneby 7, 193, 199
572 LEGUMES OF THE WORLD
Oddoniodendron De Wild 13, 103, 105 Papilionoideae L. ex DC. 1, 2, 3, 5, 7, 15, 21, 22, 34, 46, 47, Phaseoleae (Bronn) DC. 5, 9, 18, 22, 52, 355, 361, 367,
- micranthum (Harms) Baker f. 105 215, 227, 505 394, 433
- romeroi Mendes 105 Paraberlinia Pelleg1·. 105 - sens. lat. clade 9, 52, 367, 389, 393, 433, 447
Ohwia H.Ohashi 9, 18, 437 Paracalyx Ali 18, 52, 408 - sens. strict. 5, 9
Old World clade 8, 51 Paraderris (Miq.) R.Geesink 9, 17, 52, 382, 383 - subtribe Abrinae Wight & Arn. ex End!. 389
Old World group of lngeae 195 - elliptica (Wall.) Adema 383 - subtribe Cajaninae Benth. 393, 394
Oleiocaipus Dwyer 251 Paraglycine Fj.Herm. 400 - subtribe Clitoriinae Benth. 393, 394
Oligostemon Benth. 113 Paramachaerium Ducke 16, 321 - subtribe Diocleinae Benth . 367, 393, 394
Olneya A.Gray 19, 467, 469 Paramacrolobium ].Leonard 13, 99, 100 - subtribe Glycininae Benth. 393, 394, 447
- tesota A.Gray 469 - coeruleum (Taub.) ].Leonard 100 - subtribe Kennediinae Benth. 394
Onob1ychis Mill . 3, 19, 489, 494, 495 Parapiptadenia Brenan 14, 180, 181, 182, 183 - subtribe Ophrestiinae ].A.Lackey 367, 393, 394
- viciifolia Scop. 494 - rigida (Benth.) Brenan 181 - subtribe Phaseolinae Bronn 42, 52, 393, 394
Ononideae Hutch . 499 Pararchidendron LC.Nielsen 15, 193, 204 Phaseolus L. 10, 18, 31, 428
Ononis L. 19, 499, 502 - pruinosum (Benth.) LC.Nielsen 204 - acutifolius A.Gray 428
- repens L. 502 Paraserianthes LC.Nielsen 15, 193, 202, 203, 211 - coccineus L. 428
- spinosa L. 502 - lophantha (Willd.) LC.Nielsen 203 - dumosus Macfad. 428
Ophiocarpus (Bunge) Ikonn. 10, 19, 476, 481, 482 - sect. Falcataria I.C.Nielsen 193, 202, 203 - lunatus L. 428
- aitchisonii (Baker) Podlech 482 Paratephrosia Domin 9, 17, 367, 387 - polyanthus Greenman 428
Ophrestia H.M.L.Forbes 18, 400 Parkia R.Br. 14, 34, 163, 165, 178 - vulgaris L. 428
Ophrestiinae ].A.Lackey 9, 394 - biglobosa (Jacq.) G.Don 178 Philenoptera Fenzl ex A.Rich. 9, 17, 52, 378, 380
Orbexilum Raf. 19, 448 - speciosa Hassk. 178 Phylacium Benn. 9, 18, 421, 422, 433
Oreophysa (Bunge ex Boiss.) Bornm. 19, 476, 483 Parkieae (Wight & Arn.) End/. 6, 163, 184 Phyllocaipeae B1'itton & Rose 69
Ormocarpopsis R.Vig. 17, 42, 307, 333, 334, 335 Parkinsonia L. 14, 127, 153 Phyllocapus Riedel ex Tu!. 6, 73
- aspera R.Vig. 335 - aculeata L. 153 Phyllodium group 9, 433
Ormocarpum clade 31, 42 - africana Sond. 153 Phyllodium Desv. 18, 436
Ormocarpum P.Beauv. 17, 42, 50, 307, 308, 333, 334, 335 Parochetinae Chaudhaiy & Sanjappa 499 Phyllota (DC.) Benth. 17, 345
Ormosia group 227, 228 Parochetus Buch.-Ham. ex D.Don 19, 54, 496, 499, 501 Phylloxylon Baill. 17, 51, 361, 362
Ormosia Jacks. 3, 5, 15, 34, 42, 49, 235, 236 - africanus Polhill 501 phylogenetic structure 23, 41, 43
Ormosiopsis Ducke 235 - communis Buch.-Ham. ex D.Don subsp. africanus Physanthyllis Boiss. 465
OrnlthopusL. 19, 53, 455, 456,459, 460, 461 (Polhill) Chaudhary & Sanjappa 501 Physostigma Balf. 18, 426
- sativus Brot. 461 Panyella Torr. & A.Gray ex A.Gray 16, 299, 301 - venenosum Balf. 426
Orobus L. 505 - filifolia Torr. & A.Gray 301 Pickeringia Nutt. ex Torr. & A.Gray 5, 8, 16, 233, 263, 264
Orphanodendron group 127 Pauletia Cav. 58, 61 - montana Nutt. ex Torr. & A.Gray 264
Orphanodendron Barneby &].W.Grimes 14, 128, 161 Pearsonia Dummer 16, 50, 273, 281 Picretia DC. 8, 17, 42, 307, 308, 333, 334, 335
Oryxis A.Delgado & G.P.Lewis 9, 18, 430 Pediomelum Rydb. 19, 450 Piliostigma Hochst. 13, 58, 66
Ostryocarpus Hook.f. 17, 373 - esculentum (Pursh) Rydb. 450 - reticulatum (DC.) Hochst. 66
Ostryoderris Dunn 373 Peekelia Harms 427 - thonningii (Schumach.) Milne-Redh. 66
Otholobium C.H. Stitt. 19, 36, 52, 447, 448 Pellegriniodenclron ].Leonard 13, 99, 104 Piptadenia group 163, 164, 165
Otion (ined.) 8, 17, 345, 346 Peltiera Labat & Du Puy 8, 17, 42, 308, 334, 335 Piptadenia Benth. 14, 47, 163, 169, 179, 180, 181, 182, 183
Otoptera DC. 18, 414, 415 Peltogyne Vogel 13, 79 - viridiflora (Kunth) Benth. 179
Ottleya D.D.Sokoloff 9, 19, 53, 461, 462, 463 Peltophoropsis Chiov. 153 - sect. Niopa Benth. 179
Ougeinia Benth. 9, 18, 436 Peltophorum group 127, 128 Piptadeniastrum group 6, 164, 165
0Jq'lobium Andrews 17, 349, 350, 351 Peltophorum (Vogel) Benth. 14, 31, 47, 48, 127, 152 Piptadeniastrum Brenan 14, 163, 169
- lanceo/atum (Vent.) Druce 351 - dubium (Spreng.) Taub. 152 - africanum (Hook.f.) Brenan 169
O:i.'Yrhynchus Brandegee 18, 427, 428, 429, 430 - venezuelense L.C:irdenas, Rodr.-Rodr. & Varela 152 Piptadenieae Bentb. 163
Oxystigma Harms 13, 34, 42, 74, 75, 76 Pentaclethra group 6, 164, 165 Piptadeniopsis Burkatt 14, 163, 173, 184
- oxyphyllum (Harms)]. Leonard 74 Pentaclethra Benth. 5, 6, 14, 34, 47, 48, 127, 156, 157, 158, - lomentifera Burka1t 173
Oxytropis DC. 1, 3, 19, 54, 475, 476, 480 163, 166 Piptanthus Sweet 16, 263, 265
- campestris (L.) DC. 43 - macroloba (Wille!.) Kuntze 166 - nepalensis (Hook.) Sweet 265
- podocarpa A.Gray 43 - macrophylla Benth. 166 Piptomeris Turcz. 343
- viscida Nutt. 43 Periandra Mart. ex Benth. 18, 402 Piscidia group 368
Pericopsis Thwaites 15, 236 Piscidia L. 17, 52, 378, 379
Pachecoa Stand!. & Steyerm. 8, 308, 324, 325 - angolensis (Baker) van Meewen 236 - e1ythrina L. 379
Pachyelasma Harms 14, 127, 156, 157 Perlebia Spix & Mart. 58, 61 Pisum L. 19, 505, 508
- tessmannii (Harms) Harms 156 Petaladenium Ducke 15, 238 - sativum L. 10, 508
Pachyrhizus Rich. ex DC. 18, 52, 416, 417 - urceoliferum Ducke 238 Pithecellobium alliance 195
- erosus (L.) Urb. 416 Petalostemon Michx. 299, 304 Pithecellobium Mart. 15, 48, 193, 207, 209
Padbruggea Miq. 370 Petalostylis R.Br. 14, 111, 115, 116, 117, 119, 120 - dulce (Roxb .) Benth. 209
Pahudia Miq. 91 Peteria A.Gray 19, 467, 469, 472 - incuriale (Veil.) Benth. 193
Painteria Britton & Rose 7, 15, 193, 207, 208 - glandulosa (A.Gray ex S.Watson) Rydb. 472 - sect. Abaremotemon Benth. 205
Paloue Aubl.13, 96 Petteria C.Presl 16, 284, 290, 292 Pitryocapa Britton & Rose 180
- guianensis Aubl. 96 - rarnentacea (Sieber) C.Presl 290 Placolobium Miq. 235
Palovea fuss. 96 Phaca L. sens. strict. 481 Plagiocarpus Benth. 16, 253, 256
Paloveopsis R.S.Cowan 13, 96 Phaenoboffmannia Kuntze 281 Plagiosiphon Harms 13, 88
Panurea Spruce ex Benth. 15, 237, 248 Phanera Lour. 3, 13, 45, 57, 58, 62, 63, 65, 66 Plathymenia group 164, 165
INDEX TO SCIENTIFIC NAMES 573
 Plathymenia Benth. 14, 47, 170 Pseudoeriosema Hauman 18, 400 Rainforest biome 23, 24, 32, 33, 44, 46, 48 Sclerolobium Vogel 6, 150, 151, 160
- reticulata Benth. 170 Pseudoglycine Fj.Herm . 400 Ramirezella Rose 9, 18, 427, 428, 429 Sclerothamnus R.Br. ex WTAiton 347
Platycelyphium Harms 15, 239, 240 Pseudolotus Rech.f. 19, 460 Ramorinoa Speg. 16, 320 Scorodophloeus Harms 13, 86, 87, 88
- voense (Engl.) Wild 240 Pseudomacrolobium Hauman 13, 102 - girolae Speg. 320 - fischeri (Taub.) ].Leonard 86
Platycyamus Benth. 17, 34, 51, 52, 377, 407 - mengei (De Wild.) Hauman 102 Recchia Mos;. & Sesse ex DC. 45 - zenkeri Harms 86
Platylobium Sm. 17, 355, 356, 357 Pseudomelissitus Ovcz., Rassulova & Kinzikaeva 499, 503 reciprocal monophyly 31, 34, 42, 43 Scorpiurus L. 19, 53, 457
- formosum Sm. 357 Pseudopiptadenia Rauschert 14, 179 Recordoxylon Ducke 14, 127, 128, 148, 149 - vermiculatus L. 457
Platymiscium Vogel 16, 317 - pittieri (Harms) G.P.Lewis 179 Rehsonia Stritch 372 seasonally d1y tropical forests (SDTF) 23, 32
Platypodium Vogel 16, 50, 318, 322 Pseudoprosopis Harms 14, 168 Reichardia Roth 137 Securigera DC. 19, 53, 455, 458
- elegans Vogel 318 Pseudosamanea Harms 7, 15, 193, 210 Requienia DC. 9, 17, 367, 387 - securidaca (L.) Degen & Dorf!. 458
Platysepalum Welw. ex Baker 17, 375 - cubana (Britton & Rose) Barneby & J.W.Grimes 210 Retama Raf. 16, 50, 283, 284, 294, 295 - varia (L.) Lassen 458
Pleistocenic Arc 23, 24, 32, 42 - guachapele (Kunth) Harms 210 - monosperma (L.) Boiss. 294 Sellocharis Taub. 8, 16, 273, 283, 284, 287, 289
Podalyria Willd. 16, 263, 269, 270 Pseudosindora Symington 4, 84 Rhodopis Urb. 18, 51, 395, 399 Senegalia Raj 5, 187
- calyptrata (Retz.) Willd. 269 Pseudovigna (Harms) Verde. 18, 419, 420 Rhodusia Vassilcz. 499, 503 Senna Mill. 3, 5, 6, 12, 14, 31, 111, 112, 122, 123, 124, 127,
Podalyrieae Benth. 1837, emend. A. L. Schutte 5, 8, 16, 36, - argentea (Willd.) Verde. 420 Rhynchosia Lour. 3, 9, 18, 52, 408, 410, 411 134, 150
50, 253, 263, 267, 273, 283, 337 Psophocarpus Neck. ex DC. 18, 52, 414 - ferulifolia Benth. ex Harv. 410 - alata (L.) Roxb. 123
- subtribe Mirbeliinae Bentb . 339 - tetragonolobus (L.) DC. 414 Rhynchotropis Harms 17, 361, 362, 363 - alexandrina Mill. 111, 123
- subtribe Xiphothecinae A.L.Schutte 268 Psoralea L. 19, 36, 52, 299, 447, 448 Rhytidomene Ryd. 448 Serianthes Benth. 15, 202, 203
Podlecbiella Maassoumi & Kazempow· Osaloo 476 - subgen. Pediomelum (Rydb.) Ockendon 450 Riedeliella Harms 7, 8, 16, 50, 227, 247, 315 - nelsonii Merr. 202
Podocarpium (Bentb.) YC. Yang & PH.Huang 441 Psoraleeae Lowe 5, 9, 19, 36, 52, 53, 361, 393, 394, 447 Rivasgodaya Esteve 295 Sesbania Adans. 9, 12, 19, 31, 53, 452, 453, 455
Podocytisus Boiss. & Heldr. 16, 284, 291, 292 Psoraleeae sensu Hutch. 299 Robinia L. 12, 19, 53, 467, 469, 470 - bispinosa (Jacq.) W.Wight 453
- caramanicus Boiss. & Heldr. 291 Psoralidium Rydb. 19, 447, 449 - hispida L. 470 - exaltata (Raf.) Cory 453
Podolobium R.Br. ex W.T.Aiton 17, 350, 351 Psorobatus Rydb. 302 - pseudoacacia L. 470 - grandiflora (L.) Pers. 453
Podolotus Royle 19, 53, 455 , 458 Psorodendron Rydb. 8, 303 Robinieae (Benth.) Hutch. 4, 5, 9, 19, 43, 53, 299, 307, 308, - punicea Benth. 453
Podopetalum F.Muell. 235 Psorothamnus Rydb. 16, 299, 303 452, 455, 456, 467 Sesbanieae (Rydb.) Hutch. 4, 5, 9, 19, 53, 452, 455 , 456,
Poecilanthe Benth. 7, 9, 16, 49, 253, 254, 367 - scoparius (A.Gray) Rydb. 303 Robinioid clade 5, 8, 9, 22, 53, 361 467
Poeppigia group 24, 127 Pterocarpus clade 5, 8, 50, 308, 309, 433 Robynsiophyton R.Wilczek 16, 50, 273, 281 Shuteria Wight & Arn. 18, 404, 406
Poeppigia C.Presl 6, 13, 31, 46, 111, 113, 114, 127, 128 Pterocarpus]acq. 12, 16, 34, 307, 308, 317, 320, 322, 323, Rothia Pers. 16, 50, 273, 281 Sideropsis Small 208
- procera C.Presl 113 324 Ruddia Yakovlev 235 Sindora Miq. 13, 34, 69, 83
Poeppigieae Britton & Rose 111 - angolensis DC. 322 Rudolphia Willd. 399 Sindoropsis ].Leonard 13, 83
Poinciana sensu Bail/. 154 - indicus Willd. 322 Rueppellia A.Rich. 329 - letestui (Pellegr.) ].Leonard 83
Poinciana L. 140, 142 - santalinoides L'Her. ex DC. 322 Rupertia J.W.Grimes 19, 447, 449 Sinodolichos Verde. 18, 419
Poincianella Britton & Rose 6, 14, 128, 142 - soyauxii Taub. 322 Ruprechtia CA.Mey. 33 Smirnowia Bunge 19, 476, 483
Poiretia Vent. 16, 312, 313 - tinctorius Welw. 322 Russellodendron Britton & Rose 138 - turkestana Bunge 483
- punctata Desv. 313 Pterodon Vogel 15, 250, 251 Smithia Aiton 17, 329, 330, 331
Poissonia Baill. 9, 19, 467, 469, 470, 471 Pterogyne group 6, 127, 129 Sabinea DC. 469 - conferta Sm. 330
Poitea Vent. 19, 42, 467, 469 Pterogyne Tu!. 14, 111, 127, 134, 150 Sakoanala R.Vig. 15, 238, 239 - elliotii Baker f. 330
Polhillia C.H.Stirt. 16, 50, 267, 273, 283, 284, 286, 287 - nitens Tu!. 134 - madagascariensis R.Vig. 238 - sensitiva Aiton 330
Polygalaceae 1, 3, 4, 5, 22, 45 Pterolobium R.Br. ex Wight & Arn. 14, 135, 136, 137 Salweenia Baker f. 15, 244 Soemmeringia Mart. 17, 308, 329, 330
Polystemonanthus Hanns 13, 34, 101 - lacerans (Roxb.) Wight & Arn. 137 - wardii Baker f. 244 Soja Moench 421
Pomaria Cav. 6, 14, 128, 141 - stellatum (Forssk.) Brenan 136, 137 Samanea alliance 195 Sophora group 227, 228
Pongamia Vent. 9, 383, 384 Pterospartum (Spacb) K.Kocb 295 Samanea (Benth.) Merr. 15, 48, 193, 210 Sophora L. 8, 15, 36, 49, 233, 242, 243, 260, 283
Pongamiopsis R.Vig. 17, 384 Pterygopodium Harms 74 - saman (Jacq.) Merr. 210 - davidii (Franch.) Skeels 243
Possira Aubl. 218 Ptycholobium Harms 9, 17, 367, 387 Saraca L. 13, 85, 86 - flavescens Aiton 243
Priestleya DC. 8, 270 Ptychosema Benth. 355, 358 Sarcodum Lour. 17, 371 - gibbosa (DC.) Yakovlev 242
- sect. Aneisotbea DC. 268 - trifoliolatum F.Muell. 358 Sarothamnus Wimm. 292 - japonica L. 233
- sect. Eisotbea DC. 270 · Pueraria DC. 18, 52, 417, 418, 419 Sartoria Boiss. & Heldr. 19, 495 - microphylla Aiton 243
- sect. Priestleya 270 - montana (Lour.) Merr. var. lobata (Willd.) Maesen & Sauvallella Rydb. 469 - pachycarpa CA.Mey. 242
Prioria clade 71 Almeida 418 Schefflerodendron Harms 17, 376 - secundijlora (Ortega) DC. 233
Prioria Griseb. 10, 13, 34, 42, 69, 74, 75, 76 - phaseoloides (Roxb.) Benth. 418 - usambarense Harms 376 - tetraptera J.F.Mill. 243
- copaifera Griseb. 75 Pultenaea Sm. 3, 17, 339, 343, 344, 345, 346, 347, 348, 349, Schizolobium Vogel 14, 127, 151 - tomentosa L. 243, 260
Priotropis Wight & Arn. 279 350, 351, 352 - parahyba (Veil.) S.F.Blake var. amazonicum (Ducke) - tonkinensis Gagnep. 243
Prosopidastrum Burkart 14, 163, 173, 174, 184 - selaginoides Hook/ 346 Barneby 151 - sens. strict. 36, 49, 227, 243
Prosopis group 164, 165, 184 - sect. Phyllota DC. 345 Schleinitzia Warb. ex Nevling & Niezgoda 14, 34, 174, 176, - sect. Calia (Teran & Berland.) Rudd 233
Prosopis L. 12, 14, 24, 47, 163, 172 Punjuba Britton & Rose 193, 205 175 - sect. Edwardsia (Salisb.) Taub. 49, 243
- juliflora (Sw.) DC. 172 Pycnospora R.Br. ex Wight & Arn. 18, 442 Schnella Raddi 57, 65 - sect. Styphnolobium (Schott) Yakovlev 233
- pallida (Humb. & Bonpl. ex Willd.) Kunth 172 Pynae11iodendron De Wild 100 Schotia Jacq. 13, 31, 46, 73, 86, 87, 93 Sophoreae Spreng. ex DC. 4, 5, 7, 8, 15, 49, 215, 227, 250,
Pseudarthria Wight & Arn. 18, 442 Pyranthus Du Puy & Labat 9, 17, 384 - afra (L.) Thunb. 73 263, 271, 279, 307, 480
- hookeri Wight & Arn. 442 - latifolia Jacq. 73 - sens. lat. 4, 7, 8, 216, 250, 253
Pseudeminia Verde. 18, 419, 420 Quillajaceae 4, 5, 45 Scbrammia Britton & Rose 141 - sens. strict. 4, 7, 8, 227, 253, 260, 263, 267, 273
Pseudoalbizzia Britton & Rose 211 Scbranckiastrum Hass/. 183 Sopropis Britton & Rose 172
Pseudoberlinia P.A.Duvign. 105 Racosperma Mart. 187, 188 Schrankia Willd. 183 Spartidium Pomel 8, 16, 50, 273, 275, 283
Pseudocadia Harms 230 Radiata Medik. 499, 503 Sciaplea Rauschert 120 - saharae (Cosson & Durieu) Pomel 275
Pseudocopaiva Britton & Wilson 81 Rafnia Thunb. 16, 50, 276 Sclerolobieae Benth. 127 Spartiinae Benth. 284
Pseudoentada Britton & Rose 182 - amplexicaulis (L.) Thunb. 276 Sclerolobium group 127, 128 Spartium L. 16, 283, 284, 294, 296

574 LEGUMES OF THE WORLD INDEX TO SCIENTIFIC NAMES 575

- junceum L. 296 - sect. Fistuloides 217 - himalaica (Baker) H.P.Tsui 479 - eivilia (L.) Willd. 506
Spartocytisus Webb & Berthe/. 292 Swartzieae DC. 5, 7, 15, 49, 215, 227 Ticanto Adans. 140 - faba L. 10, 506
Spathionema Taub. 18, 426 - sens. lat. 215, 250 Tipuana (Benth.) Benth. 12, 16, 34, 42, 50, 319, 320, 322 - narbonensis L. 506
Spatholobus Hassk. 18, 52, 407, 408 - sens. strict. 5, 7, 215, 216, 228 - tipu (Benth.) Kuntze 320 - sativa L. 506
Sphaeridiophorum Desv. 364 Sweetia Spreng 5, 7, 8, 15, 234 Toluifera L. 232 - villosa Roth 506
Sphaerolobium Sm. 17, 340, 341, 355 - fruticosa Spreng. 234 Torresea Allemao 220 Vicieae (Bronn) DC. 10, 505
Sphaerophysa DC. 19, 484 Sylitra E. Mey. 387 Toubaoute Aubrev. & Pellegr. 103 Vicioid clade 475, 476, 477, 489, 496, 499, 505
- salsula (Pall.) DC. 484 Sylvichadsia Du Puy & Labat 9, 17, 375, 415 Tounatea Au.bl. 218 Vigna Savi 3, 9, 18, 31, 42, 52, 393, 426, 427
Sphenostylis E.Mey. 18, 422 Sympetalandra Stapf 14, 159 Tounateeae Baill. 215 - aconitifolia Qacq.) Marechal 427
- stenocarpa (Hochst. ex A. Rich.) Hanns 422 - schmutzii Steenis 159 Touniaya gossweileri (Baker f) Schmitz 62 - angularis (Wille!.) Ohwi & H.Ohashi 427
Sphinctospermum Rose 19, 467, 469, 472 Syrmatium Vogel 9, 19, 53, 460, 462, 463 Tracbylobiwn Hayne 80 - mungo (L.) Hepper 427
Sphinga Bameby & ].WGrimes 7, 15, 193, 207 Tragacantha Mill. 481 - radiata (L.) R.Wilczek 427
Spirotropis Tul. 15, 248 Tachigali group 127, 128, 129 trans-oceanic disjunctions 42, 43, 44, 309 - subterranea (L.) Verde. 427
Spongiocarpella Yakovlev & N.Ulziykh. 19, 478 Tachigali Aubl. 6, 14, 47, 127, 150, 151, 160 Trichocyamos Yakovlev 235 - umbellata (Thunb.) Owhi & H.Ohashi 427
Stachyothyrsus Harms 14, 159 - tinctoria (Benth.) Zarucchi & Herend. 150 Trifidacanthus Merr. 18, 440 - unguiculata (L.) Walp . 10, 427
Stahlia Bello 14, 145 Tachigalia Auhl. 150 Trifolieae (Bronn) Encl!. 5, 10, 19, 36, 53, 54, 455, 475, 496, - subgen. Sigmoidotropis (Piper) Verde. 427
- monosperma (Tul.) Urb. 145 Tadehagi H.Ohashi 18, 438, 439 499, 505 Vignopsis De Wild . 414
Stauracanthus Link 16, 284, 295, 296 Talbotiella Baker f. 13, 86 - subtribe Ti~foliinae Bronn 499 Viguieranthus Villiers 7, 15, 193, 197, 199, 207
Steinbachiella Harms 333 - gentii Hutch. & Greenway 86 Trifolium L. 3, 10, 19, 54, 499, 501, 505 Viminaria Sm. 17, 341, 342, 355
stem clade 45 Tamarindus L. 13, 69, 90 - pratense L. 501 - juncea (Schrad.) Hoffmanns. 342
Stemonocoleus Harms 13, 34, 79 - indica L. 90 - repens L. 501 Virgilia Poir. 16, 271
- micranthus Harms 79 Tara Molina 6, 14, 128, 138, 139 Trigonella L. 19, 499, 503 Voandzeia Tboua1·s 427
Stenoclrepanum Harms 14, 146 - spinosa (Molina) Britton & Rose 138 - foem nn-graecum L. 503 Vouacapoua Aubl. 14, 127, 128, 148, 161
Stirtonanthus B.-E.Van Wyk & A.L.Schutte 8, 16, 269 Taralea Aubl. 15, 250, 251 Trigonelleae Schulze 499 - americana Aubl. 161
Stirtonia B.-E. Van Wyk & A.I.Schutte 269 - oppositifolia Aubl. 251 Trioptolemea Mart. ex Bentb. 327
Stizolobium PBrowne 405 Taverniera DC. 19, 489, 494 Tripeta!anthus A.Cbev. 88 Wagatea Dalzell 147
Stonesiella Crisp & P.H.Weston 8, 17, 346, 347 Telestria Raf 61 Triplisomeris Aubrev. & Pe/legr. 102 Wajira Thulin 18, 42, 52, 422
Storckiella Seem. 14, 111, 115, 116 Teline Medik. 295 Tripodion Medik. 19, 459, 465 - graharniana (Wight & Arn.) Thulin & Lavin 422
- australiensis ].I-I.Ross & B.Hyland 116, 117 Temperate biome 23, 36, 49, 53, 54 Trischidium Tu!. 7, 15, 215, 218, 219 Wallaceodendron Koord. 15, 204
Stracheya Benth. 10, 489, 493 Templetonia group 7, 8, 253 Tylosema (Schweinf.) Torre & Hille . 13, 45, 58, 62, 63 - celebicum Koord. 20
Streblorrhiza End!. 19, 487 Templetonia R.Br. ex WT.Aiton 16, 253, 256, 257, 355 - esculentum (Burch.) A.Schreib. 63 Weberbauerella Ulbr. 17, 50, 308, 335
- speciosa End!. 487 - retusa (Vent.) R.Br. 256 Welwitschia Hook.f. 24
Strombocaipa (Benth.) Engelm. & A. Gray 172 Tephrosia group 368 Uittienia Steenis 6, 14, 111, 119, 120 Wenderothia Schltdl. 396
Strongylodon Vogel 12, 18, 415 Tephrosia Pers. 3, 17, 31, 52, 367, 379, 385, 386, 387 Uleanthus Harms 15, 248 Westia Vahl 103
- macrobotrys A.Gray 415 - candida (Roxb.) DC. 386 Ulex L. 16, 295, 296 Whitfordiodendron Elmer 370
- perrieri R. Vig . 415 - purpurea (L.) Pers. 386 - e uropae us L. 296 Wiborgia Thunb. 16, 50, 273, 275, 276
Strophostyles Elliott 18, 429, 430 - virginiana (L.) Pers . 386 Uliceae Webb 283 Willardia Rose 9, 380
Stryphnodendron Mart. 14, 180, 181, 182, 183 - vogelii Hook.f. 386 Umtiza clacle 4, 5, 6, 24, 31, 42, 47, 111, 127, 128, 129 Willdampia A .S. Georp,e 485
- adstringens (Mart.) Coville 182 - sect. Mundulea DC. 385 Umtiza Sim 4, 6, 14, 24, 42, 47, 69, 111 , 127, 131, 133 Wisteria Nutt. 5, 9, 17, 53, 372, 475, 478
- microstachyum Poepp. & End!. 182 - sect. Requienia (DC.) Benth. 387 - listerana Sim 131
Stuhlmannia Taub. 6, 14, 127, 135, 136 Tephrosieae (Benth.) Hutch. 367 Uraria Desv. 18, 443 Xanthobiychis Galushko 494
- moavi Taub. 136 Teramnus P.Browne 18, 420, 421 - picta Qacq.) DC. 443 Xanthocercis Bail!. 7, 15, 220 , 230
Stylobasium Desf. 45 Terua Stand/. & Ff. Herm. 380 Urariopsis Schindl. 443 - zambesiaca (Baker) Dumaz-le-Grand 230
Stylosanthes Sw. 10, 16, 31, 50, 308, 325, 326 Tessmannia Hanns 13, 34, 83 Uribea Dugand & Romero 15, 222, 234 Xerocladia Harv. 14, 24, 173
- erecta P.Beauv 325 Tethys Seaway 23, 24, 31, 34, 40, 41, 43, 44, 51 - tamarindoides Dugand & Romero 234 - viridirarnis (Burch.) Taub. 173
- fruticosa (Retz.) Alston 325 Tetraberlinia (Harms) Hauman 13, 107, 106, 108 Urodon Turcz. 17, 345, 346 Xeroderris Roberty 17, 374
- guianensis (Aubl.) Sw. 325 Tetragonolobus Scop. 9, 19, 456, 464 Xiphotheca Eckl. & Zeyh. 8, 16, 268
- humilis Kunth 325 - purpureus Moench 464 Vacbellia Wight & Arn. 187, 188 Xylia group 6, 163
- viscosa Sw. 325 Tetrapleura Benth. 14, 167 Vandasia Domin 407 Xylia Benth. 14, 163, 168
Styphnolobium Schott 5, 15, 31, 40, 49, 227, 233, 243, 260, - tetraptera (Schum. & Thonn.) Taub. 167 Vandasina Rauschert 18, 407 - dolabriformis Bentb . 168
264 Tetrapterocarpon Humbert 14, 24, 42, 47, 111 , 127, 131, Vatairea Aubl. 8, 16, 234, 307, 310 - :>..'Ylocarpa (Roxb.) Taub. 168
- japonicurn (L.) Schott 233 132, 133 - macrocarpa Ducke 310
Succulent Biome 21, 23, 24, 41, 42, 43, 44, 45, 46, 47, 48, - geayi Humbert 132 Vataireoid clade 5, 7, 8, 49, 215, 228, 307, 309 Yucaratonia Burkart 468
49, 52, 53, 54 Teyleria Backer 18, 416, 417, 418 Vataireopsis Ducke 8, 16, 234, 307, 310
Sulla Medik. 10, 19, 489, 493 Thailentadopsis Kosterm. 7, 15, 199, 207 - araroba (Aguiar) Ducke 310 Zapoteca H.M.Hern . 15, 31, 193, 196
- coronaria (L.) Medik. 493 Thermopsideae Yakovlev 5, 8, 16, 36, 49, 228, 253, 263, Vatovaea Chiov. 18, 52, 426 Zenia Chun 14, 111, 115, 118, 119, 120
supertree 5, 21, 23, 24, 45 267, 283 - pseudolablab (Harms) ].B.Gillett 426 - insignis Chun 119
Suriana maritima L. 45 Thermopsis R.Br. ex WT.Aiton 16, 263, 265, 283 Vaughania S.Moore 8, 17, 361, 363 Zenkerella Taub. 13, 93
Surianaceae 1, 4, 5, 36, 45 - rhombifolia (Pursh) Richardson var. montana (Nutt.) Vavilovia Al.Fed. 19, 505, 508 Zollernia Wied-Neuw. & Nees 15, 222, 234
Sutherlandia R.Br. ex WT.Aiton 19, 475, 476, 485 Isley 265 Verdcourtia R. Wilczek 425 Zornia ].F.Gmel. 16, 308, 312, 313, 325
- frutescens (L.) R. Br. 485 Thinicola ].H.Ross 7, 16, 253, 256, 258 Vermifruxj.B.Gillett 9, 460, 461 Zuccagnia Cav. 14, 145, 146, 147
Swainsona Salisb. 10, 19, 475, 476, 485, 486 - incana Q.H. Ross) J.H.Ross 258 Vexihia Raf 243 Zygia P.Browne 15, 34, 193, 200, 201, 202
- formosa CG.Don) Joy Thomps. 485 'fl1ornbera Rydb. 299, 304 Vexi/lifera Ducke 231 - longifolia (Willd.) Britton & Rose 201
Swartzia Schreb. 3, 7, 15, 34, 42, 49, 215, 218, 219 'fl1ylacanthus Tu/. 6, 105 vicariance 43, 44, 46, 53 Zygocarpum Thulin & Lavin 8, 17, 42, 308, 334
- sect. Cyathostegia Benth. 219 Tibetia (Ali) H.P.Tsui 10, 19, 476, 479 Vida L. 1, 3, 10, 19, 54, 505, 506

576 LEGUMES OF THE WORLD INDEX TO SCIENTIFIC NAMES 577