274

13 .
l~ .

I :' .

10.
17.
18.
In.

2,n.
2 1.
~..~.
~1.

2., .

.J epson, F. P. Depart. ngric, l<' ij i, Pall/pillet r,'o. ](j .

p. 3, Suva 1915.
'Kern er I"on Maurilaul1, A- The Na.lmuZ History of P7,MltS.
Rnglish Eel . Yol. n , 1.onelol1 1895. pp. 9(i & 99.
I,"uth, B. fla11 dbllch de'r lilldcnbiolo.r;ie. BIl. III, Lei p'l,ig.
19M, pp. 59 and ') 8 .
1\1IJlI'er , Jlsie M. , Imperfec-t Coronl1t ~-./ o'l/./"Ii . .\" e'/l) J"017t; Hal .
(:(trf/. .. 11 , 1901 , PI'. 69-7 1.
I,oyola, J. I. rl e. ('jili 'ILI'as I lIi{;ia'lI(Js, Orlim , Cioa, 1806.
I'etch, 'I'. 'l'he Flowe r 0'1" ihe Cocon ut - Trop . .'I.r;rirll/.I1I.1i.,/ .
XU , 19 1B. pp. ~4!) - -15,.5 .
Ilnngachari. K.- JI .l;a:n1l1J7 of NlclI1en/al'l/ lio/II/I,I/ /01' Iidin.
2 nd ErlitOo Marlra s. p. ~ 1 5.
~a , .'1. ~<f. de . 0 ('ol/1U'iro, :\ov>I-Uoa . 1 ~98 .
Sampson . 1-1. COo The ('OCOIl1!/ 1'01111. I,ondon, '192:, .
Smith. If. fl. & F. A. fl . 1II1'e, .('O/,O/llUS: fhe ('OIiSO/.~ of II, I'
ga.,I, 2nd E(l . London, 191,3 . .
\'an (ler I\olrl . Quoted h~' I-hlll ger , PI'. ?,:L
\\'orsd ell , \\'. COo {'rili cip/es
{'lal1/-T I'I'II/olo(J!J. \'01. I I,
Londol1 191 G, pp. 191-102.

at

BRANCHED COCONUT PALMS AND THEIR

FERTILITY .
Apropos of the note on the Fel'till:tY, of Bmnchcri ('oco'lltil
Pltllns by Mr. Burkill, published on page 1-2 of Vol. III of t hi s
Rltllelin, the following may not he without in tere. t to reaelers.
Xorma ll), one cocon ut fruit gives ri se to . one shoot an el this
in its turn to one stem. It is not infr equently, howevel', thnt one
meets with wiele deviations from this normal p henomen on. A nu t,
for in tan ce, may on germination, give ri se to more than one shoot,
each arising from a separate carpel in th e nu t. Th e writer has 110t
~onle across an in stance wher e a nut had more tlian two fer tile
cHI'pel s: but it mu st be r emembered that t he coconut has a triloPlllar ovary whereill normally two of the locules become abortive.
('a ses, the;efore. ma.' occur ;vhere all t hree ca rpels ma.,I' be stimulated i'o become fertile a.' in som e specieR Tela1'ed to Cocos 1Mwifem.
POI'heR reports of. "a nut w ith t hree cell s separated hI' leathery
"',11k" 8aml'son (I") appears to ha ve seen cases where t he septa
separntill g the ovules were hard amI not lea.thery. 'POlbes. in his
ll1'tic-le above referred t o write : " 1 ha ve seen al 0 nuts with cell~
l'angilll( from 1'0111' to eight f;lld ten . r ~end VOll a rough ouLline
sketrh 01' a tree whir-h ha s eome up from a nut of fOlll'teell celk
all of \\'h ieh germinating. prorlu('in,!); a h'pe with fomtecn stems

275
u ll.ited at the uase." If Forbes' conclu sions as to the occurrence 0-1
more than t hree cells in a nut ar e con'ect, then the ph enomenon
C'oncerned might ue a case of vegeta tive multiplication of car pels.
I f, all the othe" hand , his conclnsiol1 s wer e based mer ely on the
Illlm h<:r of shoots t hat. appea recl on germination of a nut, then
(hese alterna tI ves arc possihle (l ) t hat the numerous shoots H e
due either to t he occurrence of polyembryony as in mango, ci t rus,
oni01l , etc., or (2) to t he l'asc iation of the plumule, or (3) to the
m ono pod ia I hranc hin g wi th very mu ch abbrel,iated axis. All these
" hCIJ Ome nH Hre loosch descr ibec] as " llTancbillg " a.lthough only
in t he last two cases t he tr ue phenom enon of branching occurs.
l " Hli other cascs, thc shoots Hre iListill ct incli vicluals, thongh due to
" dosc" acl hesion among themsehes t hey may appear to star t from
si ngle point. Th e re fer ences ahout t he OCf:Ul'l'ence of polyembryon y
01' mOl'e thall t hree (arpels in eoc'onu ts are IIOt deal' . and , th er efor e,
Ihc "uhj ed ci cn","d " flll' t he,' investigHtion s.
Th e I'ill1li f"ic:ation of' t he ma in shoot ill coconuts .i s not r e. ,,(Tictecl to the yOllllg age only. Th el'e are n u merou~ instances wher e
'l cl ult co(on ut pHIlliS hHve brHn checl . Th e ph enomenoll conrerncd
ill I l'illlehillg l'almR has hee n cl iseussecl b\" Mr. Riclley, (") the
hlle [)ireetor 0(' t he Botani c Clarclens. Singapore. H e appears to
hold t hc \"iew tha t truc dichotomy never occurs in palms. If' thi s
vie w i, correct then the phenomenon of fasciation also never occur~
in i'a l m ~, sin ce fH sc i"Lion and di chotom y are esse ntially one and
t he sa me ph enomenon wit h the (li"ffer ence that, whi le i n the latter
t here ,lppeal'S on l,)' two 'boot, in the former they are always more
than hro . Iiut t he riew t hH t t ru e dichotomy never occurs ill palms
iR not absolutel." t nl e. ,Schou te (IT) has observed the phenomenon
of f'ascia tion in stems of some abnormal palm, ancl t hat of "true
dichotom~' of t he stem, as a normal fent7/'e. in Hypha,ene, an
Afriean palm . Tt (u chotomy ) arose a follow.: 'The apical
m er istem ceased to grow, and in place of it, at equal lateral
c] i. tflnces t herefr om, two llew 011es appeared.' From t his it i~
obvious t ha t ther e was no division of t he apical mer istem into two,
ancl yet tr ue di chotomy is pre. en t : because the two latera l bran ch e~
di c] not ari se owing to iujUl'y to the stem-apex ; because thev
are not ax illary to any leaves; and heca nse an 'angle-leaf' is
pr esent opposite the fork as is the case in clichotom.ni ng stems of
('r~'ptogam s .
Moreover, the two bran ches bore lateral bud s in the
axil s of every leaf'. Velenovsky (17) has also establi shed by per"anal oh ~er vati oll thflt tru e dichotomy occurs in C'hamnerlol'ecl
Jtror/"'iana ." IITor. dell (21) cite. a case wh er e the plumule of the
pa lm P.i?1(l'IIgfl 'Il1acu.lrtia in t he Kew Garclens, was branehecl clOWll
to the extr eme hase, the bi'Huches being subtendecl hy a common
hasal sheat.b. Th e ease wa one where it was not eas\" to cleeic]e
whet.her t.he phenom enon con cern ecl wa s a true di choton;y or not.
Th er e is another p hen omenon which might be confuserl
with t.he tr ue branch ing. On the ;iloakhallv Planta t ion . AI.-yah.
Burma, T was shown plants which wer e consider ed as cases of bu cl"ot and t hese plants had tl1Pil' ee nh'al leHves cl ecaving mlcl in somc

276
caRes gJVw g cl"i sagn)'lble sm ell. 'l'hough a specia l wateh was not
kept on th e plan tat ion 1'0 1' t he di sease ..),e1 it IV,I S customarv to
kill and bUnl nil su ch p lan ts t ha t cam e to t he no ti ce of t he manager
a nd sp ray t he lJ ei gh bo urin g OJ]es wit h copper s ulph ate. rt was
sa id t ha t t he disease wa s never see n in t he Ii}sta te on p[,ln ts older
t han te n 01' t weh e vea l'S a nd t hat t he cases wer e alw,lVs spoTud ie,
the aU-ac keel tl'pes ll (-lre J' oCl.: tllT iJl g ill groups 01' hecu'-ill g

HIl.\r

defill ij'c

relation to t he old ('a"es . All t hi s inl'OI"l11 ation cOllpl ed wit h nn'

obse rvation of t he I'act t ha t n um erou s plants show in g s imilar
o\'mp toIllS, eseapcd th e Ho t ice of t he Ill all in cha rge of t he phllltaho n and thai' t he disease neit her killed t hese nn treated pl ant,
nor d id it see m to spreael i" rolll these to t he neighho urin g trees
leel m e to the eon cl usion t ha t t he t ro uble ill qu estion wa s ot her
t ha n t he ' u spected bn<1-mt, genuin e cases of wbi ch (li se'lse I n ever
mot on t he plan tat ion during m y si x mo nths stav t her e. F urt hc,'
obser mtions sho,reel t hat t,he i"'ltholog icaj, conditio n ill most ca:e;;.
(o nId he tra c-ed as the c-o nseqll e ll c-e, o f tJ~njllrv to th e pa lm by the
l1111 ch-prevnlell t rh illor'eros eeeties. ""hen t he injury to t he cabbage
reached th e apex, t he latter was stimul nteCl to change its di r ection
01' g rowth lI"ith th e resul t th at t he young leaves produ cerl prev iou f;
to th.i;; ch ange II bt e sthl \ tel ,0 death tin d st,uted' to decti). X ow the
idea th at this change of direction of: g r owin g apex could be mistaken
tor t rue branching would n ever h ave occurred to m e were it not
for th e fact, th at, through th e cri ticism s by r etch and Gadd, m v
attention was drawn to ~ su cb views ellter t ain erl h" Shar ples and
L ambou rll e. 'I'h e form er wTiters h a "e discu ssed the m atter at
;;llffi'cien t len gth ill t he A nnals of 80tr1'1IY, YoL ;17, Jull' 1923, pp.
445-4~'O, aml shown that the cases which a,r e consider ed as genuin e
ea sei< of lateral hran chin g by the latter (virl e 11. 11110ls, 1'01. 36, .Jan.
'1 92~) fire most probably t he on es wher e a ch ange 'in th e directio n
of gr owth of t he a pica l burl f rom ver t.ical to hor izo nta l if; in vol ved.
' l'her e is vet anoth er phenomen on commonlv call ed ." bran ching ", though strictly spea king it is a case wh ich ough t to go und er
t he ti tle of chloran th), or proli feration. Tt con sists ill t he m etamorphos is of an. inflorescen ce shoot a.nd u sually th e tr ees start
pr odu cing th ese abllOTmaI stru r:tures when it is of t h e age to
produ ce normal inflorescen ces an rl i t continu es to produce . them
ti ll its cl eath. Ridley . ( ]O) P etch (") Tvengnr ('0) and Burkill
(' ) ha've discu ssed I;hi s abnoTll1 alitv in some detai.l\If From t he
peru sa l of t h e li te ra t ure consulted anel fr om mv own observation s
T am len to con clud e th at ~h ese " hulbil s ". 1'0" so th el' aTe term erl
hI' Rid ley . ar e n ever pers iste nt, t hough FOl'bes, fr01)1 hi s ohser vations
marl e in the ( oros K eeling T ~laJ1(ls, wa s inclill ed to heli eve th em so.
:\"0 rloubt th esp ou tgrowths hH ve u Slla lh~ a Illu ch longer li fe than
lIo rmal inflorescen ces. or t he leaves of t he trees. Tn ol'le case T
ohser ven th ese abnorm al infl Ol'escenres g rew for mOTe than five
."e>l r s. Rut eve n if th eir l ife was I)]"olongerl for mOre veaTS T elo
n ot think th ey would g ive ri se to th e peTman ent ly bran ched palm.
at lenst th e ph enomrnon co nce rn ed woulrl he ver I' different f rom
that of fhe ord in Hl'y "Hl11 iAcai:ion in palms. (cf. Burkill's pa.per ).

277
The peduncle th at bears these abnormal structmes is essentially
o'E th e type of the normal inflorescences : clean, and fibrous for a
:foot or less hom the base, while t he structure of the branches
from the poin t of fork is qf the type of the stem. 'rh e p eduncle
does n ot increase in size much more than the peduncle of normal
initoresccn ces, and , lik e t he peduncle of normal inflorescen ces, has
a tendency to form absciss layer and separate form th e main axis.
S ome of these bulbils bear minute inflorescen ces con sisting of tiny
male floret s and no femal e. In the cases obser ved by me these
itower s were born e by secondary shoots produced on t he bulbils.
The bulbils do not show any t end en cy to form roots in urtir.c-ial
medi a (r. ) & ('0).
Dr. Pulney Andy (' ) has descri bed a mOllstrosit.y which
con sisted in th e phyHody of t he greater portion of the female
itower s in the spadix, and not of the whole inflorescence itself.
H ere too " th e flower-bud cannot, by such metamorphosis, g.ive
ri se to r amificati"on in a palm without a tru e axillary leaf-bud."
Fertility.
In the above cli scussion i t becomes clear that the term
"branching" is loosely applied to various phenom ena, includin g
some that cannot be classed as bran ching. And it is possible that
the conception that the true branched palms are permanently
infertile has orginated from the con fusion which exists as to the
meaning of t he term " branched " coconut 1Jalms, that is, by tran sference to other s of th e con ception derived from some special case
or vh enomena.
Otherwi se the prevalence of the idea that th e
bran ched palms are perman en tly unproductive is r eally unfound ed.
I have seen a coconut "Inanching" at the base bearing
fruits ; and ther e are num erous refer ences which show that bran ched
coconuts are not in fertile. Morri s, (11) for instance, reports a
fruit in the K ew Museum hom a bran chp,d coconu t tree of th e
Fiji I sland s. According to Scott, ( IS) there was a t.ree in the
vicinity of C'alcutta wi th five fruit-bearing br an ches. H enry (8)
makes men tion of a coconut tree in the Marquezas I slands, which
at the age of eighteen branched in to two, both h eacIs starting to beaT
fruit s after two years from the occurrence of the f orking. Among
these cases th ere are at least two references which are of inter est
to Malaya. Th e front ispiece of the t reatise on coconut by Munro
and Brown ('" ) is of a coconut tree br anching out near the t o.p
into fi ve di stin ct stems, each of which is carrying fruit. 'l'he
"econd is of a coconut t ree in ' P enang which hall six bTan ches, each
of which wer e " loa ded with nuts." (3) Apart f rom these, the
Consois of the Ertst by Smi th and P ape (,0) and Hunger's (Jocos
nmcifem ( 0). have photographs of branched coconut palms bearing
frui ts. In f act consid ering th e presen ce of the photographs in the
above quoted standard treatises on coconu t, one cannot un der stand
the reputed prevalence in Malaya of t he view that bran ched coconut
palms aTe st erile,

278
By thi s I do not wi sh to be lUl derstood to say that all t ruly
bran ched coconut palms are productive. Cases may occur wheTe
bran ched coconuts may not start yielding for a very long period ;
but in such cases, before one can draw conclusions, it has to be ascertained whether t he unproductiveness is due to want of proper
caTe, to the fact that the t Tee has not yet fin ished bran.ching, or to
some other cause.
A veTY inteTestin g })oint in connection with t hese abnormal
palms is to Imow how then progeny behave and it is very astonishing
that n obody has yet tTied t o solve this question. It is not yet
known, fOT in stance, whether seeds of these palms Will keep true
to the abnoTmal chaTactei' 01' Teturn to the character o:f the parents
to these abnormal ones ; whether all of the seeds will do so 01: only,
a paTt of t hem, and. how large a part. I n carrying investigations
in these lin es care has to be taken t o preven t cross-fer tilisation
with the pollen from other plants. Tb is ",ill probably he an easy
matter with tbe dwarf coconuts in :NIal aya wher e normally the
male an d female flower s r ipen ill a manner as to cnSUTe selfpollination if protected by a bag fTom foreign pollen. If, however,
t he infl orescen ces of t hese abllorm al palms are protanch ous, then
the female flowers will have t o be artifi cially fertili sed wi th t he
pollen f rom the male flowers in the same tree. The brallches pf
palms su bject to heavy w:inds and, therefore, liable to be broken ,
may be induced to form aerial roots on stems by marcottage and
then detached and nlanted in a place convenient to carry on with
t he observations. The crown of a palm t hus detached has been
observed by me to geow when planted out amI this practice, it
appears, is som etimes resorted to by the gardeners in Malabar,
I n elia. (0)
I n cases wher e the infloresce)) ces foliate I have not come
across a sillgle r efeTen ce stating that such tTees or their bulbiis have
borne fruits. In th e case describecl bv Dr. Pulne~' Andy the tree
does not pTodu ce a pCJfect fru it. An d probably these aTe the only
two' phenom ena of the ones described above which are in fertile.
These cases however should n ot be confused with the ones where
coconut fru its , in stead of 'falling when matUTe and dry, remain on
the stalk aud under certain con ditions germinate on the tree.
GuPpy (') saw a case where a. seedli ng thu s germinated was about
eigh te~n in ches high.
C. X. FURTADO.
Bibl iography.
ANDY, P . S.- On Branched P alms in Southern Tndia.T1'(!lls. Linn S oc. XXVI, (1870) pp. 661-662.
See alRo ' J. Shor t.t's arti cle on the same subject in t he
JOU1. Linn. S oc. Botany, Vol XI, ( 1871) pp. 14-17.
2. BURKILL, I . J-f. - An A l moTm ~lity in the Coconut Palm.J01t1. R . .As;nt1c Soc., Stmits' 131'. No. &9, (1 915) p. 15.
3. Ceylon Obse1vc r. ~ ov. 7, ] 9lM- Quoted in A.gric. IV ews,
Barbados,
T, p. 62,
1.

'11

279
Nucifera-Jo~,'r ,

Botany,

4.

l"OHJJ I':S, H. O.-Notes on the Cocos

XVII, (1879) pp. 193-194.

5.

FUJt1'ADO, C. X . and BAlUlli'J:O, P. C.- An Abnormality of

Coconut Inflorescence-The Poona l1glic. College Magazine, XII, (19 ) No. 2, p. 81-82.

6.

FUH1'ADO, C. X.-Vegetative Reproduction in CoconutsTh e POO'I/ (' 11 g'ric. Coli. Mag . XV, 1922, pp. 144-146.

7.

Guppy, IT. B .- Obse'rv(,tions of u Natu'm list in the Pacific,

London, 1906, Vol II, p. 472.

8. HENRY, ('.- l ntelm,t. R ev , of S ci. (md Pmct. of Agl'im,ltu're,

Rome, (1920), No. 1088.
9. H UNGEH, F. W. 1'.- Cocos m,cij'era., Amsterdam, 1920, p. 218,
pI. LIX.
10.

IYENGAR, M. O. P .-Note on a Bulbiferous Coconut Tree

from M:alabar-Jowr. Ind'ian Botany, Vol. III, (1923)
pp. 2 9-291.

11.

MOlUll S, D.-On the Phenomenon Concerned in the Production of Forked and Branched Palms-Joul. Linn. Soc.
Bot. XXIX, ( 1892) pp. 281-298.

12.

MUNHO, Jl.. IV. an cl BllOIV N. JJ. C.-A. Pmctic(,l

Coco nut Planting, London , 1916.

G~,ide

to

13. l)],1'c]I", T.-Ahnormalities of the Coconut Palm-Ann. Roy.

Bot. Gwrd. Pcmdeniya" Vol VI, (191 5-17) pp. 25-29.
14.

PETCH '1'. and G,WD, C. H .- The Heplacement of the

'ferminal Bud ill the Coconut Palm. Annuls of Botany,
XXXVII, 1923, pp. 445-450.

15.

RIDLEY, H . N .-Branching ill Palms-Annals of Botuny,

Vol XXI, (1907) , .pp. 415-See also WorsdelFs Teratology, Vol 1. pp. ' 10,9-111.

16.

SAMp ON, H, C.- T he COCO'n-l,t Fulm : The Science und

Pmctice of C'oco n~, t C~,ltivation, London, 1923. pp.

17.

~()UOUTJo:

18.

SCOTT J. - (]wrden e'/"S' Ch1-onicle, Vol. L, pt. I (1874) p. 118

19.

SUAHPLES, A., and LAMBOUlu'lE, J.-Observations in Malaya

on Bud-rot of Cocouuts. Anna,ls of Botuny, . XXXVI,
1922, pp. 55-70.

20.

SMl1'lf, U. H. and P APE, F . A. G.-Coconuts: The Consols

of the East, London, 1913, p. 424.

21.

WonsDleLL, I'V. C.- '1' he Principles of Plant-Teratology, Vol.

I, London, 1915, pp, 109-111.

et V.l ': LE K OV~Ky- Quoted by WOIlSDELL, 01). Cit.

pp. 85 et 90.