Flora

Organization for Flora Neotropica
Pinus (Pinaceae)
Author(s): Aljos Farjon and Brian T. Styles
Source: Flora Neotropica, Vol. 75, Pinus (Pinaceae) (Nov. 21, 1997), pp. 1-291
Published by: New York Botanical Garden Press on behalf of Organization for Flora
Neotropica
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Brian T. Styles
26
Sepember1934 -27 Jue 199
FLORA
NEOTROPICA
MONOGRAPH75
PINUS (PINACEAE)
ALJOS FARJONAND BRIAN T. STYLES
8^^
1
PIC OP
CANCr
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FLORA(
NEOTROPICA
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r11PIC
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Pinus (Pinaceae)
Flora Neotropica
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The New YorkBotanicalGarden
Ariane L. Peixoto, Presidlent,UniversidadFederal Ruraldo Rio de Janeiro,Caixa Postal 74852, Seropedica, 23851 Itaguaf,Rio de Janeiro, Brazil (1996).
Stephan Beck, Vice-President,HerbarioNacional de Bolivia, Casilla 10077, La Paz, Bolivia
(1996).
Enrique Forero, Executive Director,Ex Qfficio, Institute de Ciencias Naturales, Universidad Nacional de
Colombia, ApartadoAereo 7495, Bogota, Colombia.
Renee Fortunato, Herbario,Institute do Recursos Biol6gicos, I.N.T.A., 1712 Castelar, Buenos Aires, Argentina (1999).
Paul J. M. Maas, Treasurer,Instituteof Systematic Botany, Heidelberglaan2, P.O. Box 80.102, 3508 TC
Utrecht, Netherlands(1999).
Ghillean T. Prance, Royal Botanic Gardens,Kew, Richmond,SurreyTW9 3AB, United Kingdom (1996).
Laurence Skog, Secretarn,U.S. National Herbarium,Dept. of Botany, NHB-166, Smithsonian Institution,
Washington,D.C. 20560, USA (1996).
Rodolphe Spichiger, Conservatoireet Jardinbotaniquesde la Ville de Geneve, Case postale 60, CH-1292
Chambesy/Geneve,Switzerland(1996).
Wm. Wayt Thomas, The New York Botanical Garden,Bronx, New York 10458, USA (1999).
Paulo G. Windisch, UniversidadeEstadualPaulista(UNESP), Depto. de Botanica,Caixa Postal 136, 15001
S. Jose do Rio Preto, Sao Paulo, Brazil (1999).
Staff Committee
EnriqueForero,Chairman
Paulo G. Windisch, Ex Officio
Board Representativeto UNESCO

Alicia Lourteig,Ex Officio
EditorialCommittee
James L. Luteyn, Editor
S. Rob Gradstein,Editor
Claudio Delgadillo M. (1996-2001)
LaurenceJ. Dorr(1996-2001)
Jose R. Pirani(1996-2001)
MariaMercedesArbo (1993-1998)
RodolpheSpichiger(1993-1998)
Olga Yano (1993-1998)
ORGANIZATION FOR FLORA NEOTROPICA
Founded and Conductedunderthe Auspices of UNESCO
The Scientific PublicationsProgramat The New York Botanical Garden

thanksthe following donors for their generous supportof this publication:
MetropolitanLife Foundation
The TinkerFoundationIncorporated
PINUS (PINACEAE)
ALJOS FARJON AND BRIAN T. STYLES
With Accounts of Wood Anatomy by IAN D. GOURLAY,

Pollen Morphology by MARIEH. KURMANN,
and Monoterpenes by JACQUELINE
S. BIRKS
CONTENTS
Abstract/Resum n ...........................................................................................................................................................
Preface (by Ghillean T Prance) ............................................................................................
Introduction ....................................................................................................................................................................
Historical Survey ............................................................................................................................................................
Morphologyand Anatomy .............................................................................................
Wood Anatomy (by an D. Gourla ) ............................................................................................................................
.......................................................................................................................................................................
KaryologyKaryology
.............................................
Pollen Morphology(by Marie H. Kurmann)........................................
The Role of Monoterpenesin the Taxonomyof Mexican and MesoamericanPines (by Jacqueline S. Birks) .........
...........................................................................
ReproductiveBiology ....................
Phylogeny and Evolution ............... ...............................................................................
Distributionand Ecology .............. .................................................................................
........................................
Uses and Conservation................ ........................................
Treatment.....................................................................................................................................................
Systematic
TreatmentSystematic
Subgenus
Pinus ..........................................................................................................................................................
PinuSubgenus
.................................................................................
Subgenus Strobus .............
Doubtful Names and ExcludedTaxa ........................................
...........................................
Acknowledgements ........................................................................................................................................................
Cited ..............................................................................................................................................................
Literature
CitedLiterature
NumericalList of Taxa ..................
............................................................................
ExsiccataeList
List of Exsiccatae ...........................................................................................................................................................
Index of Local Names ................ ................................................................................
Index of Scientific Names .......................................................................................
ABSTRACT
Farjon, A. (HerbariumDivision, Royal Botanic Gardens, Kew, Richmond, Surrey
TW9 3AB, U.K.) & B. T. Styles (Oxford ForestryInstitute, Departmentof Plant Sciences, University of Oxford, South Parks Road, Oxford OXI 3RB, U.K.). Pinus (Pinaceae). Flora NeotropicaMonograph75: 1-293. 1997.-A revision of the pines native to
Mexico, CentralAmerica, and the Caribbeanis presented.Of the 47 species recognized,
most are endemic to this region, but a number extend into Mexico from the United
States, while P. elliottii var. densa is included because it occurs on the Florida Keys in
the Caribbean.The extensive synonymy has been largely resolved by (lecto-)typification.
Some taxonomic and nomenclaturalproblems have been addressedin previous papers;
only one new combination,P arizonica var. cooperi (C. E. Blanco) Farjon,is proposed
here. All species are fully illustrated.The introductioncovers all aspects of pines that
are of interest to both taxonomists and more general readers, and includes specialist
contributionson wood anatomy (I. D. Gourlay), pollen morphology(M. H. Kurmann),
and taxonomy based on monoterpenes(J. S. Birks).
3
4
5
13
30
35
36
38
41
42
49
59
62
71
198
255
256
257
267
267
285
285
Flora Neotropica
RESUMEN
Farjon, A. (HerbariumDivision, Royal Botanic Gardens, Kew, Richmond, Surrey
TW9 3AB, U.K.) & B. T. Styles (Oxford Forestry Institute, Departmentof Plant Sciences, University of Oxford, South Parks Road, Oxford OXI 3RB, U.K.). Pinus (Pinaceae). Flora Neotropica Monograph75: 1-293. 1997.-Se presentauna revision de los
pinos nativos de Mexico, Centroamericay el Caribe. La mayoria de las 47 especies
reconocidasson endemicas a esta regi6n, pero algunas se extienden hacia Mexico desde
los Estados Unidos, y se incluye P elliotii var. densa porque se encuentraen las islas
de Florida en el Caribe. La sinonimfa muy extensiva en el genero, se ha resuelto en
gran parte por medio de (lecto-) tipificaci6n.Algunos problemastaxon6micos y de nomenclaturafueron discutidos en varias publicaciones anteriores;se propone aquf solamente una combinaci6n nueva: P arizonica var. cooperi (C. E. Blanco) Farjon.Todas
las especies estan ampliamentegrabadas.Los capitulos preliminaresincluyen todos los
aspectos de los pinos de internsparatax6nomos tanto para los lectores mas generales, e
incluyen contribucionesescritas por especialistas sobre anatomia de la madera (I. D.
Gourlay), morfologia del polen (M. H. Kurmann)y aspectos taxon6micos con relaci6n
al anilisis de monoterpenoides(J. S. Birks).
PREFACE
Many people, tending to think of pines as a genus that is largely distributedin the
forests of the temperate region, are unawareof the importanceof the tropical pines.
Mexico and Central America have a great richness of this ecologically and
economically importantgroup, and so it is vital that they be monographedfor Flora
Neotropica. Twenty-five years ago the vision of this task came from the Oxford
ForestryInstitute,and in particularfrom Dr. BrianT. Styles, who was also co-authorof
the treatment of Meliaceae for Flora Neotropica (Pennington, 1981). He began his
work on the neotropical pines in 1970, and it was his greatest ambition to finish the
monographhe considered would be his magtnumtopus. During his years of studying
pines he travelled widely in Mexico and CentralAmerica and collected a large amount
of valuable material that has helped to resolve the taxonomy of several difficult
species-complexes. He also studied a vast amount of herbariummaterial and placed
this on a computeriseddatabase.Although he publishedseveral papersabout the pines,
much of the informationhe had accumulatedremainedin his head, to be committed to
paper during the next few years. His notes and the drawings of each species which he
had had prepared by artist Rosemary Wise enabled the synthesis of his taxonomic
ideas. Unfortunately he was not to see the completion of this project during his
lifetime because of his untimely death in 1993, at the age of 58.
We are very fortunatethat a second specialist both existed and was free to devote
time to the completion of this project.All too many taxonomic projects die along with
the specialist. However, in this case, Aljos Farjon stepped into the breach and pulled
together the many loose ends to produce this fine monograph. It is interesting and
fortunatethat Farjonagreed closely with Styles about the basic taxonomic framework
of the neotropical pines. Although much of the compilation and a great deal of
additional research has been carried out by Aljos Farjon, this monograph also
representsthe taxonomy that would have been producedby Brian Styles.
This commendable, and regrettably enforced, collaboration has produced an
outstanding monograph that is a most significant contribution to our knowledge of
tropical pines. It will stand as a fitting memorialto a generous person who was always
helping other people and who gave so much of his time to teaching botany to a large
numberof students, including me during my undergraduateclasses in plant taxonomy.
There are many of us who are indebtedto Brian Styles for his help, and so we are also
grateful to Aljos Farjon for taking up the difficult challenge of completing someone
else's work.
Ghillean T. Prance
INTRODUCTION
Pinus (Pinaceae,Pinales), an exceptionally natural
genus almost entirelyconfinedto the NorthernHemisphere, is representedin Mexico, Central America,
and the Caribbeanby nearly one-half of its roughly
110 species. The constituenttaxa show more features
in common with each other (synapomorphies)than
they share with any other genus in the family Pinaceae, so the genus is distinctly monophyletic. In
fact, during the last 50 years, only one species, P.
krempfiiLecomte-a rare, localized endemic in the
high mountains of Vietnam-has been placed in a
separategenus DucampopinusA. Chev., on account
of its paired, broad, flat leaves that lack a fascicle
sheath at maturity,though otherwise it fits well into
subgen. Strobus. The mature ovuliferous cone is
typical of this subgenus, and in most recentrevisions
it has been properlyincluded in the genus Pinus.
The genus is distinguished from other members
of the Pinaceae by its needle-like secondary leaves,
borne singly or more commonly in fascicles of 2-8
on short shoots (dwarf shoots), with a fascicle sheath
of bud scales, present at least when young. Primary
leaves are retained in mature plants, but modified
into scariouscataphyllswhich are helically arranged,
as are the leaves of the other genera, subtending
the needle fascicles. The mature ovuliferous cone
consists of few to many woody seed scales with
thickened distal parts (apophyses), each of which is
subtended by a small, insignificantbract. All pines
are woody, and many form large forest trees, some
up to 70 m tall. They frequently grow in stands,
forming extensive forests composed of a single species or a mixtureof several.
Although Florin (1963) held that the Pinaceaeand
the genus Pinus originated in the Northern Hemisphere during the Jurassic, subsequent evidence
seems to point at a possible origin in the Triassic
(Miller, 1977). The genus representsthe earliest evidence of the family. However, soon after this time
the two major pine groups, Haploxylon (subgen.
Strobus)and Diploxylon (subgen. Pinus) had already
differentiated. More recent interpretationsof many
Jurassic fossil cones ascribed to Pinus have cast
doubts on this north-temperateorigin and have dated
the emergence of the genus later, perhaps at the
beginning of the Cretaceous(Axelrod, 1986; Millar,
1993). So far as the Mexican pines are concerned,
their ancestors may have existed in North America
since the lower Cretaceous,where fossil cones similar to those of P devonianahave been found(Chaney,
1954). According to Miller (1977), the pines reached
Mexico from the United States not laterthanthe MidTertiary.Mirov (1967) suggested that they entered
Mexico along the Sierra MadreOccidental from the
North American Cordillera during the Late Cretaceous or at the beginning of the Tertiary. Later
invasions may have been connected with floral retreatsfrom glaciations in the northernpartsof North
America and associated southwardshifts of biomes.
Conversely, Axelrod (1980) and Axelrod and Cota
(1993) postulated a Mexican pre-Miocene origin of
the closed-cone pines (Pinus subsect. Oocarpae),
on the basis of plate tectonic history and present
distributions. Millar (1993) assumed that Central
America may have been one of the refugia of pines
duringthe Late Eocene cooling of the climate. Unfortunately, fossil pines are virtually unknown from
Mexico, and none have been found in Central
America or on any of the Caribbeanislands (Mirov,
1967;Axelrod, 1980; Millar, 1993). The earliest finds
in Mexico are Late Oligocene pollen found in amber
deposits in Chiapas, which may themselves be of
pines but cannot be definitively determinedas such.
Pines are mostly pioneerspecies and probablyspread
quickly on the lava flows thatcame from the volcanic
activity occurringthen throughoutthe greaterpartof
Mexico.
It is not surprising, in a genus with such an
extensive worldwide distributionas Pinus and with
the diversity it displays in several of its characters,
thatthere is little agreementon the numberof species
extant. Shaw's (1914) exceptionally conservative
monographacknowledged only 66, whereas Gaussen's (1960) treatmentincluded 120. Critchfieldand
Little's (1966) geographic distribution maps show
94, while Mirov(1967) recognizedsome 103 species.
For Mexico the situation is not much better: Shaw
(1909) recognized only 19 species for Mexico, but
40 years later Martinez (1948) counted 39 species
there. Loock (1950) accepted 28 species for Mexico
and Belize, and finally Perry (1991) recognized 58
species for Mexico and Central America. There is
little doubt, given the number of new pines being
described each year, that the species concept is becoming narrower still. Taxa formerly described at
the level of varietyor subspecies have been elevated
to the rankof species. This trend is ascribed mainly
to the use of chemical information,where if a population differs from the species as a whole in the level
of a certain compound or in the absence of an
enzyme, that population must be a different taxon.
But chemical constituentscan be as variable as any
4
HistoricalSurvey
other characters,and difficulties arise in interpreting
them taxonomically,especially if such qualifications
are based on compound data (Birks, this volume).
Much of this "splitting"has been proposed by nontaxonomists, who do not seem to appreciatesufficiently that widespreadtaxa often show geographical
variationsthat may or may not be worthy of taxonomic recognition. Given the known history of migrationof many pines, it is considered-and, indeed,
observed when assessed-that the variationin many
of them is clinal in nature. Perhaps the most important reason for the disagreement about species
limits lies in the morphology of pine plants themselves; taxonomistsare forced to base theirdecisions
on the variationfound in leaves and matureovuliferous cones. Leaves (needles) of pines show comparatively little variation, compared with the leaves of
angiosperms,and may be greatly subjectto extremes
of environment.Pollen cones are extremely similar
in most species. Thus the seed cones remain as the
chief source of taxonomic information.
No othergroup of woody plantsof equivalentsize
has been so intensively studied as the genus Pinus.

The taxonomic literature alone is extensive; with
references to related disciplines added to this there
are thousandsof relevanttitles (Farjon, 1990b). The
synonymy to be dealt with is also substantial:for 47
species and 20 infraspecific taxa here recognized,
more than 400 names have been proposed. This
monographis the first critical revision of the pines
of Latin America since Shaw (1909); it is the most
comprehensiveassessmentof morphologicaland anatomical data to date. It cannot definitively solve all
taxonomic problems: in particular,the delimitation
of taxa and their interrelationshipsin the speciesP ponderosa and P
complexes of P. montezumnae,
pseudostrobus,and the relationshipsof disjunctpopulationsand taxa in P cembroidesand its close allies
merit further investigation. However, new approaches such as cpDNA variationanalysis may be
more promising than furtherinvestigations into the
apparentlypolymorphicmorphologicaland chemical
characters which have resulted in the conflicting
taxonomies.
HISTORICAL SURVEY
F. L. von Schlechtendaland A. von Chamisso (1830,
1831). A decade later,anotherGerman,C. G. Ehrenberg, collected pine specimens in Mexico; the new
The earliest period of incidentalcollecting of pines, species were again described by Schlechtendal
1785-1914, is here brokendown into two periods.
(1838) and by Gordonand Glendinning(in Gordon,
1858). A fourth German, a gardener named K. T.
Hartweg,set out for Mexico to gathernew plantsfor
I. 1785-1850
the HorticulturalSociety of London;he made further
Botanical collecting in both the Caribbean and additions,and it was throughhis specimens, widely
Mexico startedwith a few late seventeenth-and early distributed,that Mexico became known to host a
eighteenth-centurycollections, mainlyby Dutchtrav- diversity of these trees. The plant collection as a
ellers (Adams et al., 1987; Divila A. & GermanR., whole was enumeratedand described by Bentham
1991). The earliest botanical exploration started in (1840, 1842), who describedonly one new pine from
Mexico with the expeditions led by Martinde Sess6 Hartweg's gatherings, using a later homonym (P
in 1786 and 1788-1795, but no pine specimens from tenuifolia) to name it. Several more were described
these collecting activities are known. There may be by J. Lindley, Secretaryof the HorticulturalSociety,
some specimens of pines from the "Royal Botanical who was instrumentalin the distributionof some of
Expedition to New Spain" in 1790-91 in Madrid Hartweg's pine material to "subscribers"or to his
(MA). The oldest collections we have seen are two own botanical acquaintances,e.g., S. Endlicher in
specimens collected by M. A. Bonplandand A. von Vienna. F M. Liebmanncollected pines mainly in
Humboldt,now in Paris(P-Bonpl.).The earliest pine Oaxaca; B. C. Seemann collected several specimens
specimen from the Caribbeanwas collected by Olof on his second trip into the interiorfrom Mazatlanto
Peter Swartz, a pupil of Linnaeus filius, on Hispan- Durango.Among these was P lumholtzii,which was
iola in 1785. Althoughabundantin many areas,pines not recognized as a distinct species until Robinson
were apparently of little interest botanically until and Fernald (1894) described it based on Hartman
1828-1829, when C. J. W. Schiede & F. Deppe made 541 from near Coloradas, Chihuahua,collected in
several collections in Mexico, to be describedby D. May 1893. Liebmann's specimens went to various
HISTORYOF COLLECTING
AND DESCRIPTION
Flora Neotropica
herbariain the United States, and those of Seemann

went to England (BM, K); no new species were
described from any of these. J. Gregg discovered a
new species, which was named after him by G.
Engelmann and described by Parlatore(1868). He
also travelled to northernMexico. The far north of
thatcountrywas exploredfromthe UnitedStates on a
boundarysurvey,on which F A. Wislizenuscollected
specimens of Pinus strobiformis,describedas a new
species by Engelmann(1848).
These modest early collections from a vast and
largely unexplored territory nevertheless yielded a
substantial portion of the species now known to
occur in Mexico. The pine forests of Mesoamerica
(Chiapas and the Central American countries) remained virtually unknown-only Hartweg had collected a few specimens in Guatemala,described by
Bentham (1842) as Pinus tenuifolia (= P maximinoi). In the Caribbeana pine was known from
Hispaniola by two collections (P occidentalis), but
none were collected in Cuba and the Bahamas.
1. Collectors in Mexico 1785-1850, with number
of discrete collections:
1806 M. A. Bonpland& A. von Humboldt(2)
1827 F. Karwinski(3)
1828-1829 C. J. W. Schiede & F. Deppe (28)
1837-1840 C. Ehrenberg(25)
1838, 1845-1847 K. T. Hartweg(22)
1839 J. Parkinson(I)
1840 H. G. Galeotti (2)
1841-1842 F. M. Liebmann(15)
1843 A. B. Ghiesbrecht(1)
1846 F. A. Wislizenus (5)
1847-1849 J. Gregg (9)
1850 C. C. Parry(1)
1850 B. C. Seemann (6)
2. The only collector in Central America during
the period 1785-1850, was K. T. Hartwegwho made
2 discrete collections in Guatemala,1841-1842.
3. Collectors in the Caribbean 1785-1850, with
number of discrete collections and place of collection:
1785 0. P. Swartz (1, Hispaniola)
1828 B. Jaeger (1, Hispaniola)
P engelnmannii
(Wislizenus)
P greggii (Gregg)
P hartwegii (Ehrenberg,Galeotti, Hartweg, Karwinski)
P herrerae (Liebmann)
P leiophylla (Ehrenberg, Gregg, Liebmann,
Schiede & Deppe, Seemann)
P. leiophylla var. chihuahuana (Seemann, Wislizenus)
P. lumholtzii(Seemann)
P mnaximinoi
(Bonpland & Humboldt,Hartweg)
P n ontezumae(Bonpland& Humboldt,Hartweg,
Parkinson,Schiede & Deppe)
P. montezumaevar. gordoniana (Hartweg)
P. occidentalis (Jaeger, Swartz)
P. oocarpa (Ehrenberg, Liebmann, Schiede &
Deppe)
P patula (Ehrenberg,Galeotti, Gregg, Hartweg,
Liebmann,Schiede & Deppe)
P. pinceana (Ehrenberg, Ghiesbreght, Karwinski?)
P pseudostrobus(Hartweg)
P pseudostrobusvar. apulcensis (Hartweg)
P quadrifolia (Parry)
P strobiformis(Wislizenus)
P tecunumanii(Liebmann)
P teocote (Ehrenberg, Gregg, Hartweg, Liebmann, Schiede & Deppe)
By the end of 1850, a total of 21 species and 3
varieties out of 47 species and 20 infraspecifictaxa
presently recognized had been collected-that is,
nearly one-half of the species now recognized were
known in the first 50 years of botanical exploration
of the region. More were named by then, but a
numberhave been reduced to varieties (3 taxa, here
mentioned)or lost into synonymy.
Although some collectors-e.g., K. T. Hartwegtook care to collect pines purposefully,none of the
early collectors had a special interest in them, and
they were collected incidentallyby botanistsor professional collectors with other, or at least much
wider, interests in plants.
II. 1851-1914
Early in this period, the curious efforts by B.

4. Species collected 1785-1850 (in their current
Roezl
(for a more detailed account, see under P
taxonomy), with collectors names in parentheses:
montezumae)resulted in a substantialcollection of a
limited number of species from Central Mexico.
Pinus ayacahuite (Ehrenberg,Liebmann)
P cembroides(Ehrenberg,Gregg, Hartweg, Kar- However, since they were almost all presented as
new species, it was never meant to be a collection
winski, Schiede & Deppe, Seemann)
P devoniana(Gregg, Hartweg,Schiede & Deppe) to investigate the variationwithin species. The exact
HistoricalSurvey
dates of his gatheringsare unknownbut seem to fall
between 1854 and 1858. His material was sent
mostly to Europe as "vouchers" for seed batches,
which were sold to subscribersin the Horticultural
Society of London and to well-known gardenersand
plantsmenlike George Gordon in Englandand E. A.
Carrierein France.Roezl had given them names and,
in most cases, descriptionsin his catalogues (Roezl,
1857, 1858). Both Gordon and Glendinning(1862)
and Carriere (1867), ill equipped to judge on the
taxonomyof Pinus in Mexico and perhapsalso eager
to accept"noveltiesfor the garden,"acceptedmost of
them. Schlechtendal(1857-1858) translatedRoezl's
descriptionsfrom French into Latin; without having
seen a single specimen he merely commented that
so many new species in Pinus seemed unlikely even
for Mexico. F. Parlatore,when in London in August
1862, boughtRoezl's specimens from John Standish,
gardener of the Duchess of Gloucester at Bagshot
Park, Surrey: on 14 October 1862, a total of 435
specimens of conifers were entered in the registerof
accessions to the Botanical Museum in Florence.
Parlatore(1868) made short shrift of Roezl's species
and put them all into synonymy.
Very few pine collections were made until 18851886, when two professional plant collectors, E.
Palmer and C. G. Pringle, started making general
plant collections on a large and systematic scale.
Palmer's plants were enumeratedand described by
Sereno Watson at HarvardUniversity, but the pines
were entrustedto Engelmannat the MissouriBotanical Garden, who described Pinus radiata (as P insignis) var.binata from Palmer's 1875 gatheringson
GuadalupeIsland. Pringle was a most prolific plant
collector, and it was his custom to make sets of 60
duplicates whenever possible (McVaugh, 1972), but
sets of 12, 15, or 20 are also common, and there
are even (unnumbered)unicates. As a consequence,
Pringle's specimens were distributedto many herbaria. His earliest pine specimens are from Baja California (P quadrifolia, collected in 1882). Very few
collections of pines were made by him in the southern states of Mexico (one only, from Oaxaca, has
come to our attention),and he did not visit Central
America. New species were described by Shaw
(1905a, 1905b, 1905c), of which P pringlei is here
accepted. E. W. Nelson-sometimes, as in 1897,
with E. H. Goldman and J. N. Rose (the latter
numberedhis specimens separatelyas he worked for
the U.S. National Museum)-collected for the U.S.
Departmentof Agriculture,and those main sets of
specimens are kept in Washington(US). It was again
G. R. Shaw at the Arnold Arboretumof Harvard
7
University who describeda new species: P. nelsonii,
named after its discoverer. His monographon the
pines of Mexico (Shaw, 1909) was based primarily
on their collections. Shaw himself went to Mexico
in 1904, travelling with Pringle, to study his specialty, the genus Pinus (McVaugh, 1972). His is the
first trip in the history of pine collecting in Latin
America specially dedicated to the study of the genus. Because Shaw entertaineda very broad species
concept (Shaw, 1909; and especially Shaw, 1914)
there are several new species among the gatherings
of Goldman,Nelson, and Rose which were only later
recognizedand described in the next period, usually
based on new collections.
No pines had been collected from countries in
Central America other than Guatemala until after
the First World War. The collections made on the
CaribbeanIslands during this period were still few.
J. A. Shafer, gatheringin Cuba, made an exemplary
exception with 30 collections of the two species
occurring in western Cuba; for the first time in
the history of pine collecting there was a collector
gatheringmaterialof a species in sufficientquantities
to assess morphological variation among populations. Othercollectors before him had only accidentally, not purposely, made multiple gatherings of a
single species. This was to change fundamentally
in the next period: although incidental collecting
continued until the present, a more systematic approach was adopted by many collectors who now
took a special interestin the pines themselves.
1. Collectors with -10 discrete collections in
Mexico, 1851-1914, with numberof discrete collections:
1854-1858 B. Roezl (84)
(1875) 1885-1908 E. Palmer (26; 3 with C. C.
Parry)
1886-1907 C. G. Pringle (96)
1898-1908 E. H. Goldman(28)
1892-1903 E. W. Nelson (106; 3 with E. H.
Goldman)
1897-1911 J. N. Rose (48; 10 with R. Hay; 5
with W. Hough; 8 with J. H. Painter)
1904 G. R. Shaw (47)
1903-1913 C. A. Purpus(18)
1905 J. G. Lemmon (13)
1907-1912 G. Arsene (24)
1910-1911 A. Arsene (10)
2. Collectors with >5 discrete collections in Central America, 1851-1914, with number of discrete
collections:
Flora Neotropica
1876 C. G. Bernoulli & R. Cario (8)
1890-1892 J. Donnell-Smith(6)
1895-1896 E. W. Nelson (6)
1904-1907 H. von Tuerckheim(5)
1905 W. R. Maxon & R. Hay (6)
1905-1907 W. A. Kellerman(14)
3. Collectors in the Caribbean, 1851-1914, with

numberof discrete collections:
1853 R. H. Schomburgk(1)
1856-1865 C. Wright (8; I with C. C. Parry,C.
Wright& Brummell)
1887 H. F A. von Eggers (1)
1903-1912 J. A. Shafer (30)
1903-1905 G. V. Nash & N. Taylor (2)
1904-1914 HermanoLe6n [brotherLe6n] (2)
1910 H. von Tuerckheim(1)
1911 H. Clinton-Baker(1)
1914 E. L. Ekman (3)
1878-1914 e.g., L. J. K. Brace; J. K. Small &
J. J. Carter(16, Bahamas)
P pseudostrobusf. protuberans(Pringle)
P radiata var. binata (Palmer, 1875, Rose, 1911)
P tropicalis (Clinton-Baker,Shafer, Wright)
By the end of 1914 an additional 15 species and
9 infraspecifictaxa had been collected. More often
than duringthe previous period, these nomenclatural
innovations involved taxa only much later to be
recognized as distinct, e.g., Pinus douglasiana, P
durangensis, P praetermissa, and the infraspecific
taxa in P. arizonica and P cembroides.
The contemporaryperiod (1915-present) of systematic collecting of pines is broken down into two
periods.
III. 1915-1950
We characterizethe time after the FirstWorldWar

to the present as the period of systematic collecting
of pines. With few exceptions (Shaw, Shafer), specimens of the genus collected in the foregoing period
had been gathered by people who were out in the
4. Additionalspecies collected 1851-1914 (in their
field making general collections of plants, often focurrenttaxonomy), with principalcollectors' names
cusing on entirely different taxa. Even Roezl, who
in parentheses:
at one time was involved with pines, is much better
known for the great number of orchids and other
Pinus arizonica (Nelson, Palmer)
P arizonica var. cooperi (Pringle)
plants he collected, for which he undertook much
wider exploration than he did for the pines. To be
P arizonica var. stormiae (Palmer)
P. ayacahuite var. veitchii (Roezl, 1857, lost, sure, general plant collecting continued, and these
efforts brought in pine specimens as well. But inPringle)
P caribaea (original materialdescribedby More- creasinglywe encountercollectors who are interested
in the genus Pinus itself, primarilyor exclusively.
let 1851, lost; Rowlee, 1901, Le6n brothers)
P caribaea var.bahamensis(earliestBrace, 1878) Native-borncollectors are beginning to play an imP caribaea var. hondurensis (original material portantpart in the collecting; it is no longer solely
described by S6enclauze 1868, lost; Ber- the concern of Europeansor Americans.
noulli & Cario, Donnell-Smith, Kellerman,
Partly of Mexican descent and later living and
Maxon & Hay)
working in California, Y. Mexia made an extensive
P cembroides var. bicolor (Nelson, 1892)
trip to the western states of Mexico in 1926-1927
P cembroidessubsp. lagunae Nelson & Goldman, making general collections for the University of
Californiaat Berkeley (UC). She was the first female
1906)
R contorta var. murrayana(Goldman, 1905)
plant collector to make substantial collections of
P cubensis (Ekman, Shafer,Wright)
pines in Mexico. E. Matuda, a Mexican collector,
was very active in Chiapas from 1936 to 1950 and
P douglasiana (Nelson, Rose)
collected almost all of his specimens of pines there.
P durangensis (Pringle, Rose)
Yet he did not discover P strobus var. chiapensis,
P flexilis var. reflexa(Nelson)
and his collections cover the common species. One
P jeffreyi (Goldman, 1905)
of these is P tecunumanii,not yet recognized; his
P lambertiana(Goldman, 1905)
collections of this species were considered to be of
P lawsonii (Pringle, Purpus,Roezl, Shaw)
P. oocarpa or P patula.
P nelsonii (Nelson, Pringle, Shaw)
M. Martinez(1888-1964), who was Professor of
P oocarpa var. trifoliata (Rose)
P praetermissa (Palmer,Rose)
Botany at the Universidad Nacional Aut6noma de
Mexico in Mexico City and a prolific writer on the
P pringlei (Nelson, Pringle)
HistoricalSurvey
Mexican flora, did not collect pine specimens himself. They were obtained for him by other persons,
but he assigned his own numbersto the specimens
"for ease of citation and reference" (McVaugh,
1972). He rarely mentioned the person who made
the gathering,an exception being C. E. Blanco, for
whom he named a variety of P arizonica. The pine
collections date from 1937 to 1950, with a few to
1954. Their collection numbers correspond in no
way to these dates on the labels. The informationis
written in Martinez's hand and no original labels
exist; obvious errorsencounteredmake it clear that
the information is not entirely trustworthy,but it
cannot be checked. The collections (both unicates
and duplicates) are in MEXU and in part also in
herbariain the United States. Martinez described a
few new species and numerous new infraspecific
taxa based on these collections. Most of the latter
appearto be mere local forms or variantsof limited
taxonomic meaning; in part this is due to the fact
that Martinez,working during the difficult years of
the Second WorldWar,had no access to collections
outside Mexico and limited access to literatureand
few contacts with other specialists. However, his
lasting contributionsto the taxonomy of Pinus in
Mexico are several (some of which are listed below),
while others (new varieties of P arizonica and new
species P douglasiana, P durangensis, P herrerae,
and P michoacana = P devoniana) were described
by him but had in fact already been collected and
deposited in herbariaoutside Mexico well before his
time.
In Central America, it was the collecting and
floristicwork by P. C. Standleyand J. A. Steyermark
during this period, culminating in their Flora of
Guatemala (1958), published in Fieldiana, which
established the basic taxonomic knowledge of the
genus in that country and beyond. Collections were
also made in Belize, Honduras, El Salvador, and
Nicaragua,but these yielded no new taxa of Pinus.
We now know that all of CentralAmerica's pines,
even P caribaea var. hondurensis, also occur in
Mexico.
Only E. L. Ekman gathered more than a few
numbersof Pinus in the Caribbeanduringthis period.
He collected pines in Cuba and Hispaniola; they
were treated by Pilger (1926) and later by Florin
(1933), who described the distributionof the pines
on these islands.
1. Collectors with >15 discrete collections in
Mexico, 1915-1950, with numberof discrete collections:
9
1926-1938 Y. Mexia (20)
1933-1937 G. B. Hinton (21; 4 by Hinton et al.)
1933-1939 C. H. Mueller (25; 14 with M. T.
Mueller)
1935-1950 H. S. Gentry 18; 3 by Gentryet al.)
1936-1950(-1959) E. Matuda(110)
1937-1944(-1954) M. Martinez(216, not by himself)
1938-1941 S. S. White (28; 1 with R. M. Chatters)
1929-1930, 1938-1948 I. L. Wiggins (10; 6 with
D. Demaree in 1930; 3 with J. W. Gillespie in
1929)
2. Collectors with '5 discrete collections in Central America, 1915-1950, with number of discrete
collections:
1922, 1928-1950 P. C. Standley (53; 2 with P.
Chacon; 1 with L. O. Williams)
1932 J. B. Edwards(5)
1933-1934 A. F. Skutch (8)
1938-1939 H. von Schrenk(5)
1939-1942 J. A. Steyermark(30)
1943-1944 J. Valerio(3; 3 with Rodriguez)
1945-1946 A. J. Sharp(8)
1946-1950 L. O. Williams (2; 5 with A. R.
Molina)
1947-1948 L. R. Holdridge(5)
3. The only collector with 25 discrete collections
in the Caribbeanduring the period 1915-1950 was
E. L. Ekman, who made 20 collections from 1915
to 1927.
4. Additionalspecies collected 1915-1950 (in their
currenttaxonomy), with principalcollectors' names
in parentheses:
Pinus attenuata (Martinez)
P coulteri (Martinez)
P culminicola (Hinton, Mueller)
P monophylla(P. Litchey 1936, Martinez, 1939,
Wiggins, 1946, Gentryet al., 1950)
P muricata (Gentry,Martinez)
P patula var. longipedunculata(Martinez, coil.
E. E. M. Loock)
P ponderosa var. scopulorum (E L. Wynd &
C. H. Mueller, 1936, J. M. Tucker, 1952)
P remota(Mueller)
P strobus var. chiapensis (Martinez)
In this period, the additions to the Mexican flora
resulted mainly from the finding of more localized
taxa, notably the extension of four Californianspe-
10
Flora Neotropica
cies into Baja California(P attenuata,P coulteri, P

and P muricata)and the subalpinespe- Mexico, 1951-present, with numberof discrete colmonoophylla,
cies P culminicola, the latter not recognized as new lections:
until 1961. The first four were collected for Martinez
1949-1975 R. McVaugh(67; 3 with W. N. Koelz)
by anonymous collectors; of the last he had no
1951-1955 J. H. Maysilles (40)
knowledge (Martinez, 1945, 1948), even though it
1951-1964 E. Matuda(49; 3 by Matudaet al.)
had been collected years before by two independent
1952-1982 R. Moran(82; 6 by Moranet al.)
collectors. Two varietiesare taxa describedby Marti1952-1985 J. T. Marshall(24)
nez which are recognizedhere;the remarkableoccur1952-1989 J. Rzedowski (95; 2 with G. B. Hinrence of a variety of P strobus in southernMexico
ton; 7 with R. McVaugh)
and Guatemala is the more noteworthyof the two.
1955 B. Zobel (21)
The occurrenceof P ponderosa var. scopulorumjust
1957-1960 L. Kaplan(19; 4 with E. L. Davis)
inside Mexico is known from the two collections
1957-1961 J. H. Beaman (19; 9 with J. W. Anmentioned;all other collections under P ponderosa
dresen)
from Mexico studied by us appeared to be other
1959-1980 X. MadrigalS. (165; 22 by Madrigal
taxa.
S. et al.)
1960-1963 E. L. Little (170)
1960-1974 L. Vela G. (26)
IV. 1951-Present
1962-1980 D. E. Breedlove(54; 21 by Breedlove
Of ca. 8500 collections in our database, the vast
et al.)
majority, roughly 6000, has been gathered since
1963-1970 A. May Nah (64)
1950. More recently, since around 1970, they have
1967-1982 R. HerndndezM. (8; 17 by Hernandez
increasingly been accumulatedas vouchers to seed
et al.)
collections by collectors working for forest geneti1970-1971 M. A. Capo (35)
cists or by the researchersthemselves. They have
1971-1984 R. A. Bye (23; 5 by R. A. Bye et al.)
been deposited in a limited number of herbaria,
1972-1974 B. T. Styles (85; 14 with R. F. Kemp)
e.g., FHO in Oxford and NCSC in Raleigh, North
1974-1983 D. K. Bailey (25; 19 by Bailey et al.)
Carolina. One recognizes such specimens collected
1975-1984 A. A. Ventura(20)
for this purpose often by their bound-up foliage,
1977-1978 T. Eguiluz (24)
which makes them virtuallyworthless for herbarium
1977-1981 W. L. Mittak(102)
studies. Emphasis is on species and "provenances"
1978-1981 J. W. Stead & B. T. Styles (145 + ca.
of these, which have a putativepotentialin plantation
240 as "populationcollections"; 7 by Stead et
forestry.Often, 5-20 trees were sampledof a species
al.)
in a stand or otherwise limited area ("population
1979-1985 J. C. Soto N. (7; 30 by Soto N. et al.)
collections"; such collections have been enumerated
1981-1986 M. Nee (11; 65 by Nee et al.)
with serial numbersin the list of exsiccatae). Because
1982-1991 J. A. Perez de la Rosa (25)
pines extend into the tropics in our region, there
1983-1984 K. Taylor & M. Nee (22)
was much interest in collecting taxa from the more
1983-1985 P. S. McCarter (7; 24 with C. E.
southernparts of their ranges, which might be used
Hughes; 33 with B. T. Styles + ca. 100 in
in other tropical countries. The data presented here
"populationcollections")
for this period may therefore be biased, despite
1983-1993 C. E. Hughes (34; 19 with B. T.
thoroughscrutinyof collections in many of the major
Styles)
herbaria,especially by B. T. Styles.
1984 K. D. Rushforth(20)
Botanists were active, too, and they are largely
1984-1986, 1991 B. T. Styles et al. (40)
responsible for the collecting of the remainderof
1992-1994 A. Farjon(5; 33 with D. Mejia)
species and for the discovery of new ones. In the
1993 S. Higman et al. (52)
late 1980s and early 1990s the emphasis returnedto
botanical collecting, although some organizationsCentral
America, 1951-present, with numberof dise.g., CAMCORE, Raleigh, North Carolina (not increte
collections:
cluded in list of exsiccatae)-continued to make
1951 F. Schwerdtfeger(36, Guatemala)
vouchercollections of seed-samplingefforts through1956-1975 A. Molina R. (28; 12 by Molina R.
out the ranges of a number of timber-producing
et al.)
species.
History of Classification
1957 H. A. Luckhoff
1969-1972, 1977 R. H. Kemp (123)
1972, 1986 B. T. Styles (5; 82 with R. H. Kemp)
1972-1975 A. Greaves (20)
1975-1979 G. Chaplin (32; 1 with A. Greaves)
1975-1981 W. L. Mittak(308 + ca. 150 "population collections"; 5 with E. Espinosa; 13 with
J. P. Perry,Guatemala)
1977-1982 J. W. Stead (132 + ca. 250 "population
collections"; 160 with B. T. Styles)
1979, 1985-1987 A. Grijalva(7; 1 with M. Araquistain; 13 with E. Sediles, Nicaragua)
1981-1983, 1993 C. E. Hughes (68; 4 with B. T.
Styles)
1983-1988 P. S. McCarter(55; 20 with C. E.
Hughes;64 with B. T. Styles + ca. 100 "population collections";6 with Hughes & Styles)
3. Collectors with 210 discrete collections in the
Caribbean, 1951-present, with number of discrete
collections and place of collection:
1956-1957 H. A. Luckhoff (13, Bahamas)
1966-1971 J. Bisse et al. (13, Cuba)
1968 A. F. A. Lamb (30, Cuba)
1980-1988 T. A. Zanoni et al. (20, Hispaniola)
1988 W. B. H. Baylis (28, Bahamas)
4. Additional species collected 1951-present (in
their current taxonomy), with principal collectors'
names in parentheses:
Pinus cembroidessubsp. orizabensis (C. T. Hartweg, before 1850 [lost]; M.-F Robert, 1970;
Bailey, 1980)
P jaliscana (S. Carvajal,1982; Perez de la Rosa,
1983; Farjon& Mejia)
P maximartinezii (Rzedowski, 1964; Bailey,
Hughes, Farjon& Mejia)
P rzedowskii(MadrigalS., 1968; Farjon& Mejia)
Despite the enormous increase in the number of
collections made since 1950, only four additional
taxa were discovered, all in Mexico. Of Hartweg's
original collection nothing remains; this subspecies
of P. cembroides was rediscovered by Bailey, although it had earlierbeen collected by M.-F Robert
without recognizing it as distinct. Two very remarkable new species were discovered, both narrowendemics: P maximartinezii and P rzedowskii; the
latest new discovery is P jaliscana, a species allied
to P herrerae.Especially in the "pinyonpines"other
new taxa have been describedduringthis period,but
we have not recognized them as distinct from those
which had been named earlier.
I1
HISTORYOF CLASSIFICATION
The genus Pinus was established by Linnaeus
(1753, 1754) with 10 species, of which only five are
truepines (Pinus s.str.),the othersbelonging to Abies
(2), Cedrus(1), Larix (I), and Picea (I) in Pinaceae
(Farjon, 1990a). The earliest segregation of Linnaeus's species in Pinus s.str. was by Miller (1754).
Numerousproposalsto furthersubdividethe genus
Pinus have been made, the firstof these by Duhamel
du Monceau(1755) on the basis of numberof leaves
per fascicle. Mirov (1967) discussed some major
classifications and gave lists of species with each.
Little and Critchfield (1969), in their well-documented classification,gave a review of the previous
principalclassifications. A review of 10 classifications, from Shaw (1914) to van der Burgh (1973),
was furthergiven by Landry(1978). We think it is
appropriateto refer to these reviews here instead of
repeatingthem entirely, but a few salient examples,
pertainingto two majorsystems of subdivision"competing" up to the present time, may be highlighted.
These two systems differ in that one recognizes a
basic binarydivision in the genus, whereasthe other
divides the genus into more than two groups at the
highest subgeneric rank recognized. The singular
segregation of P krempfii,an aberrantspecies also
placed in a separate monotypic genus, in otherwise
binary classifications, is not counted as a deviation
from that system.
Secondly, as already accentuated by Little and
Critchfield(1969), there are nomenclaturalimplications which, despite a comprehensive treatmentby
these authors, are still being ignored in much of
the more recent literatureon pine phylogeny and
classification.The most often encounterednomenclatural error is the failure to recognize the ruling of
Article 22 of the Code (e.g., see Greuter et al.,
1994), which prescribesthe use of the genus name
("autonym")as the epithet for thatpartof any subdivision of a genus that includes the type of thatgenus.
The most widely acceptedsubdivisionof the genus
Pitnusis the binarydivision into two groupsof equal
rank,each usually subdividedfurtherinto groups of
species. The firstauthorto proposethis was Parlatore
(1868), who recognized within Pinus subgen. Pinus
the two sections ("sectio")Pinea and Cembra.Engelmann (1880a), treatingthe genus in its strict sense,
divided it (in a key to the species) into two sections,
Strobus and Pinaster. It is importantto note that
Engelmanntried to use several independentcharacters (which admittedly did not all fit equally well)
insteadof only one (see below). Section Strobuswas
12
characterizedby (terminology modernized here) a
terminalumbo, deciduous fascicle sheaths, leaves in
fascicles of 5, microsporophyllswithout a terminal
crest, and wood less dense and less resinous. Section
Pinaster had a dorsal umbo, usually persistentfascicle sheaths, a variableleaf number,microsporophylls
with a semi-orbicularcrest, and denser, more resinous wood. Notably, this led to his grouping of P
cembroides and its allies, as well as P balfouriana,
with sect. Pinaster; these taxa have been classified
with (subgen.) Strobus in most later binary classifications. This binary treatment of species groups
within the genus was accepted by the authors of
major"NaturalSystems"of plantclassification,such
as Bentham and Hooker (1880) and Eichler (1889),
as well as by Sargent(1897) in his authoritativetree
manual, which certainly stimulated its acceptance.
Lemmon (1888, 1895) was the first author to use
the rank subgenus for the two main subdivisions
designated previously as sections by Engelmann,for
which the currentcorrectcitations are Pinus subgen.
Strobus (Sweet ex Spach) Lemmon and Pinus subgen. Pinus [not Pinaster (Engelmann)Lemmon].
Koehne (1893) discovered that a binary division
of the genus was supportedby the presenceof either
one or two vascular bundles in the leaf and consequently proposed yet another set of names for two
sections: Haploxylon(I vascularbundle)and Diploxylon (2 vascular bundles). According to this single
characterscheme, P balfouriana and P. cembroides
have to be classified with Haploxylon (= Strobus).
These two "appropriatelydescriptive names" (Little & Critchfield, 1969) are still widely used, especially in non-taxonomicliteratureand at subgeneric
rank,despite their illegitimacy underthe rules of the
Code.
Non-binaryclassificationshave been proposedand
were adoptedas alternativesto the basic division into
two groupsas outlinedabove, and indeed precededit.
Early examples are Spach (1841), with a division
into four sections; Endlicher(1847), with six sections
in Pinus s.str.; and Koch (1873), with six "Gruppe,"
presumablycorrespondingwith the rank of section.
Nevertheless, all classificationsof the (entire) genus,
proposed between 1890 and 1940, recognized the
binary division of the genus, either as subgeneraor
sections, albeit undera numberof (now illegitimate)
names.
Martinez(1945, 1948) arrangedthe Mexicanpines
into nine sections, some of which were furthersubdivided into "groups."None of these were formally
proposed with a Latin description and some had
Mexican, not Latinized, names. Yet it is clear that
Flora Neotropica
Martinezabandonedthe binaryarrangementsof previous authorsand returnedto the system of the early
period of classification.
An entirely differentschool of thoughtwas developed by Campo-Duplan(1950) and de Ferre (1953,
1965), based on ideas of evolution in conifers held
by Gaussen.This led to detailedobservationalstudies
of those charactersbelieved to have evolved much
more than others, e.g., charactersof pollen (size and
shape) and seedlings (cotyledons), and to the ignoring of others believed to be primitive, e.g., the
morphologyof the female reproductiveorgan. For a
critique see van der Burgh, 1973. The genus was
accordinglysubdividedinto two subgenera(CampoDuplan, 1950) on the basis of pollen morphology,or
into four subgenera (de Ferr6, 1953, 1965) on the
basis of seedlings, the latter with further subdivisions. Landry(1974, 1978, 1994) reasoned contrary
to Gaussen(1960), who had rejectedcone characters,
and believed in a Linnaeanprincipleof classification:
since the greatmasterhad used flowers,the reproductive organs of pines must give the clue to an unambiguous classification. Although Landry has been
correct in recognizing some distinctive taxa (Pinus
leiophylla: Landry,1992; P nelsonii: Landry,1994),
his classification,like those from the school of Gaussen, must be rejectedas entirely artificial.
Few other contemporaneousinvestigations using
a limited number of charactershave proposed new
classifications for the genus. We mention Hudson
(1960) on wood anatomy (non-binary) and Jahrig
(1962, 1968) on leaf anatomy (binary). The contrasting method, phenetic classification in the sense
of Sokal and Sneath (1963), has never been applied
to the genus as a whole, althoughclassifications like
those of Little and Critchfield (1969) and van der
Burgh (1973) are phenetic in the sense that they
group taxa on overall similaritycomparing multiple
characters.
Increasingly, evaluation of special characters is
being used to assess phylogenetic relationshipsand/
or to test previously proposed "traditional"classifications. Most papersinvestigate only a limited number of species traditionallyclassified in a section or
subsection of the genus; these are not considered
here, as we are interested in the history of the
classification of the genus as a whole. Most studies
of monoterpenes,for instance,fall into this category.
Yet there are a few chemical studies that did find
differences between the two subgenera,such as Erdtman (1959) on heartwood polyphenols. Molecular
techniques,constantly improving,have made it possible to study genetic variationanalysing allozymes
13
Morphologyand Anatomy
and even more directly in the DNA itself. Examples
of allozyme studies are those by Karalamangalaand
Nickrent (1989), who looked at variationamong 14
Mexican and southwesternU.S. species in subgen.
Pinus ("Diploxylon"),and by Shurkhalet al. (1991),
who studiedEurasianrepresentativesof both subgenera. A phylogenetic analysis based on nuclear DNA
(ribosomal DNA) was carried out by Govindaraju
et al. (1992) assessing 30 species. The chloroplast
genome, paternallyinheritedin conifers, appearsin
these studies to give the most reliable results on
relationshipsbetween taxa, includinghybridrelationships (Wang, 1992). A numberof papershave studied
chloroplast-DNA polymorphisms, using restriction
fragmentlength analysis, in a (random)selection of
species assigned to several traditionalgroups (subgenera, sections), and evaluatedthese using a cladistic method (usually presenting parsimony cladograms) to investigate the phylogeny of that sample
of taxa. Examples of such cpDNA analyses are
Szmidt et al. (1988), with 20 species studied, Strauss
and Doerksen (1990), with 19 species tested, Wang
and Szmidt (1993), with 18 species, and Perez de la
Rosa et al. (1995), using 12 species. Only the last of
these studied Mexican pines.
The majorityof these investigationstends to support a binary classification of the genus; at lower
levels of hierarchy,several supportLittle and Critchfield (1969) and similar classificationsgenerally,but
give indications that some groupings, such as the
heterogeneoussect. Parryanae,are paraphyleticand
need to be revised. Malusa (1992), in a phylogenetic
study using morphologicalcharactersof the pinyon

pines, arrivedat a similarconclusion. Classifications
ought to be based on phylogeny, investigated by
means of the evaluationof a numberof independent
characters,consistent with a testable theory of the
evolution of the genus. The recent molecularevaluations have been, and promise to continue to be,
helpful test cases for existing classifications as well
as sources for useful taxonomiccharacters;a synthesis with anatomical and morphological data seems
the way forward.
A final remark on a recent(?) trend in conifer
taxonomy and classification is relevant here. There
has been a consistentnarrowingof the genus concept,
both in less well-studied families such as Podocarpaceae and in intenselystudiedones such as Pinaceae
(e.g., Page, 1989, 1990). How well this can be
justified in the light of the evolutionary history of
conifers generally is not a point of discussion here.
Whatmust be questionedis the assumptionthatsince
the (morphological)differencesbetween some of the
segregate genera appear to be small, a large and
polymorphicgenus such as Pitins ought to be broken
up as well. Divergence, caused by the "accumulation" of autapomorphies,may lead to relative differences between otherwise perfectly monophyletic
taxa. Contraryto the "rule"still maintainedby some
cladists, that each clade (sister group) should be
given equal rank or that all nodes representnatural
taxa and should consequentlybe ranked,it may well
be valid not to do so (Stace, 1989).
MORPHOLOGY AND ANATOMY

This section comprisesan accountof the morphology and anatomy of habit, shoots, vegetative structures, bark, and reproductivestructures.Wood and
pollen are described in separatechaptersby specialists, in acknowledgementof their responsibilityfor
these contributions;their treatises immediately follow this chapter.
ally, resulting in a hierarchical,symmetricalsystem

of branchingfrom the centre toward the periphery,
known as "Rauh's growth model" (Halle et al.,
1978). The characteristicsare an erect leading shoot,
assurgent higher-orderbranches, and a non-plagiotropic disposition of the branchesof all subsequent
orders, usually 4-5. Normally, shoot elongation is
uninodal, i.e., during the growing season a single
system of apical and lateral primordia of stem,
GROWTH FORM AND
leaves, and reproductiveorgans, initiatedin a previBRANCHING PATTERN
ous season and forminga terminaland severalsubterThe genus Pinus consists of perennial,arborescent minal "winterbuds," elongates and develops at the
species, many of which attain great age and size. ends of stem and branches;growththen ceases and a
The principalbranchingpatternis monopodial,of a new set of primordiais initiatedbefore the dormancy
type in which the lateral branches are (pseudo-) period. In a number( 10 in our region) of species
whorled.The apical shoot meristemdominates over of subgen. Pinus, multinodal shoots occur or are
the laterals;this patternis repeated,usually season- predominant.Multinodalsystems are likewise initi-
14
Flora Neotropica
ated in the winterbuds (see below), and development

of the vegetative organs, as well as of the reproductive organs, especially ovuliferous cones, occurs simultaneously on these internodes within a single
growing season. Shaw (1914) believed there was in
the genus an evolutionary tendency toward
multinodal growth, most prevalent in the "serotinous" pines. Multinodalshoots are not a fixed characterstate in any species; rather,they are a traitthatis
either rare (occurringonly in strong leading shoots),
common, or prevalent;this type of occurrencecould
support Shaw's view if it appears to be congruent
with the phylogeny of species inferred from fixed
derived characters.Secondarygrowth occurs primarily fromthe cambiumlayers, formingbarkand wood,
although reiterationfrom dormantbuds may occur,
especially in a few species that also coppice, e.g.,
Pinus leiophylla. A subapical meristem commonly
assumes dominance upon damage to or elimination
of the leading shoot; this, rather than reiteration,
causes irregularstem shape and branchingpatternin
older trees. In a few species, the plants remain
small and shrub-like,primarilyby limitationof apical
dominance of the main (stem) shoot; however, all
species in our region, even P culminicola, remain
monopodial, and no truly multistemmedbranching
patterns(nor layering)has been observed in Mexican
shrub-likepines.
SHOOTS
The genus is characterizedby extreme shoot dimorphism: branching shoots and dwarfed foliage
shoots. Shoot dimorphism occurs in all genera of
Pinaceaeto a greateror lesser degree (Frankis,1989),
with examples of strong dimorphism in Cedrus,
Larix, and Pseudolarix and weak dimorphism in
Abies and Picea. In Pinus, the short foliage shoots
are "extreme"in thatthey neitherelongate repeatedly
nor producemore than a first set of leaves normally,
as is the case in Cedrus, Larix, and Pseudolarix
(numberof leaves indefinite).They are thereforebest
defined as dwarf shoots (brachyblasts)to distinguish
them from the short shoots of the other genera.After
initial growth and formationof a definite numberof
green leaves, they remain on the plant for the duration of the leaves (2-20 years or more, but in our
region not longer than8 years) and thereafterbecome
deciduous with them, leaving in some cases a few
basal remnantsof sheath(= bud) scales. In the other
genera, the short or spur shoots are persistent and
only the leaves fall. The secondary leaves are also
helically arrangedin the former three genera (in a
very low spiral), but they are cyclic (whorled) in

Pinus. Primary leaves continue to develop on the
branchingshoot each season in Cedrus, Larix, and
Pseudolarix, whereas in mature Pinus only cataphylls (also termed"scale leaves" or "bracts")appear
(Thomson, 1914). Dwarf shoots cease to grow apically at a length of ca. 250 aum(Mirov, 1967) and
terminate in a dormant bud. After defoliation, this
bud may produce a second fascicle of secondary
leaves. In some cases, shoot proliferationoccurs as
a response to damage (e.g., after damage to the long
shoot by herbivory).In most pines the initiation of
long shoots from shortshoots is not the normalmode
of branching,as it is in the otherthreegenera(Farjon,
1990a). Nevertheless,the distinction is only gradual,
with the condition in Pinus as the most advanced
and specialized of a derived state in the family.
The branching shoots (internodes) develop from
single terminaland multiple subterminal,rarely lateral, buds by elongation (the whole of a season's
growth, vide Shaw, 1914) and are defined by a
leafless base and a terminal node of buds. As described above, several species in our region develop
multinodal shoots, but the prevalent mode of shoot
development is uninodal. Shoot elongation commonly concurs with elongation of secondary leaves
(see below), but in P maximartinezii,P nelsonii,
and, to a lesser extent, P cembroides and its allies,
growth of the leaves is postponed until the shoots
have reached full length. Shoot elongation, apart
from apical dominance,is influencedby environmental factors and is generally greater in species restricted to a mesic environment.A few taxa in our
region, most notably P elliottii var. densa, exhibit
arrestedapical growth of the apical meristem in the
seedling stage ("grassstage"). Instead,the hypocotyl
is greatlyenlargedand the root system develops first,
while the foliage undergoes the normal developmentaltransitions(see below). Once foliage and root
system are sufficient to sustain it, rapid growth of
the stem lifts the young tree and its foliage above
the normal zone of grass fires, where it is relatively
safe and starts lateral branching.A comprehensive
study of shoot morphologywas publishedby Thomson (1914).
Pubescence is rarein our region and of very short
duration;several species commonly have glaucous
to pruinose shoots. Shaw (1914) saw a correlation
between glaucousnessand aridity,but in the Mexican
species thatgrow in the most aridclimatic conditions
it is absent or only weakly developed. Glaucousness
is also stronglycorrelatedwith age and vigour, being
much moreobvious on shoots of young, fast-growing
15
trees and nearly absent on shoots of senescent trees primary leaves of species in subgen. Pinus is on
of most species.
average somewhat higher: respectively, 1-5 and 2The cataphyll bases (pulvini) of most species are 4. There are 2-3 resin ducts in the primaryleaves;
persistentuntil after the leaves have fallen; these are the average number is slightly higher than in the
either short decurrent(subgen. Strobus, sect. Stro- cotyledons. Primaryleaves are usually producedfor
bus), short to intermediate(subgen. Strobus, sect. only the first season and only on the main stem in
Parryanae) or long decurrent (subgen. Pinus), in most species, but in sect. Parrvanae,and more rarely
which case they form ridges between the leaf bases in other sections, they continue to be producedfor
(Fig. IA-B). A furtherdifference between the two severalyears(up to 20 years in P maximartinezii)and
subgenera is the way the pulvini disintegrate: in also on lateral shoots. They may also be produced
subgen. Strobusthey erode slightly and become al- again on injury-responseshoots from dormantfascimost imperceptibly incorporated in the growing, cle buds and adventitiousshoots of matureplants in
smooth bark (but some exceptions occur in sect. several species in sect. Parranae. De Ferr6(1952)
Parryanae);in subgen. Pinus they become scaly and published a comprehensive study of the juvenile
break away from the underlying bark, which then stages of Pinaceae, including Pinus. from which she
inferredphylogeneticlinages and a classification.
attains a rough and scaly surface.
The secondary leaves begin to appeartoward the
The buds of long shoots, as definedhere, are really
known
as
"winter
end
of the firstgrowing season or early in the second
buds"
buds")
(also
"compound
consisting of the compacted,helically arrangedbuds in many species, but in other species (notably in
of dwarf shoots with secondary leaves, each subsect. Parryanae)not untilthe thirdor fourthyear.The
tended by a bracteatecataphyll; the latter are more secondary leaves develop on dwarf shoots initially
or less imbricateand envelop the primordialleaved axillary to primaryleaves, later subtendedby catashoot (Fig. IC).
phylls, which are homologous with primaryleaves,
on (decurrent) pulvini. Thomson (1914) reported
transitionalstages between primaryleaves and cataLEAVES
phylls in several species. Cataphylls are alternate
The leaves of pines are of four types when ac- (helically arranged) non-chlorophyllous primary
counting for all developmental stages in the plant: leaves; their relative separationincreases with shoot
cotyledons, primary (juvenile) leaves, cataphylls elongation and secondary growth (girth). They are
(scale leaves), and secondary leaves (needles) (Fig. relatively small, subulatein most species, lanceolate
ID-G). Cotyledons appear epigeous in a whorl at in a few species, thin, soon scarious, and often
the apical end of the hypocotyl of seedlings and are recurved. In several species the margins are ciliate
linear,entire, and epistomatic.They vary in number and hyaline, whereas the central part is thicker and
from 4 to 24 (P maximartinezii),the highest number pale brown to reddish brown or dark brown. Cataobserved in any plant.There are (0-)2-3 resin ducts phylls are often stomatiferouson the adaxial side
in the cotyledons and either one (haploxyl) or two (epistomatic) (Florin, 1931). They are small, with
(diploxyl) vascular bundles. According to Masters erose-hyaline margins and early deciduous in most
(1904), some diploxyl pines in the maturestate are species of subgen. Strobus; they are larger, with
haploxyl in the juvenile stage (both cotyledons and ciliate margins and persisting longer in species of
primary leaves). A similar condition is noted by subgen. Pinus (Fig. IF).
Florin (1931) for the earliest secondaryleaves in the
Secondary leaves (needles) are the permanently
extralimitalP halepensis Miller. Masters'sinference green, metabolically active leaves of the evergreen
about the evolution of diploxyl pines from haploxyl pines. They are borne on dwarf shoots axillary to
predecessors ("ontogeny reflects phylogeny") is a cataphylls on terminaland lateral shoots (stem and
conclusion we must nowadays treat with caution. branches), in clusters or fascicles of 1-8 leaves.
The cotyledons are succeeded by alternate(helically Single-leaved fascicles occur in P mnonophylla
only,
or spirally arranged)primaryleaves on the growing resulting not from abortion of one of its two leaf
stem (and, in some taxa, the initial branchlets)of the primordiain the fascicle (Masters, 1904) but rather
seedling. Primaryleaves are linear-acicular,serrulate, from a single primordium(Gabilo & Mogensen,
amphistomatic, and usually glaucous. In subgen. 1973). Common numbersof leaves per fascicle are
Strobusthey have 1-3 lines of stomataon each side 2 (rare in our region and mostly confined to boreal
of a medial line abaxially and 1-7 lines on each side species), 3, 5, and occasionally 4 and 6-8. Numbers
of the medial line adaxially.The numberof lines in are variable to some extent in almost all species;
16
Flora Neotropica
?:
?' X,'
t/f
:
'I1i
H....':i'
.
.-.:,:-'l
'. K
'.
''j'
'.
FIG. 1. Morphologyof foliage in Pinus. A, Ha. P. montezumae.B. P. lambertiana.Ga. P durangensis.Gb. P. caribaea

var. bahamensis. Hb. P caribaea var. caribaea. He. P. attenuata. J. P strobus var. chiapensis). A. Branchiet typical of
Pinus subgen. Pinus. B. Branchlettypical of P subgen. Strobus.C. "Winterbud"complex, with terminaland subterminal
buds. D. Seedling with cotyledons. E. Seedling with primary leaves. F. Types of cataphylls. G. Secondary leaves in
fascicles. H. Fascicle sheaths, subgen. Pinus. I. Entire leaf margins.J. Remotely serrulateleaf margins. K. Serrulateleaf
margins.(Magnifications X 0.5.)
very constantare the 5-leaved pines of sect. Strobus,
but even in these taxa exceptions have been found
(Thomson, 1914). Distinctionscan be made between
low-numberand high-numbertaxa in the more variable group. Commonly or prevailinglyheteromerous
leaf fascicles occur in relativelyfew species, notably
in several belonging to subgen. Pinus and in some
species in subsect. Cembroidesof subgen. Strobus.
Numbers above five are frequent in taxa of the
informal "Montezumae" and "Pseudostrobus"
groups, most of which show considerable variation
indeed in these numbers. There is a north-south
trend or dine in leaf number within the group of
heteromeroustaxa in subgen. Pinus; low numbers
are predominantin northerlyspecies, but this dine
is apparentonly on the continent as a whole and
much less evident on the subcontinent(Mexico and
Central America). Clinal within this more limited
geographical range is, e.g., P leiophylla. Supernumerary leaves, i.e., numbers in a fascicle that are
above what is "normal" in the species, are also
correlatedwith growth vigour (Thomson, 1914) and
are more often found on leading shoots of young
plants.
Secondary leaves are acicular, highly variable in
length between and often within species, ranging
from (2-)2.5 to 50 cm, rigid or lax, varyingin width
between 0.5 and 2.5(-3) mm (up to 7 mm in one
extralimitalspecies), and either epistomatic(stomata
on the adaxial side(s) only) or amphistomatic(stomata on all sides). Hypostomatic secondary leaves
(stomata restrictedto the abaxial face), as in Abies,
do not occur in Pinus except in P krempfii(as an
irregularcondition). Length of leaves is generally
correlated with both annual mean temperatureand
aridity,with the shortest leaves occurringin species
with boreal or subalpinedistributionsand in species
adapted to semi-arid environments.However, there
is also a correlationbetween leaf width and seasonal
aridity, allowing long-leaved species (e.g., P
coulteri, P devoniana, P engelmannii) to occur in
areas or environmentswith long periods of intense
insolationand withoutrain. Long, slender,lax leaves
with relatively thin epi- and hypodermallayers (see
below) are confined mostly to warm temperate to
subtropicalmesic environments.
The secondaryleaves are subtendedby a number
(usually 6-9) of helically arranged,thin, oblong to
linear, chartaceous (papery), and partly hyaline
scales, homologous with primary leaves and cataphylls, which are initially imbricate and are either
deciduousor persistent,forming a basal sheath (Fig.
IH). In the species of sect. Strobus they are early
17
deciduous. In most species of subsect. Cembroides,
as well as in P rzedowskii, the distal scales are
deciduousbut the proximalscales separateand recoil
to form a basal rosette of scales before falling from
the fascicle or sometimes with them. In most species
of subgen. Pinus (and in P nelsonii) the scales
remainimbricate,and while the thinner,often ciliate
fringes and apices erode away, the remainingparts
form a tight, persistent, and sometimes resinous
sheath around the base of the leaf fascicle. Exceptional in this subgenus are P leiophylla and P lumholtzii, which have early deciduous sheath scales as
in pines of sect. Strobus.
The secondaryleaves, while still enveloped by the
fascicle scales, are connateand togetherform a terete
fascicle. The numberof leaves per fascicle therefore
determinesthe transverseshape of the leaves, being
a sector of a cylinder: circular (terete) in P monophylla, semi-circular (plano-convex) in two-leaved
fascicles, transverse-triangularin three-leaved fascicles, and (broadly) triangular in fascicles with
leaves in fours and higher (Fig. 2). The abaxial side
is always convex and the adaxial side is straightin
non-driedmaterial.The lateralmarginsof the leaves
are serrulatein all species of subgen. Pinus and are
serrulate,remotely serrulate,or entire in species of
subgen. Strobus(Fig. II-K).
Stomataon secondaryleaves are arrangedin longitudinal (grooved) lines running along the entire
length of the leaf, as indicatedabove, either on the
adaxial face(s) only or on all faces (Fig. 2). In the
species of subgen. Strobusthe leaves are commonly
epistomatic,occasionallyamphistomatic.The abaxial
face has 1-10 lines of stomata (the terete leaf of P
monophyllahas 12-25 or more lines), the two adaxial
faces 2-10 lines each. Species of subgen. Pinus have
only amphistomaticleaves. The abaxial face of 3sided leaves has 2-17 lines, the two adaxial faces
have 2-12 lines each (2-sided or plano-convexleaves
may have more lines, but truly 2-leaved pines do
not occur in our region). The position and external
structureof the stomata and their subsidiary cells
has been the subject of elaborate studies, foremost
of which is the classic work of Florin (1931), where
a representativenumberof pine species was treated.
The stomata are generally longitudinally oriented
and monocyclic, with 4-10(-14) subsidiary cells.
The subsidiary cells encircling the stomata form a
moreor less raisedring termedthe "Florinring"(Fig.
3C; see also Yoshie & Sakai, 1985). Investigating51
species, Yoshie and Sakai (1985) observed six types
of Florinrings in the genus, of which five were found
in Mexican pines. Types E and F are characteristicof
18
Flora Neotropica
S A
FIG.~~~~~~~~~'
2.
Aaoyi
rs
eto
nPnswt
fscnaylae
aclrsse
alxl(-)addpoy
E
. nlsnii C P.st,bfi~mi. DP st~bu vt. hipenis
L).A-D Psitonof esidctsexernl.A. .mnohyla.B.
E - .
.atna
.P otravt
Poiioeindctsreil
f
uryn.HJ oiino
.P
uagni.F
3--40.)
s~~~~~~~~~~~~epa.KP.jisa.L..tpiai.(Mgiiainx
FIG. 2. Anatomy in cross section of secondary leaves in Pinus with vascularsystem haploxyl (A-D) and diploxyl (EL). A-D. Position of resin ducts external.A. P monwphylla.B. P nelsonii. C. P strobiformis.D. P stn)bus var. (hiapensis.
E-G. Position of resin ducts medial. E. P durangensis. F. P uttenuata.G. P contorta var. murraiaina.H-J. Position of
resin ducts internal. H. P cubensis. I. PRcaribaea var. hondurensis.J. P praeterniissa. K, L. Position of resin ducts
septal. K. P jualiscana.L. P tropicalis). (Magnifications X 30-40.)
19
.r;i
~'"~:----------":
'L.
";;-..:
~surface
:r------~'
', stomatal plugs C.
stomatal rows. B. Abaxial surface showing
18980).A-C. Outer view. A. Abaxial
'.. " showing ~T'l
~i~I
B
...
.:,,.
~ ':~:.:.}::,.
Abaxial
and weakl :
surfae , showing stomatal apparatus
.:
..:..::
:' ..__._.~i~~
.d F
i~~~~~~~~~~~~~~~~~~~~'~"
.:!~i~i:A- e
._'.. ..
~ii
~ii::~i.i
-::
.F.-..
'.
..
D.-F.
ring. Inner?view.D.
'~-
...
surfac
'.
.........,, .-,,~ ....~.~~~~~~~~~~~~~~~~~~

i..: ~ ::,:.....
F ,.,.::,
i~~~~~~~,.l.~[1Jl~'--"~?'M~
~
~~~~~~"
.,
t l'~._..~.~.i:A -
FIG;. 3. Scanning electronmicrographs

(SEM) of cuticles ofPinus
Little~~~~~~~~~~~~~~~~~~~~~
10170:
D-F,
montezumae
(A-C,
Pringle
1880.
-C utr
ie. .Abxil ufae,shwigstmaalrws
B Aaxa srfce
hoin
somtl lus.C
Abaxial~~
surfae,
shoi
st dvlpdFoi
in ngoaa gpaatsadwal
F
n
e
r
v e .
. Aaxaurae
edge'
daiai
osadlnitui
na oienaion.E
s .
tmtlsrfe
ofsrrl te law ithoaafe
show~~~~ingsoaa
showing stomatal rows and longitudinalstomatalorientation.E. Adaxial surface edge of serrulateleaf with stomata free
zone. F. Adaxial surface, showing stomatal apparatusand six subsidiarycells. (Scale bars = 50 ,um.)
20
Flora Neotropica
parenchyma
:
bundle "xylem
/endo-rd
de r
m~ esophyll
schlerenchyma
1000,,
resin canal
~
or duct
guard
esophyllhloem
schlerenchymo
resin
canal
sculr
?
.iil~r
epidermis
duct
scherenchymo
pore
mesophyll
epidermis
schlerenchymo
FIG. 4. Cross section of a two-needled pine leaf (inset) and cellular detail of a section from the epidermis to one of two
vascular bundles.(FromT. E. Weier et al., eds. 1982. Botany: an introductionto plant biology. Ed. 6. John Wiley, New
York.)
taxa of subgen. Strobus and do not occur in other

pines; all other types are found only in subgen.
Pinus. Types E and F are characterizedby the ring
forming an abrupt, high ridge and a rectangular
stomatalopening. The other types have more-gradually rising rings, grading to virtual absence, and
irregularto circular stomatal openings. Distribution
of epicuticular wax, formed in the stomatal zones
(lines), appears to be correlatedwith the Florin ring
types, especially at subgeneric level. Type C, in
which the Florin ring overarches the stomatal pore
resultingin a "volcanic"shape, with a small stomatal
opening, may be uniqueto the genus Pinus (Yoshie &
Sakai, 1985). Although observed in other species, it
is prevalent in subsect. Oocarpae. Generally, more
wax development was found in stomatal types with
wider openings, suggesting a correlation with gas
diffusion and (alternative) strategies in relation to
waterstress by decreasingwaterloss due to transpiration.
The anatomyof the secondaryleaves, as observed
in cross section under magnification of thin slices

(microtome-cut and usually stained), has been the
subject of numerous studies. General reviews are
given by Coulter and Rose (1886), Doi and Morikawa (1929), Harlow (1931), Sutherland(1934), Jahrig (1962), and Kwei and Lee (1963), and summarized accounts are given in Shaw (1914), Martinez
(1948), and Farjon(1984), among others. The transverse section of the leaf can be divided into three
distinct structuraland functionalparts:(a) the dermal
tissues, including the stomata; (b) the green tissue
(mesophyll), containing the resin ducts; and (c)
the stelar tissues, containing the vascular system
(Fig. 4).
The dermal tissues consist of the cuticula, epidermis, and hypodermis, in which the stomata are embedded, providing openings to the underlying green
tissue. The cuticula is a layer of cutin, a highly
polymerized mixture of lipids and waxes (Mirov,
1967), covering the outer surface of the epidermal
cells. It gives most leaves a more or less glossy
surface and appears to exfoliate in a few species,
e.g., P monophylla. The epidermis consists of a
single layer of thick-walledcells. The shape of these
cells is fairly constant but varies among individual
trees and/or species. Jahrig distinguished four
classes: square, transverseoblong (rectangular),an
intermediatebetween the two former, and oblong
(either rectangularor elliptic). The hypodermis is
considerably more variable as it consists of I to
several rows of sclerenchym cells, often unequally
distributed, with concentrations in the lateral and
axial marginsof the leaf (Fig. 2). In some species,
notably P douglasiana, it forms extensions penetrating through the mesophyll and connecting with the
endodermis,often enveloping resin ducts. The cells
are also more or less thick-walled but are often
variable in this respect as well as in shape within a
single leaf. Generally, the thickest cell walls are
found in the inner rows of multi-layeredhypoderms.
Shaw (1914) classified the hypodermalcells as uniform (either thin or thick), biform (both thin and
thick), or multiform(with a gradualtransitiontoward
the thicker-walledcells). In some species-e.g., P
ayacahuite, P strobiformis,and P strobus-epidermis and hypodermisare of similar single-cell thickness and similar cell shape and size, but their different nature becomes quite apparent in the staining
process, whereby under the action of hydrochloric
acid the hypodermis is sharply differentiatedfrom
the epidermis:the formerbecomes differentlystained
(red when safranin is used, against blue or green).
The width expressed in uni- or multicellular rows
and the polymorphismof the hypodermalcells have
been used as diagnostic characters(Harlow, 1931;
Jihrig, 1962). We have not followed this distinction
for cell thickness in the characters described for
the species, but have distinguishedbetween uniform
(homomorph)and polymorphhypodermsin termsof
their overall structure. Next in complexity to the
unicellularhypodermisof the above-mentionedspecies is the uniform but double-layeredhypodermis
of, e.g., P greggii and P patula. A simple form of
polymorphism,already mentioned, is the concentration of hypodermaltissue along the marginalareas,
while the remainderof the hypodermisremains uniform with two rows of cells. All other, usually
thicker, hypodermalstructures,including intrusions
into the mesophyll, are classified as multi-layered
(polymorph)in this work.
The stomatapass inwardthroughthe dermalcells
to the mesophyllzone and are essentially of the same
structurethroughoutthe family (Florin, 1931; Farjon,
1990a). Commonly there is a substomatalchamber
21
in the mesophyll and the outwardopening is flanked
by two guardcells, which are sunkenbelow the level
of the epidermis.The guard cells are moveable and
control the stomatalopening (Fig. 4).
The mesophyllof pines consists of largecells with
thin, undulatedto invaginatedwalls, filling the entire
space between the hypodermis and the endodermis
and enveloping the resin ducts. They contain organelles, among which are plastids (chloroplasts),mitochondria, ribosomes, and the nucleus. They are the
cells from which increasingly taxonomically informative molecular data, especially chloroplast-DNA
(cpDNA) sequences, are obtained. The resin ducts
are narrowtubes usually runningthe entire length of
the leaf. They are not connected to cambium or
xylem resin ducts. They have walls of thick-walled,
sclerenchymatouscells of similar appearance and
structureas the hypodermalcells and give a similar
chemical reactionto the stainingreagent.These cells
may be laid down in single (most common) or
multiple rows; in the latter case they decrease in
thickness and size inwardly (Fig. 4). Their number
and position relative to the dermal walls of the leaf
are highly variable among species and can be used
as diagnosticcharacters.The numberof ducts within
species is generally more variablethantheirposition,
but some measure of polymorphism is present in
almost all taxa for both characters.The numberof
resin ducts in a leaf ranges from 1-12 or more in
Latin American pines, and is even higher in some
otherspecies. Since Engelmann(1880a), the position
of the resin ducts has been used as a characterto
classify the pines; since Coulter and Rose (1886),
these positions have been termed external, medial,
internal,and septal. External(as "peripheral"in Engelmann, 1880a) ducts are situatedagainst the hypodermonly. In some cases the hypodermalcells merge
with those surroundingthe resin duct and it is then
difficultto distinguishtheirorigin. Medial(parenchymatous)ducts are free in the mesophyll, touching no
dermalstructure.Internalducts touch the endodermis
only. Septal ducts, additionallyrecognized by Shaw
(1914), touch both hypodermand endoderm, sometimes interconnectedthroughintrusionsof the hypoderm, forminga septum. Most species are characterized by a single position (Fig. 2). In a comprehensive
study of nearlyall species in the genus, Jahrig(1962)
never encountereda combinationof more than two
positions, but considered medial and internalducts
as occurringwith equal frequencyin severalMexican
pines of subgen. Pinus. In several species there is a
combinationin which one positionprevailsbut where
one or a few often smaller ducts occupy one of
Flora Neotropica
22
the other positions. In many species with variable
numbers of resin ducts, certain ducts, with a fixed
position and slightly larger,are always present. We
consider these primaryducts and the others subsidiary ducts.
The stele of the secondary leaf is a tubularstructure, usually terete, but often more or less triangular
or oval in cross section, being correlated to the
transverse shape of the leaves. Its wall, the endodermis, consists of a single row of relatively thin or
partly thick-walled, globose to oval cells. The outer
cell walls in several species are thickened, and this
is often used as a diagnostic character,as it appears
mostly constant. Only a very few species are polymorphic in this respect, e.g., P leiophylla var. chihuahuana. Inside this wall is the vascular strand(s)
or bundle(s) embedded in a matrix of transfusion
tissue. Often there are strands of sclerenchym cells
which intermittentlysurroundand often divide the
vascular bundles (Fig. 2). The vascular bundles are
either single or double in pines. This distinctiongave
rise to Koehne's sections Haploxvlon and Diploxylon. The double bundle is usually obvious, even
when (partly) connate, in at least the majority of
leaves of species of subgen. Pinus. The vascular
bundle consists of two connate strands, an adaxial
xylem strand,and an abaxial phloem strand(Fig. 4).
In diploxyl pines they are either distinctly separate
or connate on the xylem side; in a few instances
they may be entirely connate and seemingly form a
single bundle,as in all haploxyl pines. An interesting
feature has been found in the haploxyl species P
maximartinezii,which has a strandor wall of sclerenchyma cells separating the phloem of the single
vascular bundle and occurring in the majority of
leaves observed (Rzedowski, 1964).
BARK
The bark of pines, as commonly mentioned in
descriptionsof species, is the outerbark(rhytidome),
which often forms characteristicbark patternsespecially on the (lower) bole of maturetrees. The bark
has a complex structureand consists of a variety of
tissues, separatedinto two zones outside the vascular
cambium: the inner bark (phloem) and the outer
bark (rhytidome). From the vascularcambium layer
outward,the barkconsists of the secondaryphloem,
the primarytissues still existing outside the secondary phloem, the periderm, and dead tissue outside
the periderm.The vascularcambiumproduceswood
(xylem) on the inside and phloem on the outside.The
first peridermcontains a cambium layer (phellogen)
which produces the phellem (-- periderm) outward
and a thin layer of phelloderm inward. Commonly

the barkof pines is divided into threezones: phloem,
cortex, and epidermis (Mirov, 1967; Farjon, 1984);
but we have followed the terminology proposed by
Junnika (1994) here, as it is more accurate. The
outer bark is thin on young trees and branches,
but increases in thickness by the accumulation of
peridermlayers added by the phellogen. Simultaneously, the outermostlayers of peridermexfoliate, so
thatthe ultimatethicknessof the outerbarkis defined
by the age of the stem and net result of these two
processes. In some species exfoliation startsearly, in
others late.
Bark patternsin pines result mainly from fissuring
and scaling (Fig. 5). Fissuring is caused by the
expansion growth of stem and (larger)branchesand
results primarilyin lengthwise cracking into fissures
separated by ridges. In most species these fissures
are irregularlyconnected. Incomplete shedding of
scales results in horizontal cracks that widen by
further wood expansion, forming tessellated or
polygonal patterns.The longitudinalfissuresare deep
or shallow, with "shallow" defined as a depth less
than one-half the total thickness of the bark. They
can be V-shaped, irregular,or square-shaped.The
ridges are usually flattened,sometimes rounded,and
often reticulate. Flakes are usually irregular. On
young trees and the upper stem and branches they
are commonly papery, on the lower trunk they are
often thick, harder, and plated or shaggy. Colour
patternsresult mostly from scaling, whereby exfoliated flakes reveal younger, less weatheredsurfaces,
or from fissuring and cracking, which may expose
varicoloured inner bark. Illustrationsof patterns of
bark have been publishedby Mirov (1967), Howard
(1971), Farjon (1984), and Perry (1991); the latter
author provides small photographsof boles for all
Mexican taxa. Although some generalizations can
certainlybe made-such as the long-retainedsmooth
bark of Pinus sect. Strobus, in which the formation
of periderm is delayed, or the often tessellated or
polygonally patternedbark with large fissures and
ridges in subsect. Ponderosae-identifying species
on the basis of bark charactersshould be practiced
cautiously. Some species (e.g., P nielsonii in our
region) have very characteristic barks that make
identificationat species level possible. In most taxa,
however,thereis considerablevariation,and similarities between taxonomicallywidely separatedspecies
are common. The anatomyof bark of 10 species of
Pinus from the southeasternUnited States (all subgen. Pinus, one of them, P elliottii var. densa, in-
23
~
!~;:j~~~~~~~~~~~~~~~~~~~~~~~~~~~a
~
~ ::i;;i;? '
'::
.
,:.,~~~~~~~~~~
~~~~~~~~~.-~
~ ~
'.'?.?.
~.:"L"....!::-."
X.-:.::'.?
... ..... ....

...'.'..'..
.~....:..:
;:...
: ::
'
...
...
:~~...
..
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.......-...:
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~~~~~~~~~~~~~~~~~~~~~~...
.a?'.'....-.'..
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:,
.........~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
.: .";';.~'~.':.
.....,;,..!~;;~,
...
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...
i'!Si~~~iii~i~i::;:::::.;..
: . . . . . . . . . '"....
. ..... .
'
,~,.
,,~,,..,:,:.;..
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.
'
'N::.; ,
:::::::::::::::::::::::::
:. : : : : :
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~ ~ ~~ ~ ~ ~ ~ ~ ~~~~~~~~~
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..".,,':'...'.:
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i
:i'~*.~~
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:"i~i;i"~::'~'":::;'"..'..
.:::.::::::
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i',;i;',,ii:::',:i/,i::;:;i~~~~i~i,::,:,',',i..,.!
..:.:i;;
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? ::.:i.
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nelsonii.lc~5Ii3:
'.%:
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ill
:i~~~~~~~~~~?:L?~~~~~~~.?
X.:':..: .
.
-
A~~~~~~~~~~~--
?-?
a---- , ~~
i.~~~~~~~~~~~~~~~
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_ ~ . ;~<.....~~~~~~~~~~~~~~~:!,......
7~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~b:
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.
i;?;;;i::i::i!
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:
:::::::::::::::::::::::::::::~~~~~~~~~~~~~~~~~~~~~~i
,.----~-:;??- i~~~~~~~~~~~~~~~~~~~i,;?:E~~~i::
tlW~.~'
~' ":'~~~
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-"~..."~-,,..';~~~_~...
~
~~~~~~;:.
i~.
....
?
C.
"gi.."
:;:::utlos?.
b.s.
'D:.:::
~ ~ tj
~.
'
:..oiina:'Z..:i::
E:::!::. n.........:'.
F.::':::':"...':.:P::.:::
::s.:i. H::. P
nwvi::.;fiez'/?.1.'
'..:;,::~'. .......~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~1
:?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
..!....,
,x~~~~~~~~~~~~~~ ::
': .':''..
:'::':;""::R:
::ii....
....:,:n~~~~~~~~~~~~~~~~~~~~~:i'
E
:~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~?
"..!.,,i!'
i:,::
;'.- '.' :
FIG.
5. Trunks
wegi. . Ppeudstrbu.
ine
pten.
A.:
PiQf~iittusih
ilrntbr
of Mex~~~icnadCtrAica
DP.deonina.E.P.
axiini. . Pocara.
. .. zedwsii.H.
leiophylt. B. hart
..maxntatiezi. I P
24
Flora Neotropica
cluded in our region) has been describedby Howard small (1-2 mm) and subpeltate (free only at the
lower end) and the uppermarginis entire. In species
(1971).
of subgen. Pinus, the microsporophyllis larger (ca.
2-3 mm diam.), peltate, with an erose-denticulate
REPRODUCTIVEORGANS
uppermargin.The outer surface in the formersubgeThe genus Pinus is monoecious. Male and female nus is smooth and in the latterradiallystriated.These
reproductiveorgans (strobili) are separatefrom each differences are true for all species studied here, but
other, arising from separate axillary buds, though perhapsnot for the entire genus. Furtherdifferences
commonly occurringon the same branchesand occa- are found in the size of the entire pollen cone, which
sionally on the same long shoot. In older literature is smaller in subgen. Strobus.In species of subsect.
(e.g., Shaw, 1914) and in many papers by foresters Cembroides,pollen cones are not more than 10 mm
on the subject, angiosperm terminology such as long and in P rzedowskii only 5 mm; they also
"flowers,""anthers,""catkins,""staminate,""pistil- extend up to more than one-half of the length of the
late," "flowering,"etc., are commonly used in de- shoot, forming spike-like clusters. In most species
scriptions of reproductiveorgans and phenology of of subgen. Pinus in our region, the pollen cones
conifers. This is to be avoided, as it not only leads form short, "capitate"clusters, but there are some
to confusion but, more importantly,is misleading exceptions and this trait is variable in many species.
since both structure and reproductive biology are The microsporophyllsof some species are initially
red or purplish,of others yellowish green or yellow,
very different from those in angiosperms.
and most turn yellowish brown to brown. There is
again uncertaintyaboutthe fixity of these differences
POLLENCONES
and, as has been demonstratedfor ovuliferous cones
Pollen cones arise from axillary buds in the proxi- in Pinaceae,the reddishcolorationmay be influenced
mal part of the compound or aggregatebud through by environmentalfactors. Due to the minimal intera modification of dwarf shoot primordia (Mirov, specific differences, or the lack of even these, pollen
1967); their subtendingbractsare homologous to the cones rarely receive attentionin descriptions.
scales of fascicle sheaths. In most species of subgen.
Pinus, buds of incipient pollen cones are slightly
SEED CONES
larger and paler than those of sterile dwarf shoots
The seed cones (ovuliferous cones) are the most
above them; in most species of subgen. Strobus
they are inconspicuous(Shaw, 1914). They are also characteristicstructureof the coniferous plant. Much
distributedover a greater length of the shoot in the has been writtenabout their ontogeny, structure,and
lattersubgenus, especially so in the Mexican species homologies. We shall not attemptto fully summarize
that literature,but we would like to recall the fact
P maximartineziiand P rzedowskii.
The pollen cone (microsporangiatestrobilus, Fig. that the studies of fossil primitiveconifers by Florin
6A-B) within the Pinaceae is much less diversified (1938-1945, 1951) established beyond doubt that
than the ovuliferous or seed cone (Farjon, 1990a). the seed cone of Pinaceae is a compound structure
It is a relatively small, weak and soft, ephemeral, homologous to a long or branchingshoot. Ontogecompoundstructure,consisting of an axis with many netic observations confirm this interpretation.Lemhelically arrangedmicrosporophylls,usually ovoid- oine-Sebastian(1975) found stomataon both apophoblong to cylindrical and up to a few centimetres ysis and umbo of all seven species (in both subgenera
long. Like the leaf fascicle, the pollen cone itself is of pines) studied. This, and a reduction in vascular
surrounded at the base by a number of helically strands from seed scale proper to apophysis, gave
arranged scales, but these are usually broader and her reason to designate the formeras "axial"and the
not ciliate-margined.The axis is thin and flexible. latteras "foliar,"i.e., homologous to a leaf. Stomata,
Dependent on their length at maturity the pollen however, are not restricted to leaves of vascular
cones of several species become subpendulousafter plants or their homologues. Comprehensivedescripshedding the pollen. The microsporophylls (Fig. tions of cone morphologyare given by Shaw (1914)
6Aa-b, Ba-b) are borne on minute stalks and termi- and Klaus (1980). The cones originate from "winter
nate in a (sub-)peltate structurebearing two pollen buds." The peduncle, varying in length from a few
sacs (microsporangia) on the lower adaxial side, millimetres to 6-7 cm in P nelsonii and up to 12
which dehisce by means of a longitudinal slit. In cm in P. lambertiana,is an axillary shoot positioned
species of subgen. Strobus, the microsporophyllis subterminally,or sometimes laterally,on a branching
25
.. .
I'
'
'" i r:
..
....
'igs
adnate,~~~~~~~~
'tfctv
ig.....
fetv
rtcuat,
K . :.....h
..~k,ruri. I~ i
i.
?hni.
." .' .-:.:"'ic?lrtui
vestigial ings remainin on the scale(a)."('..,
seeds (b)with articulae,
b = seeds. N,~ nlonc~l~lt!lkr
~~crrtunucr~cr.M,I! strr,hif~rrllis seeds

AJ
wings; a = seed scale with seed cavity tone seed

with adnate, ineffective ~bortive)'
:L.
M.b. -.
N- b:
FIG. 6. Morphologyof pollen cones, seed scales, and seeds in Pinus. A, B. Pollen cones microsporophylltype. a =
B. P (subgen. Pinn.s) maximinoi.C. Seed scale type
abaxial view. b = lateralview. A. P (subgen. Strn)bus) monoplh U11(.
of P. (subgen. Strobus. sect. Strobs)
strn)ifonms.
a = abaxial view, b = adaxial view, I = apophysis. 2 = terminal
umbo. D. Seed scale type of P (subgen. Pintos)dlevonni(a. a = abaxial view, b = adaxial view. c = lateral view. I =
apophysis. 2 = dorsal umbo. 3 = transversekeel. E-H. Increaseddevelopment of apophysis and umbo in seed scales
F. P cnribaea var. caribiena.G, H. P. ps.eudo.strobsvar. puk-ensis. 1, J. Seeds with
of P. subgen. Pinn.s.E. P.jlis-a.
adnate,effective wings. I. P cubensis. J. P lamberitun. K, L. Seeds with articulate,effective wings.. K. coultei. L.
seeds with adnate, ineffective wings; a = seed scale with seed cavity (one seed abortive).
P attenuila. M. P strohin)bifonns
b = seeds. N. P mnoplylla seeds (b) with articulate,vestigial wings remainingon the scale (a). (Magnifications x.5.
26
shoot. It is initially erect but commonly becomes
recurvedor pendulouswhile the cone grows to maturity.Cones may be solitarybut more often are placed
in pseudo-whorls,and in some species many cones
appear each growing season. There is considerable
variation in the persistence of the cones, especially
within subgen. Pinus, from deciduous shortly after
seed release to almost indefinitelypersistent(e.g., P
nuricata, P attenuata, P radiata, P greggii). Shaw
(1914) assumed an evolutionary trend in the genus
leading from symmetrical,readily opening, and deciduous cones to asymmetrical,serotinous, and persistent cones (Fig. 7). Cones of pines in subgen.
Strobus are normally deciduous, with the point of
detachmentat or near the base of the peduncle. In
several species of subgen. Pittus there is a partial
disarticulationof the cone, whereby the proximal
scales remain with the persistent peduncle; in P
praetermissa these scales even fall separately.With
P nelsonii there is articulationat the apex of the
peduncle. In this and some other species (e.g., P
cembroides and its allies, P pinceana), there is a
(very) thin attachmentof the scale to a (weak) rachis,
which often leads to early disintegrationbut usually
not before the cone has fallen. The peduncle is
covered with helically arrangedcataphylls;normally
there is no development of vegetative dwarf shoots,
though they are known in P nelsonii (M. P. Frankis,
pers. comm.). The ovuliferous scales are axillary to
small cataphylls, and helically arranged,forming a
determinate cone. Shaw (1914) has explained the
phyllotaxis of pine cones. The two subgeneradiffer
primarilyin there being a relatively low numberof
scales per unit length of axis in Strobusand a higher
number in Pinus. Due to the varying shapes of the
cones, their phyllotaxis is variouslycomplicatedand
deviates from the theoreticalarrangementin spirals
on a cylinder. Unlike in other genera (e.g., Abies and
Larix), no proliferation of a vegetative shoot has
been observed in the genus. The seed scale, with its
subtending bract, is homologous to the vegetative
dwarf shoot (and to the fertile dwarf shoot in several
fossil conifers).
At pollination, the seed scale (ovuliferous scale)
consists of a flattened structure in the shape of a
short, apiculate to acuminate spoon. The seed scale
in pines is a highly specialized organ, resultingfrom
the ontogeny of the cone and the periodical growth
phases on the apophysis as concentric differentiations. After pollinationthe seed cone (megasporangiate strobilus, Fig. 8) closes its scales by expansion.
Subtending bracts are still visible at the beginning
of this stage but soon become submerged between
Flora Neotropica
the growing scales. Growth then stops until the
following growing season, often six months or more
later. Pinus nelsonii is unique in the genus in not
having a definite resting period (Shaw, 1905a, 1909,
1914); as a consequence, its umbo is rather illdefined and poorly differentiated.The conelet at this
stage (referred to in the descriptions as "immature
cone") exposes the later umbos. The umbo is often
armed with a spine (crista) or short prickle (mucro)
in species of subgen. Pinus; it is obtuse (mucro
absent) in sect. Strobus. In a few species of sect.
Parrvanae (e.g., P cembroides and its allies), the
mucro is placed eccentrically,i.e., not on the line of
the transversekeel (Klaus, 1980). The outer edge of
the scale later forms the transverse keel, which is
continuedon the apophysis(Fig. 6Da-3). Pines show
two phases of cone expansion, in the second phase
the apophysis (the part of the scale exposed when
the cone is still closed) is formed and the seed scale
bearing the seeds grows. In this phase, the umbo
appears to be terminal in species of sect. Strobus
and dorsal in other pines occurring in our region,
belonging to both subgenera (Fig. 6Ca-2, Da-2).
Biennial cone maturationvaries among species-it
takes mostly from about 17 to 24-25 months-but
it is fairly constant within most species. In our
region, P oocarpa may have the shortest maturation
period. In a few species (P leiophylla in the Flora),
growth of the cone progresses only slowly in the
second growing season, and a thirdseason is required
to reach maturity(triennialcone development).This
results in a distinct band between the umbo (first
year of growth) and the lower part of the apophysis
(third year of growth). This concentric band was
termed the "vallum"by Klaus (1980).
The apophysis is a conspicuous subterminalstructure of the cone scale, which has attained a high
degree of diversification in the genus separating it
from other genera in the family (Shaw, 1914; Klaus,
1980; Frankis,1989; Farjon,1990a);consequentlyits
charactershave frequentlybeen used for taxonomic
delimitations.The texture of the apophysis is markedly different from the anteriorpart of the scale. It
is also more extensive on the abaxial side than it is
on the adaxial side, where it consists merely of the
outer rim of the scale (Fig. 6D). It is simplest in
species of sect. Strobus,consisting of the (slightly)
thickenedpartof the scale which was exposed in the
second growing season, terminated by the obtuse
and commonly resinous umbo. It has a terminal
orientationand growth pattern,although outside our
region some transitionalconditions toward the following type exist in subsect. Cembrae. Its abaxial
27
Morphology and Anatomy
,j
FIG. 7. Morphology of seed cone types in Pinus. A-E. Seed dispersal anemochorous.A. P. strobus var. chiapensis.
Pendulouscone type with quickly parting seed scales. B, C. Subsessile cone types with slowly partingseed scales. B. P.
montezumaevar. gordoniana. C. P durangensis. D. P pseudostrobusvar. apulcensis. Semi-persistentand subserotinous
cone type with armored,slowly parting seed scales. E. P attenuata. Persistent and serotinous cone type with armored
and closed seed scales. F. P quadrifolia. Seed dispersal zoochorous. Deciduous cone type with "wingless"seeds. (Magnifications x 0.5.)
28
Flora Neotropica
ovule
/~. ~-.-" --
"~._
...polk..
?;"
megasporo)mcte lh
ni
pltiropylt
ovufrus
'-;
- wrs....me
vuitcm
.
y't="'.
.rs"'r
sca
'
-~
.t:?':" .i
ov ueinl ,
lom
Ochp
st~~~~~~~robsius~~
uvn#v->D
ok
. uL
plmiunse
ion
t ibe_
u
vegeiaiive mmdci
sinu ci
emuinMion
--^~
SUSri6flSCf
co.yeedys'
- -- -
-- c
"Staminate" strobilus, pollen, development of the pollen tube; embryo and germination of seed. For further explanation
see text. (From T. E. Weier et al.. eds. 1982. Botany: an introductionto plant biology. Ed. 6. John Wiley, New York.)
surface is often irregularor longitudinally rugose.

In some taxa, e.g., P avyacahuitevar. veitchii, the
apophysis is distally contracted and elongated or
recurved/reflexed.
The apophysis in other pines of both subgenerais
more complex and its orientationis dissimilar: bordered in the closed cone on all sides by the margins
of other apophyses, its growth is a dorsal extension
or thickening (Shaw, 1914). The outline is rhombic,
or irregularlypentagonal,or in other cases it has a
roundedupper marginand an angularlower margin.
It can be nearly flat, conical, or gibbous, or form a
conspicuous and often curved protuberancein some
cases (Fig. 6G-H). Extreme apophysaldevelopment
occurs, e.g., in P attenuata, P coulteri, P muricata,
posed to sun and rain are most developed (see

above). Almost all apophyses with dorsal development are transversely keeled as a result of preexpansive connection and partial overlapping; mucro, umbo, and transversekeel are (retained)features
of an earlier phase in the cone ontogeny. Klaus
(1980) distinguishes three ontogenetic phases, each
reflected by its own morphological features of the
mature seed cone: I) strobilus at pollination, with
free ovuliferous scales, transverse keel, and crista
(mucro);2) firstyear's cone, with expandedovuliferous scales with umbo, transverse keel, and mucro;
3) second or third year's cone, with lignified, enlarged seed scales with keeled apophysis. umbo,
transversekeel, and mucro. From the centre to the
and P pseudostrobus var. apulcensis. On such properiphery,therefore,one encountersconcentric zones
nounced apophyses the umbo may be extremely of youngerphases. A secondaryontogenetic differendeveloped, too, which merely indicates that this ex- tiation is responsiblefor differences in size and shape
cessive growth started in the earlier stage of cone of mucro, umbo, and apophysis between proximal,
development,interruptedby the restingseason. There central, and apical scales. Whereas such differences
is invariablya longitudinaland one-sided asymmetry may be typical of certain species-e.g., the strongly
in the development of abnormallyshaped apophyses recurved proximal scales of P. avacahuite or the
on a cone, whereby those on the section most ex- straight proximal scales of P strobus var. chia-
Morphologyand Anatomy29
pensis-comparisons between species are to be made
on scales of the same zone. In general,differentiation
decreases toward the apex of the cone.
The anterior part of the seed scales of a mature
cone ("seed scales," in the species descriptions)consists of two different structuraltissues and also the
thin tissue that forms the seed wings. The abaxial
tissue has much greater lengthwise swelling and
shrinkingpropertiesassociatedwith moisturecontent
thandoes the adaxialtissue, which consists primarily
of a numberof longitudinalxylem strands(Harlow
et al., 1964). This differential provides the opening
mechanism of pine cones through parting of the
scales by recurving.There are considerable differences between species, both in the conditions required for seed scale parting and in the extent to
which they can spread.The serotinouscones of some
species usually open only after excessive heating by
tire ("closed-cone"pines), whereas in other species
the scales part only after considerable time (years)
but under more normalclimatic conditions of insolation and dehydration.Cone serotiny is correlated
with persistence. Shaw (1924) further observed a
correlation with obliqueness ("asymmetry")of the
cone resulting from a lesser or greater inequalityin
the growth of both the seed scale proper and the
posterior tissues forming the apophysis and the
umbo. The condition is indeed unique in the family
if (slight) curvatureof the cone without substantial
differential growth of scales is not considered.
Whereasthe symmetrical,cylindriccones of subsect.
Strobi are pendulous on straightor recurvedpeduncles, the cones of many of the closed-cone species
are reflexeddown the branchon very shortpeduncles
and become effectively sessile at maturityby thickening of the branch. Of the two conditions, the
formeris consideredthe plesiomorphicstate by most
authors(e.g., Shaw, 1914; Klaus, 1980). The greater
development of the apophysis on the side of the
cone persistentlyexposed to sun and weathercan be
interpretedas an adaptationto protectthe developing
seeds (Shaw, 1914), thereby enabling the species to
occupy environmentswith more extreme daily and
seasonal temperaturefluctuations.
SEEDS
The family Pinaceae is characterizedby two anatropous ovules on the proximal end of the adaxial
side of each ovuliferous scale (Fig. 8). The seed
containsan embryo,embeddedin the female gametophyte, which in turn is surroundedby the nucellus
and the seed coat or testa (Fig. 8).
Comparedto other genera in the family, the seeds

of Pinus are much more variable in size; this is in
part correlatedto variations in cone size, but Pinrs
cembroides, P culninicola, P m7onophylla, P quadri-
folia, and to a lesser extent P nelsonii and P

pinceanCa have proportionallyvery large seeds. The
largest seed in the genus is found in P maximartinezii, which reaches 28 X 12 X 10 mm, in an
exceptionally large cone. The smallest seeds are ca.
4 mm long. Shape is variable,too, but most are (ob-)
ovoid and slightly flattened(Fig. 61-N). In the pinyon pines (P cembroi(iesand allies) thereis a marked
difference in the thickness of the seed coat among
the several species. Thick seed coats are a protection against predation by mammals and select for
certainbirdsthat play a role in dissemination.Unlike
in some other genera, there are no resin vesicles in
the seed coat. The outer surface is usually smooth
and dark grey-brown to blackish, but light brown
seeds, with or without dark spots, occur in several
species.
The seed wing is not derived from tissue of or
surroundingthe seed coat, as Shaw (1914) understood it, but from tissue on the adaxial face of the
seed scale. Seed wings are, unlike those of other
genera in the family, highly diversified within the
genus Pinus (Frankis, 1989; Farjon, 1990a). This
diversificationshows adaptationsto different strategies of seed dispersal and is evolutionaryand taxonomically significant. Basically there are two types,
related to wing attachment:articulate and adnate
(Fig. 6K, J). Even in species with vestigial or "absent" wings, remnantsof basal wing tissue are present on the seed scale, on the seed, or on both. The
first author to observe this was Engelmann(1862),
in a footnote about P cembroides, but his remarks
were not acknowledged,nor the fact observedagain,
by subsequentdescribersof pine morphology(Masters, 1904; Shaw, 1914; Martinez, 1948; Mirov,
1967), who all made a primarydistinction between
wingless seeds and winged seeds instead. However,
Small (1903) and Frankis(1989) correctly observed
the situation. In the articulate seed, the wing of a
full-grownseed is loosely attachedto the seed, which
is held either by two narrow,claw-like extensions of
the wing or by these and an extension covering other
partsof the seed on the adaxial side (Fig. 6K, I). It
has been suggested that the outer layer of the integument plays a part in the formation of this latter
section of the seed wing membrane(Schnarf, 1933).
These appendages hold to the seed in all species
with well-developedwings thatdetachfrom the scale
during the drying of the cone and the partingof its
30
Flora Neotropica
seed scales. Upon dispersal they are normally still

attached(and therebyeffective) but easily separable.
Adnate wings remaininseparablefrom the seed upon
dispersal and cover the seed partially,usually more
so than articulate wings (Fig. 6I-J). All effective
seed wings are thin and membranous,though a few
species (e.g., P coulteri) have very thickened seed
wing bases, filled with light, spongy tissue. The
outline of effective wings is oblique, usually with
one more or less straightside which was adjacentto
the other wing on the scale. The colour is usually a
translucentlight brown with darkerstripesor streaks.
"Wingless"seeds occur in two ways, depending on
whether the wing remnantis adnate, in which case
it remains attached to the seed, or whether it is
articulate, in which case it usually remains loosely
attached to the scale and leaves no trace (or a very
faint one) on the seed.
There is evolutionary significance in the various
modes of seed wing attachmentand development.
The more common anemochorous dispersal would
be most effective when the seed is small relative to
the wing and the wing adnate. Indeed, this situation
is found in Pinus subsect. Strobi, in which the cone
is pendent and opens readily on maturityto release
the seeds simply by gravity. Trees of several of its
species are very tall (P ayacahuite, P lambertiaaa)
and theircones are situatedprimarilyon the extremities of the higher (main) branches. Articulate but
effective seed wings may be of adaptive value in
dispersal after fire-they help carry the seed some
distance-but with an (easy) articulation of the
highly flammable wing, chances of the seed being
burnedare probablysignificantlyreduced.Naturally,
dispersal under normalconditions requiresan effective wing only; the articulatewing is usually as good
as an adnate wing in this respect. Perhaps the loss
of the wings serves to camouflage the seeds against
predation.The reductionof the wing below effectiveness, and/or its loss before dispersal, is connected
with zoochory (Lanner, 1982). "Wingless" seeds,
occurringin our region in both sect. Strobusand sect.
Parryanae-and also in the only distantly related
Mediterraneanspecies, P pinea-evolved in coevolution with birds (jays and nutcrackers;see Tomback & Linhart,1990; Mattes, 1994). They are therefore examples of parallelismthrough selection of a
morphological potential inherent in the genus as a
whole: a trend towardcompletion of the articulation
of the seed wing. Enlargement of the seed and
strengtheningof the seed coat are othertrendsrelated
to this highly selective type of bird dispersal, which
in some cases is connected with a single bird species
(Mattes, 1994).
WOOD ANATOMY (by Ian D. Gourlay)

ANATOMICAL DESCRIPTION OF
PINACEAE (according to Greguss, 1972)
Cross section-Annual ring boundariesnormally conspicuous. Resin ducts presentor absent. Epithelial cells thin or thick walled.
Some genera possess axial parenchyma.
Radial section-Ray tracheids present or
absent. Some species have rays without transverse tracheids or resin ducts. Some species
have both thick and thin walled ray parenchyma cells. Ray tracheidsmost often characterised by different forms of dentation.Walls
of the ray tracheids smooth Pinus, group
Haploxylon = subgen. Strobusor dentate [Pinus, group Diploxylon = subgen. Pinlus].The
end walls of the ray cells exhibit sieve-like
structures.Cross fields exhibit one to six (seldom eight) pits, large oval pinoid pits filling
out the entire field, one to four small round
piceoid pits with oblique slit-like aperturesor
taxodioid pits. Some genera contain xylem

parenchyma,others lack it (Pinus). Some genera exhibit spiral thickening in their late tracheids, other genera have it in the axial and
transversetracheidsof all their species.
Tangential section-Rays one to 45 cells
tall, most of them typically uniseriate,except
those including resin ducts. End walls of longitudinal parenchymacells nowhere smooth.
GENERALINTRODUCTIONTO PINUS
The wood of pines, like that of all conifers, is a
simple structurecomposed of tracheids and parenchyma cells. The tracheidshave lignified walls characterisedby the presence of borderedpits. The rays
are usually one cell wide and rarely visible without
a lens. These vary in depth, the central rows consisting of parenchymawith simple pits on their radial
walls, the upper and lower rows consisting of tracheids with borderedpits.
31
Wood Anatomy
The genus Pinus is readily distinguished from
other genera by the presence of resin ducts distributed throughoutthe growth ring and surroundedby
thin-walledepithelialcells. It is in these ducts during
theiractive life that the productionof oleoresin takes
place. The ducts are formed as intercellularspaces
between the resin-producingthin-walled epithelial
cells, which are often damaged in the course of
sectioning. Resin ducts/canals are often thought to
be developed as a response to injury; however, in
the pines this is not necessarily so (Mirov, 1967).
The product of this response to wounding in the
pines is often exploited in commercial turpentine
extraction.
The wood of pines frequently shows a marked
transitionbetween early and late wood, togetherwith
a generally reddish-colouredheart. It is also, on the
whole, light, elastic, strongfor its weight, and widely
available. Close-grown and plantationmaterialwill
normally produce stems of straight form, ideal for
a wide range of construction uses. This structure,
therefore,can in some species (e.g., Pinus strobus)
present a fairly uniform wood, easy to work and of
reasonable durability when there is a high resin
content. Pine timber can also be used for producing
paper pulp, although it is less suitable than that of
some of the other conifers, such as spruce (Picea),
because of its resin content.
GROUPINGOF SPECIESACCORDING
TO ANATOMICALFEATURES
Several attemptshave been made to furthersubdivide the genus, most commonly into two groups, the
"hard"and "soft" pines (referred to in the United
States as the southern yellow pine group and the
white pine group).
The "hard"pines (Diploxylon, Fig. 9) possess two
vascularbundlesper leaf and their wood is generally
harder,heavier, and darker in colour, with a much
clearer demarcation between early and late wood
zones within a growth ring. The walls of their ray
tracheidsreveal thickened,uneven, tooth-likeprojections (dentation).Resin ducts are numerous.Within
this group are the common commercial northern
timber trees, Pinus sylvestris and P. nigra, together
with the majorityof NorthAmericanspecies. Almost
all of the economically importanttimber trees of
Mexico and CentralAmerica and all the Caribbean
pines also belong to this group.
The "soft" pines (Haploxylon, Fig. 10) possess
one vascular bundle per leaf, are lighter in weight
and colour, and show a more gradualtransitionfrom

early to late wood zones. Resin ducts are not as
numerous. Their ray tracheids have smooth walls
and include P cetmbraof Europeand otherAmerican
species including P. strobus and P lambertiana. In
Mexico, P avacahuite and P strobiformisare commercially importantmembersof this group.
In the "hard"pines, the bark is rough, grey, and
scaly, thickly covering the trunks of even young
trees. In the "soft" pines, bark formationis delayed,
giving the appearance in young trees of a very
smooth trunk.Scaly bark formationdevelops at the
base with age, gradually extending up the tree but
often with the uppertrunkremainingsmooth (Perry,
1991).
Jones (1924) subdivided the genus according to
needle grouping: (a) two-, (b) three-, and (c) fiveneedle pines:
a. Autumn wood of each ring is well-defined with
numerous resin ducts. Rays are uniseriate and
multiseriate(the latter usually containing a horizontal resin duct). Tracheidcells in the rays usually occur only towardthe upperand lower margins, and there is pronouncedirregularthickening
of the walls of the ray tracheids.Parenchymatous
cells occur medianly,with large simple pits.
b. Thoughsimilar in most characteristicsto the twoneedle pines, the smooth walls of the raytracheids
are in strong contrast with the jagged walls of
the two-needle pines.
c. Though similar to two- and three-needlegroups,
the rays differ in thatthe simple pits are considerably smaller and more numerous.Moreover,the
ray tracheidshave comparativelysmooth walls.
It should be noted that our currentunderstandingis
that the numberof needles in a cluster is not a good
phylogenetic marker(see Mabberley,1987).
A furtherclassificationof the genus, on the basis
of form of smooth or dentate ray tracheids, was
presentedby Phillips (1941). He stated that species
identification is not generally possible unless the
geographical origin of the material is known, but
detailed microscopicexaminationwill usually permit
a specimen being classified underone of the following seven groups (with added Mexican species given
in parentheses)recognized by Rol (1932):
I. Strobus.Ray tracheidswith smooth walls; crossfield pits 1-3, large, tangentialwall pits on latewood tracheidsnumerous,e.g. P strobus,P cembra, P. koraiensis, P.lambertiana. (P avacahuite
and P strobifornmis).
32
Flora Neotropica
.88~'~:: .:~88~88I;:::
. .. . ........
.............
..............
":?'-~
BBBBsBg:,:?::
B~B~:.:
?????~
...?.?:::?.?:-'?Bg~H~k~:r:
?????:
?????
??Y
9Bh~'P~n~:
F
vertical
res
resinducwithi..
I i
..anG. dn
show.
..........t..
eu a rc l 9.P.P.A....
y.demarc..e..zone.between.early.and..ate.wo.....B..Tangential....ion,
showing
a horizontal
Radia
l
si
cd l
...... ~~~~~~~~~~~~~~~~~~~~~~~~~~
:'::i~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
.......
... ...............~
t
tracheids and
larger
pits...........agifc
. (3-6)......cross-field
........ ... . ......
.....os0.
..
..p~
~~~~~~~~~.....
,,......
Ei~~~~~~~~~~~~~~~~~~~~~~i
~
~
:I:I~~~~~~~~~~~~~~~~..
I?.-.?.-.-~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~....
i::i~i~i.Z~i~~~P::ii'
ij~4'3
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~'..-:.7...
.
:::~~~~~~~~~~~~~~~~..
....
IAi
?II'?lli
.....
''''''''''
i:lrii
:~~~~~~~:?:?
?
...
..
:?:?:?:?:
....
...
. .
......
...
iii
ii:::i!'i:
:?:~~~~~~~~~~~~~~~~~~~.
'~i
...
j~~~~B~~~-i~~il
S8~~~~~~~~~~~D
8~...
l,l,l'l
...
..
:::
?I:
...
.
....
II
..
.............
i:::::::g~~~~~~~~~~~~~~~~~~~~~~~~~~~...
B3I:::~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.....
.........
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...
..
....
....
..
......
....
. ...
......
Ix
::::::::.::::::
....
.....
..........~~~~~~~~~.
of the "hard"pines (Diplox'i-loit subgen. Piiiiis). A. Transverse

a~~~~~~~~~~~~~~~~~~~~
FIG. 9. Piiiiis ooc(irpi,
section, showing
demarcatedzone
between early and late wood. B. Tangentialsection. showing a
verticalresinduct and a typical
horizontal:::::
hicenedtooh-lke
resi dut wihina ry. clearly
C Raialsecton,showng
pojetion (dntae) i th wals o th ra
trachidsad lager (-6)
coss-ield
its. Magnficatons x100.
Wood Anatomy
33
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~l
~~~~~~~~~~
?..........
~~~~~~~~~...
~~~~
~~~~~~~~~~~~~~~~
. :'~'"
wE
....0
"
... . .....
:...
......
?`: -
_z
_i~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
?. :i:?'-i
? "::i:i::.:
::!':.~:~
':::.
.... :~i:~:.!ii"':i"1i~?:;::!i
:iiiiiii~::~'i::~
'::.-':!:iiiiiiii~?:i:
'
i::i ii ii:ii- : !ii
crs-ied
MgA.
'
I0
C.
25._
....
::::
: ?:i:::i."
ii
FIC,.10.Pi~tltsceotbroidles.
typicalof the"soft"pines;(Halplo.-v~lo.t
= subgen.Strobtts).
A. Tralnsiverse
section.showing
two vertically
resinductsand a less clearlymarked.gradual
transition, fromearlyto late woodl.B. Tangentiall
section.
showinga horizontalresinductwithinaI ray.C. Radialsection. showingraytracheidsi
withsmoothwallsandsmall(2pits.(Malgnificaltions;:
4) cro.ss-field
A. B. x 10():C. x2:50.)
Flora Neotropica
34
2. Parryana.Ray tracheidswith smooth walls; small
cross-field pits 2-4, piceoid to pinoid; tangential
wall pits on latewood tracheidsnumerous,e.g. P
gerardiana (P cembroides).
3. Sula. Ray tracheids inconspicuously dentate;
cross-field pits 1-4, pinoid; tangential wall pits
absent,e.g. P halepensis, P canariensis, P leucodermis, P roxburghii.
4. Ponderosa. ray tracheidsdentate; cross-field pits
1-4, pinoid; tangential wall pits absent, e.g. P
ponderosa, P. contorta, P patula, P pittaster, P
radiata.
5. Taeda.Ray tracheidsdentateand reticulate;crossfield pits 3-6, pinoid; tangentialwall pits sparse,
e.g. P palustris, P banksiana, P. caribaea, P
echinata, P taeda.
6. Khasya. Ray tracheids inconspicuously dentate;
cross-field pits 1-2, large; tangentialwall absent,
e.g. P kesiya.
7. Sylvestris. Ray tracheids dentate; cross-field pits
1-2, large; tangential wall pits rare, e.g. P. sylvestris, P densiflora, P nigra, P resinosa.
A more detailed breakdowncomparingboth gross
morphologicaland wood anatomicalfeaturescan be
found in the work of Hudson (1960), who further
attempted to classify the pines by the degree of
dentation in the ray tracheids. He stated that slight
swelling and low dentationwere seen to be common
in those species that have smooth-walled ray tracheids. It is to be appreciated, however, that the
classes described are arbitraryand that some difficulty is involved in deciding into which class some
specimens should be placed. The classes attemptto
show the gradual development of the ray tracheid
dentationfrom slight swellings to reticulations.
Classificationof the ray tracheidsaccordingto the
degree of dentation (Hudson, 1960; some classes
with minor differences have been taken together
here):
1. Slight thickening of the wall adjacent to the
bordered pits of the ray tracheids, e.g., P.
cembra.
2, 3, 4. Very slight dentations most pronounced
near the borderedpits of the ray tracheids,e.g.,
P koraiensis, P strobus, P wallichiana.
5. Moderatelydentateray tracheids,e.g., P pinea.
6-10. Gradualdevelopmentfromdentateto heavily dentate ray tracheids, e.g., P merkusii, P
roxburghii,P cubensis, P pintaster,P resinosa.
I I. Heavily dentate, with some reticulate ray
tracheids,e.g., P banksiana.
12-14. Partiallyreticulateto completely reticulate
ray tracheids, e.g., P ponderosa, P taeda, P

palustris.
Frey-Wysslinget al. (1955, 1956) suggested that
pines with dentate ray tracheidshave "warts"on the
inner wall of the borderedpits of the axial tracheids
(observablewith electronmicroscope),and thatpines
with smooth-walled ray tracheids have no "warts."
Furtheranatomicaldescriptionsof individualspecies
were given by Greguss (1972).
CENTRAL AMERICAN PINES

Although few of the above charactersare observable in the field, the terms"hardpine"and "soft pine"
have significance in Mexico and Central America
because species in the soft pine group are much
sought after, as they are more easily worked by
hand and less resinous (Perry, 1991). The turpentine
industry in Mexico and Central America is quite
large and is based on the collection of resin from
species in the hard pine group.
Wood anatomicaldescriptionsof 12 Mexican timber trees including eight pines-P ayacahuite, P
P leiophylla, P monophylla,PFnelsonii,
cemnbroides,
P pinceana, P quadrifolia, and P strobus var. chiapensis-can be found in Crespo (1963). A summary
of the charactersstudied in each species is included
in the condensed tables. The form, frequency, and
dimensions of the tracheids, rays, parenchyma,and
resin canals are reported,together with photomicrographs.
Adopting the system proposedby Martinez(1948,
1953) Crespo placed P cembroides, P monophylla,
P nelsonii, P pinceana, and P quadrifolia in his
pinyon section. These species were characterizedas
having in common 1-4 piceoid pits (with the exception of P nelsonii, 1-6), small areolate pits in the
summer wood tracheids, smooth-walled ray tracheids, and ray cells with pitted horizontalwalls and
nodulartransversewalls. Interspecificdifferences included the fact that P monophylla possesses very
wide rays, whereas splits in the tracheid walls of P
pinceana and P nelsonii are visible.
Martinezplaced P. ayacahuite and P strobus var.
chiapensis in his Ayacahuite section, and Crespo
describes them as differing from the pinyon group
by having pinoid or fenestroid pits, 1-2 per field,
with slightly toothed walls in the ray tracheids.Pinus
leiophylla is also placed in this group (wood anatomy, but not Martinez'sgrouping) but differs in that
the ray tracheidsare more clearly toothed.
35
Karyology
The above-mentionedMexican species and others appearto correspondwith the general morphological/
examined at the University of Oxford (i.e., P cacri- anatomical features that attempt to separate pines
baea, P ntontezumae,P oocarpa, and P ponderosa) into hardand soft groups.
KARYOLOGY
All species of Pinus have the same haploidnumber
of chromosomes: n =
12; 2n =
24 (Mehra &
Khoshoo, 1956; Saylor, 1972, 1983; Mehra, 1988).

This is also the predominantbasic number in the
remainderof the Pinaceae, where only one species
in Pseudotsuga (P menziesii, n = 13; 2n = 26) and
the monotypic genus Pseudolarix (n = 22; 2i =

44) are known to be aberrant.Polyploidy, or even
the additionor loss of a single chromosome,does not
exist in the genus except for an occasional individual
(Saylor, 1983). Mirov (1967) reportedthat attempts
to artificially induce polyploidy in pines met with
difficulty and abnormal development in almost all
cases. Chromosomestructureis equally conservative.
Karyotypes,based primarilyon centromerelocation
and arm lengths, are constanteven within very widespreadspecies such as Pintussv,lestris (Mirov, 1967),
and differ but slightly among species. The most
comprehensiveanalysis has been providedby Saylor
(1964, 1972, 1983), covering 87 species in total.The
I I largerchromosomes of all species are similar in
length and have median centromeres,i.e., two arms
of equal length. Only the smallest chromosome [no.
12, also No. 11 in species of subsect. Sylvestres
(= subsect. Pinus)] is consistently heterobrachial
(centromeresubmedian),with a ratio of ca. 0.6 between the a and b arms.
Nevertheless, more detailed observations, using
charactersequences as summarizedby Saylor (1972,
1983), allowed species comparisonsand yielded occasional differences of karyotypeespecially at taxonomic levels above species. Some of these, such as
the additionof No. I I to the heterobrachialchromosomes in subsect. Pinus, seem to delimit such groupings of lower rank.Others,however,appearto group
species across the two subgenera. There are also
some differences between species karyotypeswhich
seem helpful in elucidating their relationships. In
general,Saylor (1983) concluded,that karyologyhas
"not provided evidence to contradict major taxonomic relationshipsestablishedpreviously,"referring
to the classification given by Little and Critchfield
(1969). A few of his observations relevant to Latin
American pines will be mentioned and discussed
here.
Morphologicaldifferences between Pinus leiophylla and P lumiholtzii,two species forming subsect.

Leiophyllae,include resinousand non-resinousbuds,
3-5 and 2-3 leaves per fascicle, medium size and
long leaves, low and high numberof stomatallines,
uniform and pluriform hypodermis in the leaves,
and three- and two-year developmentof ovuliferous
cones, respectively.The only conspicuous similarity
is the deciduous fascicle sheath, which appearsas a
uniquely shared characterin subgen. Pinus but as a
parallelismin the genus as a whole. Yet it is the only
reason for their classification in a subsection or
"group"(Shaw, 1914; Little & Critchfield, 1969).
PinusI leioph!lla and P lumholtzii have different
karyotypes.A noticeable difference was found between the karyotypes of P elliottii (including var.
densa) and P caribaea (including its varieties) in
chromosome No. 11, which links the latter to P
occidentailis with a similar pattern. This evidence
supports the classification proposed in this monograph. Another interesting observation is that the
karyotypeof P miontezu.uaemore closely resembles
P. ponderosa and its relatives than it does the morphologically similar P pseudostrobus,with which it
is also largely sympatric. Pinus lawsonii and P
teocote have similarkaryotypeswhich set them apart
from the other species classified in subsect. Ponderosae (Little & Critchfield, 1969). The species in the
rather heterogeneous subsections Ponderosae and
Oocarpae as delimitedby Little and Critchfieldhave
been rearrangedby van der Burgh (1973), who accommodatedboth species in the lattersubsection,an
arrangementthat is supported by the classification
proposedhere.A similarbreakupseparatingthe "California closed-cone pines" from Little and Critchfield's subsect. Oocarpae(van der Burgh, 1973) finds
supportin the karyotypes,which are similarfor these
pines but differ from the Mexican species originally
grouped in that subsection.
In subgen. Strobus only a few Mexican species
were studied (Saylor, 1983), some of which also are
widespread in the United States. It was found that
the karyotypesfor Pinus flexilis and P strobiformis
are rathersimilar but that P ayacahuite differs noticeably from these two taxa. Since there appearsto
36
Flora Neotropica
be a geographical dine in several morphological

characters of the first two species, with P. strobiformis approachingP. ayacahuite in its more southern populations(see discussion with the species accounts in this monograph),it would be of interestto
have known the localities from which the samples
for Saylor's studies originated.No informationabout
the origin of the samples (seedlings) is given, which
is a serious omission. Unfortunatelyalso, only three

species of Little and Critchfield's subsect. Cembroidces were studied-P
cembroides, P edullis, and
P monophylla-thereby leaving out all the more

distinct ones which could have yielded interesting
information.Not surprisingly,the three taxa investigated appearedto share similar karyotypes.
POLLENMORPHOLOGY(by Marie H. Kurmann)

Pollen grains of Pinus are bilateral,characterised
by a centralcorpus with two sacci placed symmetrically, one on each side (Fig. IA-C). They range in
size from 70 to >100 /jm. Surface sculpturingand
wall stratificationvary around the pollen grain. In
the distal area, the sculpturing ranges from psilate
(Fig. IIA) to scabrate (Fig. IIB). The distal wall
(leptoma)consists mainly of a thick, lamellatedendexine (Fig. I IF). The proximal surface is more or
less rugulateand the proximal wall (cappa) is thick,
consisting of an outer, alveolate ektexine subtended
by a narrow lamellated endexine (Fig. IG). In the
lateralregions, the ektexine is split to form the sacci.
The saccus floor (Fig. II E) consists of a lamellated
endexine and the ektexinous foot layer, and the
saccus roof is made up of the tectum subtendedby
endoreticulations(Fig. 1D).
Pine pollen is morphologicallyrelatively uniform
in most species, and the range of intraspecificvariation overlaps with the interspecificone. Furthermore,
preparationmethods may affect morphologicaland
size charactersdifferently. It is therefore important
to take preparationtechniques into account when
describing the pollen grains (Kurmann& Zavada,
1994). There seems to be some evidence that pollen
morphologicalcharactersdiffer between the two subgenera Strobus(Haploxylontype) and Pinus (Diploxylon type), but there is also a group of species in
which these are mixed. In the Haploxylon type, the
sacci are usually broadly attached,the distal area is
sculpturedand is often borderedlaterally by thickened ridges (e.g., Pinus rzedowskii,Fig. II B). In the
Diploxvlon type the sacci are contractedalong their
roots. The distal area is usually smooth and is not
borderedby a thickenedridge (e.g., Pinus leiophylla,
Fig. IIA).
In 1950, van Campo-Duplan published a study
of the Pinaceae based on pollen morphology. She
investigated ca. 60 species of Pinus pollen with
the light microscope, and this remains the most
comprehensivepalynological treatmentof the genus
to date. For each of the species, she listed the size

and shape of the corpus, shape and ornamentationof
the distal area, structureand shape of the saccus,
ratio of saccus height to width, and the proximal
attachmentof the saccus to the corpus. She concluded that it is difficult to separatepine species on
the basis of pollen morphology and considered it
impossible to draw phylogenetic conclusions. Her
results do not contradict the classification into the
two subgenera,but neitherdo they allow their recognition on the basis of pollen morphologicalfeatures
alone. Erdtman(1965) described the pollen of 25
Piins species. In most instances the characters he
used concern size and size ratios. Klaus (1975, 1978)
also investigated the pollen morphology in Piiins
species, using not only light microscopy but also
scanning and transmission electron microscopy.
Electronmicroscopy appearsto enhance species distinctions, not only by providing new charactersbut
also by clarifyingcharactersused in light-microscopical studies. Morphologicallyuseful charactersin his
account included saccus structureand variation in
corpus structuresas revealed by transmissionelectron microscopy. He concluded that in individual
cases it might be possible to differentiategroups or
even species if a large number of characters were
used. He also consideredornamentationof the distal
area to be the most useful characterin differentiating
the two subgenera.
Pinus pollen is often found in great abundancein
fossil deposits. Since differentspecies have different
climatic requirements,identification of these fossil
grains to the species level would greatly enhance
paleo-environmentalinterpretations.Several studies
have been undertakento identify modern Pinaceae
pollen grainsto species level in restrictedgeographic
areas. In an attemptto distinguishmodernspecies of
Piins in the Pacific Northwest, Hansen (1947) used
size frequency. However, this approach has been
questioned by Mack (1971). Hansen and Cushing
(1973) proposeda method for distinguishingspecies
37
Pollen Morphology
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38
Flora Neotropica
which combines quantitative and morphological

characters.Jacobs (1985) also demonstratedthat for
some geographicgroups, pollen identificationat low
taxonomic levels is possible if a combination of
structuraland linear charactersis used.
The relative uniformityof pollen in Pinus makes
it difficult to classify it at the species level. Careful
considerationof preparationtechniquesand a combination of structuraland quantitativecharactersmay

prove to be useful in distinguishing species in restricted geographic areas. Electron microscopy provides additionalcharactersand should be employed
in attemptsto identify pine species based on pollen
morphology.
THE ROLEOF MONOTERPENESIN THE TAXONOMYOF THE MEXICAN

AND MESOAMERICAN
PINES (by JacquelineS. Birks)
matography.Terpenes are the subset of terpenoids
which have exact multiples of the isoprene units
Plant isoprenoids are compounds derived from
(hemiterpenes, C5; monoterpenes, C)o; sesquiterisoprene.They are divided into primaryand second- penes, C15;
diterpenes,C20).There are thousandsof
ary metabolites.The secondary metabolitesof plant known terpenes and
many have been found in the
cells, chemicals of low molecular weight that are resins of
pines. They have a role in disease resistance
not directly involved in essential metabolic action, and defence
against insect attack.
include the monoterpenes, the sesquiterpenes, and
The biosyntheticpathways,by which the terpenoid
the diterpenes. The secondary metabolites of the constituents of resin
are formed, are difficult to
genus Pinus include a wide range of chemical prod- determine
directly (White, 1983) and were little unucts with industrial or pharmaceuticaluses. As a derstooduntil
significantadvances were made in the
result, the chemistry of many of these productshas last decade (Gershenzon& Croteau,
1990). A review
been thoroughly investigated and this in turn has
by Chappell(1995) emphasises the progressthat has
often been the precursorof more fundamentalscien- been made in
understandingthe enzymes involved
tific research.The results of this extensive research in the
biosynthetic pathways but reminds us that
generated interest in measuring and describing the there is still much that is not understood.
chemical components and using this informationas
Resin samples may be obtained from steam disa purely scientific tool to investigate relationships tillation of leaf oils
(Zavarin, 1970) or from cortical
between the different species.
tissue (Squillace, 1977; Meier & Goggans, 1978) or
Mirov (1967) reviewed the full range of chemical
xylem tissue (Franklin, 1976; Strauss & Critchfield,
products of pines, which are numerous. Although 1982). The cortical resins are easier to collect
(Squilattempts at classification and delimitation of taxa lace, 1977).
have been made with all secondary compounds for
pines, the components of the oleoresin, and particularly the monoterpenes,have attractedthe most attenGas Chromatographic Analysis and the
tion and yielded the most contributionsto the taxoQuantitative Results
nomic debate.
Gas chromatography(GC) is an efficient and precise method for determinationof resin composition
The Pine Oleoresin
(von Rudloff, 1975). Results appear as a plot of a
The mixtureof rosin and essential oils that can be series of peaks, each of which can be identified as
obtained by cutting the resin canals, of leaves or of belonging to a particularcompound, using previous
the wood, is known as the oleoresin but is usually results from known compounds.The area undereach
referredto simply as the resin. Resins are mixtures peak measures the relative proportionof each comof compounds including terpenoids,flavonoids, and pound presentin the sample. Results are reportedon
fatty acids. Essential oils are volatile oils that are a number of different bases (Squillace, 1976). For
recovered from plant tissue by steam distillation or monoterpenes three bases are commonly used: 1)
solvent extraction. The terpenoids, comprising C5 relative to all other monoterpenes (total monoterisoprene units, have been the most widely studied pene), 2) relative to all other constituents of the
class of compounds extracted from pines because oleoresin (oleoresin basis), and 3) per unit weight or
they are easily identifiedand quantifiedby gas chro- volume of tissue (tissue basis). The data defined by
INTRODUCTION
The Role of Monoterpenesin the Taxonomyof the Mexican and MesoamericanPines

the latter are usually described as absolute amounts
(White, 1983), althoughthereis nothinguniqueabout
this base as other absolute amountscould be defined.
The proportionsof compounds are independentof
the total yield of oleoresin, but data reportedon a
tissue basis are not. White and Nilsson (1984) found
that the yield of leaf oil monoterpenes in Pinus
contorta was relatedto the frequencyof resin canals
in the leaves, which was itself variable and subject
to genetic control. They concluded that the proportions of compounds should be used to study differences between populations. Shaw et al. (1982)
reachedthe same conclusion but cited the confounding effects of environment and genotype on yield.
Consequently, most studies of resin composition
have been based on the interpretationof data comprising the proportionsof individual terpenes detected in resin samples. Such data are referredto as
compositionaldata.
Variation in the Composition of Resin
The resin is synthesised in the resin ducts of the
different tissues. Each of these samples an independent system (von Rudloff, 1975), and the compositions of the oleoresin obtainedfromdifferentsources,
from a single tree, are not the same. The compositions of foliar oils and cortical resins are often similar. The composition of xylem resin is often less
complex than that of cortical resin (Squillace, 1976).
The composition varies within the same system at
differentpositions in the tree. Forexample, the xylem
resin has been found to vary with the height of
sampling (Rockwood, 1973; Roberts, 1970). There
is variation with the developmental stage of a tree
and with the season of the year (von Rudloff, 1975).
Moisture stress has been shown to affect the resin
composition in some species (Langenheim et al.,
1979), and insect infestation has been shown to
increase monoterpeneand diterpeneproductionin P
contorta (Raffa & Berryman,1983).
Because there are so many factors affecting resin
composition, apparentlycontradictoryresults are reported.Genotypicallyidenticaltrees of P monticola,
growing in diverse environments,showed negligible
differences in terpenecomposition (Hanover, 1966),
but genotypically identical trees of P sylvestris
grownon threevery differentsites in Sweden showed
moderatedifferences in terpene composition (Baradat & Yazdani, 1988).
Hanover (1966) and Squillace and Fisher (1966)
believed thattheirresearchshowed thatthe composition of resin is not only strongly inherited but the
39
production of certain monoterpenes is often controlledby single genes with majoreffects. The belief
that the non-genetic effects are minimal or easily
controlled is what made resin composition such an
interestingtrait for taxonomists.
Monoterpenes Found in Mexican and
Mesoamerican Pines
The oleoresinof Mexicanand Mesoamericanpines
is clearly differentiatedfrom that of other pines, the
main constituents being a-pinene and /3-phellandrene./-phellandrene is almost uniqueto the species
of Mexico and Mesoamerica(Mirov et al., 1965). A3-carene and longifolene are quite common constituents of the oleoresins, again rarelyfound elsewhere,
their only other occurrencebeing in the pine species
of western North America. In the north of Mexico
there is evidence of a different oleoresin. The oleoresin of Pinus jeffrevi, a species whose southern
extent is the north of Mexico, is dominated by
heptane, with minute proportions of a-pinene,
whereas P coulteri, which also just extends into the
north of Mexico, shows a more typical oleoresin
dominatedby a-pinene, plus /3-phellandrene.
THE USE OF MONOTERPENE DATA

IN TAXONOMIC STUDIES
Monoterpenedatahave usually played a role complementaryto other taxonomic charactersand therefore have been introducedto solve the more intractable problems concerned with species delimitation
in groups of closely related species alliances or
widespreadpolymorphicspecies complexes. Initially,
qualitative or discrete characters (the presence or
absence of compounds) were used to examine relationships or delimit taxa, but when the group of
species being investigated displayed the same set
of compounds, quantitative traits were used. The
possibilityof single-gene controlof some compounds
is often mentioned but not seriously addressed in
taxonomic studies.
Qualitative Characters
Occasionally the monoterpenedata have shown
discrete, qualitativedifferences between species, but
these results have often reflected inadequate sampling at one or more levels. A compound that is
declared missing in a population in one study may
be detected in small proportions,or at low frequencies, in another.An early comparisonof resincompo-
40
Flora Neotropica
sition of Piins monophylla and P quadrifolia (An-
derson et al., 1970) found that B/-phellandrene,

terpinolene, and camphene were missing from P
qucadrifoliabut were presentin P. monophylla:but a
laterstudy of these two species (Zavarin& Snajberk,
1980) shows these threecompoundspresentin small
proportionsin both species. An investigationof the
closely relatedspecies P estevezii, P. pseudostrobus,
and P montezunae (Perry, 1982) showed heptaneto
be present in the oleoresin in all samples of P
estevezii but absent in all samples of the other two
species. However,only 10 trees from one population
of each species were examined, and this interesting
observationrequiresfurtherinvestigation.
Quantitative Traits
The terpene traits are used to investigate species
relationshipsas a basis for classifications and, more
widely, for the delimitation of species (Zavarin &
Snajberk, 1973). Most studies employ monoterpene
data as though the proportionof each compound
were a continuous variable measuredindependently
of the other compounds. The means and variances
of the proportionsof each compound are analysed
between and within populationsusing univariateand
multivariatestatistical methods.
It is this wide use of quantitativeterpene traits,
nearly always in the form of proportions,that has
been questioned, and discussed in depth (Birks &
Kanowski, 1988, 1993) in relation to the known
problems inherentin the analysis and interpretation
of data describing compositions.
Many interpretationsof resin compositional data
have acknowledged but not acted upon the conceptual difference between compositional data and a
multivariatedata set, where the same constraintsdo
not apply. These proportionsare not metric traits in
the usual sense. They are not measuredindependently
of each other,and methodsof analysingmetrictraits,
such as analysis of varianceor t-tests, are invalid if
applied to single componentsof these data. Multivariate methods of analysis, such as principal components analysis, are invalid if appliedto compositional
data.
raised to species rankfrom that of variety within the

previous two decades on the basis of morphological
evidence. Zavarin and Snajberk, with others, have
conducteda series of investigationson the monoterpene traits of subsets of the pinyon pines, together
with morphologicaltraits, in order to evaluate these
taxonomic revisions. Zavarinet al. (1980) compared
P quadrifolia with P monophylla: Bailey et al.
(1982) comparedP discolor with P cembroides:and
Zavarin and Snajberk(1985, 1986, 1987) looked at
differences between populations within P cembroides, compared P discolor with P johannis, and
investigated differences among P culnirnicola, P

nelson ii, P pinceana, and P mraxilartilnezii. The
first two papers deal with the question of (putative)

hybridizationbetween these species. In both studies
there is an apparentlack of understandingof what
taxonomic characters are: "deviating values" (e.g.,
when these fall within the range of the wrong species) are left out of the equation.The same problems
are apparent in Zavarin and Snajberk (1987: table
3), where, for instance, P maximartine-iiis presented with a mean of 88.6% limonene, but a sample
measuring only 16.8% from the same population
is left out as a "deviating value." We are not informed about the percentages of the other compounds, which must of necessity be higher than in
the "mean." Since this highly distinctive species
does not hybridize with any other (to explain the
deviation),the figuresgiven in the table are meaningless as taxonomic characters.
The separationof Pinus patula and P oocarpa is
not usually disputed, but the status of other closely
related taxa has caused considerable controversy.
Monoterpenedata were used to investigate the status
of P patula var. longipeduncultata,P tectunumanlii,
and the possible hybridizationbetween P oocarpa
and P caribaea var. hondurensis(Squillace & Perry,
1992). The statistical procedures ("renormalizing"
proportions to sum to 100% and assuming 10%
limonene even where this constituent was not assessed) as well as the presentationof data in the
analysis makean assessmentof taxonomiccharacters
from this paper problematical.There appears to be
little support for the taxonomies based on morphological data from the monoterpenesanalysed in this
study.
Some Regional Examples of Taxonomic

Studies Using Monoterpene Data
CONCLUSION
The Mexican pinyon pines, Pinus subsect. Cembroides Engelmanns.l., comprise seven species and
three varieties or subspecies. Several taxa have been
Absolutedeterminationof resincompositiondefies
definition and measurement; the only assessment
possible is of the proportionsof differentconstituents
41
ReproductiveBiology
in a sample, but these cannot be defined uniquely.
The analyses of the data are all affected by some or
all of the problemsdescribedby Birksand Kanowski
(1988, 1993). Inferentialstatistical methods applied
to single components of a composition, as has been
done in the studies described here, are not correct,
and the results will be misleading. Descriptivemethods of data analysis applied to a subcomposition
may help in clustering populations, but these are
superficialmethods for taxonomic studies. The studies described here not only fail to address these
problemsbut also attemptto use quantitativedata in
ways that do not constitute meaningful taxonomic

characters.
It is possible that the presence or absence of
a constituent may provide taxonomic charactersas
defined by Davis and Nixon (1992), but there has
been little evidence so far that these exist at the
species level. The conclusion must be that,considering the combined problems of the analysis of the
data of compositions together with the unsuitability
of the traits derived from the monoterpenedata as
taxonomic traits, we are not in a position to review
the taxonomy of the pines.
REPRODUCTIVE BIOLOGY
Generalbutdetailedaccountsdescribingthe reproductive biology-including pollination,the ontogeny
of the reproductiveorgans, and embryology-in conifers are given by Strasburger(1869, 1872), Pilger
(1926), Chamberlain(1935), and Hirmer (1936); a
synopsis is presentedby Weieret al. (1982), exemplified by PinLus.
POLLINATION
The pines, like almost all gymnosperms, are entirely wind pollinated.Individualtrees produceenormous quantitiesof pollen, which can be blown great
distances from its source. Pollen dispersal(anthesis)
times vary among species in our region from sometime in November/Decemberto June/July,coinciding
with the dry season in most regions of Mexico and
CentralAmerica.The late springdispersalsare found
mostly in taxa with northerlydistributions and of
high altitudes (e.g., Pinus hartwegii). Pines, being
monoecious, are frequentlyself-pollinated, although
cross-pollinationis stimulatedby the relativeposition
of male and female strobilion the branchesof a tree.
At the time of pollen dispersal, the female strobili
have emergedfrom the buds and have slightly parted
their scales, permittingwind-blown pollen grains to
sift down to the axis of the cone, where they come
into contactwith a sticky substance(pollinationdrop)
secreted by the ovule. This fluid was first observed
by Strasburger(1872), and its role in transporting
the pollen up to the receptive surfaceof the nucellus
was definitely established by Doyle and O'Leary
(1935). The role of this micropylar fluid has been
studied in detail by McWilliam (1958). The micropyle of the anatropousovule consists of a chamber
formedby the integumentextendingbeyond the apex
of the nucellus, a canal terminating in a narrow
opening with a closing mechanism("neck")and two

appendages("arms").The sticky fluid on these parts
(Fig. 8) causes pollen grains to adhere to their surfaces. Reabsorptionof the pollinationdrop, possibly
by surface tension or partial drying, brings some
pollen grains into the micropylarchamber, where
one of them germinatesand grows a pollen tube into
the nucellus. During this process, the germinated
pollen grain slowly develops into a tubular male
gametophyte. Meanwhile, the micropylarcanal has
been closed by a swelling of the neck cells. With
this, pollinationis completed.
FERTILIZATION AND EMBRYOGENY

In Pimus,a single megasporocytelies in the centre
of each ovule; it is surroundedby the nucellus,
containing nutritive tissue. The pollen tube grows
through the nucellus and reaches an archegonium.
This development requires about one year to be
completed, when the pollen tube releases its contents
into the archegonium. Meanwhile, the megasporocyte in the ovule has been divided by meiosis into
four haploid megaspores;one of these furtherdevelops into a female gametophyte,containing archegonia surroundingan egg cell (Fig. 8). When the male
gametophyte reaches the archegonium, the female
gametophyteis fully developed and ready for fertilization (Fig. 8). One of the two sperm nuclei then
unites with the egg nucleus. Theoretically,since the
female gametophytecontains two or more archegonia, each with an egg cell, more than one egg may
be fertilized and develop into an embryo in a single
seed. However, although seeds with two embryos
occur regularly,normallyonly a single embryodevelops (Fig. 8). At the time of fertilization, the seed
cone will have reached nearly full size but is still
42
Flora Neotropica
"green"and the scales are tightly closed. The events

following fertilisationhave been described in detail
by Maheshwariand Konar (1971) and only a short
summaryof embryogenywill be given here. Mitotic
divisions (4 X) result in the formationof a proembryo with four tiers of four cells each (Fig. 8). Any
remaining nuclei (paranuclei)disappearduring this
stage. Of the four tiers of the proembryo,the two
nearest the micropyle degenerate,while the suspensor elongates, thrustingthe terminalembryoniccells
deeper into the gametophytic tissue. The proembryonalcells then divide into several proembryos,of
which one succeeds and develops further into the

final embryo. Cellular diversification leads to the
formation of a rootcap followed by root initials
(radicle)at the proximalend of the suspensor,cotyledons at the distal end of the embryo and the hypocotyl in between these structures(Fig. 8). The fully
developed embryo is embedded in the enlarged female gametophyte (often referredto as endosperm,
but its ontogeny is entirelydifferentfrom angiosperm
seed development). Both embryo and gametophyte
are enclosed in a thin chartaceous shell (all that
remainsof the nucellus) and a hard seed coat.
PHYLOGENYAND EVOLUTION
GENERICRELATIONSHIPS
popinus) has been acknowledged by most workers,

but few have had access to materialof this rare and
Pinus (Pinaceae) is considered monophyletic
Vietnamese pine. Apart from its two
endangered
(Flous, 1936; Niemann& van Genderen, 1980; Price
sometimes hypostomaticleaves, it fits
flat,
peculiar,
et al., 1987; Farjon, 1990a); it is also the largest and
well in subgen. Strobus.
most diverse genus. Its sister group is probablythe
genus Picea, from the evidence of two types of
characters(chemicaland morphological)investigated
Further Divisions
in these studies. Intergeneric relationships among
the II (sometimes 10, 12, or 13) genera currently
The subdivision of the two subgenerais considerrecognizedare not well established.Recent phyloge- ably morecontroversial,and a largenumberof classinetic studies involving genera in the Pinaceae reject fications has been proposed, dealing either with the
the traditionalclassificationbased on foliage charac- entire genus or with parts of it (see History of
ters as artificial (Niemann & van Genderen, 1980; Classification). Some, because they are based on a
Price et al., 1987; Frankis, 1989; Farjon, 1990a), but very limited numberof characterschosen for identionly the latter two, based primarilyon morphology, fication purposes (e.g., Pilger, 1926), are definitely
give Pinus a more basal position on the tree in a artificialand do not reflectphylogeneticrelationships
distinct subfamilyPinoideae.Molecularstudies using or evolution of characters.Others, though intentionDNA and comparingthe genera are yet to be under- ally phylogenetic, seem to classify species in groups
taken.
defined by unlikely combinations of homoplasies,
e.g., the reversal to deciduous fascicle sheaths in
some and the reversal to adnate seed wings in other
INTERGENERICRELATIONSHIPS
species of the same (sub)section (Shaw, 1914; Little & Critchfield, 1969). Phylogeneticanalysis based
Subgenera
on the requirementof monophyly and considering
It is widely accepted that there are two natural
only sharedderived (synapomorphous)charactersas
groups within the genus Pinus, which are often called indicative of common
ancestry (Hennig, 1966) has
Haploxylon and Diploxylon but which, under the called such classifications into
question (Strauss &
rules of nomenclature,must be named subgen. StroDoerksen, 1990; Malusa, 1992; Wang & Szmidt,
bus and subgen. Pinus, respectively (see History of
1993).
Classification). Morphological and anatomical
Little
&
Critchfield,
(Koehne, 1893; Shaw, 1914;
1969; Farjon, 1984), chemical (Lindstedt, 1951), and
Subgen. Pinus
molecular characters (Strauss & Doerksen, 1990;
Of the 60 taxa covered by this treatment, 42
Wang & Szmidt, 1993) have been analyzed,supporting this basic division. Since Little and Critchfield belong to subgen. Pinus. Speciationin Mexican pines
(1969), the subgeneric distinction of P krempfii(or is consideredto be concentratedmainly in this subgeeven its recognition as a monotypic genus, Ducam- nus, with various "species complexes" being com-
Phylogeny and Evolution

monly recognized (Martinez, 1948; Stead & Styles,
1984; Squillace & Perry, 1992). Subsections (according to Little & Critchfield, 1969) with major
representationin Mexico and Central America include Leiophyllae, Ponderosae, and Oocarpae; in
the Caribbean the taxa belong mainly to subsect.
Australes. A refinement of Little and Critchfield's
classification [which retained much of Shaw's
scheme despite claims by others that it was based
largelyon Duffield's (1952) hybridizationtrials] was
given by van der Burgh(1973) and slightly amended
by Farjon(1984). This classificationespecially subdivided Little and Critchfield'ssubsect. Ponderosaeby
splittingoff subsect. Pseudostrobi,and Oocarpae by
splitting off Attenuatae.Subsection Leiophyllae was
elevated in rank to a section, with two subsections
each accommodating one or two species. Styles
(1993a) slightly modified Little and Critchfield's
scheme as applied to Mexican taxa, with additions
of species and varieties not recognized or known in
the originalclassification.A comprehensivephylogenetic analysis of subgen. Pinus is hamperedby the
43
paucity of discrete characters,which in the genus
Pitus are scarce when compared to other plant
groups (Styles, 1993a). A cladistic analysis of all
taxa in this revision resulted in a strict consensus
tree with both unresolvedand resolved clades within
subgen. Pinus. With successive approximations
weighting, which uses the rescaledconsistency index
to calculate weights, nearly all clades (trees not
shown) were resolved. Due to the preliminarynature
of the cladistic analysis, the incomplete sample of
taxa, and the high levels of homoplasy in most
characters,the analysis is used sparinglyto influence
the classification. We therefore propose to present
the species of this subgenus tentativelyaccordingto
Little and Critchfield (1969) and van der Burgh
(1973), with such emendations as are substantiated
by the cladistic analysis of our data. The list of
charactersand a data matrix used in the analysis
(using PAUP version 3.1) are presentedin Tables I
and II, respectively.Only one cladogram(Fig. 12),
representinga phylogeny of species in subgen. Strobus, is presentedhere.
Table I
Character numbers and states used in the cladistic analysis of taxa in the genus Pinus. Characters 4, 5, 11
and 15 are represented by the first state only in species of subgen. Strobus; character 6 does not apply for
that subgenus (see Table II).
1. Winterbuds with resin, 0; without resin, I.
2. Sheaths of matureleaf fascicles early deciduous, 0; late deciduous, forming a rosette, 1; persistent,2.
3. Leaves epistomatic,0; amphistomatic,I.
4. Resin ducts (primarily)external, I; medial, 2; internal,3; septal, 4.
5. Vascularsystem in leaves haploxyl, 0; diploxyl, 1.
6. Vascularbundles in diploxyl pines connate, 0; separate, I.
7. Hypodermisuniform,0; of variablethickness, I.
8. Outercell walls of endodermisthin, 0; thickened, I.
9. Seed cones opening soon, deciduous, 0; serotinous,persistent, 1; opening slightly, disintegrating,deciduous, 2.
10. Peduncledeciduous with cone, 0; persisting with some basal scales, 1.
11. Apophyses in mid-cone nearly monomorph,0; strongly heteromorph,1.
12. Umbo terminal,0; dorsal, 1.
13. Umbo with deciduous spine, or spine absent, 0; spine persistent, 1.
14. Seed wing articulate,much larger than seed, I; seed wing adnate, vestigial, 2; seed wing absent from matureseed,
3; seed wing adnate, much largerthan seed, 4.
15. Long shoots uninodal,0; multinodal, 1.
16. Fascicle bract (cataphyll)bases virtuallynon-decurrent,0; decurrent,I.
17. Numberof leaves per fascicle 1(-2), 1; 2(-3), 2; (2-)3, 3; 3(-5), 4; 3-5, 5; 5, 6; 5-8, 7.
18. Sheaths of leaf fascicles initially long (>20mm), 1; of medium length (15-20mm), 2; short ( 15mm). 3.
19. Sheaths of leaf fascicles reduced in length at maturity(?30%), 0; persistentlylong, I.
20. Length of leaves -5cm, 1; 5-10cm, 2; 10-15cm, 3; 15-25cm, 4; 25-50cm, 5.
21. Leaves -I mm wide, rigid, erect, 1; 1-1.2 mm wide, lax, drooping or erect, 2; - I mm wide, soft, drooping or
pendent,3.
22. Peduncles of seed cones long (>2.5 cm), slender, 1; 1-2 cm long, stout, 2; very short (-I cm), 3 (for very large
cones multiply by 2).
23. Length of matureseed cones -5 cm, 1; 5-8(-10) cm, 2; (8-)10-15 cm, 3; (15-)18-25 cm. 4; -25 cm, 5.
24. Numberof seed scales per cone (average) -60, 0; 60-110, 1; 110-170, 2; >170, 3.
25. Numberof stomatal lines on abaxial face of leaves 2-5, 1; 6-10, 2; -10, 3.
44
Flora Neotropica
Table II
Characterstate matrix for taxa in Pinus subgen.
Strobus,comparing20 characters(excluding
characters4, 5, 6, 11 and 15, which are
autapomorphiesof Pinus subgen. Pinus).
Polymorphismand absence of charactersare
indicated by a question mark(?).
11111111222222
12378902346789012345
P. strobus var. chiapensis
P laihbertiana
P flexilis var. reflera
P strobiformis
P. vacahitte
P avtacahuite var. veitchii
P rzedowskii
P pincelana
P nelsoniii
P maximartinezii
P culhninicola
P remote
P cembroides var. bicolor
P cembroides subsp. orizahensis
P. cembroides subsp. lagunae
P cemtroides subsp. cembroides
P mnonophylla
P. /uadlrifoiia
Picea chihualmana
1000000004063?32131?
1010000004063?221521
00?0000002063?22231?
10?0000002061?232???
1000000004062?332?2?
1000000004062?33152?
1200000111053?23131?
1010020113143'22121?
01100201131431221211
0200020103063?23341?
0200000113063? 11310?
1010100103023?113101
1200000103043?123101
02 10000)103043?123101
1210000113033?223101
1210000113033?123101
0011000103013'?113103
0000000 103053?113101
07?0100??7?
10'?? 11?322
ayacahuite
veitchii
lambertiana
reflexa
strobifonnis
chiapensis
pinceana
nelsonii
rzedowskii
maximartinezii
lagunae
cembroides
bicolor
orizabensis
culminicola
remola
monophylla
quadrifolia
Picea
Fig. 12. Strictconsensustreeof 24 equallyparsimonious
trees(Cl = 0.586, numberof steps = 58), analyzing20
characters
(see TablesI andII),usinga heumorphological
risticsearch(randomaddition;subtreepruning/regrafting)
withFitchparsimony
(Swofford,1993).
pringlei, classified in subsect. Oocarpae s.str. by van

der Burgh, but in two different subsections by Little
Little and Critchfield(1969) classified Pinus leio- and Critchfield,appearin the cladistic analysis (strict
phylla (including var.chihuahuana)and P lumiholtzii consensus tree) in a clade with P. leiophylla. When
as a subsection within sect. Pinea Endl., which con- charactersare weighted against homoplasy, this artained two other subsections of Mediterraneanpines. rangementis refuted (strict consensus of nine trees,
Van der Burgh (1973) recognized the distinctness of not shown). The fossil recordappearsto supportthe
these Mexican pines and accorded each of them the occurrence of pines related to P leiophyllacin the
rank of section. Morphologically, these pines are Late Paleogene of Colorado. In the fossils described
unique in subgen. Pinus in having deciduous fascicle by Axelrod (1986), no mention is made of the diagsheath scales. Recent isozyme analysis (Karalaman- nostic marksof three-seasonalgrowthon the apophygala & Nickrent, 1989) and phylogenetic analysis sis of the seed scale, but if cones and leaf fascicles
using rDNA restriction fragments (Govindarajuet with lost sheaths indeed came from the same pine,
al., 1992) confirm the genetic distinctiveness of P P coloradensis is likely to be closely related.
leiophylla and show that it is ill-placed within sect.
Pinea as circumscribed by Little and Critchfield SECT.PINUS
Van der Burgh (1973) restrictedsect. Pittus sensu
(1969). This species is also distinguishedby its threeyear development of the seed cones, which is shared Little and Critchfield (1969) to a group of pines
by the MediterraneanP pinea. Both charactersap- with a primarilyEurasiandistributionand only two
pear to be examples of parallelismratherthan syna- American representatives,one of them P. tropicalis.
pomorphies. Pinus lumholtzii appears in our clado- This species is anomalous within that group for
gram (not shown) near P pseudostrobus. Its having septal resin ducts in the leaves (Farjon, 1984).
relationshipdoes not seem to be with P leiophylla, The section also contains the Southeast Asian P
but unlike that species, its place in a distinct section
nerkusii,which has an anomalouscone morphology
is not confirmed. Pinus teocote, P. lawsonii, and P and in a cladistic analysis based on cpDNA characSECT.LEIOPHYLLAE
(LOUDON)VANDERBURGH:
SECT.LUMHOLTZIAE
VANDER BURGH

ters emerges as a sister group to sect. Pinus (Wang&
Szmidt, 1993). Phylogenetic analysis comparing P
tropicalis with its putative relatives as well as with
species with nearbyCaribbeanand SE NorthAmerican distributionsmay change its classification, but
such work could not be undertakenwithin the scope
of this monograph.It differs from other Caribbean
pines in its septal resin ducts (versus internal), its
uniform hypodermis, its low leaf number and its
very short cone peduncle.
SECT.PINASTERENDLICHER
This section as recognized by van der Burghcontains the remaining species in subgen. Piiiius. Although it represents fewer species than Little and
Critchfield'ssect. Pi.nus, which included the group
of pines mentioned in the previous section, it has
still some 40% of all the species in the genus. This
reflects the difficulty in separatingthem into distinct
taxonomic units. Both Little and Critchfield(1969)
and van der Burgh (1973) have divided their largest
section into 6-8 subsections, some of which seem to
representnaturalgroups.
45
sifted species of western NorthAmerica and Mexico
to CentralAmericaof Shaw's (1914) groupAustrales
in an amendedsubsect. Ponderosae.It includes three
commonly recognized "species complexes" around
P ponderosa, P mnontezumae,and P pseudostrobus
(Mirov, 1967). Vander Burgh(1973) segregatedtaxa

considered to belong to the latter two groups in his
subsect. Pseudostrobi. Species delimitation within
these groups is often difficult and, in part as a
consequence of this, there is little agreement concerning relationshipsof the constituenttaxa. Cladistic analyses of all taxa in the region, as well as of a
limited numberwithin subgen. Pinus, permit a few
observationsabout their possible relationships:
I. Pinus montezutmaeand P pseudostrobusare species in a monophyleticgroup in most consensus

trees, commonlyjoined by P devoniana, P douglasiana, P lumiholtzii,and P miaximinoi.
2. Pinus harvtegii, placed in subsect. Pseudostrobi
by van der Burghand consideredto be a species
in the "montezumae complex" (Martinez, 1948:
Mirov, 1967) does not appearto be closely related
to it.
Subsect. ContortaeLittle & Critc/ifield.- This is 3. Pinus engelinanniimay be relatedto P arizonica,
P ponderosa, and P jeffrevi, but also to a clade
a subsection with a northerndistribution,restricted
formed by species mentionedunder 1 above.
by van der Burgh (1973) to Pintus contorta and
P bantksiana.It is representedby P contorta var.
There are a few Tertiaryfossils known that resemimurrayana,which extends along the Cascades-Sible
cones and seeds of different extant species aserra Nevada axis and has a southern disjunction in
to these subsections, most of them from the
our region (SierraSan PedroMartir,BajaCalifornia). signed
Mountains
northof the U.S.-Mexican border
Rocky
Pilius herrerae,placed in subsect. Oocarpae s.str.by
and
from
the
Eocene to the Miocene (Axelrod,
dating
van der Burgh,does not seem to fit therecomfortably.
cones
1986).
Large
resemblingthose of P montezucould be related to
Pirus contorta var. mulrravanac
this pine with very small cones, as the cladistic inae from the fossil Creede flora (Late Oligocene,
26.5 Ma) of Colorado, and others from Miocene
analysis suggests.Though fossils are scarce, P alvordensis Axelrod from the Late Eocene Bull Run flora (12.5 Ma) deposits in Western Nevada resembling
of northernNevada is believed to belong here (Axel- those of P pseudostrobus, indicate that these or
relatedspecies occurredconsiderablyfarther
rod, 1986). We tentatively group these two species closely
norththan they do at present.
the
need
for
further
research
into
together,stressing
their relationships.
Subsects.OocarpaeLittle& Critchfield,Attenuatae
van
der Burgh, and SabinianaeLoudon.- As proSubsect. Australes Loudon. In both classificaLittle and Critchfield(1969), subsect. Ooctions, this subsectioncomprisesspecies rangingfrom posed by
the eastern and southeasternUnited States through arpae includes the Mexican and Californian"closedthe Caribbean to Central America. In our region, cone" pines. Van der Burgh (1973) divided this
these are P elliottii var. densa, P caribaea with its subsection into two subsections, restrictingsubsect.
threevarieties,P cubensis, and P occidentalis.Their Oocarpae to the Mexican-Central American pines
and adding species to both. His subsect. Attenuatae
synapomorphyis a multinodallong shoot (not in P
consists
of P muricata, P radiata, P attenuata, P
is
The
at
all, repreoccidentalis).
group poorly, if
greggii, and P. patula. Cladistic analysis revealed
sented in the fossil record.
that Little and Critchfield's subsection is paraphySubsects. Ponderosae Lotuldon and Pseudostrobi letic. The first three species of van der Burgh's
van der Burgh.- Little and Critchfield(1969) clas- subsect. Attenuatae form a clade with P greggii
46
and/or P coulteri (trees not shown). Millar (1986a)
considered the only truly naturalgroup in subsect.
Oocarpae sensu Little and Critchfieldto be the one
that comprises the three Californian closed-cone
pines. Pinus coulteri is classified in both systems in
subsect. Sabinianae,togetherwith two (or one) other
Californian pines with large cones and prolonged
apophyses. It shares the characters Millar (1986a)
mentionsas distinctfor the closed-cone pines: medial
resin ducts, multinodal long shoots, and armed,
asymmetrical,serotinouscones. The cladistic analysis does indicate,contraryto Millar's (1986a) objections, the existence of two groups similar to van
der Burgh's classification, with P coulteri possibly
related to the Californianclosed-cone pines. Pinus
greggii shares clustered, asymmetrical, serotinous
cones, multinodallong shoots, and medial resin ducts
with the Californianclosed-cone pines. The position
of P patula is less clear. Pinus durangensis, P
jaliscana, P. praetermissa, and P tecunumanii,and
more distantly, P oocarpa and P pringlei, seem to
be related to it. This outcome gives some support
for an amended subsect. Oocarpae. Pinus lawsonii,
P pringlei, and P teocote are provisionally retained
in it. The closed-cone pines (subsect.Attenuatae)are
abundantlyrepresentedin the fossil record(Axelrod,
1980, 1986). Most are from the Pleistocene of California and resemble or belong to extant taxa of the
region, but some are older and date as far back as
the Miocene.
Flora Neotropica
either a pair of closely related species (Perry, 1991;
Perez de la Rosa, 1993) or a single species (Carvajal & McVaugh, 1992). Pit2uslambertiana is more
closely related to P ayacahuite in this cladogram.
The homogeneity within subsect. Strobi relative to
the subgenus as a whole seems to indicate that these
species have radiatedrelatively recently, yet fossils
very similarto P lambertianahave been found in the
Middle Eocene nearSan Diego, California(Axelrod,
1986). Interestingly,species of this subsection are
rare in Mexico and Central America. Pinus strobiformis representsthe northernelement, P ayacahuite
the southernelement in highland pine forests, while
each is accompaniedby numerousspecies in subgen.
Pinus (see also Mirov, 1967). In all, six taxa of
subsect. Strobioccur in our region, two of them only
just south of the U.S.-Mexican border. It seems
that this subsection originated elsewhere, either in
northernNorth America or, perhaps more likely, in
Asia. No species of subsect. Cembraeoccurs in our
region, and only one (P albicaulis) occurs in the
United States and Canada and is strikingly similar
in its seed cones to the Eurasian species-pair P
cembra and P sibirica. For both subsections, an
early Tertiaryorigin is supportedby palaeobotanical
evidence (Axelrod, 1986).
SECT. PARRYANAE MAYR
Nearly all recentcommentatorson pine phylogeny

and classification agree that Pinus sect. Parryanae
Mayr,as interpretedby Little and Critchfield(1969),
is extremely heterogeneous,more so than any other
Subgen. Strobus (Sweet ex Spach) Lemmon
section commonly recognized in the genus (van der
Burgh, 1973; Klaus, 1980; Farjon, 1984; Strauss &
SECT.STROBUSSWEETEX SPACH,SUBSECT.STROBI Doerksen, 1990; Malusa, 1992; Styles, 1993a). Only
LOUDON
one of its threesubsections,Cembroidess.l., is repreRecent cladistic analyses of morphologicalcharac- sented in our region,but the heterogeneitywithin this
ters (Malusa, 1992; Farjon & Styles, this volume) subgroupis nearly as great. Phylogenetic analysis of
and of molecular characters (Strauss & Doerksen, both morphological and molecular characters(Ma1990; Govindarajuet al., 1992; Wang & Szmidt, lusa, 1992; Strauss & Doerksen, 1990; Wang &
1993; Perez de la Rosa et al., 1995) agree on the Szmidt, 1993; P6rezde la Rosa et al., 1995; Farjon&
monophyly of sect. Strobus and the paraphyly of Styles, this volume) gives evidence for paraphylyin
sect. Parryanaewithin the subgenusStrobus.Within sect. Parryanae,indicatingthat it is unlikely to be a
the section Strobus, the subsections Cembrae and naturalgroup. Malusa (1992), using Wagner parsiStrobi are very similar (Strauss & Doerksen, 1990) mony analysis of mainly morphological characters
and either monophyletic (Govindarajuet al., 1992; for all species in the section, found a clade consisting
Wang & Szmidt, 1993, who looked at Asiatic spe- of P. maximartinezii,P rzedowskii, P nelsonii, P
cies) or unresolved/paraphyletic(Strauss & Doerk- pinceana, and the Asiatic pair P gerardiana and P
sen, 1990). The strict consensus tree using Fitch bungeanato form a sister group to the other "pinyon
parsimonyanalysis of morphologicalcharacters(Fig. pines" of the traditionalsubsect. Cembroides.Perez
12), thoughcomparingonly a limited numberof taxa de la Rosa et al. (1995) found P. nelsonii to be very
in subsect. Strobi, separates Pinus ayacahuite from distinct in that it possessed neitherof the two length
P strobiformis,two taxa commonly consideredto be mutationsfound in the cpDNA of 11 other Mexican
47

species of Pinus representingboth subgenera.Pinus
pinceana and P. maximartineziishare one synapomorphy in their cpDNA, but none with P cembroides.The strictconsensus tree found in our analysis (Fig. 12) seems to confirm Malusa's findings
which place these pines clearly outside a group with
P cembroides(subsect. Cembroidess.str.).This tree
supportsthe theory that the four Mexican species P
maximartinezii, P rzedowskii, P nelsonii, and P
pinceana are relict taxa, today with (very) limited
and probably naturallydecreasing distributionsand
populations. They very likely had relatives in the
geologic past which became extinct. Their treatment
in the same section or subsection with P cembroides
and its allies has no phylogenetic basis. Pinus nelsonii has a numberof autapomorphies(see Systematic Treatment)that set it apart;a separatesubsection
has been proposed for it (Carvajal, 1986). Pinus
nelsonii and P pinceana appear to be united on a
clade in the strictconsensus tree (Malusa, 1992; Fig.
12). As these two species also form a pair of related
terminal taxa in all trees generated from our data,
we proposeto include P pinceana in that subsection.
Pinus rzedowskii has a seed cone quite unlike any
"pinyon pine"; it could representa more primitive
cone type. It has also seeds with large, effective
wings, which almost certainly is the plesiomorphic
state in pines. Fossil evidence is as yet of little help:
almost all of the materialthat is known from fossils
with resemblanceto subsect. Cembroidess.l. seems
closest to P cembroidesor P edulis. Axelrod (1986)
discusses the possibility that the fossil P lindgrenii
Knowlton,from the Late Miocene in southernIdaho,
is intermediatebetween these and P pinceana. Pending a comprehensive study of morphological and
molecular characters,it seems most appropriateto
recognize subsect. Cembroidesin a strictsense in this
monograph,to recognize subsect. Nelsoniae Carvajal
with two species, and to list the two remaining
aberrantspecies as unclassifiedwithin subgen. Strobus. Section Parryanae is marked with "?" in the
classificationbelow, to indicate its paraphyly.

Based on the considerations outlined above, the
taxa of the genus Pinus accepted in this monograph
are provisionallyclassified; it must be stressed that
the ranking presented below does not necessarily
follow from the cladogramsproduced.Most subsections neverthelessformed lower-orderclades within
the consensus trees.
Pinus subgen. Pinus
Sect. Leiophyllae
la. P leiophylla var. leiophylla
lb. P leiophylla var. chihuahuana

Sect. Pinus
2. P tropicalis
Sect. Pinaster
Subsect. Contortae
3. P. contorta var. murravana
4. P. herrerae
Subsect. Australes
5a. P caribaea var. caribaea
5b. P caribaea var. bahamensis
Sc. P caribaea var. hondurensis
6. P elliottii var. densa
7. P cubensis
8. P occidentalis
Subsect. Ponderosae
9. P ponderosa var. scopulorum
O1a.P arizonica var. arizonica
lOb. P arizonica var. cooperi
IOc. P arizonica var. stormiae
I 1. P jeffreyi
12. P engelmannii
(?) 13. P hartwegii
Subsect. Pseudostrobi
14a. P pseudostrobus var. pseudostrobus
14b. P pseudostrobus var. apulcensis
15a. P. montezumae var. montezumae
15b. P mnontezumaevar. gordoniana
16. P devoniana
17. P douglasiana
18. P maximinoi
(?) 19. P lumholtzii
Subsect. Oocarpae
20a. P. oocarpa var. oocarpa
20b. P. oocarpa var. trifoliata
21. P praetermissa
22a. P patula var. patula
22b. P patula var. longipedunculata
23. P jaliscana
24. P tecunumaniii
(?) 25. P durangensis
(?) 26. P lawsonii
(?) 27. P pringlei
(?) 28. P teocote
Subsect. Attenuatae
29. P muricata var. muricata
30. P radiata var. binata
31. P attenuata
32. P. greggii
(?) 33. P coulteri
Pinus subgen. Strobus
Sect. Strobus
Subsect. Strobi
34a. P avacahuite var. avacahuite
48
Flora Neotropica
34b. P ayacahuite var. veitchii
35. P lIambertiana
36. P flexilis var. reflexa
37. P strobiformis
38. P strobus var. chiapensis
(?) Sect. Parrvanae

(?) 39. P rzedowskii
(?) 40. P maximartinezuii
Subsect. Nelsoniae
41. P nelsonii
42. P piniceana
Subsect. Cembroides
43al. P cembroides subsp. cembroides
var. cemlbroides
43a2. P cembroides subsp. cembroides

var. bicolor
43b. P. cembroidessubsp. lagunae
43c. P cembroides subsp. orizabensis
44. P culminicola
45. P remota
46. P monophyllla
47. P quadrifolia
EVOLUTION
The Mesozoic, lasting from ca. 225 Ma to ca. 65
Ma, was the era of conifers; there were many more
forms than there are at the presenttime, and most of
these became extinct. The evolution of the modern
conifer families-e.g., Araucariaceae,Cupressaceae,
Pinaceae, and Podocarpaceae-has been interpreted
as a radiation from the Permian-MesozoicVoltziaceae (Voltziales),which in turnevolved from Palaeozoic Lebachiaceae(Florin, 1951, 1954; Miller, 1982).
However, there is cladistic evidence that different
genera in Voltziaceaewere possibly ancestralto different modernconifer families, one clade leading to
Cupressaceae,anotherto the other families, and that
some genera were not ancestralto any extantconifers
(Miller, 1988). The Pinaceae must have evolved
from a common pinaceous ancestorsometime in the
Jurassic, as several distinct lineages, among them
Pinus, were already present at the beginning of the
Cretaceous.The organ genus Pityostrobus(Nathorst)
Dutt possibly representsseveral genera of pinaceous
structurallypreserved ovuliferous cones belonging
to Pinaceae. More than 20 species have been described so far. Almost all of these fossils have some
characters assigned to Pinus but also have other
features not known in modern pines (Miller, 1976).
The thickened scale apex (apophysis?) is the only
morphological character;the others are anatomical
and relate to the position of vascularstrandsor traces
and resin ducts in the bract-scalecomplexes. Despite

these peculiarities of their anatomy (how accurate
are these observationsin each case?), several of these
cones probably represent fossil species of Piiis.
These fossils date from the Lower Cretaceous(Wealden, ca. 125 Ma) as well as from youngerCretaceous
to Palaeocene (Axelrod, 1986) formations;they suggest that Pinus may have a central place in the
phylogeny of the family (Miller, 1976, 1977). Older
pinaceous cones (e.g., ComisostrobusDelevoryas &
Hope, from the Late Triassic of North Carolina)
are as yet of more doubtful affinity, due mainly to
imperfectpreservationof these remains.
Unambiguousevidence of macrofossilsdates back
to the Lower Cretaceous(Miller, 1976, 1977; Millar,
1993). Structurallypreserved ovulate cones of the
genus (Pilnus belgica Alvin) are known from the
Wealden (Neocomian) of Belgium; several other
early species date from the Lower Cretaceousto the
Eocene/Oligocene.Almost all appearto be members
of subgen. Pinu.s,having small, conical cones with
dorsal (abaxial) umbos (Miller, 1976, 1977), which
are comparable to the modern cones of subsects.
Contortae, Oocarpae, and Pinus (syn. Svlvestres).
Cretaceousfossils of anatomicallypreservedsecondary leaves reveal diploxyl vascularizationin accordance with subgen. Pinus (Miller, 1977). The only
evidence for subgen. Strobusto have occurredin the
Cretaceous is a cone described as P. magothensis
Penny from Delaware,of which the original material
has been lost (Miller, 1977). Similarly described
cones have more recently been reassigned to the
organ genus Pityostrobus.
The origin and early radiation of pines, being
xerophytes, is likely to have occurred in relation to
increased seasonality, especially aridity, of which
there is evidence in the Early Cretaceous (Mirov,
1967; Axelrod, 1986). By the Middle Eocene, a
majorshift to the spreadingof dry climate in southwesternNorthAmericacreatedconditions favourable
to furtherspeciationin Piinus(Axelrod, 1986; Millar,
1993).
Shaw (1914) based his classification of the pines
on "the gradual evolution of the fruit from a cone
symmetricalin form, parenchymatousin tissue, indehiscent and deciduousat maturity,releasing its wingless seed by disintegration[subsect. Cembrae]-to a
cone oblique in form, very strong and durable in
tissue, persistenton the tree, intermittentlydehiscent,
releasing its winged seeds partly at maturity,partly
at indefiniteintervalsduring several years." He concluded: "Thisevolution embracestwo extreme forms
of fruit, one the most primitive, the other the most
Distributionand Ecology
49
elaborate,among conifers."When the cone morphology of Pinus is compared with the other genera in
Pinaceae, neitherof these types occur outside Piinis.
The "plesiomorphic"type would seem to be the
pendulous, deciduous, and more or less cylindrical
cone with relatively thin, freely parting seed scales
releasing seeds with adnate wings. Secondly, phylogenetic analysis of many charactersclearly indicates
that the species of subsects. Cembrae and Strobi
have evolved ratherlaterthanthose with "less primitive" cone types (Fig. 12; see also Strauss& Doerksen, 1990; Wang & Szmidt, 1993). We now know
that Shaw's "most primitive"cone type in reality is
the outcome of coevolution involving seed dispersal
by corvid birds, e.g., the relationship between P
cembra and Niucifraga(carvocatactes in the Alps and
similarmutualismsbetween nutcrackersandjays and

P albicaulis and P edulis in the United States
(Mattes, 1994;Tomback& Linhart,1990). The serotinous cones (in several species of subgen. Pinuss)are
indeed a resultof "elaborateevolution,"but no more
so than those of subsect. Cem7brae. The admittedly
scant fossil recordfrom the Mesozoic gives evidence
contrary to Shaw's ideas on pine cone evolution
(Miller, 1977). From the Tertiaryof NorthAmerica,
no fossils have been ascribed to subsect. Cembrae
(Axelrod, 1986); from Eurasia there is fossil evidence from the Palaeogene only (Gaussen, 1960).
Given the probability of monophyly of the two
subgenera, the ancestral cone form of Piinuswould
have had apophyses with a dorsal umbo, a condition
which now prevails in subgen. Piinusand is present
in a part of subgen. Strobhs.The "typical"Strobus

cone with a terminalumbo is then the derived state.
The Cembrae cone, with nearly indehiscent scales
and vestigial seed wings, is a furtherspecialization
from this lineage, in coevolution with seed dispersal
by birds. Serotinouscones with asymmetricaldevelopment of apophyses are derived from the ancestral
type. Their evolution is probably linked with more
extreme conditions of climate, but defense against
predationmay also have played a part. Most species
with such cones occur at present in regions with
climatic extremes, especially prolongeddrought.
Evolutionarytrendsin foliage charactersare difficult to reconstruct.Leaf number is highly variable
in most groups; it is most consistent in subsects.
Pinus (s.str.)and Strobi.There is a very general trend
toward increased numbers from boreal to tropical
North America, apparentlyrelated to climate. Pinus
nelsonii is exceptional within subgen. Strobus in
having fascicles enclosed in persistentsheaths. This
condition and a variable (low?) number of leaves
may representthe primitive state. The fossil record
from the Cretaceous is too fragmentary to draw
conclusions, yet the earliest pines known were of
the few-leaved, diploxyl type (Miller, 1976, 1977).
Convergence or parallelism, such as the deciduous
fascicle sheaths of P. leiophylla and P. Ilutholt.ii. or
the "wingless"seeds in the MediterraneanP pinea,
seems to have occurredoccasionally. However, until
we know in more detail how these characters
evolved, we cannot be entirely certain of their homology.
DISTRIBUTION AND ECOLOGY

DISTRIBUTION
(for P ctiribaea var. bahllmensis in the Bahamas) to
The genus Pinus is the most widespreadgenus of

Pinaceae,extending across the NorthernHemisphere
roughly between latitudes 15?N(-02?S) and 66?
(-70?)N in NorthAmericaand Eurasia(Fig. 13). The
most southerly occurrence in the Western Hemisphere is at 12?N in Nicaragua, where Caribbean
coastal populationsof Pinus caribaea var. hondurensis reach as far south as Bluefelds (LUckhoff,
1964). As with other genera in the family, species
distributionsare very large in the boreal zone but
species diversity is low; the distributionsare reduced
towardthe oceanic coasts and in the southernmontane regions, and species diversity increases(Farjon,
1990a).Altitudinaldistributionranges from sea level
4000 m (for P hairtwegiiin Mexico). Other genera

in the fanily (Abies, Picea) cover similar or even
greater altitudinal ranges, but their low-elevation
species occur only in northernlatitudes,as compared
to several low-elevation pines in the tropics. Even
though the altitudinalranges of these genera reflect
their latitudinalranges, as both are correlatedwith
climate, the pines can be said to have attained the
greatestoverallenvironmentalamplitude.Special adaptations(see Ecology, below), resulting in species
radiation,have allowed the pines to spread into the
warmerand drierenvironmentsof southernlatitudes
and lowlands.
Unlike the other medium-sizedto large genera in
Pinaceae, species diversity in Piins is greater in
50
Flora Neotropica
FIG. 13. General distributionof the genus Pinus in North America and Eurasia, showing gaps in midwestern North
America and SW and CentralAsia.
North America than in Eurasia.Geographically,this

coincides with a more complete occupation of the
North American continent as compared to Eurasia.
In North America, the genus is totally absent only
in the Arctic, on the Great Plains, and in scattered
deserts. In Eurasia,it is absent in the greaterpart of
Central,Southwest, and South Asia, with only notable exceptions in the Turco-Caucasianregion and the
Himalayas. While pines occur in many areas within
the deciduous forest zone of the eastern United
States, they are virtually absent in the equivalent,
but impoverished,zone in Europe. In the subtropics
and tropics of the Western Hemisphere, pines likewise occupy an almost continuous belt and more
groundthan in SoutheastAsia, where they are mostly
scattered. Three North American regions of high
species diversity in the genus Pinus can be recognized: I) Mexico, 2) California,and 3) southeastern
United States. These are here defined in a phytogeographical sense: Mexico extends into the United
States north to the Mogollon Rim in Arizona and
New Mexico, and into CentralAmerica as far as the
distributions of pines are concerned to Nicaragua;
California includes parts of Oregon, Nevada, and
northernBaja California. Rzedowski (1993) recognises an extended Mexican region (Mega-Mexico 3)
in a similar sense to define Mexican endemism. Of
the 47 species and 20 infraspecific taxa recognized

in this monograph,42 species and 18 infraspecific
taxa occur in Mexico (all taxa in Central America
are also in the country of Mexico, including P
caribaea var. hondurensis; see, e.g., Eguiluz, 1982;
Styles & Hughes, 1983). Five species and two varieties occur in the Caribbean,one of them, P. elliottii
var. densa, on the Florida Keys and mainlandcoast.
Nine taxa are Californian;one of these, P radiata
var. binata, is an island endemic that does not occur
in Californiabut has strong affinities with P radiata
var. radiata, which is confined to that state. The
coastal and insular Californian pines are relicts of
a Madro-Tertiaryflora confined to fog-influenced
maritime and disjunct distributions resulting from
the northwestwardspreadof the Sonoranand Mojave
Deserts (Axelrod, 1976, 1980).
Rzedowski (1993) mentions the genus Pitius
among the genera for which (parts of) Mexico provided centresof speciation.The other possible source
of species diversity in a given region is migration.
In order to evaluate the relative importanceof each
of these factors,it is necessary to look at the distribution of pines on the North American continent and
its environs as a whole. The species that extend into
Mega-Mexico 3 [as defined by Rzedowski (1993)]
have distributionsthat can be grouped into six re-
Distribution and Ecology
51
lo,
:/..F`
j -t
--.?
...../
-1"-
^-/^
p<'^
_.../
-_
z 'Ji ..
.
1?.
-o
FIG. 14. Regions of North American pine species extending into Mexico and the SE United States-Caribbean-Central
American connection. A. Californianregion. B. Great Basin region. C. Rocky Mountainregion. D. N Mexican region.
E. Mexican
Mexican Gulf-Atlantic
Gulf-Atlantic coastal
coastal plain
F. Caribbean
Caribbeanregion.
plain region.
region. F.
E.
region.
gions (Fig. 14). Two of these-the Californianregion

(A) and the Mexican Gulf-Atlantic coastal plain
region (E)-are North American centres of pine diversity, as is Mexico itself. From neither of them
have species migratedvery far northinto Mexico. In
BajaCalifornia,the only truly migratoryspecies may
well be the high-elevationones, i.e., P contorta var.
murrayana,P.jeffreyi, and P lambertiana,which are
distributedon the higher portions of the block-fault
zone that runs from here far into California.Its uplift
started in the Late Tertiaryand still continues. The
pinyon pines of Baja California (P monophylla, P
quadrifolia), as well as P coulteri, occur at lower
elevations and may have spread from their centres
of origin following the desertificationof the southwestern partof the continent. Pinus monophyllais a
Great Basin element, and the other two are Californian. In the southeastern United States there is an
abundanceof pine species, six of which occur in the
lowlands of Florida. Only P elliottii var. densa extends a short way into the Caribbean;the closely
related P caribaea occurs with two varieties in the
lowland pine savannas of the Caribbean islands.
From there, a third variety (P caribaea var. hondurensis) has somehow made it across the Cayman
Trough-perhaps its seeds were carried by hurricanes.
The much less diverse Rocky Mountain pine region (C) has been of more significance in the spread
of pines into greaterMexico. Two taxa with a northerly distribution(P flexilis var. reflexa and P ponderosa var. scopulorum) reach into the country of
Mexico, and several of the pines restricted to the
northernMexican region (D) are closely related to
species occurring in C and are derived either from
these or from a common ancestor. These are P
arizonica (with 3 varieties), P durangensis,P engelmannii, P flexilis var. reflexa, and P strobiformis.
The pinyon pines common to these two adjacent
regions-P. cembroides var. cembroides. P cernbroides var. bicolor, P edulis, and P remota-are
constituentsof the widespreadpinyon-juniperwoodland ecosystem, in which the minor diversification
of the species as well as the coevolution with corvid
birds related to seed dispersal strategies (see Ecology) point to a relatively recent radiation related
52
Flora Neotropica
CARIBBEAN AMERICA
II? ??)~??
FIG 15 Nmesotaa(pcs,
counries
an Caiba
islnds
suseis
?~
,' . .
aite)i
hisi
'
tesae
~:_; ...i
fMxc.CnrlAeia
FIG. 15. Numbers of taxa (species, subspecies, and varieties) in Pinui.sin the states of Mexico. Central American
countries, and Caribbeanislands.
to climatic change. The common ancestor of this

monophyletic group of pines (see Phylogeny and
Evolution) probably originated somewhere in the
Mexico-United States borderarea.
Eguiluz (1985) recognized six principalregions of
distributionof Mexican pines. These regions with
majordiversity are I) the Sierra Madre Occidental,
2) the Sierra Madre Oriental, 3) the Eje Volcdnico
Transversaland Mesa Central, 4) the Sierra Juarez
and Sierra San Pedro Martir in Baja California, 5)
the Sierra Madre del Sur and Macizo de Oaxaca,
and 6) the highlands of Chiapas, which extend into
Guatemala.These were termed "centres of species
diversity of pines in Mexico" by Styles (1993a). The
Sierra Judrez and Sierra San Pedro Martir in Baja
California, as mentioned above, have only Californian pines. Mappingthe numberof taxa in each state
of Mexico roughly confirms Eguiluz's regions, but
the Mesa Central, being a distinct morphotectonic
province (Ferrusqufa-Villafranca,1993) is conspicuously less diverse than the Eje VolcanicoTransversal
(Fig. 15). High diversity in northeasternMexico is
partly explained by past migrationsfrom the Rocky
Mountainregion. Anotherquite different component

is local endemism, with species such as P culminicola, P nelsonii, and P pi,nceana.
As noted above,
this region is also the most likely centre of origin of

the species of Pinus subsect. Cembroides. Toward
the southeast,diversity decreases, with the numberof
taxa in Chiapas (along with Oaxaca. the botanically
richest state of Mexico; see, e.g.. Graham, 1993)
alreadysignificantlyless, until the distributionof the
pines reaches its terminationwith only four species
in Nicaragua. Many of these southern pines are
widespreadspecies, pointingto migrationas the most
likely cause of pine diversity in Mesoamerica.South
of the Eje VolcanicoTransversal,local endemism of
pines is absent, in line with the scarcity of local
endemics in most other plant groups (Rzedowski,
1993). This, too, is an indication that speciation
has not occurred in southern Mexico and Central
America, possibly with the exception of P tecunumanii, which may be a relatively recently derived
taxon.
From this evidence, two centres of pine diversity
anldevolutionemerge: the Eje VolcdnicoTransversal,
53
Uses and Conservation

with extensions to the northwestand southeastalong
the Sierra Madre Occidental and Sierra Madre del
Sur, and northeasternMexico. Here we find 14-18
taxa in most states (only the very small states around
Mexico City have fewer taxa, as expected). The
species of the first centre are almost all Mexican
taxa sensu stricto. This centre is also the major
area for the three polymorphicspecies complexes P
and P pseudostroblts(for
devoniana, P monntezumae,
the first two see also Styles, 1993), which may have
radiated from here over much of montane Mexico
and CentralAmerica. The geological history of two
of its main mountainchains is similar (FerrusquiaVillafranca,1993). The SierraMadreOccidentaland
the Eje VolcanicoTransversalare both volcanic complexes-the first with a long history of volcanism
beginning in the Upper Cretaceous and encompassing the Tertiary,the latterof more recent origin.
The SierraMadredel Sur has a much more complex
geology, but (high) mountainshave existed here at
least since the early Tertiary.The second centre of
pine diversity and evolution is characterizedby 1)
adaptationto aridity, 2) disjunctionon high "island
mountainranges" surroundedby (semi-)desert, and
3) local endemism. Speciation as well as migration
from the Rocky Mountainregion are the main causes
of pine diversity here. A more ancient geological
historyof this partof Mexico-with Late Cretaceous
to Early Tertiaryuplift of mostly Mesozoic marine
sediments, forming mountain ranges separated by
intermontanebasins (Ferrusquia-Villafranca,1993),
and climatic change involving desertificationin the
Miocene (Millar, 1993)-mostly explains these patterns.
A somewhat different approachis to analyze the
distributions, both geographical and altitudinal, of
all taxa and classify these. Altitudinal ranges have
been computed from the specimen database for this
monograph; for altitudinal ranges of species occurring both in Mexico and the United States, see
also Yeaton (1982). The geographical distributions
(see maps in SystematicTreatment)can be classified
into seven regions:
1.
II.
III.
IV.
V.
VI.
VII.
California
northwesternMexico
western Mexico
northeastern& eastern Mexico
central & southernMexico
Mesoamerica
Caribbean
The altitudinalranges can be classified into four

categories:
1.
2.
3.
4.
lowland [1-300(-700) ml
foothills [(100-)300-1200 m]
montane [(700-)1000-2600(-2800) ml
high montane [(2000-)2500-4000(-4300) ml
In Table III all taxa have been tabulatedagainst

these two parameters.In region I, the Californian
taxa are predominantlymontane,but two taxa occur
at lower elevations. Their altitudes have been given
only for Mexico here, but these do not greatly differ
from the distributionsfound in California.
In northwesternMexico (region II), most distributions are confined to the Sierra Madre Occidental.
Nine taxa are montane and four are high montane.
Most of these populationsoccur some distance from
the coast, and the lower and middle elevations tend
to be much drier than similar altitudes nearer the
coast, especially on the continentalside of the Sierra
Madre. Here, the drought-resistantP cembroides
formsan intermittentbelt below the mainpine forests
(Perry, 1991). Pinus cembroides var. bicolor occurs
at higher altitudes but in dry regions with cold
winters.
WesternMexican species (region III) have distributions predominantlyin the southernSierra Madre
Occidentaland westernmostpartof the Eje Volcanico
Transversal,partly overlappingwith those of northwestern Mexican taxa. Most taxa in this region are
montane; the higher precipitationlevels allow pine
forest to grow at lower altitudes.This is particularly
evident on the oceanic slopes of the mountainranges.
Here we find the greatest diversity of pines in
Mexico.
In northeasternand eastern Mexico (region IV),
distributionsare concentrated in the Sierra Madre
Oriental,with extensions to the northwestand in the
easternmost part of the Eje VolcainicoTransversal.
Most taxa are montane;several are scatteredalong
an intermittentbelt below the main pine forests on
the continental side of the Sierras, e.g., P c-embroides, P nelsonii. and P pinceana.
A few species are restrictedto southern Mexico
(region V): P lawsonii, P.pringlei, and P strobusvar.
chiapensis, the latterextending into the highlandsof
Guatemala.They occupy montaneregions in a section of Mexico where the distance to the coasts is
much reduced, with the ameliorating influence on
climate as mentionedabove.
Othertaxa in centraland southernMexico (region
V) have very wide distributions.Most of them have
very wide altitudinalranges, too. These are due in
part to an upward shift of the lower (and upper)
altitudinallimits from southeast to northwestgoing
Table III
Distribution of Latin American pine species and infraspecific taxa according to geographical regions (II (California)
I (lowland:
1-300[-700] m)
II (NW Mexico)
III (W Mexico)
IV (NE & E Mexico)
2 (foothills:
P. attenuata
[100-]300-1200 m)
3 (montane:
[700- 1000-2600
[-2800] m)
4 (high montane:
[2000-]2500-4000
[-43001 m)
V (C & S M
P. muricata
P. contorta var.
nurravana
P. coulteri
P. jeffreyi
P. lambertiana
P. monophylla
P. quadrifolia
P. radiata var.
binata*
P. devonian
P. maximino
P. oocarpa
P. arizonica + var.
cooperi
P. cembroides
P. devoniana
P. engelmannii
P. leiophvlla var. chihuahuana
P. leiophylla var. leiophylla
P. oocarpa
P. teocote
P. cembroides
subsp. lagunae*
P. devoniana
P. douglasiana
P. durangensis
P. herrerae
P. jaliscana*
P. Ieiophylla var.
chihuahuana
P. leiophylla var.
leiophylla
P. lumholtzii
P. maximartinezii*
P. maximinoi
P. montezumae
P. oocarpa
P. praetermissa
P. pseudostrobus +
var. apulcensis
P. rzedowskii*
P. teocote
P. arizonica var.
stormiae
P. cembroides +
subsp. orizabensis*
P. devoniana
P. engelmannii
P. greggii
P. leiophylla var. leiophylla
P. montezumae
P. nelsonii*
P. patula + variety
P. pinceana
P. pseudostrobus
P. remota
P. teocote
cembroid
devonian
douglasia
herrerae
lawsonii
leiophylla
leiophylla
P. maximino
P. montezum
var. gordoniana
P. oocarpa +
P. patula + v
P. pringlei
P. pseudostr
+ var. apu
censis
P. strobus va
chiapensis
P. tecunuma
P. teocote
P. cembroides var.
bicolor
P. flexilis var. reflexa
P. strobiformnis
P. teocote
P. hartwegii
P. montezumae
P. pseudostrobus +
var. apulcensis
P. strobiformis
P. teocore
P. avacahuite + var.
veitchii
P. culminicola*
P. flexilis var. reflexa
P. hartwegii
P. montezumnae
P. pseudostrobus
P. teocote
P. strobiformis
P. avacahuit
P. hartwegii
P. montezum
P. pseudostr
+ var. apu
censis
P. teocote
* Narrow endemic taxa; for furtherexplanation, see text.
P.
P.
P.
P.
P.
P.

from Mesoamerica further into Mexico. With the
widening of the Mexican landmass, interior forests
are blocked from oceanic weather and only receive
sufficient precipitation at higher altitudes. Longer
summer day length away from the equator enables
trees to reach higher altitudes as growing-season
temperaturesare higher, and they can become dormant in the colder season. However, even within
a relatively restricted area the range can be very
is found in Oaxaca
considerable,e.g., P nmontezumae
between 1200 and 3000 m. Most of these widespread
species are morphologically highly variable, with
several varieties being recognized here. Pinus hartwegii is found at the tree line, to an upper limit of
ca. 4000 m, on the volcanos of the Eje Volcanico
Transversalwhich rise above 5000 m. Yet this species occurs down to 2300-2500 m in the highlands
of Oaxaca and elsewhere.
With the one exception of P. culminicola, all
narrowly restrictedtaxa (indicated with an asterisk
in Table III) have a montane distribution with a
narrowaltitudinalrangenot exceeding 1000 m. Pinus
culminicola is a high-montanespecies occurring at
3000-3700 m. The term "endemic"has been variously interpretedin the literaturebut is here restricted
to taxa with a localized distribution (narrow endemic). Rare but scatteredspecies which occur over
a large area-e.g., P muricataand P. pinceana-are
thus excluded from this category; their distributions
are more aptly described as disjunct or interrupted.
No narrowendemics occur in the southernregions;
most are western and northeasternand eastern Mexican.
The Caribbeantaxa are mostly lowland pines, with
P cubensis ascending to a maximum of 1200 m in
the mountainsof eastern Cuba. Several occur only
just above sea level, but this could be due in part to
a limited topography,such as on the Bahamas and
the Florida Keys. One notable exception is the only
pine on mountainous Hispaniola, P occidentailis,
which, with an altitudinal range of (200-)9002700(-3175) m, exceeds thatof all other LatinAmerican pines (Darrow& Zanoni, 1991).
ECOLOGY
The geographical distributionof the Pinaceae in
the Northern Hemisphere reflects the climatically
dominatedecological conditions under which these
conifers thrive and compete successfully. Relatively
low temperaturesduring the growing season are a
limiting factor to deciduous broad-leavedtrees, and
evergreenconifers take advantageof this; as a result
55
they form the dominant forest types northwardas
well as at higher altitudes in mid-latitudemountain
ranges. The duration of the growing season (the
periodwith meandaily temperature> 10?Cfor coniferous trees; Schmidt-Vogt, 1977) shortens progressively with increasing latitude and altitude, until it
reaches the 10?C July-isotherm (Northern Hemisphere) which is the climatic tree line. At the other
end of the temperaturedine, two factors limit successful growth of conifers: droughtand competition
by broad-leaved trees. Moisture stress is usually
related to lack of precipitationat critical periods of
the life cycle, e.g., the seedling and sapling stages.
Competitionby broad-leavedtrees occurs underconditions where limitationsto their success are absent
and one or more potentially successful species are
present.This means that an undisturbedsuccession
of the vegetation under these favourableconditions
will lead to dominanceof broad-leavedtrees.
As can be surmisedfrom its greatergeographical
and altitudinalrange,the genus Pinus has a considerably greater ecological amplitude than any other
genus in the family Pinaceae (Farjon, 1990a). This
is especially true toward the warmerand drier ecosystems of scrubland/woodland bordering semideserts and in pine savannas with prolonged dry
seasons and naturalfire regimes.Adaptationsto these
conditions, involving specializationof vegetative as
well as reproductiveorgans, have led to radiations
of taxa and colonization of environmentswhere few
other trees can survive. Such specialization is especially true for the pines of California and Mexico.
Both regions have experienced great geological and
climatic changes in relatively recent (Miocene
throughPleistocene) geological time; it has therefore
been suggested that much of this diversity reflects
relatively recent evolution (Little, 1962; Axelrod,
1980).
Mexico is a predominantlyupland country, with
more than one-half of its area above 1000 m. The
Central American countries where pines occur are
also mountainous.Extensive lowlands occur on the
Yucatan Peninsula, the Atlantic coastal plains of
Central America, and, on a smaller scale, in the
Isthmus of Tehuantepec.Although P oocarpa descends into the lowlands of the isthmus, only the
Atlantic coastal plain from Nicaraguato Belize has
significantpine forests with P caribaea var.hondurensis. "Pine savannas"also occur on several Caribbean Islands,but here, too, most islands with natural
pine forest are mountainous. Because of Mexico's
extraordinarilyvariedtopographyand climate, which
range from wet, lowland tropical rainforests and
56
Flora Neotropica
montanetropicalcloud forests to hot scrublandsand

deserts and snow-capped mountains, often in close
proximity to each other, virtually all ecotopes in
which pines are found worldwide are also present in
this region. Due to the great elevation of some of
the mountainsand their situation between 12?N and
33?N latitude,climatic zonationis pronouncedwithin
short distances and isolation of populationsand taxa
is commonplace. Several authorshave tabulatedclimatic zones for Mexican pines (e.g., Loock, 1950;
Eguiluz, 1982, 1988), using altitudinal,temperature,
and precipitationgradientsas criteria.
Plant Communities
The vegetation of Mexico has been classified by
Mirandaand Hernndez (1963). Pines occur in chaparral,juniper woodland, pine forest, oak forest, deciduous forest, and fir forest, and in transitional
vegetationas well as mixed types of these. Chaparral,
best developed in northwesternBaja California and
north into Californiaon the lower Pacific slopes of
the mountainranges, is a semi-evergreenvegetation
dominatedby sclerophyllousshrubs.There is a counterpartin parts of the foothills and lower slopes of
the Sierra Madre Oriental, referred to as montane
chaparral. Pinus attenuata, P muricata, and, to a
lesser extent, P monoplhyla and P quadrifolia are
associated with the Pacific variant;P cembroides,P
culmniicola, P nelsonii, and P pinceana with the
interiortype. There is a climatic difference between
the two types: the Pacific coast type (chaparralin
the strict sense) has winter rains (and summerfog in
its lower reaches) and is largely frost-free;the interior type has summer rains (thunderstorms),occasional frost, and, at high elevations, snow. Both are
subject to natural fires by lightning. Adaptations
are diverse and specialized. Prolonged drought and
intensive radiation have selected for short, rigid
leaves. Along the Pacific coast, where fires are both
relatively frequent, with 20-50-year intervals in
closed-cone pine forests, and intense,the closed-cone
type of seed dispersal, requiringextreme heating of
the outer partsof the persistentand serotinouscones
to open, has evolved in a number of species. The
seeds have articulatebut effective wings, which enable the seeds to be transportedsome distance and
be scattered sufficiently to land in places where fire
has eliminated the chaparralundergrowth.
Juniper woodland is usually an open arborescent
vegetation type with well-interspersedsmall trees, in
which P. cembroides and/or its close relatives may
occur in various proportions;the other components
are species of Juniperusand often of Quercus.This

type is the equivalentof the pinyon-juniperwoodland
of the United States, especially of the variants in
southernArizona and New Mexico. This woodland
forms the lower limit of the upland forests in much
of the interiorpartof northernMexico, especially on
the semi-arid slopes and mesas bordering the two
Sierra Madres. Studies (papers in Everett, 1987) of
succession after disturbance, as well as of stand
structuresof mature woodland types, indicate that
Juniperus would dominate the vegetation, perhaps
even to the exclusion of Pinus, in the prolonged
absence of majordisturbancessuch as fires or (over)grazing. In all pinyon pines and some white pines,
seed is bird-dispersed.In researchcarriedout in the
United States, the pinyon jay (Gymnorhinuscvanocephalus) and Clark's nutcracker(Nucifraga columbiana) are the bird species involved in pine seed
dispersal (Tomback & Linhart, 1990). No detailed
studies on these mutualisms have been carried out
for Mexican pinyon pine species, but it has been
suggested that two species in the genus Apheloconma
and the raven (Corvus corax) perform a similar
function (Peterson& Vargas-Barajas,1993). The extreme similarity and homology of the cones and
"wingless" seeds of P cenbroides and its (close or
more distant) allies-including P edulis, for which
mutualism with Clark's nutcrackerand the pinyon
jay has been demonstrated(VanderWall & Balda,
1977; Ligon, 1978)-make it very likely that such a
bird connection exists in Mexico as well. The scrub
jay (Apheloconmacoerulescens) is largely dependent
on pinyon seeds where it lives in this type of woodland and stores them, yet this bird also occurs in a
variety of habitatswhere no pines exist (Peterson &
Vargas-Barajas,1993); furtherstudies on the bird's
ecology seem to be necessary to determinethe relationship between tree and bird. On the other hand,
similar but larger seeds occur in P mnaximartinezii,
which has developed an extremely large, hard-scaled
cone from which the seeds are not easily removed
as they are in P cembroides and its allies. It seems
unlikely thatjays can handlethese cones as they can
the small, soft-scaled cones of the other pinyon
pines. Is the raven responsible for dispersal here?
Differences in thicknessof the seed coat and content
of starch and fatty oils in the seed among various
species of pinyon pines may also be of as yet unknown ecological significance.
Pine forests ("Pinares")are a heterogeneous assembly of forest types dominatedby species of Pinus
(Miranda & Hernandez, 1963). They extend from
the lowlands to the tree line and from xeric pinyon
Distributionand Ecology
pine woodland to mesic montane mixed pine forest
with a tall, closed canopy. Foothilland lowlandtypes
phase into "pine savannas"on the Atlantic coastal
plains in Central America, where trees are widely
spaced and the groundcover is dominatedby grasses,
with scattered fire-resistant(resprouting)shrubs. In
the Caribbean,P. caribaea, P. elliottii var.densa, and
P tropicalis occur in pine savannas, but in Central
America only P. caribaea var. hondurensisoccupies
this type of vegetation. Seedlings have an elongated
stem (P caribaea var. hondurensis) or develop a
"grassstage";both aredefences againstvery frequent
grass fires which usually occur every few years.
Annual precipitationcan be high, but it is unevenly
distributed and there is a prolonged dry season.
Edaphic conditions prevent moisture retention on
most pine savanna sites, and in the tropical climate
total evaporationis extremely high. In the adjacent
upland areas, precipitationincreases rapidly, especially on sites exposed to moisture-ladenwinds from
the Gulf of Mexico. Here P. cubensis and P. occidentalis are the pines of the Caribbeanuplands,but both
occur as the only species of the genus in their
respectivedistributions.On the mainland,it is mainly
P oocarpa that occupies this zone above the pine
savannas,and more locally also P tecunumanii.The
canopy is more often nearly closed and there is a
predominanceof shrubs in this type of pine forest,
among which are many legumes, but grasses are still
abundantin most places. Fires (many of them manmade) are still frequentbut occur at longer intervals.
Especially P. oocarpa has developed persistentand
semi-serotinouscones, which enables the species to
be opportunisticin its dispersalof seeds after fire.
The more mesic pine forests of the montanezone,
generallybetween (XX)and 2600 m above sea level,
are more diverse. In much of Mexico and Central
America,more thanone species, and not infrequently
as many as 5-8, may occur in a single forest, with
usually some altitudinal zonation involving some
species but with others ranging throughoutthe pine
forest. On one hand, sympatry is common among
pines in these forests, to a greaterextent than anywhere else. On the other hand, there are few if any
other conifer genera present, except at the higher
altitudes. In this they contrast with the equally diverse conifer forests of Californiaand Oregon,which
are "mixed,"i.e., composed of membersof different
families and genera but with only a few representative species of each. Within the montanepine forest
belt, there is a climatic zonation from south to north,
with warmtemperateconditionsprevailingin Central
America and southern Mexico and cool temperate
57
conditions in northernMexico. In the southernparts,
seasonality is dominated by precipitationlevels; in
the north, low winter temperaturesare more important.Adaptationsto these differences are primarily found in the foliage, with long, slender, often
pendulouslax leaves occurringespecially in pines of
subgen. Pinus in the warm temperate regions and
shorter, thick and rigid leaves predominantin the
winter-cold Sierra Madres of North Mexico. The
predominantcone type is thatof Pinus subgen.Pinus,
showing moderateto extreme diversificationin development of the apophyses (see Figs. 6-7), protecting the inner parts of the scales with the seeds
from intenseradiationby thicker,protrudingapophyses on the sun-exposed side. Cones open gradually
and are persistentuntil the seeds with articulatebut
fully effective wings are dispersed. Several species
range from the foothills to high mountainsbut are
nevertheless frequent constituents of the montane
pine forest belt. Their heterogeneity(see above) may
reflect genetic variation as much as environmental
adaptationto the variousclimatic and edaphicconditions they occupy. Pinus devoniana, with large,
woody cones and long, thick leaves, remains a medium-sizedtree and seems to occupy open secondary
woodland more often than its close relative, P mo,ntezurnae,in which these featuresare less pronounced.
Intermediateforms between both species are often
difficult to classify, and variationsof the cones have
often been interpretedas representingdistinct taxa
or hybrids. Investigation of the possible effects of
environmental conditions on the morphology of
leaves and cones in this complex could help delimit
genetically based, fixed characterstates more clearly.
There is a very gradual transition between pine
forest and oak forest at montane altitudes: most
forests classified as pine have a mixtureof Quercus
and most oak forests harbour some Pinus. Many
oaks are deciduous, shedding leaves during the dry
season, i.e., roughlyfrom Novemberto June. Natural
disturbances,such as fire, now increasinglybrought
about by man, tend to promote Pitus, but selective
logging of pines in many areas has given Quercusa
chance to close the canopy, preventing pines from
regeneratingin the absence of calamitousevents.
A few species of Piins are commonly associated
with broad-leavedtreesof warmtemperatedeciduous
forests. For instance, P. strobus var. chiapensis and
P tecunumaniiare almost always growing with LiquiMamnbarstyracifua and various other associated
broad-leaved trees characteristic of the moisturecatching slopes of the mountainsborderingthe Gulf
of Mexico in Mesoamericafrom Chiapasto Hondu-
58
ras. Precipitationis very high, often exceeding 3000
mm annually, and fog occurs year round. Other
species of Pinus thatcommonlyoccur in these forests
are P. patula and P. teocote. The pines often occupy
the steepest slopes and crags at higher elevations
within the temperatedeciduous forest belt. A gradual
transitiontowardoak forest occurs especially toward
the drier eastern slopes, where pine species more
tolerant to drought appear, such as P montezumae
and P pseudostrobus.
On cool, cloudy, wet mountain summits up to
3000 m, or in belts on largersierras or taller mountains, species of Abies-mainly A. guatemnalensis
in Mesoamerica and A. religiosa further north in
Mexico-dominate the coniferous forest. Locally,
stands of Cupressuslusitanica may replace the firs,
or both genera may occur in mixed stands. In this
mixed coniferous forest Pinus avacahuite, P maximinoi, P pseudostrobus, and, often at the upper
limit, P hartwegii are common; in northwestern
Mexico, P strobiformis replaces P ayacahuite. In
Baja California on the Sierra San Pedro Martir,a
Californianmixed coniferous forest occurs with Abies concolor. Pinus lambertiana, P jeffreyi, and P
contorta var. murrayana. This northern type has
warm summers and snowy winters, while the southern equivalents have a more equitableclimate dominated by fog and rain. In these high-montaneforests,
where competitionamong tall-growingconifers (and
fewer broad-leaved trees, mainly oaks) is severe,
effective seed dispersal is of utmost importanceto
reach distant gaps. The species all have cones that
open readily upon drying at maturityto release the
winged seeds (P avacahuite,P lambertiana,P strobiformlis);especially those of P strobiformis, with
reduced wings, are also bird-dispersed.The cones of
these conifers are placed high in the treetops, as in
Abies, or toward the end of higher branches in
the pines, to facilitate long-distancedispersal of the
seeds.
Finally, at the highest elevations in Mexico and
Guatemala, some pines grow at or near treeline.
Foremost among these is P hartwegii, which is
scattered on the highest mountain summits from
Honduras to northeasternMexico. In Mexico and
Guatemala it often forms pure stands, in Mexico
encircling the snowy summitsof the tallest volcanos
such as Pico de Orizaba,Popocatepetl,and Iztaccihuatl, generally to about 4000 m altitude. In northeastern Mexico, the summits of several mountains, all
of which are considerablylower than 4000 m, have
only scattered stands of P hartwegii, and of some
more or less extensive populations of a dwarfed
species closely related to P cembroides, P culmini-
Flora Neotropica
cola. This shrub-likepine may cover extensive areas
exclusively, as on Cerro Potosi, or in association
with other conifer species (Abies, Picea, Pinus) in
"parklandcommunities" or open woodland. Apart
from the populationon Cerro Potosi (3713 m), it is
not a treeline species (Riskind & Patterson, 1975)
but, rather,a species associated with an open, montane chaparral,merging with P hartwegii forest at
more favourablesites, where the calcareous rock is
covered with deeper soils and the slopes are less
exposed to wintry,snow-laden gales from the north.
Mycorrhizae
Most plants form a symbiotic association of fungi
with their roots creating mycorrhizae.All species of
Pinus form ectotrophicmycorrhizaewith the hyphae
of basidiomycetousand ascomycetous fungi; in this
type the hyphae form a mantle aroundthe fine roots
and root hairs and rarely penetratethe cortex cells.
Roots affected by mycorrhizaecease to grow longitudinally; instead the mycelium to which the mycorrhiza is attachedextends the root system, increasing
the absorbingcapacityof the roots. The fungus dislocates the outer cortex cells of the root and forms a
network of hyphae called the "Hartig net." The
symbionts (plant and fungus) cooperate in growth
and function. The role of the fungus is primarily
absorptionof soluble nutrients;that of the host, to
provide the fungus with its assimilation products
(organic carbon compounds). The increased intake
of nutrients-e.g., phosphates-enables the pine tree
to subsist on soil types otherwise too poor for tree
growth. Many differentspecies of fungi are involved
in mycorrhizal symbiosis, both in temperate and
tropicalclimatic zones (Barnes et al., 1996).
Diseases and Predators
Under naturalconditions in pine or pine-oak forests with mixed species and/or mixed age classes,
both diseases and phytophagous predators have a
more or less chronic but low-level negative effect
on tree health.Mistletoes, such as Arceuthobiumspp.
(Hawksworth& Wiens, 1972), can be abundantin
some partsof the forest but rarelypreventtrees from
long-termsurvival and reproduction.Fungal attacks,
if not limited to physically damaged trees, rarely
spread far and wide. Unlike these pathogens, some
insects-e.g., the pine bark beetle (Dendroctonus
mexicanus)-are capable of large-scale destruction,
especially in single-species pine forests. Enormous
tracts of Pinus oocarpa forest in Honduras were
defoliated in 1963-1966, and thousandsof hectares
59

of pine forest in Mexico were similarly affected
in 1949-1950 (Perry, 1991). Many areas are soon
naturallyreseeded,and the overall effect undernatural conditions may be rathersimilar to a forest fire.
However,if regenerationis somehow prevented(e.g.,

by livestock grazing), a more permanent shift to
other types of vegetation may be the ultimate result
of insect epidemics.
USES AND CONSERVATION

PINES AS RENEWABLE RESOURCES
to the attentionof foresters, with the shift to multipurposeforestrygoals, especially in the UnitedStates
(Everett, 1987) but also in Mexico, where three
recentsymposia were held on the subject.Their yield
of edible seeds representsa type of sustainableuse
that was already practised by prehistoric tribes in
pre-Columbiantimes (Passini et al., 1988). It is
probablyan importantsource of cash locally. Yet the
collecting of the seeds, mostly in September, is
tedious labour carried out mostly by an itinerant
labourforce of women and children.Seed crops vary
considerablyfrom year to year (Styles, 1993b), with
bumperharvestsoccurringapproximatelyevery five
years, and the methodof harvestingdiminishescone
production(Morales & Camacho, 1988). Although a
considerablevolume of seed is marketedas far away
as Mexico City, most of it is sold locally, and this
type of use can at best be considered a cottage
industry(Styles, 1993b). Largerseeds are produced
by P maximartinezii,a species with a very limited
distribution;it was discovered by the botanist J.
Rzedowski by way of the exceptionally large seeds
he saw being sold in the market at Juchipila in
southernZacatecas.
A commerciallymore importantsecondaryproduct
of pines is resin, mainly tapped from "hard"pines
(Pinus subgen. Pinus). Some 200,000 ha of pine
forest in Mexico are exploited in this way, according
to the CamaraNacional de las IndustriasDerivadas
de la Silvicultura(CNIDS). In CentralAmerica,it is
also a major secondary product, derived from the
extensive forests of P oocarpa, covering large parts
of Hondurasand Guatemala,and of P pseudostrobus
and other species in the highlands.Pine resin forms
the basis for the turpentine industry, which is a
majorsource of revenue in several Mexican statesparticularlyChiapas, Oaxaca, Michoacan, Jalisco,
and Nuevo Le6n-and in Hondurasand Nicaragua.
If the process of resin tappingis carriedout carefully,
the tree does not appearto suffer any ill effects for
many years. Frequently,however,large stripsof bark
are crudely removed from the bole, allowing entry
of pathogensthat graduallyweaken and kill the tree.
The preferred species for resin tapping appear to
The pines collectively constitute one of the most

valuable natural resources in Mexico and Central
America (Carvajal& McVaugh, 1992); in Mexico
they produced nearly 7.5 million cubic meters of
pine logs in 1986. The most importantproducers
were the states of Durango,Chihuahua,and Jalisco,
in that order. Hondurasproduces more pine timber
than any other Central American country (Perry,
1991), and a substantialportion of its economy is
based on it. Exploitationof the extensive pine forests
on Hispaniola in the Caribbeanonly began in 1920
(Darrow & Zanoni, 1991). Wood production for
lumberis most important,followed by pulped wood
for kraft paper and cardboardand local uses for
firewood, fence posts, etc. Pine's importancelies in
the fact that the xylem of most "hard"pine species
produces long fibres, giving the final productextra
strength."Soft" pines, less abundant,are sought for
their less resinous, evenly grainedwood. In Mexico,
currentlythe most heavily exploited species are Pintuspatula, P oocarpa, P pseudostrobus,P herrerae,
P leiophylla, and P arizonica (Perry, 1991; Styles,
1993a).However,frequentlymixturesof pine species
are harvested without regard to species. In Central America, P caribaea var. hondurensis and P
oocarpa arecommerciallythe most importantspecies
at the lower and middle altitudes,where they are the
dominantcomponentsof the forest, and are similarly
exploited. In the Guatemalanhighlands, mixed pine
forests are the rule and exploitation largely does not
discriminateon the basis of species. In that country,
but also in other densely populatedruralregions of
Central America and Mexico, pine trees with
branches lopped off from low on the bole upwards
are a common sight. Often very little crown is leftinsufficient to sustain the tree for long, as most
species of pines do not coppice. The branches are
used for fuel.
In northernMexico, P cembroides is widely exploited for firewood and charcoal burning,and to a
limited extent locally for lumber.Most trees are too
small and heavily branchedto yield long, straight
lumber, and they are also less desirable for pulp be P oocarpa, P montezumae, P teocote, and P
wood. The pinyon pines have recently been drawn p.seudostr.obl.s.
60
Several other countrieshave or have had an interest in Latin American pines, especially the (sub-)
tropical species, introducingthem for plantationforestry schemes in their own territoriesor in other
(sub-)tropicalcountries. Pines are virtuallyabsent in
the Southern Hemisphere.The indigenous conifers
there do not have an equivalentproductionpotential
or adaptabilityfor fast-growing,even-aged monocultures. South Africa was one of the first countries to
researchthese potentialsand implementintroduction
programs(Loock, 1950; Liickhoff, 1964). Pinus patula was introducedin South Africa from Mexico as
early as 1903. Loock, in 1947, was the first to
conduct a systematic search for suitable species,
followed in 1957 by Liickhoff, who expanded the
efforts to P caribaea var.hondurensisand, to a lesser
extent, P pseudostrobus.The Food and Agricultural
Organization(FAO) of the United Nations has recognized the potentialof the CentralAmericanpines for
afforestation in tropical countries since the early
1950s and has cooperatedwith governmentalforestry
departmentsin Mexico and severalCentralAmerican
countries (Styles, 1993a). Concernsabout deforestation in tropical countries (e.g., at the Eighth Commonwealth ForestryConference, held in Nairobi in
1962) focused on the loss of timber(not on biodiversity) and led also to suggestions of afforestationwith
(neo-)tropicalpines. This in turngeneratedresearch,
funded by the British Government,at the Commonwealth Forestry Institute,now the Forestry Institute
(OFI) of the University of Oxford, beginning in
1963, focusing on explorationand seed collection of
P caribaea var. hondurensis. Later, other species
were added, e.g., P oocarpa, P patula, P tecunumanii (discovered by F. Schwerdtfeger,investigating
insect attacks on pines on behalf of FAO in Guatemala), P maximinoi (then known as P tenuifolia),
and P pseudostrobus(Styles, 1993a). All these species were introducedto several tropical countries in
South America, Africa, and Asia. More recently,the
CentralAmerica and Mexico Coniferous Resources
Cooperative(CAMCORE),operatingfrom the North
Carolina State University and financed largely by
government as well as private-sectorsubscribers,is
continuing these programs,expanding them to still
more species (Dvorak & Donahue, 1992). Seed is
being collected from many locations and established
in two types of field plantings: gene conservation
banks and genetic tests especially in Central and
South America. Many provenanceshave been tested
and the most productive identified, and these are
eventually to be planted in tropicalcountriesaround
the globe on a commercialscale. Genetic differences
Flora Neotropica
of growth, form, and wood properties,especially in
P caribaea, are preferentiallysought out. However,
such selections may also turn out to be the more
"aggressive" invaders in the event of (unwanted)
naturalization(Hughes, 1994). After Pinus radiata, P.
patula has now become one of the most troublesome
invasive species of pine threateningnaturalvegetation and biodiversity in the highlands of southern
Africa. In many cases, it will be only a matter of
time for the newer introductionsto follow suit, the
apparenttime lag between introductionand invasion
being a thresholdrecently overcome by a numberof
species (Richardsonet al., 1994).
CONSERVATION
The importance of pines to the forestry-based
industriesin the region means that timber exploitation is increasing,and in many areas it is indiscriminate. Areas formerlycovered with pine forests have
now been depleted, and the resultingeroded hillsides
and mountainsidesare a common featurein all countries of the region. Exploitationis almost exclusively
of naturalstands, where regenerationis wholly dependenton the remainingseed trees in the area and
the conditions of soil, grazing, and fire that prevail
after clear-cutting. Hundreds of small to mediumsized sawmills have penetrated most forested regions. With the building of roads, logging trails
furtheropened up the mountainsto broaderuse by
man. Sawmills require and use only trees with the
best form, so forests are frequently"creamed"of the
straightest,small-crownedindividuals,a practicethat
will ultimately lead to the genetic impoverishment
of a species (Vela, 1976; Styles, 1993a). Plantation
forestry, although begun, is still practised on too
small a scale to be a substitute for exploitation of
naturalforests. Populationpressuresare increasingly
high, especially in alreadydensely populatedregions
such as Hispaniola, El Salvador, Guatemala, and
many locations in Hondurasand Mexico. Indiscriminatedestructionof the forestenvironmentfor agricultural production,especially on richer volcanic soils,
is widespreadin these regions.
Species with restricteddistributionsare most vulnerable, but the actual threatof extinction of these
taxa depends on the situation locally. Some of them
grow on sites that are under little exploitation pressure, as is the case with Pinus nelsonii, and unless
grazing and burning were to increase dramatically
on these marginal lands, they seem relatively safe.
Farjon et al. (1993) have listed all conifer taxa
presentlygiven a conservationcategoryby the IUCN

Species Survival Commission's Conifer Specialist
Group. Ithis list, as a continuously updatedworking
document,has recently been evaluatedin connection
with the draftingof an IUCN Action Plan for conifers; see Farjon,19961.Seventeen pines in our region
are currentlylisted; the 10 listed below are of most
urgentconcern.
Pitnusculminicola: critically endangered
Pinus rzedowskii:critically endangered
Pilnusnmaximartinezii:
endangered
Pinus pinceana: endangered
Pinus radiata var. binata: (critically) endangered
Pinus cembroidessubsp. lagunae: vulnerable
Pinus cubensis: vulnerable
Pilnusjaliscana: vulnerable
Pinus nelsonii: vulnerable
Pinlus strobus var. chiapensis: vulnerable
A furtherseven taxa are categorizedas "low risk,"

meaning that although present trends are leading to
a decrease in population sizes in most areas, they
will not lead to foreseeable genetic depletion and
extinction of the taxon. They are nevertheless listed
because the trendsobserved may lead them into one
of the higher categories when current conditions
persist for a long time.
Garcia and Gonzales (1991) have demonstrateda
dramaticdecrease in area covered by P. culminicola
on Cerro Potosi since Beaman and Andresen (1966)
investigatedthe same population.The principalthreat
is fire, to which the species is not resistant(see also
Perry, 1991). There is as yet no official protection
for any of the areas (mountainsummits) where this
species occurs. Pinuts rzedowskii occurs in three
small populationssurroundedby a largebutexploited
pine forest. Two are on inaccessible talus slopes,
and although one is situated right underneatha fire
lookout, they can potentiallybe wiped out by a single
wildfire. The third and largest population is in part
mixed with other species. Although foresters and
conservationists are aware of these pines, none of
the populations enjoys conservation status at the
currenttime. A similar situationapplies to the single
but larger population of P maximartinezii(Farjon,
1994). The woodland is privately owned by a few
villagers who use the area for cattle grazing and pine
nut collecting at the same time. Both may have
adverse effects on natural regeneration. Fire is a
hazard,but awareness is growing and effective land
managementin cooperation with local people may
prevent disastrous calamities until adequate in situ
61
conservation management (regeneration schemes
foremost) is in place. Although P. pinceana is more
widespreadthan P nelsonii, the latteroccurs on soils
(often almost bare calcareous bedrock) that support
neitherforest nor forage for grazing.Transformation
of the vegetation cover related to livestock grazing
appears to be a major threat to P. pinceana, at least
in areas we have seen (see also Perry, 1991). Both

yield edible pinyon nuts ("pinones"), but in low
quantitiesas comparedto the largerand more abundant P cembroides,and exploitationis minimal.The
island populationsof P radiata, recognized as var.
binata, are under threatfrom grazing of feral goats.
The problemis especially acuteon GuadalupeIsland,
the farthestout in the Pacific (Libby et al., 1968).
Eradicationof goats would restore regeneration,but
this is moreeasily proposedthanpracticallyeffected.
and is not likely to become a priorityfor a relatively
poor state like Baja California Norte, unless aided
by external funding.
The vulnerabletaxa are all being depletedmore or
less rapidlyand are likely to fall into the endangered
categories in the near future if present trends continue.An exemplaryaccount is given by Zamoraand
Velasco (1977) for P strobus var. chiapensis in the
state of Chiapas, Mexico. A preferentialpine for
lumbermenwith the same wood quality as its near
relative (P strobus var. strobus) of eastern North
America, it has been excessively exploited over
much of the state. Regenerationhere, as elsewhere,
is hampered by livestock grazing. At the time of
Zamoraand Velasco's study, some small exclosures
had been establishedto observe regenerationwithout
grazing pressure, and plantation forestry with the
species had been started up. However, as Perry
(1991) pointed out, with the currentgrowth of the
populationin Mexico and CentralAmerica,the need
for food productionincreasingly becomes the overwhelming factorand nullifiesmost efforts to preserve
the pine forests. The situation on the Caribbean
Islands is similar. The best strategy for preserving
the biodiversity of Pinus in this part of the world
where it is still the greatest is probablyto establish
a numberof nationalparksand preserves.Countries
like Honduras and Mexico already have a good
number of these, but without financial support for
protectionand managementfrom the wealthierparts
of the world where conservation of biodiversity is
professed to be valued so greatly, we predict that
most of these areas, too, will succumb sooner or
later to the more basic needs of a growing human
population.
Flora Neotropica
62
SYSTEMATIC TREATMENT
Pinus Linnaeus,Sp. PI. 2: 1000. 1753; Gen. PI., ed.
5, 434. 1754; Miller, Gard. Dict., abr. ed. 4, 3.
1754; Jussieu, Gen. PI. 414. 1789; L. C. Richard,
Comm. bot. Conif. Cycad. 145, 147. 1826; Zuccarini in Endlicher,Gen. PI. suppl. 2: 26. 1842; Endlicher, Syn. Conif. 81. 1847; Carriere,Tr. Gen.
Conif. 291. 1855; Parlatorein Alph. de Candolle,
Prodr. 16(2): 378. 1868; Shaw. Gen. Pinus 24.
1914; Pilger in Engler & Prantl, Nat. Pflanzenfam.,ed. 2, 13: 331. 1926; Little & Critchfield,U.S.
Dept. Agric. Forest Service Misc. Publ. 1144: 7.
1969; Price, J. ArnoldArbor.70: 262. 1989; Carvajal & McVaugh in McVaugh, Fl. Novo-Galiciana
17: 32. 1992; Kral in Flora of North America
Committee(ed.), Fl. N. Amer. 2: 373. 1993. Lectotype species (vide Britton,N. Amer.Trees 5. 1908).
Pintts sv',estris Linnaeus, Sp. PI. 2: 1000. 1753.
Pinus sect. Petce Grisebach, Spicil. Fl. Rumel. 2: 347.
1844.
StrobusOpiz, Lotos (Prague)4: 94. 1854.

Cairyopitys Small, Fl. s.e. U.S. 29. 1326. 1903.
ApinusNeckerex Rydberg,Bull. TorreyBot. Club32:

597. 1905.
Amer.Midi.Nat.3: 69. 1913.
Nieuwland,
Leucopitys
Ducanlpopinus Chevalier, Rev. Bot. appl. & Agr. trop.
24: 30. 1944.
Derivation:classical Latin name pinus. English pine,

Spanish pino.
Trees,or less commonly shrubs, evergreen, resinous in wood, bark,leaves, and often cones, aromatic;
1-75 m tall, diameter at breast height (dbh) 5-320
cm. Trunkmonopodial, or shrubs in some species
multistemmed,branchingin pseudowhorls.Bark of
older stems variously (deeply) furrowed or plated,
to thin and scaly or thin and smooth, ridges and/or
plates layered or scaly, more or less conspicuously
exfoliating in often intricate patterns. Wood with
normally conspicuous annual ring boundaries;resin
ducts present or absent; ray tracheids present or
absent, xylem parenchymaabsent.Shoots dimorphic,
with long shoots and dwarf shoots; long shoots
uninodalor less commonly multinodal;dwarf shoots
axillary to spirally arranged primary leaves (cataphylls). Catapllh!ls on long or short decurrentleaf
bases (pulvini) on long shoots, non-chlorophyllous,
enclosing "winter buds" (primordiallong shoots or
ovuliferous strobili), subtendingdwarf shoot buds or
pollen strobili,early deciduousor persistent.Secondary leaves (needles) borne in fascicles of (1-)2-5
(-8) on dwarf shoots; the fascicles surroundedat
base by an early deciduous or persistent sheath of
bud scales or their remnants,persisting 2-30 years

and falling as fascicles; length 2.5-50 cm, width
0.5-2.5(-7) mm, acicular (one species lanceolate),
plano-convex or triangular(rarely terete or flat) in
cross section, entire or serrulate;epistomatic or amphistomatic(one species occasionally hypostomatic);
resin ducts (1-)2 to several, variously positioned in
the mesophyll; vascular bundles single or double,
enclosed in a stele. Pollen cones spirally arranged
near the proximal end of new long shoots, ovoidoblong to cylindrical;consisting of a thin axis with
numerousspirally arranged,(sub-)peltatemicrosporophylls, each bearing two longitudinally dehiscent
sporangia;pollen bisaccate. Seed cones (ovuliferous
cones) subterminal or sometimes appearing to be
lateral, borne singly or more commonly clustered,
pedunculate,maturingin the second year or rarely
in the third year, shed early or variously persistent,
initially erect; maturecones pendulousor spreading,
opening soon or variously serotinous, (obliquely)
ovoid to cylindrical, 2-60 cm long. Seed scales
(ovuliferous scales) persistent, obovate to oblong,
thin or thick woody, attached to a slender to very
thick axis; the exposed portion(apoplhysis)variously
thickenedand/orelongated; bearing in a terminalor
dorsal (abaxial) position the remnantof the exposed
portion (iimbo) from the first year's development,
which may be terminatedin a (deciduous) spine or
prickle. Seeds obovoid, slightly flattened;seed coat
thin or thick, without resin vesicles; seed with an
adnate or articulatemembranouswing derived from
the adaxialpartof the seed scale; several times larger
than the seed and effective, or (greatly) reduced
and in articulate wings remaining with the scale.
Cotvledons3-24, denticulateor entire. Chromosome
number 2ii = 24 (x = 12).
Glossary of Botanical Terms Used in the Keys

adnate seed wing: wing not easily separable from
the seed, usually tearing instead (Fig. 61-J).
amphistomatic: with stomataoccurringon all faces
around the leaf; there are usually more lines or
rows of stomataon the convex abaxial (outer) face
than on each of the two adaxial faces.
apophysis: part of the seed scale that is exposed in
a mature, closed cone, representing the second
phase of growth (Fig. 6Cb-1, Dc-i).
articulate seed wing: wing easily separable from
the seed, it is held by two claw-like appendages
(Fig. 6K).
63
Systematic Treatment
attenuate: widest near the base, gradually tapering
to the apex; here referringto the shape of ovoidoblong closed cones.
cataphyll: scale leaf or primaryleaf, which initially
covers the vegetative buds (Fig. 1C); later they
are more or less persistenton the shoot (Fig. 1A,
F).
endodermis: layer of large cells surroundingthe
stele with the vascularbundle(s) in a leaf, in cross
section seen as a ring of cells, of which the outer
walls are thin (Fig. 4D) or thickened(Fig. 4E).
epistomatic: with stomata occurring on the 1-2
adaxial (inner) leaf faces only.
fascicle sheath: Aggregationof scales enveloping a
leaf fascicle; it can be deciduous or persistent;in
some species the scales recoil before becoming
deciduous (Fig. lHa-c).
leaf: secondary,chlorophyllousleaf; commonly referredto as "pine needle."
leaf fascicle: A group of leaves (needles) growing
from a dwarf shoot, initially or persistentlyenveloped at its base by a sheath of scales (Fig. IGab).
multinodal shoot: a long shoot elongating in more
thanone flushof growth,this detectedprimarilyby
noting the interruptedarrangementof leaf fascicles

and some weaker lateralshoots between the main
nodes of branching.
pulvinus (pi. pulvini): small, raised section on a
long shoot at the base of a cataphyll,either forming a ridge downward (decurrent, Fig. IA) or
ending abruptly(not decurrent,Fig. IB).
resin ducts: tubulartractscontaining resin and situated in the wood as well as in the leaves of pines;
diagnostic of several species is the position of
these resin ducts in the leaf in cross section:
external (Fig. 2A-D), medial (Fig. 2E-G), internal (Fig. 2H-J), septal (Fig. 2K-L).
serotinous cones: seed cones that open their scales
only afterextreme heatingor many years of dehydration; normally found to be closed or nearly
closed on the tree.
umbo: part of the seed scale that is exposed in the
first phase of growth, distal to the apophysis (Fig.
6Ca-2, Da-2).
vegetative bud: the terminaland subterminalcomplex of shoot primordiumandbudsof dwarfshoots
with young leaves, covered by cataphylls (Fig.
IC).
Key to the subgeneraof Pinus

1. Pulvini decurrent(Fig. IA); leaves with 2 vascularbundles and variously positioned resin ducts (Fig. 2AL), with persistent(rarely deciduous) fascicle sheath; seeds with an articulate(Fig. 6K, L). rarely adnate,
wing..............................................................................................................................................subgen. Pinus (spp. 1-33)
1. Pulvini not decurrent(Fig. IB); leaves with a single vascularbundle and external resin ducts (Fig. 2A-D),
with deciduous (rarely persistent)fascicle sheath; seeds with an adnate (Fig. 61, J), rarely articulate,wing
or wingless when free from the seed scale (Fig. 6Nb).........................................................subgen. Strohus(spp. 34-47)
Key to the sections and subsectionsof subgen. Pinus

I. A conspicuous narrowband visible aroundumbo on seed scale; sheath of leaf fascicle deciduous...............
........................................................................................................................................sect. Leiophyllae(I. P leioph!lla)
1. No band visible around the umbo, differing from other parts of the apophysis; sheath of leaf fascicle
persistent.
2. Seed cones serotinous (remaining closed), sessile in tenacious whorls, or in pairs but then large and
with elongated, hooked spines ......................................................................................................... subsect. Atte
2. Seed cones opening (relatively) soon after maturity,(short) pedunculate,solitary or in whorls (open
cones may seem to be sessile).
3. Long shoots multinodal;vegetative buds often resinous;resin ducts internal.......................................
....................................................................................................................subsect. A str les (excl. P occide utlis)
3. Long shoots uninodal;vegetative buds usually not resinous;resin ducts variously positioned.
4. Seed cones asymmetrical,oblique at base or curved. 4.5-35 cm long; seed scales spreadingto 90?
or more; apophyses nearly flat to prominentlyraised (Fig. 6E-H).
5. Leaves relatively short, erect or spreading,or if up to 25 cm long. 1.5-2 mm wide, in fascicles
subsect. Polerosae
of (2-)3-5(-6) (count 15-20 fascicles) ..............................................................
5. Leaves 20-50 cm long, drooping to pendulous,in fascicles of 5 or more ................ subsect. Pseudostrobi
4. Seed cones symmetricalor slightly oblique, ovoid or ovoid-oblong, (2-)3-10(-12) cm long; seed
scales spreading<90?; apophysis flat or slightly raised (Fig. 6E-F).
6. Seed cones up to 5.5 cm long; leaves in fascicles of 2-3 (count 15-20 fascicles)......... subsect. Contortae
6. Seed cones (4-)5-10(-12) cm long; leaves in fascicles of 2-5.
7. Leaves in fascicles of 2(-3) .......................................................................subsect. Pi s (2. P tropicalis)
Flora Neotropica
64
7. Leaves in fascicles of 3-5.

8. Vegetativebuds resinous;umbos of seed cone scales with a persistentprickle......8. P occidentalis
8. Vegetative buds not resinous; umbos of seed cone scales without a prickle in mature
sct. Oocarpae
cones .........................................................................................................................subse
Key to the species of subsect. Attenuatae

1. Leaves in fascicles of 2 (rarely 3) (count 15-20 fascicles).
2. Umbo of seed scales with a persistentspine or prickle..................................................29. P. uricata var. Imuricata
2. Umbo of seed scales without a spine or prickle................................................................ . . P radita var. bi t
I. Leaves in fascicles of 3 (rarely 2).
3. Seed cones very large, 20-35 cm long; apophyses and umbos of seed scales very strongly developed;
leaves 1.9-2.2 mm wide .......................................................................................................................... 33. P coulteri
3. Seed cones smaller; leaves <1.6 mm wide.
4. Vegetative buds resinous;apophyses of seed scales strongly developed on one side of the cone (Fig.
7E).......................................................................................................................................................3 1. P. a ttenu
at a
4. Vegetative buds not resinous;apophyses flat or only slightly raised...................................................32. P. greggii
Key to the species of subsect. Australes

1. Leaves (6-)10-15(-18) cm long; seed cones (3.5-)4-7 cm long; seeds with adnate wing (Fig. 61) ......7. P cubensis
i. Leaves (I -)14-28(-30) cm long; seed cones 5-13 cm long; seeds with articulatewing (Fig. 6K, L).
2. Vegetative buds not resinous; leaves in fascicles of 2(-3) (count 15-20 fascicles) .............. 6. P elliottii var. dcensa
2. Vegetative buds resinous; leaves in fascicles of (2-)3-5.
3. Long shoots multinodal;fascicle sheath of latest leaves 15-20 mm long: leaves in fascicles of (2-)
P. (carihbae
3-4, very rarely 5 .............................................................5.
3. Long shoots uninodal; fascicle sheaths of latest leaves (8-)l0-15 mm long; leaves in fascicles of
3-5 .....................................................................................................................................................8. P occilentt lis
Key to the species of subsect. Contortae

1. Leaves in fascicles of 2; vegetative buds resinous; umbos on seed scales with a persistentprickle...............
u1lt i ana
................................................................................................................................................3. P contorta var. m
1. Leaves in fascicles of 3; vegetative buds not resinous; umbos on seed scales with a minute, deciduous
4. P iherrerue
prickle................................................................
Key to the species of subsect. Oocarpae

I. Cones broadly ovoid to subglobose when closed.
2. Cones semi-serotinous (only the distal seed scales parting), remainingon the tree; leaves 0.8-1.6 mm
P.
wide; leaf resin ducts septal...............................................................20.
oocarpa
2. Cones opening completely when mature,falling after 1-3 years; leaves 0.5-1 mm wide; leaf resin ducts
medial or internal.
3. Cones falling with the proximal seed scales missing; fascicle sheaths 10-14 mm long; leaves (8-)
10-16 cm long, 0.5-0.8 mm wide; leaf resin ducts internal......................................................21. P. praetermissa
3. Cones falling intact: fascicle sheaths 12-18(-25) mm long; leaves (14-)16-18(-25) cm long, 0.7-1
tectunumoatii
(-1.3) mm wide; leaf resin ducts medial..................................................................................... 24..
1. Cones ovoid-attenuatewhen closed.
4. Leaves in fascicles of (4-)5-7, rarely 8 ............................................................................................ 25. P. durangensis
4. Leaves in fascicles of 3-5, rarely 2.
5. Leaves with 9 or more lines of stomataon the abaxial face; leaf resin ducts internal.
6. Cones semi-serotinous,persistent,leaving some basal scales on the branchwhen falling; umbo of
seed scales flat or depressed ..........................................................................................................27. P pringlei
6. Cones opening at maturity,falling soon with peduncles attached;umbo of seed scales pyramidal.
.........................................................................................................................................................26. P tawsonii
5. Leaves with 3-7(-10) lines of stomataon the abaxial face; leaf resin ducts medial or septal.
7. Leaves in fascicles of (4-)5, rarely3; fascicle sheaths 8-10(-15) mm long; leaf resin ducts mostly
septal..............................................................................................................................................23. P jaliscan
7. Leaves in fascicles of (2-)3-4(-5); fascicle sheaths initially long (ca. 20 mm or longer); reduced
in matureleaves to 10-15 mm; leaf resin ducts medial.
65
8. Leaves (7-)10-15(-18) cm long, 1-1.4 mm wide..................................................................... 28. P. teocote

8. Leaves (11-)15-25(-30) cm long, 0.7-0.9(-1) mm wide ..........................................................22. P patul
Key to the species
of subsect. Ponderosae
I. Leaves in fascicles of 3-5(-6). 5 usually present,often predominating.

2. Seed scales of cones relatively thin, flexible; leaf endodermiswith thin-walledcells; leaf vascularbundles connate...................................................................
13. P hartwegii
2. Seed scales of cones relatively thick, rigid; outer walls of leaf endodermalcells thickened;leaf vascular
bundles mostly separate......................................................................................................................... 10. P ri onic
I. Leaves in fascicles of (2-)3(-4), rarely 5.
3. Leaves in fascicles of 3, rarely 2..........................................................................................................
.
jeffre i
3. Leaves in fascicles of 2-3 or more.
4. Seed cones 5-10 cm long; fascicle sheaths of leaves 10-15(-20) mm long; leaves in fascicles of 2mm
mw
wide
ide.............
9.. P
................................................................................................
P. pon(lerosa
3, 1.3-1.6 m
po,derosa var. scoplrlorul
scoptdorum
;........
4. Seed cones 8-15 cm long; fascicle sheaths of leaves (15-)25-35(-40) mm long; leaves in fascicles
of (2-)3(-4), rarely 5, 1.5-2 mm wide ....................................................................................... 12. P
el nnii
Key to the species of subsect. Pseudostrobi

I. Fascicle sheaths deciduous at maturity;leaves pendulous ....................................................................... 19. P luholtii
I. Fascicle sheaths persistent;leaves drooping or spreading.
2. Cones 5-10(-12) cm long; falling intact (peduncle deciduous with the cone).
3. Seed scales of cones thin, flexible, in open cones usually strongly recurved;leaves often very slender,
drooping to pendulous,0.6-1 mm wide.......................................................................................... 18. P a i i oi
3. Seed scales of cones not flexible, not strongly recurvedin open cones; leaves droopingor spreading.
0.7-1.2 mm wide...........................................................................................................................17. P
gl si
2. Cones (7-)10-35 cm long; leaving a few proximalscales on the branchwhen falling.
4. Fascicle sheaths (20-)25-35(-40) mm long, resinous; leaves 1.1-1.6 mm wide; outer walls of leaf
endodermalcells thin; seed cones 15-35 cm long. ....................................................................... 16. P. levoia
4. Fascicle sheaths (15-)20-30(-35) mm long, usually not resinous; leaves 0.8-1.3 mm wide; outer
walls of leaf endodermalcells thickened;seed cones 7-20 cm long.
5. Leaves with connate vascular bundles;seed cones usually obliquely ovoid when closed. 14. P pseudostrohus
5. Leaves with separatevascular bundles; seed cones usually ovoid-oblong to ovoid-attenuatewhen
closed.. ......................................................................................................................................15. P. onlte
e
Key to the sections and subsections of subgen. Strobus

1. Umbo of seed scale terminal(Fig. 6Ca); leaves in fascicles of 5 ..............................................................subsect. Strohi
1. Umbo of seed scale dorsal (Fig. 6Da, c); leaves in fascicles of 1-4(-5).
2. Seed cones with -60, very widely spreading, very flexible scales; cones not longer than wide; seeds
relatively large, wingless when detached from seed scale ...........................................................subsect. Cembroiles
2. Seed cones with >60 scales spreading-70?; cones longer than broad;seeds large or small, wingless or
winged.
3. Leaves in fascicles of 3, rarely 4 (count 15-20 fascicles); seed cones oblong, irregular,pendulous.
up to 10-12 cm long; seed scales easily flexible......................................................................... subsect. Nelsoi e
3. Leaves in fascicles of (3-4-)5; seed cones regularly shaped, usually >10 cm; seed scales rigid to
inflexible.
4. Seeds small (ca. 8 mm long), winged; seed cones 10-15 cm long, with thin, rigid scales....39. P rz.e(dowskii
4. Seeds large (20-28 mm long), wingless; seed cones (15-)17-25(-27) cm long, with thick, inflexible scales...............................................................................................................................40 . P
. i rt inezii
Key to the species
of subsect.
Cembroides
. Leaves in fascicles of 1, rarely 2.............................................................................................................46. P ono lla

1. Leaves in fascicles of 2 or more.
2. Leaves in fascicles of 5 (very rarely4 or 6); a low, spreadingshrub..............................................44. P cull,lilicola
2. Leaves in fascicles of 2-4(-5); a shrubor (small) tree.
3. Scales of fascicle sheaths recoiling strongly before falling, forming a rosette; leaves usually <1 mm
wide..................................................................................................................................................43. P ce broides
3. Scales of fascicle sheaths not forming a rosette; leaves usually 1-1.5 mm wide.
66
Flora Neotropica
P. remota
4. Leaves in fascicles of 2(-3), 0.8-1.1 mm wide........................................45.
4. Leaves in fascicles of (3-)4(-5), 1-1.5 mm wide ...................................................................47. P. quadrfolia
Key to the species of subsect. Nelsoniae

I. Leaves in fascicles of 3, rarely 4, but connate, appearingas one (Fig. 60B); seed cones on thick, 2.5-6.5
cm long and curved, persistentpeduncles..................................................................................................... 41. P. nelsonii
i. Leaves in fascicles of 3, rarely4, separate(Fig. 61C); seed cones on slender, easily breakingpeduncles....
......................................... ..............................................................................................................................4 2 . P pinceanu
Key to the species of subsect. Strobi

I. Leaves amphistomatic,with 2-4(-5) lines on the abaxial face; seed scales with broad, straight and flat,
smooth apophyses....................................................................................................................................35.P. a bertian
I. Leaves epistomatic (on adaxial face), or rarely with a few stomata on the abaxial face; seed scales with
tapering,recurvedor concave, often grooved apophyses.
2. Seed wing shorter than seed or vestigial (Fig. 6Mb); leaves (5-)6-11 cm long, 0.8-1.2 mm wide (on
sterile shoots).
3. Vegetative buds resinous; fascicle sheaths up to ca. 15 mm long; seed cones 10-15 cm long.............
..............................................................................................................................................36. P. flexilis var. reflex
3. Vegetativebuds not resinous;fascicle sheaths20-25 mm long; seed cones 12-30 cm long (or longer).
. strobiformis
.........................................................................................................................................................37..
2. Seed wings (much) longer than seed (Fig. 6J); leaves (5-)6-15(-18) cm long, 0.6-1 mm wide (on sterile
shoots).
4. Seed cones large, (10-)15-40(-50) X 7-15 cm when open; seed scales with recurved or reflexed
apophyses (at least the proximal scales recurved)....................................................................... 34. P avacahuite
4. Seed cones (6-)8-16(-25) X 4-8 cm when open; seed scales with straight, ? concave apophyses..
.......................................................................................................................................38. P strobus var. chiapensis
Additional Keys by Regions I-VII

Key to the species in Baja CaliforniaNorte (I)
1. Leaves in fascicles of 5; fascicle sheaths deciduous; seed cones >25 cm long, pendulous; umbo of seed
scale terminal (Fig. 6Ca)..................................................................................................................... 35. P lambertiana
1. Leaves in fascicles of 1-4, rarely5; fascicle sheaths persistentor recoiling and deciduous;seed cones small
or large, not pendulous;umbo of seed scale dorsal (Fig. 6Da, c).
2. Fascicle sheath persistent.
3. Leaves in fascicles of 2 (count 15-20 fascicles, especially on leading shoot).
4. Leaves 4-7 cm long, persisting 5-8 years; seed cones opening gradually,(3-)4-5.5 cm long........
.................................................................................................................................3. P contorta var. murravana
4. Leaves (7-)10-14(-16) cm long, persisting 2-3 years; seed cones serotinous,5-7(-8) cm long....
.................................................................................................................................29. P muricata var. nm
uricata
3. Leaves in fascicles of 2-3 (count 15-20 fascicles, especially on leading shoot).
5. Seed cones usually in whorls of 2-5, serotinous,(5-)8-15 cm long.
6. Vegetativebuds resinous; leaves in fascicles of 3, rarely 2; seed cones ovoid-oblong to ovoida ttenuata
attenuate............................................
......................................................
P..........31.
6. Vegetative buds not resinous; leaves in fascicles of 2, sometimes 3 on leading shoots; seed
cones ovoid or obliquely ovoid. .............................................................................. 30. P. radiat var. binata
5. Seed cones solitary or in pairs, rarely in whorls of more than 2, opening graduallyor soon after
maturity,10-35 cm long.
7. Seed cones very large and heavy, 20-35 cm long, very resinous;apophyses and umbos of seed
scales very strongly developed ..................................................................................................33. P. coulteri
7. Seed cones smaller, lighter, 10-17 cm long, not resinous;apophyses and umbos of seed scales
slightly or moderately raised. .......................................................................................................1 . P. effreyi
3. Fascicle sheath deciduous.
8. Leaves in fascicles of , rarely 2..............................................................................................46. P. monoph la
47. P quadrifoia
8. Leaves in fascicles of 3-5 usually 4 ..........................................................................
67
Key to the species in northwesternMexico (I1)

(Sonora, Chihuahua,Sinaloa, Durango,Zacatecas)
1. Bases of cataphylls (pulvini) not decurrent(Fig. IB); leaves with a single vascular bundle and external
resin ducts (Fig. 2A-D); sheath of leaf fascicle deciduous.
2. Leaves in fascicles of 2-3, rarely 4 or 5; seed cones small (3-5 cm), not longer than wide when open;
seeds wingless when detached from the scale ...................................................................................43. P. ce roies
2. Leaves in fascicles of 5, rarely 6; seed cones 10 cm long or longer; seeds with a short (sometimes
vestigial) wing.
3. Vegetativebuds resinous;fascicle sheaths up to ca. 15 mm long; seed cones 10-15 cm long.............
..............................................................................................................................................36. P fl exilis var. reflex
3. Vegetativebuds not resinous;fascicle sheaths20-25 mm long; seed cones 12-30 cm long (or longer).
.........................................................................................................................................................37. P strol iformis
I. Bases of cataphylls (pulvini) decurrent(Fig. IA); leaves with 2 vascularbundles and variously positioned
resin ducts (Fig. 2A-L); sheath of leaf fascicle persistentor deciduous.
4. Sheath of leaf ascicle deciduou
. P eioph ll
...........................................................................................................
4. Sheath of leaf fascicle persistent.
5. Seed cones persistent,semi-serotinous,ovoid to globose when closed; leaf resin ducts mostly septal.
................................................................................................................................................................20.P. ooc rpa
5. Seed cones falling, at least after a few years, opening when mature,obliquely ovoid to oblong when
closed; leaf resin ducts medial.
6. Seed cones obliquely ovoid-oblong, often curved, 15-35 cm long; leaves in fascicles of 5, rarely
4 or 6 (count 15-20 fascicles) ................................................................................................. 16. P. devo ia
6. Seed cones ovoid or obliquely ovoid, less than 15 cm long; leaves in fascicles of 3-5, rarely 2.
7. Peduncledeciduous with the seed cone, which falls intact....................................................... 28. P teocote
7. Peduncle persistent;the cone leaves some of the basal scales on the branchwhen it falls.
8. Seed cones (4.5-)5-10(-14) cm long; apophyses of seed scales moderatelyraised (<'/2 as
high as wide); leaves rarely longer than 20 cm................................................................. 10. P. arizonica
8. Seed cones 8-15 cm long; apophyses strongly raised (>/2 as high as wide); leaves (18-)
20-35 cm long............................................................................................................... 12. P. engel annii
Key to the species in western Mexico (III)

California
Sur,
Nayarit,southernZacatecas,Aguascalientes,Jalisco, Colima, Michoacan)
(Baja
1. Leaf fascicle sheaths deciduous.
2. Seed cones with '60, very widely spreading,very flexible scales; cones not longer than wide....43. P. cemnbroides
2. Seed cones with '60, inflexible or at least rigid scales; cones longer than wide.
3. Leaves epistomatic, rarely a few stomataon the abaxial face; one vascularbundle in the leaf.
4. Fascicles sheaths with distinct, individuallyfalling scales; seed cones cylindrical, 12-30(-60) cm
long; apophyses of seed scales, at least the proximalones, recurvedor reflexed ................37. P. strobiformis
4. Fascicles sheaths with basally connate scales recoiling to form a rosette before they fall; seed
cones not cylindrical;apophyses not recurvedor reflexed.
5. Leaves in fascicles of (3-)4-5; seed cones 10-15 cm long, with thin, fixed but flexible scales;
seeds small (ca. 8 mm long), winged................................................................................. 39. re o skii
5. Leaves in fascicles of 5, rarely3-4; seed cones (15-)17-25(-27) cm long, with thick, inflexible
inartineii
scales; seeds 20-28 mm long, wingless ........................................................................40. P.
3. Leaves amphistomatic;2 vascularbundles in the leaf.
6. Leaves (4-)6-15(-17) cm long, spreading:seed scales with a conspicuous narrow band visible
aroundthe umbo........................................................................................................................... . P. eiophll
6. Leaves (15-)20-30(-40+) cm long, pendulous;seed scales lacking a band aroundthe umbo.........
.......................................................................................................................................................19. P lui holtzii
I. Leaf fascicle sheaths persistent.
7. Leaves in fascicles of 3, sometimes a few 2-5 (count 15-20 fascicles); seed cones (2-)3-6(-7) cm long.
8. Seed cones ovoid to subglobose when closed, shorterthan wide when opened......20b. P oocarpa var. trifblicata
8. Seed cones ovoid or obliquely ovoid when closed, longer than wide when opened.
9. Leaves slender, lax, (10-)15-20 cm long, 0.7-0.9 mm wide; seed cones (2-)3-3.(-4) cm long.4. P. herrerae
9. Leaves rigid, (7-)10-15(-18) cm long, 1-1.4 mm wide; seed cones (3-)4-6(-7) cm
28. P teocote
long ......................................................
7. Leaves in fascicles of (4-)5-6, rarely3 or 7-8; seed cones (4-)5-10(-12) cm long or much
larger.
10. Seed cones (4-)5-10(-14) cm long; seed scales with flat or slightly raised apophyses
(Fig. 6E-F).
i1. Leaves (6-)10-25(-30) cm long.
68
Flora Neotropica
12. Seed cones ovoid to subglobose when closed, shorterthan wide when opened.
13. Cones falling with the proximalseed scales missing; leaves (8-)10-16 cm
long, 0.5-0.8 mm wide; leaf resin ducts internal ........................... 21. P praetermissa
13. Cones persistent,intact when falling; leaves (I -)14-25(-30) cm long; leaf
resin ducts septal ..............................................................20a. P oocarpa var. oocarpa
12. Seed cones ovoid-oblong to ovoid-attenuateor obliquely ovoid when closed.
longer than wide when opened.
14. Proximal seed scales not parting when the cone opens; leaf resin ducts
septal. ...................................................................................................... 23. P jaliscana
14. Proximalseed scales partingwhen the cone opens; leaf resin ducts medial.
15. Seed cones with ca. 150-200 scales; apophyses of seed scales more or
less flat, weakly keeled, light brown to purplishblack ................. 13. P hartfwegii
15. Seed cones with ca. 90-120 scales; apophyses of seed scales (slightly)
raised, prominentlykeeled, ochraceous to light (reddish-)brown..........
..................................................................................................... .. P duranger
nsis
II. Leaves 20-35 cm long.
16. Leaves 0.6-1 (-1.1) mm wide, drooping to pendulous;seed scales thin, spreading 90? or reflexed when the cone is open; leaf hypodermis without or with
few intrusionsinto the mesophyll................................................................18. P. taximinoi
16. Leaves 0.7-1.2 mm wide, spreadingor drooping; seed scales thick, spreading
<90? when the cone is open; leaf hypodermis with numerous intrusions into
the mesophyll, some connecting with the endodermis ........................... 17. P douglasiana
10. Seed cones (7-)8-35 cm long; at least several seed scales with prominently raised
apophyses (rarely all nearly flat) (Fig. 6G-H).
17. Fascicle sheaths 30-40 mm long, resinous: leaves 1.1-1.6 mm wide; outer walls of
leaf endodermalcells thin; seed cones 15-35 cm long .................................... 16. P devoaniana
17. Fascicle sheaths (15-)20-30(-35) mm long. usually not resinous; leaves 0.8-1.3
mm wide; outer walls of leaf endodermalcells thickened;seed cones 8-20 cm long.
18. Leaves with partly connate vascular bundles; seed cones usually obliquely
ovoid when closed............................................................................. 14. pseu ostrob s
18. Leaves with separate vascular bundles: seed cones usually ovoid-oblong to
ovoid-attenuatewhen closed .................................................................... 15. P montezt
umae
Key to species in NE & E Mexico (IV)

(eastern Chihuahua,Coahuila, Nuevo Le6n, Tamaulipas,northernZacatecas, San Luis Potosi,
Guanajuato,Queretaro,Hidalgo, Tlaxcala, northernVeracruz,northernPuebla)
I. Leaf fascicle sheaths deciduous.
2. Seed cones with -50. very widely spreading,very moveable scales; cones not longer than wide.
3. Leaves in fascicles of 5 (very rarely4 or 6, count 15-20 fascicles); a low, spreadingshrub...44. P culininicola
3. Leaves in fascicles of 2-4(-5); a shrub or (small) tree.
4. Scales of fascicle sheaths recoiling strongly before falling; leaves in fascicles of (2-)3(-4), rarely
5, < mm wide .......................................................................................................................... P cem ro es
4. Scales of fascicle sheaths not recoiling; leaves in fascicles of 2(-3), 0 .......................................45. P remote
2. Seed cones with -50 scales, spreading -90?, rigid, or if flexible, spreading very little; cones longer
than wide.
5. Leaves in fascicles of 3, rarely4 (count 15-20 fascicles); seed cones ovoid-oblong or short cylindrical, irregular,5-10 cm long; seed scales very moveable; seeds wingless when detached from the
scale.....................................................................................................................................................42. P pinceana
5. Leaves in fascicles of (2-)3-5(-6); seed cones ovoid or long cylindrical, mostly longer than 10 cm,
regular;seed scales rigid; seeds winged.
6. Leaves in fascicles of 5, rarely 6: seed cones cylindrical.
7. Seed wing shorterthan seed or vestigial (Fig. 6Mb); leaves 6-11 cm long, 0.8-1.2 mm wide.
8. Vegetativebuds resinous;fascicle sheaths up to ca. 15 mm long; seed cones 10-15 cm long.
...............................................................................................................................36. P fl exilis var. reflexa
8. Vegetative buds usually not resinous; fascicle sheaths 20-25 mm long; seed cones 12-30
cm long (or longer).........................................................................................................37. P s ro ifo is
7. Seed wings longer than seed (Fig. 6J); leaves (6-)8-15(-18) cm long, 0.6-1 mm wide. 34. P. !avacahuite
6. Leaves in fascicles of (2-)3-5(-6); seed cones ovoid.................................................................. . P leiophylla
9. Bases of cataphylls (pulvini) not decurrent(Fig. IB); leaves in fascicles of 3, rarely 4, but
connate, appearingas one; seed cones on thick, long and curved, persistentpeduncles;seeds
P. nelsoii
wingless when detached from the scale ...............................................................41.
69
9. Bases of cataphylls (pulvini) decurrent(Fig. IA); leaves in fascicles of (2-)3-5(-6), separate; seed cones on (relatively) short peduncles;seeds winged.
10. Leaves (7-)9-15(-18) cm long; seed cones (semi-)serotinous;apophyses of seed scales
flat or only slightly raised (Fig. 6E).
II. Seed cones conspicuously pedunculate, in whorls of 1-3, (3-)4-6(-7) cm long;
leaves in fascicles of 3, but occasionally 2-5 (count 15-20 fascicles) ................ 28. P teocote
II. Seed cones appearingsessile, in whorls of 3-8, (6-)8-13(-15) cm long; leaves in
fascicles of 3..............................................................................................................32. P greggii
10. Leaves usually (except in high-altitudeP hartwegii) longer than 15 cm; seed cones
opening soon after maturity;apophyses of seed scales raised, or at least prominently
keeled (Fig. 6F-H).
12. Leaves very slender, drooping in 2 rows, 0.7-0.9(-1) mm wide, in fascicles of 34(-5 )............................................................................................................................22. pa t l
12. Leaves >1 mm wide, if 0.8-1 mm, not drooping in 2 rows and predominantlyin
fascicles of 5.
13. Leaves in fascicles of (2-)3-4(-5) (count 15-20 fascicles).
14. Umbo of seed scales with a persistent spine or prickle; seed cones 8-15
cm long; leaves (18-)20-35 cm long ............................................... 12. P. engelannlii
14. Umbo of seed scales with a tiny, deciduous prickle; seed cones (4.5-)510(-14) cm long; leaves 14-25 cm long....................... 1c. P arizonica var. storntilae
13. Leaves in fascicles of (3-)4-5(-6), predominantly5.
15. Leaves (6-)10-17(-22) cm long; seed scales of cones relatively thin.
13. P h rtwegii
slightly flexible ...................................................................
15. Leaves (15-)20-40(-45) cm long; seed scales of cones relatively thick,
mostly inflexible.
16. Fascicle sheaths 30-40 mm long, resinous; leaves 1.1-1.6 mm wide;
outer walls of leaf endodermalcells thin; seed cones 15-35 cm long..
........................................................................................................ 16.
16. P devonia
16. Fascicle sheaths (15-)20-30(-35) mm long, usually not resinous;
leaves 0.8-1.3 mm wide; outer walls of leaf endodermalcells thickened; seed cones 7-20 cm long.
17. Leaves with connate vascular bundles; seed cones usually
obliquely ovoid when closed ......................................... 14. P pseudostrobhs
17. Leaves with mostly separatevascularbundles;seed cones usually
ovoid-oblong to ovoid-attenuatewhen closed..................... 15. P. montezuettle
Key to the species of central& southernMexico (V)

(eastern Michoacan,Mexico, Distrito Federal,Morelos, Hidalgo, Tlaxcala, Puebla, Guerrero,
Oaxaca, southernVeracruz,Chiapas)
2. Seed cones with -50, very widely spreading,very moveable scales; cones not longer than wide...........
................................................................................................................................................................43.P cembroides
2. Seed cones with 250, rigid scales, spreading-90?; cones longer than wide.
3. Leaves in fascicles of (4-)5(-6) (count 15-20 fascicles); seed cones ovoid, (4-)5-7(-8) cm long;
umbo on seed scales dorsal (Fig. 6Da, c) ...............................................................la. leiophll var. leiophl
3. Leaves in fascicles of 5, rarely 6; seed cones cylindrical, (6-)8-40(-50) cm long; umbo on seed
scales terminal(Fig. 6Ca).
4. Apophyses, at least those of proximalseed scales, recurvedor reflexed;seed cones usually 15-40
cm long. ..34. P avuacahuite
4. Apophyses of seed scales not reflexed; seed cones usually 8-16 cm long, rarely longer................
..................................................................................................................................38. P strobus var. chiapensis
1. Leaf fascicle sheaths persistent.
5. Seed cones asymmetrical,oblique at base or curved,4.5-35 cm long; seed scales opening >90?; apophysis nearly flat to prominentlyraised (Fig. 6E-H).
6. Leaves relatively short, (6-)10-17(-22) cm long, spreading;apophyses of seed scales nearly flat or
slightly raised (Fig. 6E-F), light brown or purplishblack............................................................... 13. P hartwegii
6. Leaves very long, (15-)20-40(-45) cm, spreadingor drooping;apophyses of seed scales nearly flat
or prominentlyraised (Fig. 6E-H).
7. Cones 5-10(-12) cm long; falling intact (peduncle deciduous with the cone).
8. Seed scales of cones thin, flexible, in open cones usually stronglyrecurved;leaves very slender,
ininoi
drooping,0.6-1(-1.1) mm wide .......................................................................................... 8. P
70
Flora Neotropica
8. Seed scales of cones not flexible, not strongly recurved in open cones; leaves drooping or
17. P douglasiana
spreading,0.7-1.2 mm wide..................................................................................
7. Cones (7-)10-35 cm long; leaving a few proximalscales on the branchwhen falling.
9. Fascicle sheaths 30-40 mm long, resinous;leaves 1.1-1.6 mm wide; outer walls of endodermal
cells thin; seed cones 15-35 cm long................................................................................... 16. P devoniana
9. Fascicle sheaths (18-)20-30(-35) mm long, usually not resinous; leaves 0.8-1.3 mm wide;
outer walls of endodermalcells thickened;seed cones 7-20 cm long.
10. Leaves with (partly) connate vascular bundles; seed cones usually obliquely ovoid when
closed ........................................................................................................................ 1. P. pseudostrobus
10. Leaves with usually separatevascular bundles;seed cones usually ovoid-oblong to ovoidae
attenuatewhen closed...................................................................................................15. P rnontezum
5. Seed cones symmetrical,ovoid or ovoid-oblong, sometimes slightly oblique, (4-)510(-12) cm long; seed scales usually spreading<90?; apophysis flat or slightly raised
(Fig. 6E-F).
11. Cones broadlyovoid to subglobose when closed.
12. Cones semi-serotinous (only the distal seed scales parting), remaining on
the tree; leaves 0.8-1.6 mm wide; resin ducts in the leaves septal..........20. P oocarpa
12. Cones opening completely when mature,falling after 1-3 years; leaves 0.71(-1.3) mm wide; resin ducts in the leaves medial............................24. P tecunumanii
I11.Cones ovoid to ovoid-attenuatewhen closed.
13. Leaves with (6-)9 or more lines of stomata on the abaxial face; leaf resin
ducts internalor internaland medial.
14. Cones semi-serotinous, persistent, leaving some basal scales on the
branchwhen falling; umbo of seed scales flat or depressed.............. 27. P pringlei
14. Cones opening at maturity,falling soon with peduncles attached;umbo
of seed scales pyramidal,prominentlyraised ................................... 26. P iawso.nii
13. Leaves with 3-7(-10) lines of stomataon the abaxial face; leaf resin ducts
only medial.
15. Leaves (7-)10-15(-18) cm long, 1-1.4 mm wide............................... 28. P teocote
15. Leaves (11-)15-25(-30) cm long, 0.7-0.9(-1) mm wide .....................22. P patula
Key to the species of Mesoamerica(VI)

(Chiapas, QuintanaRoo, Belize, Guatemala,El Salvador,Honduras,Nicaragua)
2. Apophyses, at least those of proximal seed scales, recurvedor reflexed; seed cones usually 15-40 cm
long........................................................................................................................................................ 34. P avacahuite
2. Apophyses of seed scales not reflexed; seed cones usually 8-16 cm long, rarely longer...........................
............................................................................................................................................38. P strobus var. chiapensis
3. Long shoots usually multinodal;vegetative buds usually resinous; leaves in fascicles of 3 (rarely 2 or
c. P caribaea var. hondurensis
4); leaf resin ducts internal ......................................................
3. Long shoots uninodal;vegetative buds usually not resinous;leaves in fascicles of (3-)4-5, rarely6; leaf
resin ducts variously positioned.
4. Seed cones asymmetrical,oblique at base or curved, 4.5-35 cm long; seed scales opening to >90?;
apophysis nearly flat to prominentlyraised (Fig. 6E-H).
5. Leaves relatively short, (6-)10-17(-22) cm long, spreading;apophyses of seed scales nearly flat
or slightly raised (Fig. 6E-F), light brown or purplishblack .................................................... 13. P hartwegii
5. Leaves very long, (15-)20-40(-45) cm, spreadingor drooping; apophyses of seed scales nearly
flat or prominentlyraised (Fig. 6E-H).
6. Cones (4-)5-10(-12) cm long; falling intact (peduncle deciduous with the cone); seed scales
thin, flexible, in open cones usually strongly reflexed; leaves very slender,drooping, 0.6-1
(-1. ) mm wide ..................................................................................................................... 18. P maximinoi
6. Cones (7-)10-35 cm long; leaving a few proximal scales on the branch when falling; seed
scales thick, rigid, usually not recurvedor reflexed; leaves 0.8-1.6 mm wide.
7. Fascicle sheaths 30-40 mm long, resinous;leaves 1.1-1.6 mm wide; outer walls of endoderP. 16. devoniana
mal cells thin; seed cones 15-35 cm long ........................................................
7. Fascicle sheaths 20-30(-35) mm long, usually not resinous; leaves 0.8-1.3 mm wide; outer
walls of endodermalcells thickened;seed cones 8-20 cm long.
8. Leaves with (partly)connate vascularbundles;seed cones usually obliquely ovoid when
closed. .................................................................................................................... 14. P. pseudostrobus
8. Leaves with usually separatevascularbundles;seed cones usually ovoid-oblong to ovoidattenuatewhen closed................................................................................................ 15. P montezumae
71
4. Seed cones symmetrical, ovoid or ovoid-oblong, sometimes slightly oblique,

(3-)5-10(-12) cm long; seed scales usually spreading <90?; apophysis flat or
slightly raised (Fig. 6E-F).
9. Cones broadlyovoid to subglobose when closed.
10. Cones semi-serotinous (only the distal seed scales parting), remaining
on the tree; leaves 0.8-1.6 mm wide; leaf resin ducts septal .............20. P oocarpa
10. Cones opening completely when mature,falling after 1-3 years; leaves
0.7-1(-1.3) mm wide; leaf resin ducts medial..............................24. P tecunumanii
Key to the species of the Caribbean(VII)

(southernFlorida,Bahamas,Turks-CaicosIslands, Hispaniola,Cuba)
1. Leaves in fascicles of 2, rarely 3; resin ducts in the leaves septal (Fig. 2L); umbos on seed scales without
a prickle. ili.........................................................................................................................................................2.
P tropicalis
1. Leaves in fascicles of 2-4(-5); resin ducts in the leaves internal(Fig. 2H-I); umbos on seed scales with a
deciduous or persistentprickle.
2. Leaves (6-)10-15(-18) cm long; seed cones usually 4-7 cm long; seeds with adnate wing (Fig. 61). 7. P cuhensis
2. Leaves (11-)14-28(-30) cm long; seed cones 5-13 cm long; seeds with articulatewing (Fig. 6K-L).
3. Vegetativebuds not resinous; leaves in fascicles of 2-3 (count 15-20 fascicles) ............ 6. P elliottii var. densa(
3. Vegetativebuds usually resinous;leaves in fascicles of (2-)3-5.
4. Long shoots multinodal;fascicle sheath of newest leaves 15-20 mm long; leaves in fascicles of
c ri ea
(2-)3(-4 ), very rarely 5....................................................................................................................
4. Long shoots uninodal;fascicle sheaths of newest leaves (8-)10-15 mm long; leaves in fascicles
of 3-5 ...........................................................................................................................................
occident lis
I. Pinus subgen. Pinus
(pulvini). Cataphylls usually persistent, becoming

recurvedon shoots, falling with the leaf fascicles or
shortlyafterwards.Fascicle sheaths mostly persistent
Pinus I. Pinus MUnchhausen,Hausvater 5: 215. 1770; (deciduous in two
species). Leaves amphistomatic,
Pinus [group] Pinus Sweet, Hort. Brit., ed. 2, 475.
with serrulate margins; resin ducts variously posi1830, nom. nud.; Pinus Isubgen.] B. Pinus Endlicher,
stele with outercell walls of endodermisthin
Syn. Conif. 137. 1847; Pinus subgen. Pinus Parlatore tioned;
in Alph. de Candolle, Prodr. 16(2): 364, 378. 1868; or thickened,containingtwo vascularbundles,which
Pinus sect. Diploxylon Koehne, Deut. Dendrol. 30. are separateor (partly)connate. Pollenlcones (long)
1893; Pinus subgen. Pinaster Lemmon, Cal. State
when mature,crowded near the proximal
Board Forest. Bienn. Rep. 2: 71, 85. 1888; Lemmon, cylindrical
Cone-Bear.Trees Pacif. Slope, ed. 3, 24. 1895; Pinus end of new long shoots; microsporophyllsradially
div. CrassisquamaeMasters,J. Linn. Soc. London,Bot. striatedistally and with erose-denticulateuppermar35: 570. 1904; Pinus subgen. Diploxylon (Koehne) gin. Seed cones often persistent, symmetrical or
Rehderin Bailey, Cult. Evergr.311. 1923; Pinus subgen.
oblique. Seed scales numerous,with complex phylloEupitys (Spach) Rehder, Biblio. Cult. Trees 35. 1949; taxis;
apophysis usually transverselykeeled; umbo
Pinus subgen. Diplopinus Campo-Duplan,Trav. Lab.
Forest.Toulouse T. 2, sect. 1, 4, art. 1: 94. 1950, nom. dorsal, with or without a spine or prickle. Seeds
nud.; Pinus subgen. ScleropitysIshii, Kochi Univ. Nat. commonly with an articulate(rarely adnate) but efSci. Reports 2: 113, 125. 1952, sine diagn. lat.; Pinus fective wing which is largerthan the seed.
Pinus subgen. Pinus (spp. 1-33)
subgen. Eupinus Gaussen, Trav. Lab. Forest. Toulouse

T. 2, sect. 1, I, part 2: 94. 1960, sine descr. lat.
Lectotype species (vide Britton, N. Amer. Trees 5.
1908). Pinus sylvestris Linnaeus,Sp. PI. 2: 1000. 1753.
Pinus subgen. Pinea (Endlicher) Landry, Nat. Canad.
101(5): 774. 1974. Holotype species. Pinus pinea Linnaeus, Sp. PI. 1000. 1753.
Pinus subgen. SabinianaE. Murray,Kalmia 13: 18. 1983.
Holotype species. Pinus sabiniana Douglas ex D. Don
in Lambert,Descr. Pinus, ed. 8, 2: p. s.n. inter 144 et
145, t. 80. 1832.
Bark thin and scaly on branches and stems of

young trees, soon variously furrowed or plated and
Long shoots uninodal or
scaly, exfoliating.
multinodal, rough, with long decurrent leaf bases
1. Pinus leiophylla Schiede ex Schlechtendal &

Chamisso, Linnaea6: 354. 1831.
Tree, medium to tall, height to 15-30(-35) m,
dbh to 20-85 cm. Trunkmonopodial, erect, terete,
sometimes forked higher up, often clear of branches
for 3/5 of total height. Bark very thick, rough, scaly,
divided into elongated, irregular,scaly plates and
deep, mainly longitudinalfissures,dark grey-brown,
on young trees and branchesscaly, flaking, reddish
brown to grey-brown. Branches long, slender,
spreadingor ascending,the lower ones curveddown,
forming an open, rounded crown. Shoots more or
72
less scaly, with short, decurrent pulvini, initially
reddish brown or glaucous, soon grey-brown. Cataphylls small, ca. 6 mm long, lanceolate-subulate,
soon recurved,light orange-brownwith hyaline-ciliate margins,early deciduous. Vegetativebuds ovoidconical, obtuse, the terminalbuds 10-15 x 5-7 mm,
the laterals smaller, not or slightly resinous, light
brown; the scales imbricate, subulate, with ciliate
margins and free apex. Fascicle sheaths 12-20 mm
long, with 5-6 initially imbricate, ligulate, ciliatemargined, early deciduous scales, leaving the fullgrown fascicles devoid of a sheath. Leaves in fascicles of (2-)3-5(-6), spreading in lax or rigid tufts
near the ends of branches, persisting 2-3 years,
sometimes subpendulous,straight,variablein length
on the same branch,(4-)6-15(-17) cm X 0.5-1.3(1.5) mm, the margins (minutely) serrulate,acute or
acute-pungent, light green or glaucous-green. Stomata on all faces of leaves, in (3-)4-8(-9) lines on
the convex abaxial face, in (2-)3-4 lines on each
adaxial face, generally correlatedwith the width of
the leaf. Leaf anatomy: Cross section triangular,
with a convex abaxial side; hypodermis 1-2-layered,
rarelythickerand intrudinginto the mesophyll;resin
ducts (1-)2-6(-7), medial, occasionally 1-2 internal;
stele terete or slightly oval in cross section; outer
cell walls of endodermisnot thickenedor thickened;
vascularbundles2, distinctbut connateor with partly
merged xylem strands. Pollen cones crowded near
the proximal end of a new shoot in a short, dense
cluster,subtendedby large, subulate,scariousbracts,
cylindrical, 10-15(-20) x 4-6 mm, yellowish pink,
maturingto light brown. Microsporophyllswith relatively large, conspicuous pollen sacs. Seed cones
subterminal,solitary or in whorls of 2-5 on stout,
10-20 mm long, spreading or recurved peduncles,
persistinga few years after seed dispersal.Immature
cones broadly ovoid to subglobose, with pungent
spines, initially pink, later purplishgreen to purplish
brown,maturingin threeseasons. Maturecones (narrowly) ovoid when closed, nearly symmetrical,
(broadly) ovoid when opened, (4-)5-7(-8) X (3-)
4-5.5 cm when open. Seed scales ca. 50-70, parting
gradually,eventually spreadingwide, straightor recurved nearthe base of the cone, oblong, with nearly
straightmargins,darkpurplishbrownturninggreyish
on the abaxial side, purplish brown with lighter,
greyish marks of seed wings adaxially. Apophysis
mostly uniformaroundthe cone, raised, transversely
keeled, with the central section of the second season's growth distinctly marked as a narrow band
around the umbo, rhombic in outline, up to 15 mm
wide at mid-cone, dull brown to grey-brown. Umbo
Flora Neotropica
dorsal, pyramidal, transverse-rhombic in outline, up
to 5 mm wide, with a blunt prickle, darker than the
apophysis. Seeds obliquely ovoid, slightly flattened,
small, 3-4(-5) mm long, dark grey-brown with black
spots. Seed wings articulate, effective, held to the
seed by two claws, the membrane partly covering
the seed on one side, obliquely oval, 10-18 X 4-8
mm, yellowish brown, translucent, with a dark tinge.
Cotyledons or seedlings not observed. Adventitious
shoots sprout from the trunk or stump after fire or
cutting and may develop into a small tree.
Distribution (Map 1). SW United States (SE Arizona, SW New Mexico); in Mexico along the Sierra
Madre Occidental, the "Eje Volcanico Transversal"
and in the Sierra Madre del Sur as far SE as the
highlands of central Oaxaca.
Key to the varieties of Pinus leiophylla
1. Leaves in fascicles of (4-)5(-6), 0.5-0.9 mm wide,
with (3-)4-6(-7) lines of stomata on the abaxial
face; numberof resin ducts (1-)2-3(-4)................
.............................................................
a. va .lla ph
eio
1. Leaves in fascicles of (2-)3(-4, rarely 5), 0.91.3(-1.5) mm wide, with (4-)5-8(-9) lines of stomata on the abaxial face; number of resin ducts
na
(3-)4-6(-7) ................................... Ib. var. chiiuahuah
la. Pinus leiophylla Schiede ex Schlechtendal &
Chamisso var. leiophylla. Type. Mexico. Veracruz:
Cruz Blanca, 28 Nov 1828, Schiede 1109 (holotype,
HAL)
Fig. 16A-F
Pinus gracilis Roezl, Cat. Grain.Conif. Mexic. 21. 1857;
Pinus comnonfortii
Roezl var. gracilis (Roezl) Carriere,
Trait6G6n. Conif., ed. 2, 2: 565. 1867. Type. Mexico.
"Habite le revers des cordilleres du c6t6 du Pacifique,
a une altitude de 9500 a 10.000 pieds," 1857, Roezl
s.n. (lectotype, Fl, here designated).
Pinus comonlfortiiRoezl, Cat. Grain. Conif. Mexic. 22.
1857. Type. Mexico. Mexico: Huixquilucan, 1857,
Roezl s.n. (lectotype, Fl, here designated).
Pinus decandolleana Roezl, Cat. Grain. Conif. Mexic.
22. 1857. Type. Mexico. Locality not stated, 1857,
Roezi s.n. (lectotype, P, here designated; isolectotype,
Fl).
Pinus huisquilucaensisRoezl, Cat. Grain. Conif. Mexic.
22. 1857. Type. Mexico. Mexico: Huixquilucan. 1857,
Roezl s.n. (lectotype, FI, here designated).
Pinus monte-allegri Roezl, Cat. Grain. Conif. Mexic.
22. 1857; Pinus comonfortii Roezl var. monte-allegri
(Roezl) Carriere, Trait6 Gen. Conif., ed. 2, 2: 566.
1867. Type. Mexico. Michoacan: Near Xochiltepec at
Zitacuaro, 1857, Roezl s.n. (lectotype, Fl, here designated).
Pinus verrucosa Roezl, Cat. Grain. Conif. Mexic. 22.
1857; Pinus decandolleana Roezl var. verrucosa
(Roezl) Carriere,Trait6G6n. Conif., ed. 2, 2: 570. 1867.
Type. Mexico. Huixquilucan ("Huisquiluca"), 1857,
73
Ma1Pu
nvrcieahaa(rage)
IepIavrleo/ila(ice)Plcol
A~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~'S
tth
4
lelll (I
Map
Map
3
.
8.*
1.
-..
. I
Pinus cedrus Roezl, Cat. Grain. Conif. Mexic. 23. 1857.

Type. Mexico. Mexico: On a hillside nearthe road from
Mexico to Cuernavaca, 1857, Roezl s.n. (lectotype. P,
here designated: isolectotype, Fl).
Pinus dependens Roezl, Cat. Grain. Conif. Mexic. 23.
1857. Pinus decandolleana Roezl var. dependens
(Roezl) Carriere, Traite Gen. Conif., ed. 2, 2: 569.
1867. Type. Mexico. Mexico: Near rd. from Mexico
to Cueravaca, 1857, Roezi s.n. (lectotype, P, here
designated;isolectotype, Fl).
Pinus lerdoi Roezl, Cat. Grain. Conif. Mexic. 23. 1857.
Pinus comonfortii Roezl var. lerdoi (Roezl) Carriere.
Traite Gin. Conif. ed. 2, 2: 567. 1867. Type. Mexico.
Mexico: Sierra de Ajusco, 1857, Roezl s.n. (lectotype,
P, here designated; isolectotype, FI).
Pinus lumholtziiB. L. Robinson& Fernaldvar.microphyIla Carvajal,Phytologia 59: 135. 1986. Type. Mexico.
Jalisco: San Martinde Hidalgo, Rancho El Cobre, 16
Sep 1983, Carvajal 4031 (holotype, CREG).
Tree, usually tall, height to 20-30(-35) m, dbh to
50-85 cm. Trunk monopodial, erect, straight. Shoots
is;
(a
reddishbrown, sometimes glaucous. Leaves in fascicles of (4-)5(-6), 4 more often than 6, (6-)8-15
(-17) cm X 0.5-0.9 mm. Stomata in (3-)4-6(-7)
lines on the convex abaxial face, in (2-)3-4 lines on
each adaxial face. Resin ducts in the leaves (I-)23(-4), medial, occasionally I internal.
Distribution and ecology. Mexico: In NE Sonora,
W Chihuahua,Durango, Nayarit,Zacatecas, Jalisco,
Michoacan, Mexico D.F, Hidalgo, Morelos, Tlaxcala, Puebla, Veracruz,Guerreroand Oaxaca. In the
north it is gradually replaced by var. chihuahuana,
which is presumablythe only taxon occurringin the
United States (see Perry, 1991). Pinus leiophylla var.
leiophylla is a widespreadconstituentof montaneto
high-montane pine and pine-oak forests on deep,
well-drainedsoils derived from various, but usually
volcanic or metamorphic,rock. Its altitudinalrange
is (1500-) 19002900(-3300) m, graduallyincreasing
74
Flora Neotropica
'i)1,
Padilla
&Stles
A.Shoot
with
100EF,F
Pinus
FIG.
16.A-F.
var.'
,Alman
38).
(A-D,
Higman,
leiophlla
eiphylla
'
Ia
xtwy^f^.
F ,
young leaf fascicles. B. Shoot with leaf fascicles and pollen cones. C. Leaf fascicle. D. Cross-section of leaf. E. Ovuliferous cones. F. Seeds. G-J. P leiophYllavar. chihuahuana(S. S. White3409). G. Branchwith leaf fascicles and ovuliferous
cone. H. Leaf fascicles with sheaths. I. Leaf fascicles. J. Cross-section of leaf. K. Seed scale (three views). (Magnifications: A-C,
A-C. E---,
XO.5;
K, x
D, J,
H, x
E-G, 1, K,
x2.)
J, xX40;
tions:
0.5; D,
40; H,
2.)
from north to south. It occurs most frequently with
Quercus spp. and/or with Pinus patula, P pringlei,
P teocote, P lawsonii, P pseudostrobus, P. montezumae, P. douglasiana, P durangensis, and, at lower
altitudes, P oocarpa. In the NW of its range, P
arizonica, P engelmannii, and R leiophylla var. chihuahuana grow commonly with it. Locally, Juniperus spp. or Cupressus lusitanica are found with it.
Annual precipitation varies greatly with locality and
altitude, from a low of ca. 700 mm to 1950 mm. In
the north and at high altitudes, frost and snow are
common in winter. It is one of the few pines with
a capacity to coppice. Phenology: Time of pollen
dispersal not recorded, it is probably variable; a few
herbarium specimens suggest March in the south of
the range. The ovuliferous cones take three seasons
to mature, which is exceptional in pines.
Representative specimens examined. MEXICO. CHIHUAHUA:Madera,Rio Negro, 12 Jun 1958, Matuda32677
(FHO, MEXU); San Vicente, 30 Jun 1892, Hartman 538

(A, NY, UC, US); Arroyo Ancho, 7 Jun 1929, Mex(a
2607 (P); Mt. Mohinora, I Sep 1898, Nelson 4907 (F).
DURANGO: 13 km N of Coyotes railway sta., 4 Nov 1970,
Breedlove 18726 (CAS, ME); El Salto, 30 Apr 1987,
Hughes 978 (FHO, MEXU); Lagunadel Progreso,46 km
N of Coyotes railway sta., 8 Aug 1955, Ma.silles 8380
(FHO, MICH);Quebradade San Juan,34 km N of Coyotes
railway sta., 9 Aug 1955, Mavsilles 8435 (FHO, MICH).
2 km E of Malinaltepec,2 Jun 1981, Almazdn
GUERRERO:
100 (FHO); along rd. from Chilpancingo to San Vicente,
25 Mar 1993, Higman. Padilla & Styles 60 (EAP, FHO,
HEH, MO). JALISCO:
Tapalpa,9 Oct 1892, Ellis 30 (MO);
10 km from CiudadG6zman, 21 Feb 1980, Stead & Styles
528 (FHO, MO); 12 km from Ciudad Guzman, 16 Mar
1974, Styles 89 (FHO). Mexico: Desierto de Le6n, 2 Apr
1950, Matuda 19495 (FHO, MEXU, MO); Juchitepec, 15
Apr 1982, Lorea 1969 (FHO);Mt. Ixtaccihuatl,9 Jan 1894,
Nelson & Goldmtans.n. (A, E, US); Sierra de Ajusco, 2
Nov 1895, Pringle 6180 (A, BM, E, F, GOET, JE, K, M.
MO, NY, S). MICHOACAN:
along Hwy. 37 to Uruapan,ca.
5 km S of Carapan,6 May 1994, Farjon & Mejia 349 (E,
FHO, HEH, K, MEXU, U); Cerro Patzcuaro,Nov 1890,
Pringle 3377 (F, GH, MICH), Nov 1890, Pringle 4171 (F,
MICH, NY, UC, US), 11 Oct 1892, Pringle 4358 (BM, F,
GH, GOET, JE, K, M, MO, P, S, UC, US); Tajimaroa
("prope Tajimaroaprovinciae Michoacensis"), Dec 1829,
Schiede 1117 (HAL); San Felipe, near Zitacuaroalong rd.
to Angangueo, 8 Feb 1980, Stead & Styles 462 (FHO); 5
km N of Uruapan, 22 Feb 1974, Styles I (FHO, K).
OAXACA:Along rd. from Oaxaca to Tuxtepec (Hwy. 175),
near San Juan Atepec, 23 Apr 1994, Farjon & Mejia 315
(E, FHO, HEH, K, MEXU, U); La Carbonera, 18 Mar
1993, Higman, Padilla & Styles 38 (EAP, FHO, HEH,
MO); National Park Benito Juarez,20 Mar 1993, Higman,
Padilla & Styles 41 (EAP, FHO, HEH, MO); Miahuatlan,
22 Mar 1993, Higman, Padilla & Styles 50 (EAP, FHO,
MO, ESNAC, HEH); 1841, Liebmann 14791 (MO, NY);
Llano Verde,E of Oaxaca, 4 Mar 1985, McCarter& Styles
344 (FHO). PUEBLA:
Paso Cortez, 17 Jul 1977, Mittak8507
75
Conchatera,Aug 1901, Rose & Ha\
(FHO, M). TLAXCALA:
5968 (A, K, NY,US). VERACRUZ:

LasVigas,nearPerote,
20 May 1899,Pringle8182 (A, BM, E, F, GOET,JE, K.
"SanJose del Oro,"1839,
M, MO, NY, P). UNKNOWN:
Ehrenbergs.n. (P).
lb. Pinus leiophylla Schiede ex Schlechtendal &

Chamisso var. chihuahuana (Engelmann) G. R.
Shaw, [Pines Mexico] Publ. Arnold Arbor. 1: 14.
1909.
Fig. 16G-J
Pinuschihuahuana
in Wislizenus,
Mem.Tour
Engelmann
N. Mexico 103. 1848; Pinus leiophyllaSchiedeex
Schlechtendal
& Chamissosubsp.chihuahuana
(EngelKalmia12:23. 1982.Type.Mexico.
mann)E. Murray,
Chihuahua:
Oct 1846, Wislizenus232
Cosiquiriachi,
(holotype,MO 1817023,andcone MO 1635446).
Tree, medium size, height to 15-25 m, often not
more than 10 m, dbh to 20-60 cm. Trunkmonopodial, erect, often forked higher up. Shoots reddish
brown, often glaucous. Leaves in fascicles of (2-)
3(-4, rarely 5), (4-)6-12(-14) cm X 0.9-1.3(-1.5)
mm. Stomata in (4-)5-8(-9) lines on the convex
abaxialface, in 3-4 lines on each adaxial face. Resin
ducts in the leaves (3-)4-6(-7), medial,occasionally
1-2 internal.
Distribution and ecology. United States: SE Arizona, SW New Mexico. Mexico. Along the Sierra
MadreOccidentalin NE Sonora,W Chihuahua,Durango, Nayarit,N Jalisco, and Zacatecas.Pinus leiophylla var. chihuahuanawas included with the species in the distributionmap given by Critchfieldand
Little (1966); it has been recognized as a distinct
species by Perry (1991), who gives a generalized
map roughly in accordance with our findings. Althoughlargely sympatric,the altitudinalrangeof var.
chihuahuana is narrower- 500-2700(-2950) mbut this is in partdue to the fact that it does not occur
on some of the high volcanos of central Mexico. In
the Sierra Madre Occidental its lower limit usually
is below the level of var. leiophylla. Here it may
be found in semi-arid habitat with P. cembroides,
Juniperus spp., and xerophytes like Opuntia and
Arctostaphylos.Higher up, its habitat is similar to
that described above. Its poor growth in some areas
is related to lower precipitation and rocky, poor,
shallow soils, especially at lower and middle altitudes. Phenology:The time of pollen dispersalis not
recorded.
Representative specimens examined. MEXICO. CHIHUAHUA:
Along rd. betweenCreeland La Bufa, 12 Sep
410 (MEXU);Cerrode la Ventana,16km

1957,Knobloch
NWof Minaca,10May1929,Mexia2514(CAS,E, MICH,
MO,NY, P); aboveArroyoAnchoon Mesade Gonzalez,
8 Jun 1929,Mexia2620 (CAS, E, MICH,MO, NY. P):
76
Flora Neotropica
YerbaBuenaaboveBatopilas,Nov 1885,Palner301 (GH); titious ("epicormic")shoots along the trunkof mature

SierraLa Silla (SierraMapula),5 Nov 1886,Pringle959 trees.
(BM,F, MEXU,NY);CascadaBasaseachic,11Nov 1989,
Carvajal (1986) described Pinus lumlholtziivar.
Spellenherget al. 10064(FHO,MEXU,NMC,ESAHE),
12 Nov 1989, Spellenberget al. 10081 (FHO, ID, MEXU, microphylla based on Carvajal 4031 (CREG!) but
NMC); Cascada Basaseachic, 12 Nov 1989, Spellenberget also referringto materialcollected tentatively as P
al. 10090 (FHO, NMC);ColoniaGarcia,1 Oct 1899, chihuahuana. His conclusion, that it resembles P
Townsend& Barber359 (BM, E. F, GH, K, MO,NY, P, lumholtziimore,
appearsto have been withdrawnin
NW of El Salto, 12 Jul 1898, Nelson 4558
UC). DURANGO:
& McVaugh, 1992: 71, where its type is
Carvajal
83
Palmer
1896,
(A, K, US); SantiagoPapasquiaro,
Apr
(K, MO);nearDurango,
Apr1896,Palmer774(K),Palher synonymizedwith P leiophylla var.leiophylla. Pinus
775 (A, BM, F, K, MO,NY, US); Durango.2 Apr 1991, lumholtzii has sometimes short, non-pendulous
SantaTeresa, 1897, leaves and can in those cases become confused with
Stles, Favela et al. I (FHO). NAYARIT:
Rose 2123B (A). SONORA:Region of the Rio de Bavispe,
P leiophylla var. chihuahuana(Martinez, 1948; see
20 Aug 1940,Phillips659 (GH);
LasTieritasdel Tamblor,
SierraEl Tigre, 18 Aug 1940, White3409 (FHO,GH, also the treatmentof P lumholtzii,this volume).
The numerous "species" described or named by
MICH);regionof the Rfode Bavispe,Ranchode la Nacha
SE of El Tajo, 15 Aug 1941, White3942 (GH). ZACATECAS: Roezl, most of which were synonymizedby Parlatore
25 Jul 1982,Diggs& Nee 2997
16 kmSE of Chalchihuite,
(1868), have been typified for the first time from
(F. FHO, XAL); Sierrade los Morones,near Plateado. Roezl's materialin Fl and P. A
general comment on
1897. Rose 2742A (GH).
Roezl and his pines is given under the treatmentof
Uses. Along with many other species of pine in Pinus montezumae.
In the Botanical Appendix to Wislizenus (1848),
the Sierra Madre Occidental, Pinus leiophylla has
been heavily exploited for timber in the latter half Engelmann described three new species of pines
of this century. Because of its high resin content, it "entirely different from those found farther north,
does not provide high-qualitywood but at the same but perhaps identical with some species from the
time is a producer of good resin, for which it is Pacific coast." One of them is Pinuischihuahuana.
tapped.The variety chihuahuana,an equally prolific No comparisonswere made with other pines already
described from the Sierra Madre Occidental and
producerof resin, is less exploited for timber.
Under Coniferae, Schlechtendal and Chamisso occurringfarthersouth. In his revision of the genus,
(1831) describedtwo new species of Pinus, P patula Engelmann(1880a) classified P leiophylla in a difand P leiophylla, based on Schiede (& Deppe) 1108 ferent subgroup from P chiuahuahuana,
although he
sheaths.
fascicle
its
deciduous
observed
and 1109 respectively, and both from (approxi[Hartveg
mately) the same locality. Schiede (& Deppe) 1109 441 does indeed have persistent sheaths, but it beconsists of a single sheet with two sterile branchlets longs to P teocote, not to P leiophylla, where Engelat HAL, which is the holotype of P leiophylla. mann (1880a) and Parlatore (1868) cited it]. Not
Deppe, mentioned as the second collector in the until Shaw (1904b, 1909) recognized the prolonged
protologue, does not appear on the labels. Another phenology as well as the sharedcharacterof adventicollection in HAL, Schiede 1117, bears a small cone, tious shoots and coppicing capacity were P. leiophybut in the protologue it is claimed no cones were lla and P chihuahuanacorrectly interpretedas conobserved ("nobis deficiente"). Although meagre, specific.
Schiede 1109 is unambiguousand need not be supEngelmann(in Wislizenus, 1848) did not indicate
plemented with Schiede 1117 as an epitype; but as which of the collections of Wislizenus pertains to
another specimen annotated by Schlechtendal, the his new species, and neither does Parlatore(1868),
latterhelps to interprethis taxon more fully. Lambert who received some materialon loan from Engelmann
(1832, t. 21) had the species drawn for his third (see Carvajal& McVaugh, 1992: 69), as is annotated
edition from a specimen with a cone received from
by Engelmannon the sheet of Wislizenus232 (MO
the collector(s).
1817023). This sheet and a cone (MO 1635446),
from the loan to Parlatore,are to be considthe
returned
describe
the
first
author
to
Shaw (1904a) was
a
the
feature
ered
ovuliferous
of
the
cone,
holotype of P chihuahuana.
prolonged growth
Martinez (1948), who maintained both taxa at
shared by P pinea of the Mediterraneanbut unique
in Latin American pines. It shares this feature with the rank of species, nevertheless admitted that P
its variety, P leiophylla var.chihuahuana.Similarly, chihuahuanain the southernpartof its range merges
the ability to coppice, shared by P rigida of the with P leiophvlla to such an extent that it becomes
eastern United States, is an uncommonphenomenon very difficult to distinguish them. We, too, have
presentin both varieties.A relatedfeatureare adven- found this to be the case: and since there seem to
77
be no other than quantitative foliar characters to lines on the convex abaxial face and in 6-8 lines on
differentiatethem, while they sharetwo very distinct the adaxial face (3-leaved specimens not observed).
and uniquequalitativefeatures,these combinedargu- Leaf anatomy: Cross section semi-circular (rarely
ments stronglysupporttreatmentat varietalrank.On transverse-triangular);
hypodermisuniform in thickthe other hand, in certain areas in Durangothe two ness with 1-2 rows of cells but thicker in marginal
taxa retain their differences over large areas, where areas; resin ducts 6-10, very large, septal, or sometypical P leiophylla with fascicles of five, slender, times a few external; stele semi-terete, large; cell
pendulous leaves grow in proximity to "pure" P walls of endodermisthin; vascular bundles 2, sepaleiophylla var. chihuahuana with fascicles of three, rateor the xylem strandsmore or less connate.Pollen
thick, spreadingleaves (Styles, 1992). Furtherassess- cones crowdednearthe proximalend of a new shoot,
ment of the relationshipof these two pines and their ovoid-oblong to cylindrical, 20-30 x 5 mm, pink
putative relatives, including molecular genetics, is to yellowish, turning yellowish brown. Seed cones
desirable.
subterminal,erect on short, thick peduncles, solitary,
in pairsor whorls up to 6, remainingerect or spread2. Pinus tropicalis P. M. A. Morelet, Rev. Hort. ing obliquely. Immaturecones (narrowly)ovoid, 10
C6te d'Or 1: 106. 1851. Type. Cuba. Isla de la x 5-7 mm, purplishred, maturingin two seasons.
Juventud[Isla de Pinos]: La Caniada,16 Feb 1916, Mature cones narrowly ovoid to ovoid-attenuate
Brittonet al. 14416 (neotype, NY, here designated). when closed, ovoid with a flattened base when
Fig. 17 opened, then 5-8 x 4-5.5 cm, persistent for some
Pinus cubensisGrisebachvar. terthrocarpa
Grisebach, years but falling with the peduncle attached. Seed
Cat.PI.Cub.217. 1866;Pinulsterthrocurpa
(Grisebach) scales ca. 100-120, parting to release the seeds
G. R. Shawin Sargent,Trees& Shrubs1: 149,t. 75. except those at the base, oblong, straightor strongly
1903.Type.Cuba.Localitynotindicated,1860,Wright recurved, dark brown, with light brown marks of
marked
3189(lectotype,GOET.secondarily
3189a,here seed
wings on the adaxial side. Apophysis flat or
GH.JE, K, S).
isolectotypes.
designated:
slightly raised, transverselykeeled, rhombicto penTree, medium to large, height to 30 m, dbh to tagonal in outline, up to 12 mm wide, radially stri100? cm. Trunk monopodial, erect, terete, usually ated, light brown or reddish brown. Umbo dorsal,
slender, in forest stands with clear bole for /3-3/4 of
flat or slightly raised, rhombic, ca. 3 mm wide,
height. Barkthick, rough, scaly, breakinginto irregu- without a prickle, greyish brown to grey. Seeds
lar, polygon plates, divided by deep longitudinal obliquely ovoid, slightly flattened,5 x 4 mm, light
fissureson the lower partof the trunk,reddishbrown, grey-brown.Seed wings articulatebut effective, held
weatheringgrey, on young trees and branchesscaly, to the seed by two claws, obliquely obovate-oblong,
flaking, reddish brown. Bratches of first and higher 12-15 x 5-6 mm, yellowish with a grey or black
orderssparse, thick, spreadingto ascending, forming tinge. Seedling: Numberof cotyledons not observed.
an irregular,open crown. Shoots uninodal, thick, Suppressedterminalgrowth and a thick radicle provery rough with large, prominent and persistent, duce a "grassstage"as an adaptationto groundfires.
decurrentpulvini, lustrous orange-brownin Ist and Leaves are normalbut less straight.
2nd year, then grey, scaly with flat bases of pulvini.
Distribution and ecology (Map 2). W Cuba: in
Cataphylls subulate, recurved, scarious, with erose- Pinar del Rio and on the Isla de la Juventud(often
ciliate margins and caudate apex, brown, caducous. still referred to as Isla de Pinos (Isle of Pines) in
Vegetativebuds ovoid-oblong, acute, the terminal botanical literature).Pinus tropicalis is a lowland
bud 15-25 mm long (on vigorous leading shoots of pine, occurringon the coastal plains and low foothills
young trees larger),the lateralssmaller,not resinous; at 1-150(-300) m altitude,on nutrient-poorsandy or
the scales more or less spreading, reddish brown gravelly alluvial soils which are dry due to rapid
with ciliate, hyaline margins. Fascicle sheaths of drainage.The climate is tropical,with annualprecipijuvenile leaves ca. 20 mm long, papery, yellowish tation of ca. 1200 mm and a prolongeddry season.
brown with ciliate, whitish margins, persistent but It is in partsympatricwith P caribaea var.caribaea,
reduced to ca. 10 mm on mature leaf fascicles, which has a greateraltitudinalrange. Pine savannas
weatheringgrey. Leaves in fascicles of 2 (rarely 3), are open, grass-dominatedlowland areas which burn
spreadingobliquely,persisting2 years, very uniform, frequently; P tropicalis has an advantage over P
straightand rigid, (15-)20-30 cm X 1.5 mm, with caribaea in that it has a "grassstage" which enables
serrulate margins, acute, light or yellowish green. the seedling to survive successive fires. Thus it beStomataon all faces of leaves, in 6-8 inconspicuous comes frequently the only pine in this vegetation
78
Flora Neotropica
FIG. 17. Pinus iropicalis [a, C, D, Little 14048; B, E-H, Lambs.n. (div. coll.)]. A. Branch with leaf fascicles and two
immatureovuliferous cones. B. Seedling in "grass stage." C. Leaf fascicle. D. Cross section of leaf. E-F. Ovuliferous
cones. G. Seed scale (three views). H. Seeds. (Magnifications:A-C, E-H, X0.5; D, X25.)
79
Map 2. Pinus tropicalis (circles); P. cubensis (triangles);P. occidentalis (squares).
used in plantationforestry,mainly in Cuba, but also

to a limited extent elsewhere.
Morelet (1851) recognized from his field observaRepresentative specimens examined. CUBA. ISLADE
that there are two distinct species of pines in
tions
LAJUVENTUD
[ISLADE PINOS]:Loma Vista, near Nueva
Gerona, 26 Nov 1920, Ekman 12385 (K, S); 25 km WSW W Cuba, both on the "Isla de Pinos" and on the
of Nueva Gerona, 22 Jan 1951, Little 14048, 14052 (US); mainland. They were not to be equated with P
locality not indicated, 22 Mar 1905, Shaw s.n. (M). PINAR occidentalis Swartz, the only pine previously deDELRfo: San Vicente, 12 Sep 1910, Britton et al. 7319
scribedfrom the CaribbeanIslands,of which Morelet
(NY); Herradura,12 Feb 1906, Hermann 2653 (NY, US);
Vinales, between Pinar del Rio and San Vicente, 13 Jul said that it occurs on Hispaniola("Saint-Domingue")
1968, Lamb s.n. (FHO); Galalon, 15 Jul 1968, Lamb s.n. and E Cuba (Sierra de Cobre).
(FHO); Sandino, 30 Jun 1990, Maas et al. 7750 (U); near
Grisebach(1866) describeda (tentative)varietyof
city of Pinar del Rio, 19 May 1951, Rutten-Pekelharing P cubensis as var. terthrocarpa,a name coined by
431 (U); Herradura,30 Apr 1903, Shafer 425 (K, NY);
C. Wrightat species rank on the label of his collecLaguna el Jovero, 12 Dec 1911, Shafer 10905 (NY, U,
16
Dec
11017
el
1911, Shafer
(NY, tion no. 3189. Wrightonly mentioned "Cuba"as its
US); Laguna Bufeo,
U, US); Herradura,9 Feb 1903, Shaw s.n. (K, MO, P, US); origin, but apparentlyit is from easternCubabecause
21 km N of Pinar del Rio, 18 Jun 1955, Schultes et al. 4
P cubensis is restricted to E Cuba. Shaw (1903),
(NY). WITHOUTLOCALITY:1856, Christy s.n. (K); 12
the GH duplicate of Wright3189 and other
citing
s.n.
Clinton-Baker
(K).
May 1911,
collections, took up the specific epithetfromWright's
label and published it as a new species. He mainUses. Pinus tropicalis is an important regional
source of timber mainly used by local sawmills. Its tained this at first (Shaw, 1904a), but later (Shaw,
wood is dense and durable but resinous. Despite
1914) synonymized it underP. tropicalis. Engelmann
extensive exploitation, it is still common. Regenera(1880a), who, like others, seems not to have known
of Morelet's repeated publication (Morelet, 1851,
tion seems to be good in many areas, especially
where it is protected from grazing pressures. It is
1855) of two new Cuban pine species, mentioned
type. Phenology: Time of pollen dispersal not recorded; presumably later than P caribaea.
80
Flora Neotropica
Pinus murrca'anaBalfour in Murray,Bot. Exped. Oregon
8: 2. 1853; Pinus contorta D. Douglas ex J. C. Loudon
subsp. murravana (Balfour) W. B. Critchfield, Publ.
Maria Moors Cabot Found. Bot. Res. 3: 106. 1957.
Type. United States. California: Siskiyou Mts., s.d.
[1852]. J. Jeffrey 740 (holotype, E; isotypes, Fl, K).
s~..
\..
Tree, in Mexico of medium size, occasionally tall,

height to 25-30(-33) m, dbh to 100-150(-200) cm.
Trunkmonopodial, erect, massive, some trees may
be forked higher up, scrubby on summit ridges,
slender in dense stands. Barkthin, scaly, with numerous small, flaking plates, colour variable from light
orange- or pink-brownto greyish brown, on young
trees thin, papery. Branches of first order relatively
short, spreading to ascending; branches of higher
orders slender and flexible, spreading or ascending,
forming a narrowly conical or pyramidal,dense or
more open crown. Shoots multinodal (especially
leading shoots), lateral shoots on older trees mostly
uninodal, rough, ridged between bases of leaf fascicles, more or less glaucous; pulvini persistent.Cataphylls small, ligulate, curved, with erose margins,
early deciduous. Vegetative buds ovoid-conical,
Map 3. Pinus contorta var. murravana.
Wright's name as published by Grisebach and, apparently thinking that its "grass stage" was the reason for
its distinction, which he considered taxonomically
insignificant, dismissed it.
Previous classifications (e.g., Little & Critchfield,
1969; Farjon, 1984) group P tropicalis (and P resinosa of the eastern United States and Canada) with
the predominantly Eurasian subsect. Sylvestres on
grounds of morphological features of the seed cones
and leaf numbers. A phylogenetic approach using all
available information, preferably including molecular data, should be used to determine if P. tropicalis
indeed belongs in this rather homogeneous subsection. Unfortunately, Wang and Szmidt (1993) did
not incorporate this species in their cpDNA-based
phylogeny assessing many of the purportedly related
Asian species.
3. Pinus contorta D. Douglas ex J. C. Loudon var.
murrayana (Balfour) Engelmann in Watson, Bot.
California 2: 126. 1880.
Fig. 18
the
terminal buds ca. 15 x 6 mm, the laterals smaller,

resinous; the scales imbricate, appressed, acute, orange-brown. Fascicle sheaths short, 5-8 mm long,
reduced to ca. 5 mm in maturefascicles, persistent,
light reddish brown with hyaline-whitish margins,
weathering grey. Leaves in fascicles of 2, in dense
tufts spreading obliquely from the shoot, persisting
5-8 years, rigid, usually curved and contorted, 4-7
cm long (variable on a single branch, decreasing
with altitude), 1.2-2 mm wide, entire or sparsely
serrulate,acute-acuminate,pungent,light (yellowish)
green. Stomata on all faces of leaves, in 10-14
grooved lines on both the convex abaxial and the
slightly concave adaxial face, conspicuous. Leaf
anatomy:Cross section semi-circularto semi-elliptic;
cuticula and epidermis thick; hypodermis uniform,
2-layered; resin ducts (1-)2(-?), medial toward the
leaf margins;stele elliptic in cross section, with outer
cell walls of endodermisthickened;vascularbundles
2, distinctly separate.Pollen cones crowded near the
proximal end of a new shoot, subtended by 5-8
subulate bracts, ovoid-oblong to cylindrical, 8-14
mm long, yellowish to yellowish orange. Microsporophylls ca. I mm diam. with a radiallystriateabaxial
surface; pollen slightly largerthan in other varieties.
Seed cones subterminal,usually in pairs or whorls
of 3-5, on short, recurvedpeduncles, persisting only
1-3 years after release of seeds. Immature cones
ovoid, with prominent,erect spines, ca. 10 mm long,
reddish purple, maturing in two seasons. Mature
81
A
FIG. 18. Pinus contorta var. inurravana (A-C, E-G, Hughes & Styles 174; D, Jeffrey 740). A. Branch with leaf
fascicles and ovuliferous cone. B. Leaf fascicle. C. Cross section of leaf. D. Ovuliferous cones, closed. E. Ovuliferous
cones, opened, with lateralview of seed scale. F. Seed scale (two views). G. Seeds, with articulatewings. (Magnifications:
A, B, D-G, x 0.5; C, x20.)
cones (narrowly) ovoid when closed, symmetrical,

broadlyovoid with a more or less oblique base when
opened,(3-)4-5.5 x 3-4 cm when open. Seed scales
ca. 90-1 10, partinggraduallyafter cone maturation,
oblong, straight or recurved, thin woody, dark purplish brown on the abaxial side, dark brown with
light brown marksof seed wings on the adaxial side.

Apophysismore or less symmetricalaroundthe cone,
slightly raised, transversely keeled, somewhat gibbous on the basal scales, irregularlypentagonal in
outline, ochraceous,lustrous.Umblodorsal, rhombicpyramidal, 2-3 X 3 mm, with a prominent. thin,
Flora Neotropica
82
curved and persistentprickle, greyish brown. Seeds
obliquely ovoid, 4-5 mm long, greyish brown. Seed
wings articulate but effective, held to the seed by
two oblique, thin claws, obliquely ovate, 8-12 X 45 mm, yellowish brown, semi-transparent.Cotyledons 4-5 (number on average higher by I than in
other varieties).
Distribution and ecology (Map 3). United States:
Cascade Range of SW Washingtonand W Oregon;
southward in the Sierra Nevada of California and
extreme W Nevada, disjunct in high mountainsof S
California.Mexico: in Baja CaliforniaNorte, Sierra
San Pedro Martir.In southernCaliforniaand Mexico
it occurs in the upper zone of the Mixed Conifer
Forest, with Abies concolor and Pinusjeffreyi as the
most common conifers, and also Pinus lambertiana
and Populus tremuloides on deeper soil and more
mesic sites, respectively.The SierraSan PedroMartir
is a graniticbatholithwith a steep easternescarpment
and extensive western plateaus.The altitudinalrange
of Pinus contorta var. murravana is here 2300-
3000+ m; there is no climatic tree line, but due to

steepness of the highest summit, Picacho del Diablo
(3095), only a few erraticsaplings of Abies concolor
occur there (Minnich, 1987). Annual precipitationis
a moderate500 mm, of which 50% may be snowfall.
Frosts are common during the winter months. Phenology: Pollen is dispersed during May-June.
Representative
specimensexamined.MEXICO.BAJA
Sierra San Pedro Martir, at
CALIFORNIA NORTE:
"Vallecitos," 29 Nov 1987, Franklin & Chandler 5673
(NY), 26 Jan 1972,HallerJr. 3427 (UC), 25 May 1987,

Hughes & Styles 174 (FHO), Sep 1961, Lindsay s.n. (SD),
Sep 1940, M. Martinez 3464 (INIF. MEXU), 2 Jun 1968,
Moran15076(MEXU,SD);SierraSanPedroMartir,near
3 May 1987.TenorioL. & RomeroT 13296
Observatory,
(MEXU).
yet recognized as a distinct species, it was later

described by Loudon (1838) from a collection made
by David Douglas in 1825 near the mouth of the
Columbia River in Washington, U.S.A. (Mirov,
1954). The specimen at K! (D. Douglas s.n., ex herb.
Bentham)-annotated by Douglas as "P contorta
Sabine. ...
In swampy ground near the sea coast.
Abundantly near Cape Disappointment and Cape

Lookout. 1825.6.7."-is here designatedas the lectotype of Pinus contorta Douglas ex Loudon.
Based on collections sent by J. Jeffrey under
no. 740 to Edinburgh, Balfour (in Murray, 1853)
describedP murrayanafrom the Siskiyou Mountains
in southernOregon or northernCalifornia,accompanied by a figure done by Greville (who is often
erroneouslycited as one of the authors;in fact, the
new species are ascribed by Murrayto the "Oregon
Committee"of which Balfour was chair).
Engelmann(1880b) reducedthis species to a variety of P contorta (in which he included P contorta
var. latifolia Engelm. from the Rocky Mountains).
Apart from stressing the taller habit (especially in
the Sierra Nevada, where most conifers are indeed
taller), he describedthe longer and wider leaves and
the less oblique, often opening and deciduous cones
as the differentiatingcharactersagainst P contorta
s.str., which he confined to the Pacific coast. Critchfield (1957) changed the rank of this and other
infraspecifictaxa to subspecies, based on allopatric
distributionand growth performancerelatedto geographyand climate ("provenance").Many of his arguments (Critchfield,1957, 1963, 1980) are of ecological consequence, rather than taxonomic, and the
morphologicaldifferencesappearto be clinal in areas
where these taxa meet. Some of them, such as nonserotiny of cones ascribed to var. murrayvaa, also
occur in certain individualsof populationsbelonging
to other infraspecifictaxa. Distinctionat varietalrank
seems the most appropriatetaxonomic approach in
such cases.
Uses. Pinus contorta is one of the most important

timber trees of North America (Lotan & Critchfield,
1990), but since it is in Mexico restrictedto the high
plateaus and ridges of the Sierra San Pedro Martir,
it is of local importancethere. Much of this disjunct
4. Pinus herrerae Martfnez,Anales Inst. Biol. Univ.
population, including the best-developed stands
Nac. Mexico 11: 76. 1940.
Fig. 19
aroundhigh-montanemeadows such as "La EncanPedro
National
Martir
tada,"is now in the SierraSan
PitnusteocoteSchiedeex Schlechtendal
& Chamissovar.
Park and enjoys protection at least from lumber
herrerae(Martinez)Silba, Phytologia68: 63. 1990.
harvesting.
Type.Mexico.Jalisco:SierraLalo,Feb 1940,M.Martnez 3427 (lectotype, MEXU; isolectotypes, F, US).
Pinus contorta is highly variable genetically and
morphologically throughout its vast range, and is
Tree, usually tall, height to 30-35 m, dbh to 75only second in geographic extent to P sylvestris
As
a
several
forms
100
cm. Trunkmonopodial, straight or sometimes
(Critchfield,1980).
consequence,
of different regions have been described as species. tortuous,terete, often with a branch-freebole 2/3 or
Although first collected in Alaska (at Sitka), but not /4 its length. Bark thick, rough, with scaly plates
83
B3B
FIG. 19. Pinus herrerae (A-C, Stles 69; D-G, Hughes & St'les 137). A. Branchwith leaf fascicles and pollen cones.
B. Leaf fascicle. C. Cross section of leaf, with enlarged endodermal cells showing thick outer walls. D. Immature
ovuliferous cones. E. Matureovuliferous cones. F. Seed scale (two views). G. Seeds, with articulatewings. (Magnifications: A, B, D-G, x0.5; C, x60.)
84
and shallow, longitudinalfissures, reddish brown to
grey-brown, in young trees and on branches thin,
exfoliating in thin flakes, reddish brown. Branches
long, slender, arching or spreadinghorizontally,the
higher-orderbranches flexible, slightly pendulous.
Shoots smooth, with well-developed, long decurrent
pulvini, orange-brown.Cataphyllssubulate,scarious,
recurving,brown, weakly ciliate-hyaline at margins,
early deciduous. Vegetative buds ovoid-acute to
ovoid-oblong or cylindrical, the terminalbud 10-15
mm long, the lateralssmaller,not resinous;the scales
subulate,reflexed, weakly ciliate-hyalineat margins.
Fascicle sheaths of flushing leaves up to 20 mm
long, torn by elongating and spreading leaves; the
scales chartaceous,yellowish white; sheaths persistent but reduced to 8-15 mm in mature fascicles,
light brown to grey-brown.Leaves in fascicles of 3,
in open or dense tufts persisting 3 years, slender,
lax, drooping or spreading,(10-)15-20 cm X 0.70.9 mm, serrulate at margins, acute, light green to
yellowish green. Stomata on all faces of the leaves,
in 6-8 lines on the abaxial face and ca. 3 lines
on each adaxial face. Leaf anatomy: Cross section
transverse-triangular,with a convex abaxial side;
hypodermis2-3-layered, or thickerin marginalareas;
resin ducts 1-3(-4), internal,rarelyseptal; stele oval
in cross section; cells of endodermiswith thickened
outer walls; vascular bundles 2, (widely) separated.
Pollen cones crowded along the proximal end of a
new shoot, subtended by short bracts, spreading,
ovoid-oblong to cylindrical, 15-18 x 5 mm, yellowish green, tinged red, turninglight brown.Seed cones
subterminal,solitary or opposite, rarely in whorls of
3, on 10-15 mm long peduncles, at first erect, soon
on recurved peduncles which fall with the cones.
Immaturecones (broadly) ovoid, 8-13 mm long,
purplish red, maturing in two years. Mature cones
narrowlyovoid when closed, nearly symmetricalor
slightly curved, (broadly) ovoid when opened, (2-)
3-3.5(-4) x 2-3.5 cm when open. Seed scales ca.
50-80, partingexcept those at the base of the cone,
thick woody, oblong, straight or recurved, on the
abaxial side purplish, greyish at base, with light
brown marksof seed wings and a darkerbrowndistal
part adaxially. Apophysis slightly raised, mostly so
on a few proximal scales on one side of the cone,
transverselykeeled, the apical marginentire or crenate, radiallystriate,light brown. Unbo dorsal,pyramidal, irregularlyrhombicin outline, 3-4 mm wide,
with a minute,deciduous prickle,greyish. Seeds (ob)ovoid, slightly flattened, 2.5-4 X 2-3 mm, dark
grey-brown.Seed wings articulate,effective, held to
the seed by two small, thin claws, obliquely ovate,
Flora Neotropica
5-8 x 3-5 mm, translucent, yellowish with a dark
tinge. Number of cotyledons not observed.
Distribution and ecology (Map 4). Mainly in
disjunct populations in the west and south of Mexico,
along the Sierra Madre Occidental and more abundantly along the Sierra Madre del Sur; in SW Chihuahua, Sinaloa, Durango, W and S Jalisco, Michoacan,
and Guerrero. Its altitudinal range is (1100-) 15002600 m; its lowest altitude apparently is reached in
the Sierra de Cuale in W Jalisco. It occurs in the
mesic forest belt, with annual precipitation of ca.
900-1600 mm but with a dry season lasting from
November to May. From N to S annual precipitation
increases. Mixed pine and pine-oak forest is its usual
habitat, in which it is associated with many other
pines according to their ranges, further with Pseudotsuga locally. Other broad-leaved trees-e.g., Arbutus, Alnus, Clethra, Juglans, Persea, Clusia, and
Tilia-are sometimes common. In many areas the
forests are greatly influenced by man-made changes
in their composition, e.g., by selective felling of
larger trees of Pinus or by recurrent fires and grazing.
Phenology: Pollen dispersal has been observed in
March in Jalisco and Michoacan.
Cascada Basaseachic National Park, I km S of
the falls, 12 Nov 1989, Spellenberg et al. 10086 (FHO,
ChavarriaNuevo, 3 Apr 1991,
MEXU, NMC). DURANGO:
4 km
Styles, Favela et al. 6 (FHO, MEXU). GUERRERO:
SW of Puerto El Jiguero, 25 Oct 1987, Aquino 153 (FHO,
MEXU); 3 km NE of Cruz de Ocote, 23 Mar 1985, Lorea
3483 (FHO); Sierra Madre del Sur, Las Lumbreras,3 Jan
1938. Mexial9088 (B, K, MO, NY, S, UC); Omiltemi, Las
Trinceras,2 km N of Pablado, 6 Apr 1985, Millan Espin
582 (FHO, MEXU); Carrizalde Bravo, 8 Feb 1981, Stead,
Styles et al. 731 (FHO). JALISCO:
Loc. not stated, det. by
M. Martfnezas "P herreraisp. nov.," 1938-1939, collector
not specified, (MEXU No. 2526); SierraManantlin, 16 Apr
1987, Hughes & Styles 137 (FHO); Pihuamo, Mar 1939,
M. Martinez3599 (NY); Sierra de Cuale, along rd. to the
mines E of El Tuito, 16-19 Feb 1975, McVaugh 26346
(MICH);Tecatitlan,64 km from Ciudad Guzmain,13 Mar
1974, Styles 67 (FHO, K), 13 Mar 1974, Styles 68, 69
Dos Aguas, 24 Sep 1960, Courbas(FHO). MICHOACAN:
son & Mahieux s.n. (P); ca. 3.5 km S of Paracho de
Verduzco,at km 35 along Hwy. 37, 6 May 1994, Farjon &
Mej'a 351 (E, FHO, HEH, K, MEXU, U); Dos Aguas,
along rd. to Coalcoman, 11 Apr 1985, Soto N. et al. 8144
(EAP, NY); Dos Aguas, forest station W of Aguililla, 2
Mar 1974, Styles 41 (FHO). SINALOA:
Mun. Rosario, ca. 6
km W of El Palmitoalong Hwy. 40, 31 Jan 1962. Breedlove
1704 (CAS); Mun. Badiraguato,5 km N of Los Ornos
along rd. to Ocurahui, 2 Oct 1970, Breedlove & Thorne
18361 (CAS); Los Loberos,along main rd. between Mazatlin and Durango,30 Apr 1987, Hughes 970 (FHO, MEXU).
Uses. Along with other tall growing pines, it is
(heavily) exploited for timber throughout almost all
85
.I
;'~~
''
1..
~ ~ ~ ~ ~ .8
S..,.
""
-?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
*
..,...
iiii
....
ii
Map
4.Pinushe~~~~~~;rrre(ice)P.pedsrbsvraplesstings.
a; .
?" Z'
p4 .
....
of its range. It is considered to be of good wood

quality.Provenancesampling has been startedand is
recommended by foresters (Dvorak & Donahue,
1992) to be conducted on a largerscale.
This species was, according to Martinez (1940),
first collected in Jalisco in 1931 and again in 19381939 as P. teocote and P.patula, respectively.Critchfield and Little (1966) did not recognize it and
mappedit with P. teocote. Pinus herrerae is in many
characterssimilar to P teocote, yet distinct in a few
constant characters, most notably the very slender,
lax, and (slightly) longer leaves and the smaller
ovuliferouscones which are distinctly narrowerwhen
still closed. Other differentiatingcharactersare the
resinducts in the leaves, which are fewer and internal
in P. herrerae and medial in P teocote, and the less
ciliate cataphylls (margins hyaline) of the former
species. Perry(1991) noted virtuallythe same differences between the two taxa. Molecular evidence
(cpDNA) may providenew, non-morphologicalcharacters to more firmly establish (or refute) species
delimitations, but, apart from P patula (Perez de la
Rosa et al., 1995), no molecular data are as yet
available for these species.
5. Pinus caribaea Morelet, Rev. Hort. C6te d'Or 1:
107. 1851.
Tree, medium to tall, height to 20-35(-45) m, dbh
to 50-100 cm. Trunk monopodial, erect, slender,
terete. Bark rough, scaly, on lower part of trunk
breaking into irregularly square plates divided by
shallow or deep fissures (in part influenced by fire
damage, see Hunt, 1962), grey-brown, inner bark
86
reddish brown. Branches of first order slender,
spreadingor ascending;of second and higher orders
similar, or drooping. Crown broad conical, open or
Flora Neotropica
in outline, transverselykeeled, to 15 mm wide, ochraceous, light or dark brown, lustrous. Umnbodorsal,

flat or slightly raised or pyramidaland curved upfor
2/3
or
-/4
of
tree
3-4 mm wide, up to 3 mm high, with a minute,
a
clear
bole
ward,
usually
irregular;
height. Shoots multinodal,very rough, resinous;with persistentprickle,dull brown,weatheringgrey. Seeds
large, short decurrentand persistent pulvini. Cata- obliquely (narrowly)obovoid, slightly flattened,obphylls large, 15-20 mm long, subulate, strongly re- tuse or angular, 5-7 x 2.5-3.5 mm, light greycurved, scarious, with hyaline-ciliate margins, dark brown with dark spots, or dark brown to blackish.
brown, apex caudate, persisting some time after the Seed wings articulateor adnate,effective, held to the
leaf fascicles have fallen. Vegetative buds ovoid- seed by two claws or more commonly enveloping
oblong to cylindrical, 20-25 mm long, the laterals the seed on one side (adnate), obliquely ovate or
smaller, ovoid-acute, (slightly) resinous; scales oblong, 10-20 X 5-8 mm, thin, semi-transparent,
spreadingor recurved,scarious, with ciliate margins, yellowish grey or light brown with dark stripes.
brown. Fascicle sheaths 15-20 mm long, retaining Cotyledons(4-)5-8(-9), (12-)20-35 mm long. Seedtheir length or reducedto ca. 10 mm at maturity;the lings with an elongated stem, primaryleaves green
scales papery, with erose-ciliate margins, yellowish or glaucous, more or less ascending or spreading,
brown,turningdarkred-brownor grey-black.Leaves soon replacedby secondaryleaves or these delayed.
Distribution
in fascicles of (2-)3(-4, very rarely 5) in dense,
(Map 5). Caribbean: Bahamas,
irregularlyspaced tufts, persisting3 years, spreading, Turks-CaicosIslands, W Cuba [including Isla de la
Juventud(Isla de Pinos)]. Mesoamerica:Mexico (S
straight, (slightly) twisted, rigid, (12-)15-26(-28)
cm X (1.2-)1.4-1.8 mm, margins serrulate, apex QuintanaRoo), N Guatemala,Belize, Honduras(incl.
acute-pungent,light or darkgreen (occasionallyglau- Islas de la Bahia), Nicaragua.
cous-green), moreor less lustrous.Stomataconspicuous on all faces of leaves, in 8-11(-14) lines on the
Key to the varieties of Pinus caribaea
convex abaxial face and 4-6 lines on each adaxial
in fascicles of 2-3, mostly 3, fascicle
Leaves
i.
face. Leaf anatomy:Cross section transverse-triangusheathsreducedin lengthat maturityfrom15-20
lar; hypodermis multi-layered (3-5 rows of cells);
mm to 10 mm ................................. 5b. var. hlhamensis
resin ducts 2-5(-8), internal,rarely 1-2 medial;stele I. Leaves in fascicles of 3-4, mostly 3, very rarely2
or 5, fascicle sheaths persistently long, ca. 15-20
elliptic in cross section; outer cell walls of endomm.
dermis not thickened; vascular bundles 2, distinctly
2. Seedlings with green, ascendingprimaryleaves,
clustered
near
Pollen
cones
the
proximal
separate.
in a few years replaced by secondary leaves....
end of a new shoot, sometimes more spreading by
..........................................................5a. var. caribaea
multinodal habit, subtended by imbricate, scarious,
2. Seedlings with glaucous, spreading primary
leaves, development of secondary leaves dereddishbrown bracts,cylindrical,often curved when
layed ...........................................5c. var. hondurensis
mature, 20-30 x 5-6 mm, pink or yellow, turning
yellowish or reddishbrown.Seed cones subterminal,
often in several whorls (nodes) in one season, mostly 5a. Pinus caribaea Morelet var. caribaea. Type.
Cuba. Isla de la Juventud [Isla de Pinos]: Santa
in pairs or whorls of 3-5(-8), on 2-2.5 cm long,
Barbara,Jun 1957, Luckhoff11608 (neotype, PRF
deciduous
curved,
peduncles (sometimes breaking
Fig. 20A-F
designated by Luckhoff, 1964).
off at the base); immaturecones ovoid to narrowly
Pinus recurvataRowlee, Bull. Torrey Bot. Club 30: 107.
ovoid, 15-20 x 10-15 mm with pyramidal or
1903. Type. Cuba. Isla de la Juventud [Isla de Pinos]:
curved, acuminateumbos terminatingin an erect or
Santa Rosalie, Camp Columbia. 29 Dec 1901, Rowlee
incurvatespine, ochraceous, lustrous, maturingtime
232 (holotype, CU).
two seasons. Maturecones pedunculate,ovoid-coniPinus cubensis Grisebach var. anolmala Rowlee, Bull.
cal or ovoid when opened, more or less symmetrical
TorreyBot. Club 30: 108. 1903. Type. Cuba. Isla de la
Juventud[Isla de Pinos]:JucaroLanding,Dec 1901,
but the base obliquely flattened, (4-)5-12(-13) x
Rowlee 233 (holotype, CU).
(3-)4-6(-7) cm when open. Seed scales ca. 120200, partingto release the seeds except at the base,
Tree,medium to tall, height to 20-35(-45) m, dbh
thin or thick, oblong, straightor recurved,dark pur- to 60-100 cm. Fascicle sheaths 15-20 mm long,
plish or blackish brown on the abaxial side, dark retainingtheir length at maturity;the scales papery,
brown with faintly visible marks of seed wings on with erose-ciliate margins,yellowish brown, turning
the adaxialside. Apophysisslightly or stronglyraised dark red-brownor grey-black.Leaves in fascicles of
or nearly flat on basal scales, rhombicto pentagonal 3(-4), very rarely 2 or 5, spreading, straight,
|| e
87
?X A
...........
^...I..............
......
Map 5. Pinus carib(ea
....
..
var. caribaea (squares); P. caribaea var. bahatwensis (circles): P caribaea var. hotndurensis
(triangles);P. elliottii var. clensa (stars).
(slightly) twisted, rigid, (13-)15-26 cm X (1.2-)

1.4-1.8 mm, light or dark green, more or less lustrous. Stomata on all faces of leaves, in 8-11 lines
on the convex abaxial face and 4-6 lines on each
adaxial face. Seed cones (4-)5-10(-12) x (3-)46(-7) cm when open. Seed scales ca. 120-170, more
or less thin woody. Apophysis slightly raised or
nearly flat on basal scales, rhombicto pentagonalin
outline, ochraceous or light brown, lustrous. Umbo
dorsal, flat or slightly raised. Cotyledons (4-)6-7
(-9), 12-25 mm long. Seedlings with an elongated
stem, primaryleaves green, more or less ascending,
soon replacedby secondary leaves.
Distribution and ecology. W Cuba: Pinardel Rio,
Isla de la Juventud (Isla de Pinos). Forming pure,
open, dry fire-climax forest or open woodland with
undergrowthof grasses or scatteredshrubson sandy
or gravelly, well-drained, acidic soils. Altitudinal
range from 1-700 m, with most extensive stands
below 400 m. The growing is season continuous in

a warm tropical climate with long dry spells. Annual
precipitation varies mainly with altitude, ca. 10001800 mm, with a winter dry season. Frost does not
occur. Phenology: Pollen dispersal is in January and
February.
Representative specimens examined. CUBA. ISLADE
LA JUVENTUD[ISLA DE PINOS]: Lanier, 20 Nov 1920,
Ekman12279 (S): Nueva Gerona,22 Jan 1951. Little 14047
(US). PINARDELRio: San Diego de los Bahos. 31 Aug
1910, Brittonet al. 6766 (NY); Bahia Honda,9 Mar 1920.
Ekiman10440 (NY. S); Cortez. 27 Mar 1921, Ekman12802
(S); Topes de Collantes, Las Villas, 12 Jun 1968, Lambs.n.
(FHO); San Diego de los Bafios, at El Luiar, 14 Apr 1904,
Le6n 4250 (NY); Guane, 28 Dec 1911. Shrfer 11209 (NY,
U, US); Los Palacios. 10 Jan 1912, Shafer 11697 (B, NY.
US); Pinal de la Catalina, 18 Jan 1912, Shafer 11867 (NY,
U); San Marcos, 27 Jan 1912, Shafer 11968 (K. NY. US).
WESTERNCUBA:Loc. unknown, 1860, Wright 1462 (S);
loc. unknown, 1863, Wright1462B (GOET, K).
88
Flora Neotropica
:i@i
,I
I,
t
tzyI
FIG. 20. A-F. Piols carihaea var. carilbea (A-C. Lmnhb

34; D-F, Lal)mbs..). G-I. P. caribhte var. bahal ensis (GH. Brlis 15). J-L. P caribhea var. hiondurensis(Farjon & D. Meji4 356). A. Branch with leaf fascicles and immature
cones. B, G. Leaf fascicles. C, H, J. Cross sections of leaves. D, K. Ovuliferous cones. E. Seed scale (three views). F.
Seeds with adnate wings. I, L. Seeds with articulateand adnate wings. (Magnifications:A, B, D. F, G. 1, K. L. XO.5; C.
H. J. x30: E. x I.)
89
SystematicTreatment
Remarks on the species. Morelet (1851) described this species as one of two new pines from
the Isla de la Juventud, Cuba, with rather small
(ca. 6 cm) cones but otherwise giving an adequate
descriptionof its characters.Although Morelet'sCuban collections were deposited at P, no original
materialof P. caribaea could be found there. Luckhoff (1964) designateda neotype from SantaBarbara
on Isla de la Juventud.
Probablydue to the obscurityof Morelet'spublication in regional horticultural(1851) and naturalhistory (1855) periodicals,the name Pittuscaribaea has
been erroneouslytaken up for the slash pine of the
SE United States, P. elliottii Engelm. (e.g., de Vail,
1941), a closely related, but nevertheless distinct
species. Subsequently,Loock (1950) compared the
Mesoamerican(Belize) populations of P. caribaea
with "P. caribaea" from the United States, not from
Cuba or Isla de la Juventud.He concluded that the
pines in Belize "show . . . some distinct botanical
wings more frequentlyarticulate,also adnate.Cotyledons (4-)6-7(-9), 12-25 mm long. Seedlings with

an elongated stem, primary leaves green, more or
less ascending, soon replacedby secondary leaves.
Distribution and ecology. Bahamas: on Grand
Bahama, Great Abaco, New Providence, and North
and South Andros Islands in the north; on Great
InaguaIslandin the south.Also on the Turks-Caicos
Islands. On flat, eroded limestone rock with pockets
of sandy soil (pH >7), on sandy spits and old
beaches, forming pure, open fire-climaxstands, usually with scatteredor dense undergrowthof shrubs
(often Sabal palnetto); also invadingopen scrubland
forming secondaryforest. Altitudinalrange 1-10 m.
Annual precipitationdecreases from ca. 1500 mm
on GrandBahamato ca. 1000 mm on Andros in the
northernBahamas, mean annual temperatureis ca.
25?C and frost does not occur. Hurricanes are a
recurringdestructive factor. Phenology: Pollen dispersal occurs from Januaryto March.
Representative
specimensexamined.BAHAMAS.AN-
differences, which are considered importantenough

to warrantthe separationof the Belize plants from DROS:San Andros,25 Aug 1988, Baylis 20 (FHO); Duncan
P. caribaea, Morelet, under the name, P. hondur- Point, 30 Aug 1988, Baylis 21 (FHO); Conch Sound. 22
ensis." One can hold the view, as Shaw (1914) did, May 1890, Northrop& Northrop84A (NY); Lisbon Creek,
16 Jan 1910, Small & Carter 8497 (K, NY, US); Nicholl's
that there is just one species incorporatingpopula- Town, 4 Feb 1910, Small & Carter 8973 (K. NY, US).
tions from SE United States to Mesoamerica, but GRAND BAHAMA: McLean's Town, 16 Aug 1988. Baylis
such a broadconcept is not now generally accepted. 10 (FHO); Little HarbourKey, 20 Aug 1988. Baylis 14
The pines from Isla de la Juventud, for lack of a (FHO); Freeport, 19 Aug 1988, Baylis 15 (FHO): Garden
Cay, Neat End, 16 Apr 1905. Brace 3670 (NY). GREAT
holotype specimen, are as near as we can get to a ABACO:Sandy Point, 1 Aug 1988, Baylis 6 (FHO);Cedar
referencepoint for Morelet's Pinus caribaea.
Harbour,12 Aug 1988, Baylis 7 (FHO);MarshHarbour,11
Dec 1904, Brace 1634 (NY). NEWPROVIDENCE:Nassau.6
5b. Pinus caribaea Morelet var. bahamensis Sep 1988, Baylis 27 (FHO); Blue Hills, 7 Apr 1904. N. L.
(Grisebach) W. H. Barrett & Golfari, Caribbean Britton25 (NY); loc. unknown, 12 Mar 1907. E. G. Britton
6589 (NY, US); loc. unknown.22 Feb 1888, Eggers 4172
Forest. 23: 69. 1962.
Fig. 20G-I
(E, M, US).
Pinusbahtimensis
TURKS-CAICOS ISLANDS. Pine Cay. 25 Aug 1975,
Grisebach,Fl. Brit.W.I.:503. 1864.
Feb 1978,
20 Nov 1960,Golfari Correll43153 (NY); GrandCaicos,Banberra.
NewProvidence,
Type.Bahamas.
77571(neotype,designated
by Barrett& Golfari,1962, Correll 49463 (NY); Conch Bar. E of Conch Bar airstrip,
14 Jun 1974, Gillis & Proctor 12301 (FHO): Pine Cay. 17
BAB).
Dec 1907. Wilson7694 (K, NY).
Tree, medium size, height to 20 m, dbh to 50 cm.

Fascicle sheaths 15-20 mm long, reduced to ca. 10 5c. Pinus caribaea Morelet var. hondurensis (Sencm at maturity; the scales imbricate, with eroseeclauze) W. H. Barrett& Golfari,CaribbeanForest.
ciliate margins, dark red-brownturning grey-black.
23: 65. 1962.
Fig. 20J-L
Leaves in fascicles of 3(-2), on GrandBahama2-3,
Pinushondurensis
S6enclauze, Conif. 126. 1868.Type.
spreading,straight,(slightly) twisted, rigid, (13-)15Belize.El Cayo,5 Nov 1960,Barrett77582(neotype.
or
dark green,
26 cm X (1.2-)1.4-1.8 mm, light
designatedby Barrett& Golfari,1962,BAB).
more or less lustrous.Stomataon all faces of leaves,
PinushonthrensisLoock,[PinesMexicoBrit.Honduras]
S. Afr. Dept. Forest.Bull. No. 35: 210. 1950, non
in 8-11 lines on the convex abaxial face and 4-6
Seneclauze.Type.Belize:StannCreek,Loock10280
lines on each adaxial face. Seed conles (4-)5-10
(holotype,PRF).
(-12) x (3-)4-6(-7) cm when open. Seed scales ca.
Tree,mediumto tall, height to 20-35(-40) m, dbh
120-170, moreor less thin woody. Apophvsisslightly
raised or nearly flat on basal scales, rhombic to to 60-100 cm. Fascicle sheaths 15-20 mm long,
pentagonal in outline, ochraceous or light brown, retainingtheir length at maturity;the scales papery,
lustrous. Umbo dorsal, flat or slightly raised. Seed with erose-ciliate margins,yellowish brown, turning
90
dark red-brownor grey-black.Leaves in fascicles of
3 (rarely 2, 4, very rarely 5), spreading, straight,
(slightly) twisted, rigid, (12-)16-28 cm x (1.2-)
1.4-1.8 mm, light green, occasionally glaucousgreen, more or less lustrous.Stomata on all faces of
leaves, in 8-14 lines on the convex abaxial face and
4-6 lines on each adaxial face. Seed cones (4-)512(-13) X 3.5-7 cm when open. Seed scales ca.
120-170, moreor less thin woody.Apophysisslightly
or strongly raised, but less so on basal scales, rhombic to pentagonalin outline with uppermarginirregularly undulate,chestnut-brown,lustrous. Umbo dorsal, slightly raised or curved upward,up to 3 X 34 mm. Cotyledons 5-8, 20-35 mm long. Seedlings
with an elongated stem, primary leaves glaucous,
more or less spreading, development of secondary
leaves delayed.
Distribution and ecology. Mexico (S Quintana
Roo), Belize, N Guatemala,Honduras(includingthe
Islas de la Bahia), Nicaragua.Mainly distributedon
the lowlandcoastal plains withinthe Atlanticclimatic
influence, from the edges of mangrove swamps to
lower upland "bunch-grass/pinesavannas."In Nicaragua it is the southernmostpine in the Americas. It
occurs on well-drained, sandy or gravelly, acidic
soils, forming pure (or sometimes mixed with P.
oocarpa and/or P tecunumanii) fire-climax forest,
or pine-oak forest, with undergrowthof grasses,
Pteridium aquilinum, or Sabal palmetto nearer the
coast, possibly replacing broad-leaved rain forest
underhumaninfluencein many areas.The altitudinal
range is 1-700(-1000?) m. Annual precipitationvaries greatly, from the highest measurement,ca. 4000
mm, at Lagunadel Pinarin Nicaraguato the lowest,
ca. 660 mm, at Los Limones in the interior of
Honduras; it generally increases from the interior
valleys to the coastal plain and from north to south
along the coast. The dry winter season decreases in
durationin a similar pattern.Mean annual temperature is 21-27?C and no frosts occur. Phenology:
Pollen is shed in the early part of the dry season.
Styles et al. (1982) cited reportsof pollen dispersal
in November and Januaryin different areas.
Flora Neotropica
along rd. to El Volcan, 14 Mar 1972, Styles & Kemp 104
Just E of El Lobo, 8 Mar 1978, Stead &
(FHO). ZACAPA:
Styles 234 (EAP, FHO, K, M), Stead & Styles 235 (FHO).
BELIZE. BELIZE:Stoney Camp, II Aug 1929, Lundell
677 (F, MO, NY, S, US), Lundell678 (K, NY, US). CAYO:
Mountain Pine Ridge, Augustine, 18 Dec 1959, Hunt 62
(BM, P); MountainPine Ridge, Cooma Cairn, 19 Jul 1966,
Lamb 32 (FHO), 25 Jul 1966, Lamb 34 (FHO); Mountain
Pine Ridge, ThousandFoot Falls, 5 Nov 1960, Little 17985
(US); Augustine, 31 Nov 1960, Little 17986 (F, NY, US).
ORANGEWALK:4 km from San Antonio, 4 Mar 1977,
Stead & Styles 116 (FHO, K). STANNCREEK:All Pines,
29 Jan 1930, Schipp 709 (BM, F, K, MICH, MO, NY, S);
Cabbage Haul Creek, 14 Jan 1988, Stevens 1128 (NY).
TOLEDO:Las Lomitas, 18 Jul 1974, Chaplin c60, c61
(FHO).
HONDURAS. COMAYAGUA:
N of Siguatepeque, 11 Feb
1977, Stead & Styles 78 (FHO). COPAN:Santa Rosa, 6
Mar 1970, Kemp 38 (FHO). CORTES:Lago de Yojoa,
Canaveral,on hillside near NE shore of lake, 14 May 1994,
Farjon & Mejia 356 (E, FHO, HEH, K, MEXU, U); Lake
Yojoa, on shore of lake, 26 Nov 1983, Herndndez &
Yuscaran,Danli-YuscCornejo s.n. (HEH). EL PARAISO:
aran jct., 3 Jan 1951, Kemp 57 (FHO); Ojo de Agua, 22
May 1957, Molina R. 8506 (US); Los Limones, 17 Feb
1972, Styles & Kemp 8 (FHO); Yuscaran, 17 Feb 1972,
MORAZAN:
El
Styles & Kemp 14 (FHO, K). FRANCISCO
Zamorano, 15 Feb 1972, Styles & Kemp 42 (FHO, K).
19 Feb 1981, Stead & Styles 725 (FHO); Otoro,
INTIBUCA:
19 Feb 1981, Stead & Styles 726 (FHO). ISLASDE LA
BAHIA:Good Winter,"IslandW of Guanaja,"26 Jul 1988,
Cornelius 36 (HEH); Roatan I., Port Royal, 1 Sep 1988,
Cornelius38, 42 (HEH);Guanaja1., 30 May 1982, Hughes
165 (FHO); Guanaja I., n.d., Molina R. 20786 (EAP, F,
US). OLANCHO:
Agalta National Park, Catacamas, 1954,
Johannessen470 (IFGP);Aguacate,7 Feb 1965, Johannessen 969 (UC). SANTABARBARA:
Pinalejo, near Quimistan,
16 Mar 1977, Stead 22 (FHO).
NICARAGUA. NUEVASEGOVIA:
Santa Clara, 3 Mar
1972. Styles & Kemp 78 (EAP, FHO). ZELAYA:
Near Tala
Has and Puente Mango (over Rio Kisalaya), 18 Apr 1978,
Stevens 7638 (ENCB, FHO, MO. NY); between El Empalme and Limbaika,24 Feb 1979, Stevens 12787a (FHO.
"Froma sawmill."no location given,
MO, NY). UNKNOWN:
10 Oct 1934, Prettymans.n. (NCU).
Uses. Pinus cubensis, and especially the Mesoamerican variety hondurensis, is an important timber
tree forming easily accessible, extensive stands in
level to moderately hilly terrain. As such it is heavily
exploited, and although not in danger of extinction
in any of its major areas, many local and especially
Representative specimens examined. MEXICO.
populations that are likely to be genetically
outlying
QUINTANA ROO:Ejido Caobas, 7 Feb 1981, Avila s.n.
distinct are now severely degraded (Dvorak & Do(ENCB, MEXU).
GUATEMALA. ALTAVERAPAZ:Secanquim, 3 Jan nahue, 1992). One of these is Ejido Caobas, Quintana
1905, Maxon & Hay 3141 (GH, NY, US). BAJAVERAPAZ: Roo, Mexico, which is the northernmost occurrence
Sierra de las Minas, Mittak 8283 (FHO). CHIQUIMULA: of P caribaea var. hondurensis.
Loc. unknown, Bernoulli & Cario 1090 (B, GOET). EL
Three varieties of Pinus caribaea are recognized,
PETEN:Poptun, 21 Jun 1971, Harmon & Fuentes 5848
(NY); 1.5 km NW of Poptun, Lickhoff 11940 (PRF). as is customary also in forestry literature and pracIZABAL: Montana del Mico, Stevermark38548 (F, US);
tice. We have not found sufficient evidence (i.e.,
91
discontinuous,fixed characters)to separatevar. hondurensis from P caribaea as a distinct species, despite its greatervariability.
Rowlee (1903) was apparentlynot awareof Morelet's species, which had been publishedin a comparatively unknownperiodicaland was not mentionedin
later nineteenth-centurygeneral treatmentsof conifers. His P recurvatais a taxonomic synonym of P
caribaea var. caribaea, not of P tropicalis as it is
sometimes cited. Rowlee also described P cubensis
var. anomala from near "Jucaro Landing" on the
eastern side of Isla de la Juventud. Pinus cubensis
does not occur on this island, and this variety is
also synonymous with P caribaea var. caribaea.
Its "anomalous"leaf-like cataphylls described from
young shoots appearto be not different from those
in the species generally. Liickhoff (1964) considers
the adnateseedwing of P caribaea from Cuba(Pinar
del Rio), "as opposed to the clearly articulateseed
wings from all other areas," the most outstanding
differencebetween seeds of the variousprovenances.
His observationsare based on examinationof large
samples and methodsof "seed handlingor cleaning."
Adnate seeds occur in the Bahamas as well as in
Honduras,as has been pointed out by Loock (1950)
for Hondurasand observed in this study. This casts
serious doubts on the "infallible means of identifying" P caribaea var. caribaea by this character
state, as Lickhoff (1964) suggested.
Grisebach (1864) described Pinus bahaimensisas
a new species from the Bahamas,based on Grisebach
s.n. (holotype, K, lost). Grisebach, too, was apparently not aware of Morelet's descriptionof P caribaea. Barrettand Golfari (1962) designated a neotype. They mention several differential character
states from P caribaea and conclude that the pines
of the Bahamas should be classified as a variety of
it. Apartfrom lower leaf numbers,they find a greater
size range in the cones and predominantlyarticulate
seed wings. From our observations, it appears that
such differences are present,but less pronouncedas
suggested by Barrett and Golfari (1962) and on
the whole with considerably overlapping character
states. Both the lower average numberof leaves per
fascicle and the more often articulateseed wings are
charactersthat P caribaea var. bahamensis shares
with P elliottii (var. densa) of the SE United States.
Seneclauze (1868) describeda new species of pine
based on material he had received in 1854 for his
nurseryat Bourg-Argental(Dept. Loire, France)with
the name Pinus hondurensis.His description,though
very concise and based only on foliage, is not at
variancewith the presentconcept of the species and

cannot be confused with any other taxon in Belize
or coastal Honduras.There is no reason, therefore,
to reject the name in favour of Loock's later homonym. No original materialappearsto be extant (the
description may have been based on live material
with no specimens preserved) and a neotype has
been designatedby BarrettandGolfari(1962). Loock
(1950), like other writers before him (e.g., de Vail,
1941) comparedP. caribaea auct. from Mesoamerica
with P elliottii (then still commonly known, especially among foresters, as P caribaea Morelet) and
concluded that he had a case for treating them as
different species. Unaware of both Morelet (1851,
1855) and Seneclauze (1868), he proposedthe obvious epithet hondurensis.There is no reason to doubt
that Loock referredto the same taxon as Seneclauze
had in mind. Morphologicalcharactersare especially
variable in Mesoamerican Pinus caribaea,
which
has the greatest geographical range and ecological

amplitude of the species as a whole (Styles &
Hughes, 1983). These authors,as well as Barrettand
Golfari (1962) and Luckhoff (1964) report differences in colour and growth pattern of seedlings
between P caribaea var. hondurenlsisand the two
Caribbean varieties. We have used these, besides
mature foliage characters,to key out the varieties.
Additionally,the more stronglydeveloped apophyses
of the cone scales in P caribaea var. hondurensis
are usually a reliable characterstate.
Hybridizationbetween P caribaea var. hondurensis and P oocarpa has often been inferred from
intermediatemorphology(see Styles et al., 1982, for
review) and is possible in areas where both taxa are
sympatricand their phenologies overlap in time, as
is the case in NW Honduras.Putative hybrids with
P. tecunumanii(as P oocarpa var.ochoterenae,vide
Perry, 1991) as well as with P oocarpa are reported
from MountainPine Ridge in Belize and other areas.
6. Pinus elliottii Engelmann var. densa Little &
K. W. Dorman,J. Forest. (Washington)50: 921, f.
1, 2. 1952.
Fig. 21
Pinus elliottii Engelmannsubsp. densal(Little & K. W.
Dorman)E. Murray,Kalmia 12: 23. 1982: Pinus densa
(Little & K. W. Dorman) Silba, Phytologia Mem. 7:
50. 1984; Type. United States. Florida:HendryCo.. SE
of La Belle, 19 Jan 1951, Little & Dorman 14033
(holotype, US; isotypes, F, NY).
Pinus densa (Little & K. W. Dorman)Silba var. austrokevsensis Silba, Phytologia 68: 49. 1990. Type. United
States. Florida:Little Pine Key, I Mar 1911, J. K. Small
Flora Neotropica
92
I,
2ii
itA
FIG. 21. Pinus elliotii var. lenst (A, C, D, Hoffinan296; B, Killip 43267; E, G, H, J. K. Small & Nash s.n.; FJ. K.
Small et al. 3635). A. Branch with leaf fascicles and immaturecones. B. "Grass stage" seedling. C. Leaf fascicle. D.
Cross-section of leaf. E-F. Ovuliferous cones. G. Seed scale (two views). H. Seed with wing. (Magnifications:A-C, EH. xO.5; D, x30.)
et al. 3635 (holotype, NY; erroneously cited in the
as 3655).
protologue
Pinuscaribaeaauct.,nonMorelet(1851),proparte,e.g.,
Small,Floridatrees2. 1913;Shaw,[genusPinus]Publ.
ArnoldArbor.5: 70. 1914;De Vail,Proc.FloridaAcad.
Sci. 5: 121-132. 1941; Loock, [Pines Mexico Brit.
S. Afr.Dept.Forest.Bull.No. 35: 210-211.
Honduras]
1950.
Pinusheterophylla
(S. Elliott)Sudworth,non K. Koch
Bull.
(1849),nec Presl(1849),proparte,e.g. Sudworth,
TorreyBot. Club 20(1): 45. 1893;Sargent,Silva N.
Amer.11: 157-159,t. 591-592. 1897.
Tree,small to mediumor occasionally tall, rapidly
growing at early age (ca. 20 years), height to 1525(-30) m, dbh to 50-(X) cm. Trunkmonopodial,
erect, straight or somewhat tortuous. Bark thick,
scaly, breaking into large, irregularplates, divided
by deep fissures,reddishbrown,weatheringgrey; on
young trees rough, scaly, reddish brown. Branches
of first order few, long, spreading or ascending,
tortuousin older trees; of higher orders spreadingor
ascending, the ultimatebranchesassurging,forming
an open, roundedor flat-toppedcrown. Shoots often
multinodal,on lateral shoots uninodal, thick, rigid,
orange-brownto brown, with small, decurrent,caducous pulvini. Cataphylls small, subulate, scarious,
recurved, brown, with hyaline-erose margins, early
deciduous. Vegetativebuds ovoid-conical, the terminal buds 15-25 mm long, the laterals smaller, not
resinous;the scales imbricate,linear-acuminate,reddish brown with white, ciliate margins. Fascicle
sheaths initially 10-15 mm long, persistent but
slightly reduced to ca. 8 mm in mature foliage;
scales reddish brown with white, ciliate margins,
weathering grey. Leaves in fascicles of 2(-3), in
dense tufts near the ends of branches, spreading
obliquely,rigid, straightor slightly twisted, (16-)1825(-30) cm x ca. 1.5 mm, margins serrulate,apex
acute-acuminate,pungent, dark green, slightly lustrous. Stomataon all faces of leaves, in 10-12 lines
on the convex abaxial face and ca. 10 lines on the
adaxial face. Leaf anatomy: Cross section semicircular, with three leaves in a fascicle transversetriangular;hypodermis with (2-)3-4(-5) layers of
cells; resin ducts 3-9(-11), internal, sometimes 12(-4) medial, variablein diam.; stele elliptic in cross
section, outercell walls of endodermisnot thickened;
vascular bundles 2, distinctly separate, sometimes
connate. Pollen cones crowded near the proximal
end of a new shoot in short clusters, subtendedby
triangular,reddish brown bracts, ovoid-oblong, then
cylindrical,30-50 x 5-8 mm, at first purplish,then
reddish brown; microsporophyllspeltate, the distal
part radially striate, with erose-denticulatemargins.
93
Seed cones subterminal,commonly in whorls of 25 or more, sometimes 2(-3) whorls on multinodal
shoots, on 10-20 mm long, stout, scaly peduncles.
erect at pollination but soon spreadingor recurved,
deciduous. Immaturecones ovoid to ellipsoidal, II13 x 6 mm, pinkish to pale purple, with greenish
apophyses,maturingtime two seasons. Maturecones
ovoid-attenuateto ovoid-conical when closed, ovoid
to broad cylindrical when opened, symmetrical,
(5-)7-12(-15) X (4-)6-8.5 cm when open. Seed
scales ca. 120-150, readily parting except near the
base, thin woody, flat, oblong, straight or slightly
recurved, nearly symmetrical, dark brown on the
abaxial side, with lighter coloured marks of seed
wings on the adaxial side. Apophysis transversely
rhombicand keeled, moreor less flattenedor slightly
raised, ochraceous to (reddish) brown, lustrous.
Umbo dorsal, raised, with a persistent,stout, curved
1-2 mm long prickle, darker than the apophysis.
Seeds obliquely ovoid, acute at proximal apex, or
angular,6-7 X 4 mm, mottledgrey to nearlyblack.
Seed wings articulatebut effective, held to the seed
by two claws, oblong or obliquely obovate. size
variable with cone(scales): 15-30-(35) x up to 10
mm, light brown usually with darkertinge. Seedlings
with a very thick hypocotyl, cotyledons 5-8; developing into a "grassstage"with delayed shoot elongation, a short stem near the ground, a thick taproot
and maturefoliage.
In S Florida and north along the coasts to central
Florida; also on eight of the Lower Florida Keys.
Formingpure fire-climaxforest or scatteredin grassland on sandy lowland plains or on calcareous rock
SW of Miami and in the FloridaKeys. There is often
a dense undergrowthof the palm Sabal palmettoand
it can grow mixed with taller palms locally. Its
altitudinalrangeis 1-20 m. The climate is subtropical
to tropicalon the Lower FloridaKeys: annualprecipitation is 100-150 mm, with a six-month dry winter
period. It is especially well-adapted to frequent
ground fires due to its "grass stage" strategy,which
is found in the seedlings up to sapling age (Little &
Dorman, 1954). Phenology: Pollen dispersal takes
place from late FebruarythroughMarch,depending
on weatherconditions (Little & Dorman, 1954).
Representative specimens examined. UNITED
STATES. FLORIDA: Near Miami, 21 Mar 1904. Britten
101 (NY); Florida Keys, Big Pine Key. 4 Jul 1956. Brizickv& Stern378 (NY);Lee County,SanibelIs.. E partof
island, 29 Dec 1953. Cooley 2485 (NY); Dade County,
"edge of Everglades." 30 Dec 1940. Gifford et al. s.n.
(NCU); Florida Keys. Big Pine Key. near Blue Hole at
94
intersection of Key Deer Blvd. & Miami Ave., 16 Mar
1982, Hill 10974 (NY); MyakkaRiver State Park,Myakka.
28 Mar 1987, Hoffman296 (NY); Florida Keys, Big Pine
Key, along newly cut rd. direct from Inn to Palm Willow,
25 Apr 1953, Killip43267 (NY); EvergladesNationalPark,
between Coconut Grove and Cutler, Oct 1903, Small &
Carter 1249 (NY); near Miami, Oct 1901, Small & Nash
Flora Neotropica
as he considered the characteristicsof the seedling

"evolutionarilyimportant,"but by failing to cite the
basionym he enabled Silba (1984) to publish the
combination validly under the Code. A specimen
with small cones and relatively long (ca. 30 cm)
leaves, preservedin NY (for the cone, see Fig. 21F),
s.. (NY);FloridaKeys,Big PineKey,27 Feb 1911,Smaill was described
by Silba (1990) as a distinct variety
et al. 3575 (NY).
occurring in the Florida Keys. There is no such
Uses. Pinus elliottii var. densa, the South Florida distinction occurringas fixed charactersin the Florslash pine, is a regionally important timber tree ida Keys, as other collections from there amply
prized for its dense wood. Substantial areas are testify.
logged regularly,but in most cases regenerationis
rapidand successful. Substantialvirgin stands (natu- 7. Pinus cubensis Grisebach, [PI. Wright.] Mem.
rally subject to cyclic fires) are protected in EverAmer.Acad. Arts, ser. 2, 8: 530. 1862.
Fig. 22
glades National Park;on the other handsuburbanisaPints occidentalis Swartzvar.cuhensis (Grisebach)Silba,
tion around Miami has been a factor in its decline
PhytologiaMem. 7: 55. 1984. Type. Cuba. Eastern
there.
Cuba:Localityunknown,1856,Wright598 (lectotype,
For a considerable time botanists and foresters
GH, K, NY).
GOET,heredesignated;
isolectotypes,
PinuswrightiiEngelmann,
Trans.St. LouisAcad.Sci. 4:
have applied the binomial Pinus caribaea to the
185. 1880. Type. Cuba. EasternCuba:Localityunslash pine (P elliottii) of the SE United States, and
known.1860,Wright3190 (lectotype,MO.heredesigas
to
in
southern
the
Florida,
species
particularly
nated;isolectotypes,
GH,GOET,K, S).
well as to pines growing on the islands in the CaribPinusmaestrensisBisse, Ciencias(Havana),ser. 10, 2:
bean and in Mesoamerica. However, Engelmann
2. 1975; Pinus occidentalis Sw. var.maestrensis(Bisse)
Silba, Phytologia 68: 57. 1990. Type. Cuba. Eastern
(1880a) already had renamed, fully described, and
Cuba:SierraMaestra,nearLa Francia,28 Mar 1969,
(beautifully) illustratedP taeda L. var. heteroplhlla
Olivia& Herrera13904(holotype,HAJB-n.v.;isotype,
Elliott as P elliottii, realizing that he could not use
JE).
the epithet heterophylla at species rank. The lectoTree, medium to large, height to 25-30 m, dbh to
type of P elliottii: United States, South Carolina,20
Mar 1873, Mellichamp s.n. (MO no. 3941460) was ca. 1() cm. Trunk monopodial,erect, usually slender,
designatedby Farjon(1993). Sudworth's(1893) rais- in forest stands with clear boles for 2/3or /4 of total
ing of Elliott's varietal epithet to species rank was height. Bark thick, rough, scaly, on the lower partof
both superfluouswhen publishedand illegitimate,as the trunk breaking into irregular, squarish plates
it created a later homonym.
divided by deep fissures, or forming vertical ridges
Little and Dorman (1952) restrictedthe concept between fissures, grey-brown, weathering grey, on
of P caribaea to Caribbeanand Mesoamericanpines, young trees and branchesrough and scaly. Branches
recognizing a single variabletaxon rangingfrom the of first orderslender,spreadingirregularly;branches
Bahamas to Honduras, and further recognized P of higher orders similar, forming a small, irregular,
elliotrii with two varieties in the United States. Con- broadlydomed and open crown. Shoots more or less
sequently, they named the southern Florida taxon multinodal,rough with small, short-decurrent,semiPionuselliottii var. densa. The botanical differences persistent pulvini, glaucous to pruinose in the first
with P elliottii var. elliottii include a prolonged, and second years. Cataphylls small, subulate, scariwell-defined "grass stage" of the seedling which ous, early recurving, with ciliate margins, brown,
results from apical growth inhibition of the stem. deciduous shortly after the leaves or falling simultaThis is apparentlyan inherited trait (Little & Dor- neously. Vegetativebuds ovoid-oblong, the terminal
man, 1954) as it is retainedin seedling experiments buds 10-15 mm long, the lateralssmaller, acute, not
outside naturalconditions. Initial growth is invested resinous or sometimes slightly resinous; the scales
in a thick hypocotyl; subsequentlya large taprootis imbricate,more or less appressed, scarious, narrow,
formed and the hypocotyl grows to a short "stem" with ciliate margins, brown. Leaffascicles initially
up to 3 cm thick and 4 cm long (Fig. 21B). Other 9-12 mm long, on mature leaves reduced to 7differences appear to be more clinal (Price, 1989), 9 mm, persistent, with imbricate, silvery-brown to
such as the fewer leaves per fascicle and the thicker greyish brown scales. Leaves in fascicles of 2(-3),
in dense, spreadingtufts, persisting 2-3 years, more
hypodermisin the leaves of var. densa.
Gaussen (1960) raised the variety to specific rank, or less rigid, or flexible, straight or slightly curved,
95
1.1
~~~~~~~~~~~~"?i
w...
ZE;~~~~"
~'
FIG. 22. Pinus cubensis (A-C. Marie-Victori & Cleiment 21428; D, E. Lamib .sn. (Moa Mine); F-H. Shafer 3216).
A. Branch with leaf fascicles and immatureovuliferous cone. B. Leaf fascicles. C. Cross section of leaf. D. Immature
cones on recurved peduncles. E. Growing cones. F. Mature cone. CG.Seed scale (three views). H. Seed (two views).
(Magnifications:A, B, D-F, O.5;C, x40; G. H. x 1.)
(6-)10-15(-18) cm X 0.8-1.3 mm, margins serru- the convex abaxial face and 4-5(-7) on the adaxial
late, acute, light green, occasionally glaucous-green. face(s), weakly correlatedwith width of leaves. Leaf
Stomata on all faces of leaves, with 5-10 lines on anatoiny: Cross section semi-circularor transverse-
96
triangular(3-leaved fascicles), with convex abaxial
side; hypodermis multi-layered with 2-3 rows of
cells; resin ducts 3-6, internal; stele more or less
elliptic in cross section, outer cell walls of endodermis not thickened; vascular bundles 2, distinctly
separate. Pollen cones clustered near the proximal
end of a new shoot, cylindrical,often recurvedwhen
mature, 15-20 X 5 mm. Microsporophyllsdistally
peltate, radially striate on the abaxial side, with
erose-denticulate margins. Seed cones subterminal,
solitary or in pairs, less commonly in whorls of 34, on 1-2 cm long, bracteate, recurved or straight
peduncles, deciduous. Immature cones (narrowly)
ovoid or oval, 10-15 x 6-9 mm, ochraceous,often
pruinose;umbos rhombic, with a minute spine. Mature cones (narrowly)ovoid to ovoid-attenuatewhen
closed, (broadly) ovoid when opened, (3.5-)4-7 x
(2-)3.5-5 cm when open. Seed scales ca. 90-110,
partingreadilyexcept the basal ones, oblong, straight
or slightly recurved,thin woody, darkpurplishbrown
to nearly black on the abaxial side, dark brown with
faintly distinct marks of seed wings on the adaxial
side. Apophysismore or less symmetricalaroundthe
cone, slightly raised, transverselykeeled, transverserhombic to pentagonal in outline, with curved or
irregularupper margin, up to 11 mm wide, radially
striate,darkochraceous to dull brown. Umibodorsal,
depressed or slightly raised, transverse-rhombic,34 mm wide, brown, weathering grey; the minute
prickle early deciduous. Seeds obliquely ovoid, flattened, 5-6 x 3-3.5 mm, mottled grey-brown.Seed
wings usually adnate (with membranecovering the
seed on one side), or articulate (wing held to the
seed by two claws), obliquely ovate-oblong, 12-16
x 5-6 mm, grey-brown, with a graphite-liketinge.
Numberof cotyledons not observed.
Distribution and ecology (Map 2). EasternCuba:
From the Sierra Maestra and the Sierra de Nipe E
into the highlands terminatingthe eastern partof the
island. It occurs in foothills and highlandsas well as
in "pinebarrens"along the coast. Its altitudinalrange
is 100-900(-1200) m. Pinus chbensis forms mostly
pure but open stands or is invasive in disturbedsites
on serpentine or serpentine-derived,often ferruginous soils ("Nipe latosol" or "Nipe clay") or on
alluvial sediment near the coast. In the highlands
there is abundantrainfall (1800 mm or more annually), but a dry winter season is a typical aspect of
the tropical to subtropicalclimate. Phenology:Time
of pollen dispersal not recorded.
Flora Neotropica
s.. (FHO, NY); SierraMaestra,N of Pico Bayamesa,May
1968, Bisse & Duek 9118 (JE); Santiago de Cuba, E of
Gran Piedra, 26 Apr 1969, Bisse & Lippold 15040 (JE);
Santiago de Piedra, Gran Piedra,Jun 1967, Bisse & Rojas
3762 (JE); Sierra Maestra,28 Jun 1914, Ekman 1533 (K,
S); Sierra Maestra. near Reuni6n, 30 Mar 1915. Ekman
7026 (NY, S); Sierra de Nipe, 6 Dec 1919, Ekman 10137
(K, NY, S): Sierra Maestra, along trail to Pico Turquino,
11 Jul 1985, Gentry & Lavin 50962 (MO); Monte Cristi, 5
Aug 1968. Limb s.n. (FHO); Moa Mine, 6 Aug 1968,
Lambs.,. (FHO); Braza de Cauto, Jul 1927, Le6n & Roca
10326 (P); Rio Yao. Mar 1943, Marie-Victori, 600/0 (US);
vic. of Moa. 27 May 1943, Marie-Victorin & Clement
21428 (NY, US); MonteVerdenearYateras.29/30 Apr
1907, Matxon 4341 (MO); Sierrade Nipe, Apr 1966, Sanmek
s.n. (FHO);Sierrade Nipe, 19 Dec 1909, Shaifr 3216

I Mar1910,Shaftr
(US);nearmouthof Rio Yamaniquey,
4254 (NY); CampLa Gloria,S of SierraMoa, 24 Dec
1940, Shlafrr8128 (K, NY, P, US); along rd. from Arriba
to Bermejal,31 Jan 1911,Shafer8453 (K, NY,U. US);La

Perla.19Feb1911,Shafer8842(K,NY);localityunknown,
1860-1864,Wright1462B(MO);localityunknown,1863,
Wright1462C(GOET);nearMonteVerde,1859, Wright
1462D(GOET,K, MO),Wright1462(MO).
Uses. Pints cubensis is the only pine occurringin
the eastern partof Cuba (see below); no pines occur
naturallyin Cuba between Eastern Cuba and Pinar
del Rio in the extreme west of the island, and as
such the pines are of importanceeconomically as a
timber source to this partof Cuba.
Grisebach (1862), in his enumeration of plants
collected by CharlesWrightin Cuba,describedPinus
cubensis as a new species, based on Wright 598
and 1462. Number 1462 being an ambiguous
collection in part mixed with P caribaea (GOET,
seen by B. T. Styles), no. 598 (GOET) is here
designatedas the lectotype; it has been annotatedby
Grisebach.
Engelmann (1880a) described as a new species
No. 45 Pinls
wrightii, citing Wright 1462B and
Wright 3190. The specimen subsequently marked

1462C (GOET) belongs here; it is also a syntype
of P cubensis. The lectotype of P wrightii is designatedhere to be no. 3190 (MO). Engelmannappears

to have confused his species no. 44 P cubensis
with P caribaea Morelet ("Isle of Pines, Bahama
Islands"); and his P wrightii, which describes P
cubensis from "Mountains in Eastern Cuba" is a

superfluousname.
Sudworth (1893), erroneously equating P taeda
var. heterophvlla Elliott, described from Georgia,
United States, with P cubensis, proposedon nomenclaturalgrounds (American rules) the "older"binomial P heterophylla(Elliott) Sudworthfor the "Cuban
examined.
CUBA.
EASTERN
pine." Under the present Code, no name has
Representative specimens
CUBA:Sierra de Nipe, Mayari. 13 Apr 1969, Betancol,rt
priorityoutside its own rank. Shaw (1914) synony-
Systematic Treatment97
mized P cubensis and P wrightiiwith P occidentalis
Sw. by applicationof a much wider species concept
than one would accept today.
Bisse (1975) provideda key for the pines of Cuba
in which the first couplet divides them into 2- and
3-leaved pines. Fromthe SierraMaestrahe described
a new species: P maestrensis,with leaves in fascicles
of 3. The leaves are -I1 mm wide, glaucous-green,
and very flexible. The reportedthin leaves (0.5-0.7
mm), predominantlyin fascicles of 3, are the only
different feature of this taxon, which was said to
occur in various places between 300-1700 m together with P cubensis. The measurementsof Bisse
(1975) have not been observed on the isotype (JE),
which has leaves 0.8-0.9 mm wide, nor on any other
materialat JE determinedby Bisse as P mnaestreesis
(with up to 1.1 mm wide leaves). One of his specimens (Bisse & Duek 9118) has all leaves in fascicles
of 2, and another(Bisse & Rojas 3762) primarilyso.
Silba (1984) reducedP cubensis to a varietyof P
occidentalis, and P maestrensislikewise to a variety
of P occidentalis, repeating the original diagnosis
but without having seen any of the original material
or giving an explanation. As Darrow and Zanoni
(1991) have discussed, the pines of the SierraMaestra in E Cuba are not P occidentalis, for which they
have long been mistaken (e.g., map 46 in Critchfield & Little, 1966; Mirov, 1967)-that species is
native to Hispaniola.Neither do they belong with P
caribaea. Their placement with P cubensis is in
agreement with Florin's determination of Ekmian
1533, with morphologicalcharacterstates found in
that species east of the Sierra Maestra, and finally
with overall geographic patternsof Caribbeanpine
taxa.
8. Pinus occidentalis Swartz, Prodr. 103. 1788.
Type. Haiti. "Hispaniola: aux Pins, quartier des
Nippes, du c6ot septentrional,"1785, Swartz s.n.
(lectotype, BM, designatedby Farjon, 1993).
Fig. 23
Pinus occidentalis Swartz var.baorucoe.nsis Silba, Phyto-
Republic.Pederlogia58: 368. 1985.Type.Dominican

nales:Sierrade Baoruco,nearAceitillar,14 May1976.
Judd 1487 (holotype. A).
Tree, medium to tall, height to 30-40 m, dbh to

100-120 cm. Trunkmonopodial,erect, terete,usually
slender, in forest stands with clear bole for
2/3--4
of
height. Barkthick, on lower trunk30-40 mm, rough,

scaly, breakinginto irregularlysquare plates divided
by deep fissures, longitudinally fissured toward the
base, on young trees and branches light brown,
turning grey-brown to grey. Branches of first and

higher orders slender, spreading or ascending, in
the lower crown curved and drooping, forming an
irregular,open crown. Shoots uninodal,glaucous to
pruinosein the firstyear, later very rough with large,
short-decurrent,
persistentpulvini,brown.Cataphylls
slender, subulate to caudate, straight or recurved,
scarious,with erose margins,brown,persistingsome
time after the leaf fascicles have fallen. Vegetative
buds ovoid to ovoid-oblong, acute, the terminalbud
10-15 mm long, the lateralsshorter,usually slightly
resinous; the scales imbricate, more or less appressed, scarious, with ciliate margins, lustrous
brown. Fascicle sheaths persistent, (8-)10-15 mm
long, scales lustroussilvery-brownwith erose-ciliate
margins,weatheringgrey-brown.Leaves in fascicles
of 3-5, in dense, spreading tufts at the ends of
branches, persisting to the third year, straight or
slightly curved, more or less rigid, (I -)14-18(-20)
cm x 1.2-1.4 mm, with serrulate margins, acute.
pungent,light green. Stomata on all faces of leaves,
in 6-8 lines on the convex abaxial face and ca. 4
lines on each adaxial face. Leaf anCatomyv:Cross
section triangularor transverse-triangular,
with convex abaxial side; hypodermis multi-layered (3-4
rows of cells); resin ducts (2?-)3-5, internal;stele
elliptic to transverse-obovatein cross section; outer
cell walls of endodermis not thickened; vascular
bundles 2, distinctly separate.Pollen cones clustered
near the proximal end of a new shoot, cylindrical.
often recurvedwhen mature,15-25 X 5 mm, pinkish
yellow, turningyellowish brown. Seed cones subterminal, solitaryor in pairson 10-20 mm long, straight
or recurved, bracteatepeduncles, persisting several
years after seed dispersal, falling with the peduncle
attached. Immaturecones ovoid to ovoid-attenuate
when growing, 10-15 x 8-10 mm, with inflexed
spines, ochraceous,maturingin two seasons. Mature
cones ovoid to ovoid-conical, straight or curved.
nearlysymmetrical,largecones with a flattenedbase,
(4-)5-9(-11) x 3.5-6.5 cm when open. Seed scales
ca. 110-150, partingto release the seeds except those
at the base, oblong, straightor recurved,thin woody,
dark purplishbrown abaxially,dark brown with indistinct lighter brown marks of seed wings on the
adaxial side. Apoplhsis slightly raised, transversely
keeled, rhombic to pentagonal in outline, up to 12
mm wide, on the proximal scales more or less gibbous, (lustrous)darkbrown,radiallystriate,weathering dull grey. Umbo dorsal, raised and often curved,
usually armedwith an inflexed, 2-3 mm long spine,
grey-brown.Seeds obliquely obovoid, flattened,5-6
98
Flora Neotropica
~E
1,
FIG. 23. Pinuusoccidentalis (A-Ba, Hughes 481; Bb, C, Ea, Ga, Zanoni et al. 40761; D, Eb, Gb, Zanoni et al. 41212).
A. Branch with leaf fascicles. B. Leaf fascicles. C. Cross-section of leaf. D. Growing immatureovuliferous cone. E.
Ovuliferous cones. F. Seed scales. G. Seeds. (Magnifications:A, B, D-G, x0.5; C, x 30.)
X 3-4 mm, light, mottled grey-brown. Seed wings
articulate but effective, held to the seed by two
claws, obliquely ovate or oblong, 12-18 X 4-6
mm, ochraceous with black or grey tinge or stripes.
Numberof cotyledons not observed.
Distribution and ecology (Map 2). Hispaniola:
Both in Haiti and the Dominican Republic, where
the largerpart of its range is; reportsof the species
from EasternCuba (Oriente)are probablyall erroneous and likely pertain to P cubensis. It occurs in
diverse habitatsfrom the lowlands, at about 200 m,
to the highest mountainridges (Pico Duarteand Pico
La Pelona) on the island, at almost 3200 m. The
more extensive and pure stands occur at 900-2700
m (Mirov, 1967; Farjon, 1984), but in more accessible areas these are much depleted. Soils are either
derived from limestone at lower altitudes, or more
acid, clay-like, and shallow in the CordilleraCentral.
Pinus occidentalis consequentlyis found in a variety
of vegetation types, mostly occupying the shallow,
nutrient-poorsoils and rock outcrops, where it may
occur in open or dense, pure stands or mixed with
various broad-leavedtrees and shrubs. In disturbed
(grazed)areasPteridiumaquilinumcan dominatethe
groundcover; in frequentlyburntareas,grasses (e.g.,
Danthoniadomingensis,Andropogonspp.) and Pteridium replace shrubs and small trees. Thick bark on
the lower trunks of pines are a protection against
quickly passing fires, and regenerationcan be rapid
from seed. Annual precipitationvaries greatly with
exposition, but is in the range of 1200-1600 mm
where most pine forests occur and exceeds 2300 mm
in the N and E of the CordilleraCentral.There is a
3-5-month dry season during winter, which may
bring frost, but rarely snow, at the higher altitudes
above 1600-1800 m. Yet there are also pines in
tropical and subtropical regions at low altitudes
where no frostoccurs. A detailed accountof distribution and ecology is given by Darrow and Zanoni
(1991) from which much of the above information
is obtained.Phenology:Time of pollen dispersalis in
early spring,probablyvaryinggreatlywith altitude;a
second period has been reported in October (see
Darrow& Zanoni, 1991).
99
32443 (NY). LA VEGA:Constanza, 23 Apr 1976, Juddl
1166 (NY); Constanza, I Jul 1964, Lamb 10 (FHO); Rio
Jimenoa, 7 km from Jarabacoa,22 May 1986, Zanoni et
al. 36453 (NY): 2.5 km SE of Aguacate. 26 May 1988,
Zanom
i et cl. 41212 (FHO). PEDERNALES:Sierrade Baoruco, 26 Jul 1981, Fisher-Meerow687 (NY); island W of
Hoyo de Pelempito,NE of Cabo Rojo, I I Feb 1981, Zanoni
et al. 10889 (F, NY): Sierrade Baoruco,33 km N of Cabo
Rojo, 10 Apr 1988, Zatnoti et al. 40760, 40761 (FHO).
PERAVIA:
42 km NW of San Jose de Ocoa, 30 May 1984,
Mejia et al. 608 (NY), 30 May 1984. Mejia et al. 612
Los Cacaos, 3 May 1984,
(MO, NY, US). SANCRISTOBAL:
Rio Bao, Dec
Zcaoni et al. 29734 (FHO). UNKNOWN:
27625(P,S); Rio Bao,locationpresumably

1952,Humtbert
fartherupstreamthan Humbert27625 (P), Dec 1952.
Humbert27699 (P); loc. not recorded, 1853. Schomlburgk
43 (P).
Uses. Being the only species of Pinis on Hispaniola and formerlyabundantover much of the island,
it has been heavily exploited for timber.According
to Darrow and Zanoni (1991) it has been depleted
from an estimated3 million ha of primeval,more or
less pure pine forests to perhapsless than 5% of that
area, but accurate estimates of even the present
forestextent are lacking. Protectionin the Dominican
Republic is inadequatebut existent, contraryto the
virtually uncontrolledsituation in Haiti (Darrow &
Zanoni, 1991).
This species has been describedby Swartz (1788)
from "Hispaniola,"where he explored the western
part of the island (now Haiti) from December 1784
throughJune 1785 (Steam, 1980). He gives a concise
description of a 5-leaved pine and a reference to
"Plum.t. 161,"which is plate 161 in Burman(1758).
Darrowand Zanoni (1991) refer to a collection in S,
but this specimen,althoughfrom Swartz'sherbarium,
is a 4-leaved branchletof unknownorigin. A specimen in BM, which is a 5-leaved branchletin accordance with the protologue, bears on the reverse
of the sheet the inscription (in L'Heritier'shand):
"Hispaniola:aux Pins, quartierde Nippes, au c6ot
septentrional.Dr Swartz."This specimen has been
designatedas the lectotype (Farjon, 1993).
Shaw (1914) considered the pines of E Cuba
conspecific with those indigenousto Hispaniolaand
thereforecited P cubensis Griseb. in synonymy with
P. occidentalis Swartz. His observationof an "erect
Representative
specimensexamined.HAITI.Artibon- conelet and a reflexedcone"
may have been accurate
ite: Marmalade,
1903,Nash825 (NY). NORD:Briquis,26
Nov 1985, Zanoni & Mejia 35801 (NY). OUEST:Furcy,21 in the specimens he studied,but conelets on recurved
Aug 1917, Cook, Scofield & Doyle 25 (FHO, US). PORT- peduncles occur as well, and this "character"is
AU-PRINCE:Morne la Visite, 35 km S of Port-au-Prince,15
thereforeof little value taxonomically.Many collecJan 1985, Hughes 480, 481 (FHO). UNKNOWN:
"Sourcail," tions from E Cuba had been determined
by B. T.
20 May 1828, Jaeger 155 (S, US).
as
P.
occidentalis
because
of
Shaw's
treatment,
Styles
DOMINICAN REPUBLIC. AZUA:Sabana de Miguel
Martin,3 May 1988, Zanoni et al. 40832, 40833 (FHO). which seems to have been based on a mixture of
DAJABON:Sierra Chacuey, 27 Nov 1984, Zanoni et al.
taxa (Darrow & Zanoni, 1991). No authentic P
100
occidlentalis collected in Cuba from the wild has
come to the attentionof the first author(A. Farjon),
who agrees to restrict P occidentallisto Hispaniola
in the absence of positive evidence to the contrary.
Silba (1985b) described as new to science P occ-ideitalis var. baorucoensis, with leaves in fascicles
of 5 (rarely4), and with "swollen, sometimes protuberant apophyses with a fine, upcurved, prominent
spine" from the Sierrade Baoruco in the Dominican
Republic and other localities both in the Dominican
Republic and Haiti. Similar cone characters have
been observed in specimens with 3-leaved fascicles
(e.g., Zanoni et al. 40760, 40761 from the Sierrade
Baoruco, FHO), whereas 5-leaved specimens may
have cones with more-flattenedapophyses and less
prominentspines. The holotype specimen (A), determined by Zanoni Dec 1981 as P occidentalis, has
3-, 4-, and 5-leaved fascicles. As with virtually all
of Silba's "new"taxa, this one seems to be based on
odd observationsof only a few specimens (a scan of
the relevant materialin the herbariawhere the types
of his taxa are deposited usually reveals this), and it
was proposed without consultation of the botanist
(Zanoni, in this case) who knew intimatelyboth the
area and its pines.
9. Pinus ponderosa D. Douglas ex C. Lawson var.
scopulorum Engelmannin S. Watson,Bot. California 2: 126. 1880.
Fig. 24
Piins ponderosa D. Douglas ex C. Lawson subsp. scopu-
E. Murray,Kalmia12: 23. 1982;

loruni(Engelmann)
Pinus scopultorum (Engelmann) J. G. Lemmon, Gard.
Forest10: 183. 1897.Type.UnitedStates.Colorado:

"ColoradoTerritory,
lat. 39-41?. Alpineand Subalpine,"Jun 1862,Parrys.n. (lectotype,MO,heredesignated).
in Wislizenus,Mem.Tour
PilnusbrachYptera
Engelmann
N. Mexico89. 1848.Type.UnitedStates.New Mexico:
534
"RockCreekto SantaFe,"5 Jul 1846,WislizeYnus
(holotype,MO).
Tree, usually tall, height (Mexico) to ca. 40 m,
dbh to 80-120 cm. Trunkmonopodial,straight,erect,
usually slender.Bark thick, roughand scaly, dividing
into large, slightly elongate plates separated by
broad, shallow fissures, clear orange-pinkishor reddish brown with dark fissures on the lower part of
the bole, on young trees and branches thin, scaly,
brown or darkreddishbrown. Branchesof firstorder
relatively short,spreadingor curveddown, numerous
and persistent;branches of higher orders ascending
or spreading, the lower branches subpendulous,assurgent at tips. Shoots stout, rough, with persistent,
decurrentpulvini which leave prominentmarksafter
the fascicles have fallen, reddish brown, often glau-
Flora Neotropica
cous. Cataphylls 10-15 mm long, subulate, scarious,
recurved,with hyaline-erose to ciliate margins, reddish brown, weathering blackish, mostly abscising
with the leaf fascicles. Vegetativebuds ovoid-acute
to broad-ovoid, the terminal bud 15-25 x 10-15
mm, the laterals smaller, not resinous; the scales
spreadingor recurvedat base of bud, subulate-caudate, lustrousred-brownwith darkerapex and white
or brown ciliate margins. Fascicle sheaths initially
ca. 20 mm long, consisting of ca. 10 imbricate,
appressed,orange- to red-brownscales with hyaline
margins, in mature fascicles persistent but reduced
to 10-15 mm, with a thickened base, weathering
blackish grey. Leaves in fascicles of 2-3 (in some
trees predominantly2 or 3), in dense, spreadingtufts
toward the ends of upturnedbranches,persisting 22.5 years, straightor curved, rigid, (10-)15-25(-27)
cm X 1.3-1.6 mm, with serrulate margins, acutepungent to acuminate, light or dark green. Stomata
on all faces of the leaves, in 8-10 lines on the convex
abaxial face, in 3-4 lines on each adaxial face.
Leaf anatomy:Cross section transverse-triangular
(3leaved fascicles), or semi-circular (2-leaved fascicles), with a convex abaxial side; hypodermis uniform, with 2-3 layers of cells; resin ducts 2-6,
medial;stele oval in cross section; outercell walls of
endodermisthickened;vascularbundles 2, distinctly
separated. Pollen cones densely clustered near the
proximalend of a new shoot, subtendedby subulate
bracts with white ciliate margins, ovoid-oblong to
cylindrical, 15-30 X 5-7 mm, yellowish, turning
yellowish brown.Seed cones subterminal,solitary or
in whorls of 2-3 on short, stout, recurvedpeduncles,
persistent some time after seed dispersal, leaving a
few basal scales on the pedunclewhen falling. Immature cones broadly ovoid to subglobose, with sharp,
spreading or curved spines, (15-)20-25 x 12-18
mm, purplish brown, glaucous, maturing in two
years. Mature cones ovoid to subglobose when
opened, symmetrical or slightly oblique toward the
base, 5-10 X 4.5-7 cm when open. Seed scales ca.
90-120, partingwhen mature,thin, flexible, oblong,
with thin margins,straightor recurved,darkpurplish
brown with a light brown edge below the apophysis
abaxially,light brownwith faint marksof seed wings
adaxially. Apophysis slightly to prominentlyraised,
transverselykeeled, rhombic or irregularlypentagonal in outline, ochraceous to reddish brown. Unbo
dorsal, flat to prominently raised and transversely
keeled, 5-7 mm wide with a persistent, curved, to
3 mm long spine. Seeds obliquely ovoid, slightly
flattened,5-7 x 4-5 mm, light grey-brown, sometimes with dark spots. Seed wings articulate,effec-
101
III-o
FIG. 24. Pinus ponderosa var. scopulorun (A, Ba, C, Wynd& Mueller 650; Bb, D, E, Parry s.n. (lectotype); F-H,
MacKelveY194, FHO, from Arizona, not cited). A. Shoot with leaf fascicles. B. Leaf fascicles. C. Cross section of leaf.
D, E. Shoots with leaf fascicles and growing ovuliferous cones. F. Matureovuliferous cone. G. Seed scale (three views).
H. Seeds, with articulatewings. (Magnifications:A, B, D-H, X0.5; C, x30.)
Flora Neotropica
102
"-r~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~i....
I
A --I
...
i~.._~l,,,,; ',,. : / ,,, u ,,-~
i~
~ _. ,, , ....???...
-':
I,
. I~
~~.---?--I
..
Map 6. Pinus pondem.sa var. scopulorumn(squares); P. arizonica var. cooperi (triangles); P arizonica var. .torniae
(circles).
tive, held to the seed by two narrow,flimsy claws,

obliquely oblong, 15-20 x 6-10 mm, translucent,
light yellowish brown. Cotyledons6-9(-1 I).
Distribution and ecology (Map 6). The range of
Pinus ponderosa extends from SW Canada to the
Mexican border. Commonly, the species is recognized to occur in two wide-ranging varieties, var.
ponderosa and var. scopulorum(with P. arizonica as
a third variety in the United States, which is here
recognizedas a species). Pinus ponderosavar.scopulorumis the interioror "Rocky Mountain"form, and
it is this taxon that is found in two separatelocalities
just S of the Mexican border, in Chihuahua and
Coahuila. In the region of the SW United States
adjacentto the Mexican border,P. ponderosa occurs
in the "yellow pine belt" above a zone characterised
by Pinus edulis, Juniperus monosperma,and Quercus, on mostly isolated mountainsseparatedby broad
intermontanevalleys of sufficient elevation (above
1500 m) to receive a minimum of ca. 200 mm

precipitationduringthe growing season.
The vegetation on the two mountains in Mexico
where P. ponderosa has been found is similar to that
N of the border,with open, park-likestands of Pinus
interspersedwith Quercus, Arbutus, shrubs such as
Arctostaphylos, and grassland. Phenology: Pollen
dispersaloccurs in May-June,a dry period.
Representativespecimensexamined.MEXICO.Chihuahua:W sideof theSanLuisRange,in the firstcanyon
S of DiabloCanyon,3 Sep 1952, Tucker2596 (A, UC).
Sierradel Carmen,highpeakadjoiningCanon
COAHUILA:
Sentenela,10 Jul 1936, Wynd& Mueller650 (MO, NY,
P).
Uses. Ponderosapine is one of the most important
timbertrees of the United States. Due to the remoteness of the areas where it is found in Mexico, no
commercial exploitation is likely to occur; its use
SystematicTreatment
would otherwise be comparablewith that of P arizonica.
Pinus ponderosa var. scopulorum was distinguished from P. ponderosa of Oregon and California
by Engelmann (1880b), who also referred to it as
"Pinus ponderosa of the Rocky Mountain flora's."
The materialupon which Engelmannbased this variety included a collection by C. C. Parry,made in
June 1862 and bearing a label imprinted "Rocky
MountainFlora,"which is in Engelmann'sherbarium
(MO), with two small brancheswith leaves up to 10
cm long in fascicles of 2-3 and immatureto halfgrown seed cones. Since Engelmanndescribed also
maturecones and seeds, he must have had additional
material,but it has not been found in MO. The Parry
sheet is neverthelessdesignated as the lectotype because Engelmann used to send out other bits to
Parlatore and other botanists, which may yet be
found. Engelmann (1880a) distinguished P ponderosa var.scopulorumfrom both var.jeffreyi (= P
jeffreyi) and var.ponderosa by its smaller cones and
shorterleaves, "often in pairs."This lattercondition
is, according to Conkle and Critchfield(1988), not
found in the "Southwesternregion,"i.e., Arizonaand
New Mexico, where only 5-15% of leaf fascicles
are foundto be 2-leaved and where 3-leaved fascicles
predominate.The extension of usage of the epithet
scopulorumto the Ponderosapines of this region fits
the clinal patternof this trait from north to south in
the interiorpopulations(Conkle & Critchfield,1988:
fig. I) and is accepted here.
Shaw (1909, 1914) used a broad circumscription
of Pinus ponderosa, which included P arizonica, P
engelmannii, and P jeffreyi as varieties in Mexico
(1909), and later also P scopulorum (1914). In the
United States, P arizonica is still widely considered
a variety of P ponderosa (Little, 1979; Burns &
Honkala, 1990; Kral, 1993), but that view seems
to be based primarily on the populations in the
southwesternUnited States and not on an evaluation
of the complex throughout northern Mexico. The
only authorof these who has a broad knowledge of
Mexican pines is Little, but he favoureda conservative approachto species recognitionin his checklists,
which is of U.S. trees only (Little, 1953, 1979).
There may be naturalhybridizationin Arizona and
New Mexico where the two taxa occur together
(Conkle & Critchfield, 1988), complicating recognition of these taxa there. Monoterpenecomponents
seem to be as variable between P arizonica and P
ponderosa as within the latter species (Conkle &
Critchfield,1988) and provide inconclusive evidence
for species delimitation.
103
Pinus ponderosa s.str. is recognized here as occurringin two wide-rangingvarieties:var.ponderosa
and var. scopulorum (Kral, 1993). Trees with morphologicalcharacterstates as given in the description
above occur in the San Luis Range in extreme NE
Chihuahua,close to the United States-Mexico border, and in the Sierra del Carmen, just south of
the Rio Grande in Coahuila. They belong to var.
scopulorum. Bailey and Hawksworth(1983) anticipated such occurrencesnear the border.
Martfnez(1948), when referringto P. ponderosa
in Baja California, confused it with P jeffreyi, a
species he also recognized as distinct. The southern
California"race"of P ponderosa (Conkle & Critchfield, 1988) has its southernmostextension in the
Laguna Mountainsin San Diego County (Griffin &
Critchfield,1972;A. Farjon,pers. obs., 1992), where
it is sympatricwith P jeffrevi.
10. Pinus arizonica Engelmann,in Rothrock,Rep.
U.S. Geogr. Surv., Wheeler,6, Bot.: 260. 1879.
Tree, usually tall, height to 30-35 m, dbh to 100120 cm. Trunkmonopodial,erect, straight,tereteand
slender, often clear of branches 2/- to 3/5 its length.
Bark thick, rough, scaly, breaking into large plates
divided by broad, shallow or deep fissures, reddish
brown to dark brown, outer bark grey; on young
trees and branches thin, rough and scaly, reddish
brown.Branchesof firstorderheavy, long, spreading
horizontallyor curved down; of second and higher
orders ascending or spreading, the lower branches
subpendulous,slender.Crownin young trees pyramidal, open, in old trees roundedor flat-topped,open
or dense. Shoots at first orange brown or glaucous,
rough with persistent,decurrentpulvini. Cataphylls
up to 15 X 3-4 mm, subulate, scarious, soon reflexed, with erose-ciliate marginsand caudate apex,
dark brown. Vegetativebuds ovoid to ovoid-acute,
the terminalbuds 15-30 X 10-15 mm, the laterals
smaller,not resinous, brown;the scales spreadingor
recurvedat apex, with long-ciliate, white or brown
margins.Fascicle sheaths initially long, 15-30 mm,
with ca. 10 imbricate, orange-brownto red-brown
scales with ciliate margins,persistentbut reducedto
ca. 10 mm, with a thickenedbase, weatheringgreyblack. Leaves in fascicles of 3-5, spreadingat a wide
angle from the shoot, persisting 2-3 years, rigid to
slightly lax, straightor slightly curved and twisted,
(8-)10-20(-25) cm X 0.9-1.8 mm, margins serrulate, apex acute to pungent,light yellowish green or
glaucous-green. Stomata on all faces of leaves, in
(3-)4-8(-12) lines on the convex abaxial face and
104
Flora Neotropica
?~~~~9
'.~i~
7.
.~ ~?~~
Ma Pin
~~~~~~~~~~~'
'
'
~" i' .,- I..
'
"
arznc
???."~'-N~
~.
. .z
i
'-??
va.
arizonica?-?
in (3-)4(-5) lines on each adaxial face (the abaxial

numberof lines correlatedwith width of leaf). Leaf
anatomy: Cross section (transverse-)triangular;hypodermis multi-layered, with intrusions into the
mesophyll; resin ducts (3-)5-10, medial, usually 3
larger(external) ducts present;stele elliptic in cross
section; outer cell walls of endodermis thickened;
vascularbundles 2, more or less connateor distinctly
separate. Pollen cones densely clustered near the
proximalend of a new shoot, subtendedby subulate
bractswith ciliate margins,ovoid-oblong to cylindrical, 15-20 X 5 mm, yellowish to yellowish brown.
Seed cones subterminal,solitary, in pairs or whorls
of 3-5, on short, curved, persistent peduncles retaininga few scales on the branches.Immaturecones
ovoid-globose, ca. 15 X 10-12 mm, with sharp,
curvedor spreadingspines, purpleto purplishbrown,
maturingtime two seasons. Maturecones seemingly
sessile, ovoid or broadly ovoid with parted scales,
asymmetrical at obliquely flattened base, often
(
gr
c(Iranges
rle)
P~..?U
e 1 ggii
slightly curved, (4.5-)5-10(-14) X 3.5-6(-8) cm

when open. Seed scales ca. 90-140, most numerous
at the base of the cone, partingto release the seeds
except at the base, thick, rigid, broadlyoblong, nearly
straight or recurved especially near the base, dark
purplishto blackish brown on the abaxial side, light
brown with faint marksof seed wings on the adaxial
side. Apophysisraised in the central part, rhombicto
pentagonal in outline, mostly symmetrical around
the cone, transversely keeled, ochraceous to light
reddish brown. Umbo dorsal, raised and slightly
recurvedor nearly flat, rhombic,transverselykeeled,
4-8 mm wide, grey, with a minute,deciduous prickle.
Seeds obliquely ovoid, slightly flattened,4-7 x 35 mm, light brown with black spots. Seed wings
articulate but effective, held to the seed by two
flimsy claws, obliquely ovate, 12-20 x 4-8 mm,
semi-transparent,greyish brown with black tinge.
Numberof cotyledons 10-12.
Distribution (Maps 6, 7). SW United States: Ari-
105
zona, New Mexico. Mexico: Mainly in the Sierra

Madre Occidental, south to S Durango, scatteredin
Coahuila, NE Zacatecas, and Nuevo Le6n.
Remarks on the species. Pinus arizonica was
based on a collection made by J. T. Rothrockwhile
on one of the U.S. Army Survey expeditions in the
Santa Rita Mountains of southern Arizona. Engelmann (1879) stressed the difference in leaf number
with P ponderosa (5 vs. 3), but found it otherwise
very similar to certain forms of that species. Shaw
(1909, 1914) reduced it to a variety of P. ponderosa
when it became known that the "leaf-fascicles (are)
heteromerous,with the largernumberin the southern
partsof its range"(referringto P ponderosa s.l.). As
discussed under P ponderosa var. scopulorum, P.
arizonica has often been treated under that species
as a variety,but more recent treatmentsrecognize it
as a distinct species, with possible introgressionin
Arizona and New Mexico. Table IV compares and
contrasts the morphological characters of the two
taxa.
The cones, otherwise very similar, are generally
somewhat smaller in P arizonica, but here again
considerable overlapping is found and with no observed clinal trend.
Arbor.1: 24. 1909;PinusponderosaD. Douglasex P.

Lawsonsubsp.arizonica(Engelmann)
E. Murray,
Kalmia 12:23. 1982.Type.UnitedStates.Arizona:Santa
RitaMts., 1874,J. T Rothrock
652 (holotype.MO).
Tree, usually tall, height to 30-35 m. Leaves in
fascicles of 3-5 (predominantly3-4), rigid to slightly
lax, straightor slightly curved and twisted, (8-)1020(-23) cm x 0.9-1.4(-1.6) mm. Stomata on all
faces of leaves, in (3-)4-8 lines on the convex
abaxial face and in (3-)4(-5) lines on each adaxial
face (the abaxial number of lines correlated with
width of leaf). Seed cones ovoid to broadly ovoid,
often slightly curved, (4.5-)5-7 x 3.5-6 cm when
open. Seeds obliquely ovoid, slightly flattened,4-6
X 3-3.5 mm. Seed wings obliquely ovate, 12-15 x
4-6 mm.
Remarks. There seems to be clinal variation in
the numberof leaves per fascicle, with trees having
predominantly3(-4) leaves more frequentin Sonora,
Chihuahua,and Coahuila;those with predominantly
5 leaves are more frequent in Durango. Exceptions
do occur: 3-leaved in Sinaloa: H. S. Gentry 7242
(MICH);(4-)5-leaved in Sonora:J. T. Marshall 134
(UC), S. S. White3360 (MICH).
Distribution and ecology. SW United States:Arizona, New Mexico. Mexico: Mainly in the Sierra
Madre Occidental south to S Durango, scatteredin
Key to the varieties of Pinus arizonica
Coahuila, NE Zacatecas, and Nuevo Le6n. Forming
1. Leaves long (14-25 cm), thick (1.4-1.8 mm), with
pure stands or more commonly mixed with Quercus
8-12 lines of stomataon the abaxial side, 3-4(-5)
spp., other pines (e.g., Pinus engelmannii,P strobiper fascicle......................................... IOc.var. stormiae
formis), and occasionallyJuniperusflaccidaat lower,
1. Leaves short or variable,<1.4(-1.6) mm thick.
or J. deppeana at higher, altitudes. On various sub2. Leaves variable in length (8-)10-20(-23) cm,
in fascicles of 3-5, predominantly3-4............
strates, but the best stands in valleys and on mesas
..........................................
10a. var. arizonica with
deep soil, in moderately dry to mesic forest
2. Leaves short, (5-)6-10(-12) cm, in fascicles of
with
light winter frost occurring, at (1300-)20005................
lOb.
var.
(3-)4-5, predominantly
cooperi
2700(-3000) m. Annual precipitationis low to moderate, 700-900 mm, mostly falling duringthe winter
10a. Pinus arizonica Engelmannvar. arizonica
months. Phenology: Pollen is dispersed in springFig. 25
time, dependenton altitude and latitude;exact time
Pitnus ponderosa D. Douglas ex P. Lawson var. arizonica
not recorded.
(Engelmann)G. R. Shaw, [Pines Mexico] Publ. Arnold
Representative specimens examined. MEXICO. CHI-
HUAHUA:
SW of San Juanito,10 Aug 1954,
Mojarachic.
Table IV
Characterstates in the genus Pinus
Character
Leaves per fascicle
Widthof leaves
Stomatallines on abaxial
side of leaf
Resin ducts in leaf
Seed scales at maturity
Knobloch 1164 (MICH); El Vergel, 5 Oct 1960, Little

17899 (US); Barrancade Cobre, 7 Oct 1960, Little 17914
(US); MesaHuracan,8 Oct 1960,Little17936(US); ca.

65 km SW of CasasGrandes,2 Jul 1952. Marshall54
P arizonica P ponderosa
(UC);Madera,nearthe Rio Colorado,7 Jul 1958,Matuda
var. scopulorum 32667 (MEXU);vic. of Madera,May-Jun1908,Palmer
449 (MO);Ocampo,CascadaBasaseachicNationalPark,
3-5
2-3
0.9-1.8 mm
1.3-1.6 mm
(3-)4-8(-12)
(3-)5-10
Thick, rigid
8-10
2-6
Thin, flexible
12 Nov 1989, Spellenberget al. 10083 (FHO, NMC);Cerro
Mohinora,16 km S of Guadalupe
y Calvo,11 Aug 1960,
Straw & Forman 1968 (MICH);Cerro Mohinora, 16 km S
of Guadalupey Calvo, 12 Aug 1960,Straw& Forman

2003 (MICH);63 km W of CasasGrandes,29 Aug 1952,
Tucker2582 (UC). COAHUILA:
Cuatro Ci6negas, Canon
106
Flora Neotropica
Z
';
'
'
rai', ~'"'
~:(::::'''"
Os
Cb
"';''~";"';'":;~"as~
9~~~~~~~~~~~';M,'':'i
FIG. 25. Pinus arizonica var. arizonica (A-C, Straw & Forman1968; D, Pringle 10139). A. Branchwith leaf fascicles.
B. Leaf fascicles. C. Cross sections of leaves in (a) fascicle of 5 and (b) fascicle of 3. D. Ovuliferouscones: (a) immature
cone, (b) mature cone, (c) seed scale (three views), (d) remnants of cone bases on branch. E. Seed with wing (two
views). (Magnifications:A, B, D, E, X0.5; C, x30.)
del Agua, 8 Sep 1939, Mueller 3208 (MO, UC, US);
"mountainsnear General Cepeda," 7 Oct 1905, Pringle
10139 (E, GOET, MEXU, MO, NY, UC). DURANGO:
50
km WSW of El Salto at trail S to "Mexiquillo,"31 Aug
1951, Mavsilles 7580 (MICH);Cerro Huehueto,S of Huachicheles, 23 Jul 1955, Maysilles 7981 (MICH); El Salto,
4 Aug 1955, Mavsilles 8311 (MICH);Lagunadel Progreso,
58 km N of railroadat Coyotes, 10 Aug 1955, Maysilles
8337 (MICH);ChavarriaNuevo, 3 Apr 1991, Styles, Favela
et al. 9 (FHO, MEXU). NUEVOLEON: Unknown loc..
1900, Nelson 4502 (A). SINALOA:Sierra Surotato, Los
Pucheros,canyon below Los Pefiascos, 17 Mar 1945, Gentry 7242 (MICH, NY); "SantaLucia," Sierra MadreOccidental, 16 Nov 1926, Mexia 496 (MO). SONORA:Sierra
Huachinera,nearsummiton E slope, 18 Jun 1953, Marshall
134 (UC); NW of Aribabi, Puerto de Hu6pari,in curve of
Rio de Bavispe, 7 Sep 1939, White2748 (MICH); Puerto
de los Asseraderos,4 Aug 1940, White 3176 (MICH); El
Temblor,in the Sierra El Tigre, 18 Aug 1940, White3360
(MICH).
0Ob. Pinus arizonica Engelmann var. cooperi
(C. E. Blanco) Farjon, comb. et stat. nov.
Fig. 26A-D
Pious cooperi C. E. Blanco, Anales Inst. Biol. Univ. Nac.
Mexico 20: 185. 1950; Pinus lutea C. E. Blanco ex
Martinez,Anales Inst. Biol. Univ. Nac. Mexico 16: 233.
1945, non Walter,1788, nec G. Gordon& Glendinning,
1858. Type. Mexico. Durango:El Salto, Mar 1940 by
C. E. Blanco, M. Martinez 3442 (no original material
found in MEXU; lectotype, A, here designated).
Pinus lutea C. E. Blanco ex Martinezvar.ornelasi Martinez, Anales Inst. Biol. Univ. Nac. Mexico 16: 237.
1945; Pinus cooperi C. E. Blanco var.ornelasii (Martinez) C. E. Blanco, Anales Inst. Biol. Univ. Nac. Mexico
20: 185. 1950. Type. Mexico. Durango: El Salto, M.
Martinez3441 (holotype, MEXU).
Tree, usually tall, height to 30-35 m. Leaves in
fascicles of (3-)4-5,
predominantly 5, rigid to
lax,
curved,
(5-)6-10(-12) cm X 1usually
slightly
1.3 mm. Stomata on all faces of leaves, in 4-7 lines
on the convex abaxial face and in (3-)4 lines on
each adaxial face (the abaxial number of lines correlated with width of leaf). Seed cones ovoid to broadly
ovoid, often slightly curved, 5-10(-12) X 4-6 cm
when open. Seeds obliquely ovoid, slightly flattened,
5-7 x 4-5 mm. Seed wings obliquely ovate, 15-20
X 6-8 mm.
Distribution and ecology. Mexico: Mainly in Durango, but scattered northward in the Sierra Madre
Occidental into Chihuahua. At least in part sympatric
with var. arizonica. Similar habitat and altitudinal
range (mainly 2000-3000 m); associated pines are,
e.g., P. durangensis, P. leiophylla, P. strobiformis, P.
teocote.
Representative specimens examined. MEXICO. CHI.
HUAHUA:
Bocoyna, SW of Creel near San Ignacio, 17 Oct
107
1977, Bye & Weber8298 (E, MEXU, NCU). COAHUILA:
Serraniasdel Burro, 7 Jun 1983, Marshall 83-28 (FHO).
DURANGO:
La Ciudad, 30 Sep 1960, Courbasson & Mahieux s.n. (P); El Salto, 23 Nov 1961, G6mez P. 612
(MEXU); El Salto, 18 Sep 1961, G. Guzmdns.n. (MICH.
SD); 5 km E of El Salto, 8 Jul 1958, Hendricks513 (MO);
58 km W of Durangoalong Hwy. 40, 7 Mar 1966, Hess &
Hall 554 (MICH); 10 km E of La Ciudad, 30 Apr 1987,
Hughes 977 (FHO, K, MEXU); Cerro Huehueto, 2 Jul
1950, Mavsilles 7262 (MICH);Cerro Prieto, ca. 35 km W
of Otinapa, 10 Jul 1950, Mavsilles 7375 (MICH);Coyotes,
10 Aug 1952, Mavsilles 7697 (MEXU, MICH); Cerro
Chupadero,W of Pueblo Nuevo, 28 Jul 1955, Mavsilles
8079 (MEXU, MICH); Lagunadel Progreso, 55 km N of
railroadat Coyotes, 10 Aug 1955, Maysilles 8152 (MICH);
WSW of El Salto, between Las Adjuntas and Rucias, 4
Aug 1955, Maysilles 8312 (MEXU, MICH); El Salto, 24
Aug 1957, Solbrig & Ornduff4637(NY); ChavarriaNuevo,
5 Apr1991,Styles,Favelaet al. II (FHO);Coyotes.5 Apr

1991, Styles, Favela et al. 17 (FHO); El Madrofo, 5 Apr
1991, Styles, Favela et al. 18 (FHO, MEXU).
lOc. Pinus arizonica Engelmannvar.stormiae Martinez, Anal. Inst. Biol. Univ. Nac. Mex. 16: 284.
1945.
Fig. 26E-I
PinusponderosaDouglas ex Lawsonvar.stormiae(Mart-
inez) Silba, Phytologia68: 59. 1990.Type. Mexico.

Coahuila:Sierrala Corcordia
de los Patos")S
("Sierra
of GeneralCepeda[locationis not in accordance
with
thatgivenin Martinez
Coahu(1948)forthisspecimen:
ila, Mun.Arteaga,Las Margaritas;
probablyM's numbershavebeenmixedup],Jul 1941,M. Martinez3455
(lectotype,MEXU,heredesignated).
Tree of medium size, height to 15-20 m. Leaves

in fascicles of 3-4(-5), thick, coarse, rigid, usually
curved and twisted, 14-25 cm X 1.4-1.8 mm. Stomata on all faces of leaves, in 8-12 lines on the
convex abaxial face and in (3-)4 lines on each
adaxial face (the abaxial numberof lines correlated
with width of leaf). Resin ducts in the leaf 3-5(-8).
Seed cones ovoid to ovoid-oblong, often curved,
(4.5-)5-10(-14) x 3.5-6(-8) cm when open. Seeds
obliquely ovoid, slightly flattened,5-6 x 3.5-4 mm.
Seed wings obliquely ovate, 12-16 x 6-8 mm.
Distribution and ecology. Mexico: Mainly in S
Nuevo Le6n, a few localities in S Coahuila, Zacatecas, and possibly San Luis Potosi (dot map probably incomplete, but materialfrom several localities
attributedto var.stormiaehave here been determined
as var. arizonica). Habitat similar to var. arizonica,
but often recorded from more arid sites; forming

pure stands or mixed with Quercus,Juniperus,and/
or other pines, e.g., P cembroides (especially dry
sites) and P pseudostrobus. Altitudinal range ca.
1500-3000 m.
108
Flora Neotropica
,,{
j,,
arizonca var.
& Favela 11). E-.
A-D. Pinus arizonica var. cooper (A-C, Masilles 8152;
stormiae (E-G, Zobel 18; H, 1, Zobel 20). A. Branch with leaf fascicles. B, F. Leaf fascicles. C, G. Cross sections of
leaves. D. Ovuliferous cone. E. Branch with bud, immaturecones and leaf fascicles. H. Ovuliferous cone: (a) mature
cone, (b) seed scale (two views). 1. Seeds with wings. (Magnifications:A, B, D-F, H, 1, x 0.5; C, G, x 30.)
P
FIG. 26.
D. Styles
109
Representative specimens examined. MEXICO. is therefore more in accordance with the evidence
NUEVO
LEON:3 km S of Pablillo,20 Jul 1958,Correll& presentlyavailable.
Johnston19914(NY);Canondel Aserradero,
Miquihuana, Martinez (1948) recognized Pinus arizonica var.
Nov 1939,M. Martinez3455 (numberprobablyincorrect,
see above under type) (NY); Galeana, 20 km E of San stormiae, which he described as different from P
Roberto,4 Nov 1983, McCarter& Hughes105 (FHO); arizonica in its larger (longer and wider) leaves,
Puertode Reyna,14Jul1977,Mittak8492(FHO);Galeana, fewer resin ducts in the leaves, and largercones. The
14 Jul 1977, Mittak 8493 (FHO); I km from Iturbide, 12 other characterstates mentioned seem to be closer
Mar 1980, Stead & Styles 614 (FHO, MEXU); 30 km from
to P ponderosa var. scopuloruinthan to P arizonica
Linaresalongrd.to Iturbide,13 Mar1980,Stead& Styles
628 (FHO);Ascensi6n,Dec 1955,Zobel18 (UC);Galeana, var. arizonica, but Martinez did not consider P
Dec 1955, Zobel 20 (UC).
ponderosa as occurringin Mexico. Unlike var. cooperi, var. stormiae has a more disjunct distribution
Uses. Pinus arizonica is an importantconstituent from the mainly western range of the species.
of the pine and pine-oak forests of northernMexico,
especially in the Sierra Madre Occidental. As such 11. Pinus jeffreyi J. H. Balfour in A. Murray,Bot.
it is heavily exploited as a timbertree in much of its
Exped. Oregon 8: 2. 1853.
Fig. 27
range, especially in the more accessible regions, and
PinusponderosaD. Douglasex P. Lawsonvar.jeffreyi
stands with large trees in these localities are now
(J. H. Balfour)Vasey,Cat.ForestTreesU.S. 31. 1876
rare (Perry, 1991).
[& U.S. Commr.Agric.Rep. 1875:179. 18761;Pinus
ponderosaD. Douglasex P. Lawsonsubsp.jeffreyi(J.
Three varieties of Pinus arizonica have been recH. Balfour)E. Murray,Kalmia12: 23. 1982. Type.
ognized, based mainly on a combination of leaf
UnitedStates.California:
ShastaValley,("Chastey
Valcharacterswhich are quantitativeand mostly continuley,Lat.41?30'N"),24 Oct 1852,Jeffrey731(holotype,
ous. We have not found any distinct charactersto
E; a poorspecimenwith4 leaf fascicles,a budanda
brokenseed anda few seed wings).[Epitype:United
separate P. cooperi C. E. Blanco from P arizonica
States.California:
TrinityCo., ScottMt.,2 Sep 1958,
Engelm. to consider at the rankof species.
Alava et al. 2439A (E), here designated].
Martinez(1945) gave a Latindescriptionof a new
Pinusdefle.raJ. Torreyin Emory,U.S. Mexic.Bound.
Surv. Rpt. 2(1): 209, t. 56. 1859; Pinus jeffreyi J. H.
species of pine from Durango (El Salto). It was
Balfourvar.deflexa(J. Torrey)J. G. Lemmon,Calif.
based on a specimen, accompaniedby a description,
StateBoardForest.Bien.Rep.2: 74, 100. 1888.Type.
sent to him by C. E. Blanco under the suggested
UnitedStates.California:
"Mountains
E of SanDiego,"
name Pinus lutea, presumablynamed for its yellow
1850,Parrys.n. (lectotype,NY,heredesignated).
wood. This name is a later homonym and for this
Pinus jeffreyi J. H. Balfour var. baja-californica Silba.
reason the new species was renamed P. cooperi by
68:52. 1990.Type.Mexico.BajaCalifornia
Phytologia
Norte:BetweenOjosNegrosandNeji Rancho,16 Sep
Blanco (1950). A search by A. Farjon for the type
1929, 1. L Wiggins & Gillespie 4123 (holotype, NY;
specimen in MEXU has been unsuccessful.Martinez
isotypes, F. MEXU. MICH, SD).
P
this
with
rudis
(=
(1945, 1948) compared
species
P hartwegii) and P. montezumnae, from which it is
Tree, medium to large, height to 20-30 m, dbh to
more distinct morphologically (especially in cone 100 cm. Trunk monopodial, straight, terete. Bark
characters)than from P arizonica. Martinez(1945) thick, rough and scaly, divided by deep, longitudinal
also proposedthe variety P lutea var.ornelasi, being and anastomosing fissures into thick, elongated
a 5-leaved form with dark green, longer leaves and plates, light buff to light brown, the fissures dark
largercones. We found these charactersto be incon- grey-brown,on young trees and branchesrough and
sistent (e.g., Styles, Favela et al. 17 (FHO) with scaly, orange-brown.Branches of first order long,
leaves 6-8 cm and a maturecone 8 cm long; Styles, slender, spreading horizontally or assurgent at the
Favela et al. 18 (FHO) with leaves 11-13 cm and a ends, persistent; branches of higher orders sparse,
cone 5 cm long). This taxon was later recombined curved or assurgent,forming an open pyramidalor
by Blanco (1950). Unlike Perry(1991: 109, t. 3.10), flat-topped crown. Shoots stout, assurgent, very
we do not think that the bark between these taxa rough and scaly with persistent, decurrentpulvini,
differs consistently in the manner he described, light orange-brown,often glaucous. Cataphyllslarge,
which follows from comparison of his published ca. 20 mm long, triangular-subulate,
reflexed, scariphotographswith those given by Martinez (1948: ous, with erose-ciliatemargins,light brown,weatherfig. 195) alone. Other differences, such as those ing greyish black. Vegetativebuds ovoid-acute or
mentionedfor several cone characters,have not been subglobose, the terminal buds 15-20 mm long, the
confirmedin the collections we have studied.Treat- laterals smaller, not resinous; the scales imbricate,
ment of P cooperi as a variety of P arizonica subulate,with free apices and ciliate, white margins,
110
Flora Neotropica
\,s/
t?
aC
2D
FIG. 27. Pinus jeffreyi (A, B, Hughes & Styles 170: C-E, Hughes & Styles 181). A. Branch with leaf fascicles and
cataphylls. B. Leaf fascicle and cross section of leaf. C. Ovuliferouscone. D. Seed scale (three views). E. Seed. (Magnifications:A, C-E, x 0.5; B. x25.)
light orange-brown.Fascicle sheaths of young leaves

20-25 mm long, with 8-10 pale orange-brown,imbricate scales, persistent but reduced to 10-15 mm
in older fascicles, weatheringgrey-brownto blackish.

Leaves in fascicles of 3, sometimes a few fascicles
of 2, in dense, spreading tufts on thick branchlets,
persisting(3-)4-5(-6) years, thick, rigid, straightor
slightly curved, sometimes twisted, (12-)15-22(-25)
cm x 1.5-1.9(-2) mm, with serrulate margins,
acute-pungentto acuminate,light yellowish green to
greyish green. Stomata on all faces of leaves, in 711 grooved lines on the convex abaxial face, in 3-5
conspicuous lines on each adaxial face. Leaf anatwith a conomy: Cross section transverse-triangular,
vex abaxial side; epidermis thick; hypodermismultilayered, slightly intrudinginto the mesophyll; resin
ducts (2-)3-5, medial, the three marginalducts larger
than any subsidiaryducts; stele oval in cross section;
the outer walls of endodermalcells thickened;vascular bundles 2, distinctly separated. Pollen cones
densely clustered near the proximal end of a new
shoot, subtendedby subulatebracts,ovoid-oblong to
cylindrical, 20-35 x 6-7 mm, yellow or purplish
yellow, maturinglight brown. Seed cones subterminal, solitary or in pairs on short, stout, persistent
peduncles, spreadingand seemingly sessile at maturity, leaving a few basal scales on the branch when
falling. Immaturecones broadly ovoid, ca. 30 X 20
mm, with spreading spines, purplish, soon light
brown, maturing in two seasons (16-18 months).
Mature cones broadly ovoid to subglobose, with a
slightly oblique, flattenedbase, (10-)12-17 X 9-14
cm when open. Seed scales ca. 150-175, in a low
spiral, parting soon and wide, thin woody, straight
or slightly curved, up to 20 mm wide, light or dark
brown abaxially, with distinct, lighter marksof seed
wings on the adaxial side. Apophysis slightly raised,
transverselykeeled, broadly rhombic in outline, on
the proximal scales more gibbous, often resinous,
ochraceousto light brown. Umbo dorsal, moderately
raised, transverselykeeled, terminatingin a distinct,
curved, 3-5 mm long spine. Seeds obliquely ovoid,
slightly flattened, 9-12 mm long, light yellowish
brown with faintly darker spots. Seed wings articulate, effective, held to the seed by two thin claws,
oblique, with a straightand a curved side, 20-25 X
10 mm, light yellowish brown to light brown. Number of cotyledons 7-13, usually 10.
Remarks on morphology. The above description
pertainsto Pinus jeffreyi in Mexico only. Generally,
both tree size and sizes of leaves and cones can be
larger in the United States, especially in the Sierra
Nevada of California. There trees up to 50 m or
more occur, to which figures given by Martinez
(1948) and Perry (1991) must be referring,although
theirbooks deal with Mexican pines. Similarly,cones
in the Sierra Nevada can be up to 25-30 cm long,
but we have not seen such large cones on our visits
to the populationsof this tree in Mexico. From more
111
Q
I
^V^
\ o
Map 8. Piniis jeffreyi.
northernregions, the persistence of leaf fascicles is

also reported(Bradshaw, 1941) to be longer, up to 8
years. The almost brick-redto red-brownbark seen
by A. Farjon in parts of the Sierra Nevada has not
been observed in the Mexican populations.
Primarily in California, extending into SW Oregon
and extreme W Nevada; its main distributionis in
the Sierra Nevada; there are disjunct populationson
mountainsin S California.Mexico: In Baja California Norte, scattered in the Sierra Juarez, but more
common in the northernpartof the SierraSan Pedro
Martir,where it reaches its southernmostlimit at ca.
30?40'N. Its altitudinalrange in the Sierra Juarez is
(1 100-)1400-1800 m; in the SierraSan Pedro Martir
it occurs considerablyhigher, 1800-2500(-2700) m.
Here it forms an open, mixed coniferous forest with
other conifers. Common are also Abies concolor,
Pinus lambertiana, and P contorta var. murravana
in the upper zone. In the Sierra Juarez it is often
associated with Quercus spp. Both mountainranges
112
Flora Neotropica
view (e.g., Kral, 1993) that the two taxa are sufficiently distinct to be recognized as species. Intermediate forms have been recognized as naturalhybrids
(Haller, 1962), but these are infrequent.Under controlled conditions, crossability is relatively low
(Burns & Honkala, 1990). As it has been established
that P. ponderosa is absent from Baja California
specimensexamined.MEXICO.BAJA (Duffield & Cumming, 1949), no such hybrids are
Representative
NORTE:Sierra Juarez, San Faustino, 15 km
CALIFORNIA
S of RanchoEl Compadre,off Tecate-Rumorosa
rd., 26 to be found there.
The more or less disjunct populations in Baja
181
Mun.
Ensenada.
Styles
&
(FHO);
May 1987,Hughes
La Cienega, Jan 1942, M. Martinez 2148 (NY); Sierra California have been described by Silba (1990), on
Juarez,80 km S of Tecate,Sep 1939,M. Martinez3451
narrow evidence, as P.
var.
are graniticand provide acid soils. Annual precipitation is moderate, up to 500 mm, of which about
half comes as winter snow at the higher altitudes.
Phenology: Pollen is dispersed in June-July, depending on altitude and related climatic circumstances.
(MEXU); 1.5 km N of San Faustino, Moran 16605 (SD);
baja-califorjeffreyi
very
nica. His observation that the leaves are "mostly
SierraSan PedroMartir,20 May 1893, Brandegees.n.

(UC); 2 km N of Vallecitos,1 Feb 1971, Echenique46 two per fascicle" is contradicted by the holotype
(MEXU);SierraSanPedroMartir,
alongrd.to observatory, specimen (/. L. Wiggins & Gillespie 4123, NY) and
1790m, 25 Apr 1987,Hughes& Styles170 (FHO),2320 by all other materialwe have seen in and from Baja
m, 25 Apr 1987,Hughes& Styles175 (FHO,K); Sierra California, in which
only an occasional fascicle has
s.n.(US);LaEncantada,
SanPedroMartir,1889,Townsend
leaves.
Also
the leaves not wider than in
2
are
4920
Demaree
&
18 Sep 1930, Wiggins
(F, NY, US).
specimens from California.Cone size appearsto be
Uses. Although not an importanttimber tree in smaller on average, but this character is of little
Mexico, its geographical isolation from other Mexi- taxonomicvalue (Duffield & Cumming, 1949). Vidacan pine forests makes it of local or regional impor- kovic (1991) mentions a variety"peninsularis"from
tance. Especially in the Sierra Juarez, exploitation Baja California with "darkergrey and deeply furmay pose a threat to the limited and more scattered rowed bark"but does not cite a valid publicationof
populations. Many of the stands in the Sierra San this name.
Pedro Martirare within a national park and reason12. Pinus engelmannii Carriere,Rev. Hort., ser. 4,
ably well-protected.
Pinus jeffrevi was found by the Scottish plant
3: 227. 1854.
Fig. 28
collector John Jeffrey in northern California, in
Pinusmacrophylla
Engelmann,non Lindley(1839), in
Shasta Valley, on 24 October 1852. He had been
Wislizenus, Mem. Tour N. Mexico 103. 1848; Pinus
the
of
Botanical
sent out by the Committee
Oregon
ponderosa D. Douglas ex P. Lawson var. macrophylla
Expedition, on behalf of its subscribers,to collect
(Engelmann)G. R. Shaw, [Pines Mexicol Publ. Arnold
Arbor. 1: 24. 1909. Type. Mexico. Chihuahua:Cosiquirseeds of exotic plants. When the first batch of mateiachi, Oct 1846, Wislizenus 233 (lectotype, MO
rial finally arrived in Edinburghin the summer of
3377671, here designated; isolectotypes, MO 3377672,
1853, a list of its contents, including new species
MO 3377673, MO 3534934).
with descriptions,was publishedas a pamphlet(No.
Pinus latifolia C. S. Sargent, Gard. Forest 2: 496, fig.
135. 1889; Pinus lmavrianaSudworth, U.S.D.A. Div.
8) "by order of the committee," by its secretary,
Forest. Bull. 14: 21. 1897; Pinus ponderosa D. Douglas
Andrew Murray.Although Murray(1853) ascribes
ex P. Lawson var. mavriana (Sudworth)C. S. Sargent,
the new conifer taxa to Balfour (who was the chair;
Silva N. Amer. II: 81. 1897. Type. United States.
Greville, often cited as co-author, did the illustraArizona: S slopes of Santa Rita Mts., 1887, H. Mavr
tions), the authoritywith the new species is cited as
s.n. (holotype, A).
"Oreg. Corn.," which has led to the interpretation Pinus apacheca J. G. Lemmon. Erythea 2: 103, t. 3.
1894. Type. United States. Arizona: Chiricahua Mts.,
that Murray,not Balfour, was the validating author
Jun 1892, Lemmons.n. (lectotype, A, here designated).
in
here
revised
view
is
accorThis
1993).
(Farjon,
Pinus macrophyllaEngelmannvar.blancoi Martinez,Andance with Art. 46.2 of the Tokyo Code (Greuteret
ales Inst. Biol. Univ. Nac. Mexico 15: 345. 1944,
al.. 1994). The holotype, Jeffrey 731 (E), is such a
"'macrofila";Pinus engelnannii Carriere var. hlancoi
(Martinez) Martinez, Pinos Mexic., ed. 2, 288. 1948.
poor specimen that use has been made of the new
Mexico. Durango:(Near) El Salto, Aug 1940, M.
Type.
to
in
the
an
Code
(Art.
9.7)
designate
provision
Marlinez 3450 (lectotype, MEXU 2509, designated by
epitype to accompany it.
Carvajal& McVaugh, 1992: 59).
Formerly classified as a variety under P ponderosa, more detailed research(Bradshaw,1941; JoTree, medium to large, height to 20-25(-27) m,
hansen, 1953) has led to the now generally accepted dbh to 70-90 cm. Trunk monopodial, straight, erect,
113
\\
\\\\
\\
FIG. 28. Pinus engelmrannii(A-C, Mueller 3436; D, Pringle 13614; E, F, Mexia 2515; G, Styles, Far'elaet al. 3). A.
Branchwith leaf fascicles. B. Leaf fascicles. C. Cross section of leaf. D. Leaf fascicles and immatureovuliferous cone.
E. Ovuliferous cone. F. Seed scale (three views). G. Seed (two views). (Magnifications:A, B, E-G. x0.5; C, x 20; D.
x 1.)
114
usually free of branchesonly in the lowest part('/4'I/) or to '/2 of height. Bark thick, rough, scaly,
divided into long, irregularplates by wide, shallow,
dark fissures, reddish brown on branchesand young
trees, then dark brown, weathering grey. Branches
of first order thick, long, spreadingor assurging;of
second and higherorderssparse,stout, spreadingand
assurging, curved, forming a broad, rounded, open
crown. Shoots stout, very rough, with large, decurrent, persistentpulvini. Cataphyllslarge, 15-20 mm
long, subulate, recurved, with scarious lamina and
erose-ciliate margins, dark brown to blackish, often
caducous. Vegetativebuds large, the terminal buds
20-30 x 15-20 mm, ovoid-conical, the laterals
smaller, not resinous; the scales spreading or reflexed, subulate-caudate,with long-ciliate margins,
brown. Fascicle sheaths persistent and remaining
long, (15-)25-35(-40) mm, consisting of 10-12 imbricate, distinct bracts, reddish brown with greyish
ciliate margins, weathering brown to blackish.
Leaves in fascicles of (2-)3(-4), rarely 5, in dense,
spreading tufts toward the ends of branches, persisting 2-3 years, thick, rigid, straight or slightly
drooping, (18-)20-35 cm x 1.5-2 mm, margins
serrulate, apex acute-pungent, light (yellowish)green. Stomata on all faces of leaves, in (6-)8-15
lines on the convex abaxial face and 4-8 lines on
each adaxial face. Leaf anatomy:Cross section trianwith a convex abaxial
gular or transverse-triangular,
side; hypodermis multi-layered, slightly intruding
into the mesophyll; resin ducts (4-)5-8(-14), medial,
the 3 marginalducts usually largerand always present; stele oval in cross section; outer walls of endodermalcells not or slightly thickened;vascularbundles 2, separate or the xylem strands more or less
connate, always distinct. Pollenlcones clustered near
the proximal end of a new shoot, subtended by
subulate bracts, ovoid-oblong to cylindrical, up to
40 mm long and 12 mm wide, yellowish pink, turning
yellowish brown. Microsporophyllswith a 1.5 mm
wide peltate, more or less cordate distal part, which
appearsradially striated.Seed cones subterminal,in
pairs or in whorls of 3-5, on thick, short, curved
peduncles,persistingsome time, leaving a few scales
on the branch when falling. Immaturecones ovoid,
with prominent, spreading or slightly recurved
spines, light brown, often glaucous, maturingin two
seasons. Maturecones seemingly sessile, ovoid-oblong, curved, with an oblique base when opened, 815 x 6-10 cm when open, often resinous. Seed
scales ca. 100-140, parting to release the seeds
except at the base, oblong, straightor recurved,thick
woody, with irregularlyundulating,thin margins,up
Flora Neotropica
to 20 mm wide, dark purplishor blackish brown on
either side, with light brown marks of seed wings
on the adaxial side. Apophysis prominently raised,
transverselykeeled, often recurvedand slightly larger
on one side of the cone, rhombic or pentagonal in
outline, ochraceous or light brown, often with dark
radial lines or fissures. Umbo dorsal, large, rhombic
to transverse-rhombicin outline, with a persistent,
curved spine up to 3 mm long, slightly darkerbrown
than apophysis, weathering grey-brown. Seeds
obliquely ovoid, slightly flattened,keeled, 5-8 X 45.5 mm, light grey-brown, often with dark spots.
Seed wings articulate,effective, held to the seed with
two claws or (partly)covering the seed on one side,
ovate-oblong to obliquely ovate, 18-25 x 7-10
mm, translucentlight-ochraceous,often with a darker
tinge nearthe apex. Seedlings: Numberof cotyledons
8-10; a conspicuous "grass stage" is reported for
this species (Bailey & Hawksworth, 1983).
SE Arizona and extreme SW New Mexico. Mexico:
Extending south from the populations in the SW
United States through the Sierra Madre Occidental
in Sonora, Chihuahua, NE Sinaloa, Durango, and
more scattered in Zacatecas. It has been cited from
the SierraFriabetween ZacatecasandAguascalientes
(see Critchfield & Little, 1966: map 49), but no
collections from there have come to our attention;
Perry (1991) could not find it there. It also appears
to occur in Nuevo Le6n, in the mountains near
Galeana, from where two collections could be verified. Other reportsof disjunct populationsappeared
to be based on specimens of Pinus durangensis or
could not be verified and have been omitted. Pinus
engelmannii occurs on moderately dry, summerwarm open mountainslopes or plateaus at (1200-)
1500-2700(-3000) m, most abundantly at 20002500 m. It occurs on poor rocky (volcanic) soils as
well as on alluvial coarse sand/gravelor loamy sand.
The climate is temperate,with annual rainfall 400700 mm, increasingsouthward.Above 2000 m, frost
and snow are common in winter. It is a constituent
of open pine and pine-oak woodland, sometimes of
mixed pine forest, with, e.g., P leiophylla, P lumholtzii, and P pseudostrobus, on drier sites with P
cembroidesand Juniperussp., and usually with various species of Quercus present. Phenology: Pollen
dispersal is reportedto occur in May (Arizona); the
time is likely to be dependenton altitudeand can be
some weeks later at the highest elevations.
Guasarachic, 26 Sep 1898, Goldman 163 (A,
115
SystematicTreatment
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~?,
?
t~~~~~~~~~~~~~?13
1~~~~~~~~~~~~~~~~~~~~~~~~~~~'
15~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
?
:..' .:
'i~?
IVr
'l
'
,i .,,,,.UIL
''
...I..
?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
..
I:
r
r
?......
"~
F1~
N
'
rl??L!???1-?T-?
,.
.......
~ ~
,......,,..,.....
~~~~~~~~~~~~~~~~~~~~~~~~~~.'
'~
~ ~
~~~~~~~~~~~~~~~.......~
"?~~~~~~~~~~~~~~~~~~
''''
t ~~~~..
...
?~~~~~~~~~~~~':...
..
?t?
r~
...
."
-.
SCUM~~~~~~ i~~~~~~~~~-.
?
i
,~~~~~~~~~~~~~~~~~~~~~~~~~~~~
?r~~~~~~~~~ru~~~~~
' ld~a~l
'I"~~~
.......
-.I
'
'
'
I
I
"
~I
enelani
Map 9. Piu (crls;Pd?lsaa(rage)
i
'
......
.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~;
~~
~~~~~~~~~N'
'
rrrar ~~i...
..
?'''
(triangles).
Map9. Pin.rse.lgehmna.ii(circles);P:douglaksia.al
US); in Sierra Madre SW of Colonia Judrez, Dec 1906,
Goodding 2108 (A, GH, MO), Dec 1906, Goodding 2109
(UC); W of Chuhuichupa,26 Aug 1936, LeSueur551 (UC);
Sierra San Luis, I I Jul 1892. Mearns 424 (US); Basuchil,
near Cerro de la Ventana, 16 km NW of Miifaca, 10 May
1929, Mexia 2515 (A, E, MICH, MO, NY, P. UC); Arroyo
Ancho, 7 Jun 1929, Mexia 2608 (A, BM, MICH, MO, NY,
UC); Canon Huhuatan,13 km SE of Madera,23 Sep 1939,
Mueller 3436 (GH, MICH, MO, UC); Guadalupe, 12 Sep
1898, Nelson 5013 (K); vic. of Madera, May-Jun 1908,
Palmer330 (GH, MO, NY); "gravelly plains at the base of
the Sierra Madre." Sep 1887, Pringle 1448 (A, MEXU,
MO, NY, US); Sierra Madre, 15 Oct 1888, Pringle 1673
(P, S, UC); ca. 43 km S of Creel along rd. to La Bufa, 6
Jul 1969, Soule 513 (MO); Cascada Basaseachic, 11 Nov
1986, Spellenherg et al. 10069 (FHO, NMC); Colonia
Garcia, 26 Sep 1899. Townsend & Barber 356 (FHO,
MICH,MO, NY, P); Majalca(Pilares), 11 Aug 1939, White
Sierra de la Madera, Canon
2344 (MICH). COAHUILA:
Charretera,I I Sep 1941, Johnston 8935 (MO). DURANGO:
43 km W of Durango, along Hwy. 40, 24 Jul 1958, Correll & Johnston 20120 (NY); Tepehuanes, 30 Jul 1944,
Fisher 44236 (MO, NY); along rd. from Durango to El

Salto, nearjct. to Otinapa.25 Mar 1978, Klaus s.n. (WU);
La Punta,Nov 1940. M. Martinez2346 (US); near El Salto,
Nov 1941, M. Martinez 2363 (NY); Cerro Prieto, 30 km
W of Otinapa, 10 Jul 1950, Maysilles 7376 (MICH); 10
km W of Durango, 5 Jan 1983, Neill 5314 (MICH. MO):
El Salto, 12 Jul 1898, Nelson 4568 (A, K, US); near
Durango at Santiago Papasquiaro,Apr 1896. Palmer 776
(A, B, F, K, MO, NY); E slope of Sierra de la Ulama,
Cienega de los Canelos, 1911, Patoni 22 (MEXU): Sandia
railway sta.. 13 Oct 1905, Pringle 13614 (MICH. US);
Agua Bendita, 1932, RamirezL. 22 (MEXU); 37 km SW
from Durango, 2 Apr 1991. Styles. Favela et al. 3 (FHO).
NUEVOLE6N: Near Galeana, 1 km N of 18 de Marzo. 29
Oct 1984, Rushforth495 (E); Pablillo, 8 Aug 1936, Tarlor
104 (MO). SINALOA:
SierraMonterrey,II Mar 1940. Gentr' 5904 (NY). SONORA:Cailon de Santa Rosa, E of El
Tigre, 16 Jul 1938, White 528 (MICH); Cation de los
Metates, 24 Jun 1940, White2829 (GH, MICH);Puertode
los Asseraderos, 4 Aug 1940, White 3175 (GH, MICH);
Rancho de la Nacha, 15 Aug 1941, White 3943 (MICH,
NY); Rancho del Roble. NE of El Tigre, Sep 1941. White
116
4266 (MICH, NY); summit of pass between Imuris and
Cananea,8 May 1948, Wiggins11684 (MICH. UC, US).
Flora Neotropica
either are inconsistent, with the leaves "por lo general
algo menas robustas" but still 2 mm wide (the maximum found in the species), or fall within the range
of character states found in material from widely
separated locations, such as cone colour and shape
of seed scales and apophysis. We agree with Perry
(1991) and Carvajal and McVaugh (1992) that Martinez's variety is insufficiently distinct for taxonomic
Uses. Harvested but apparently not specifically

selected as a timbertree in most of its range together
with other pines; in some areas depletion of larger
trees has been observed (by B. T. Styles).
Engelmann (1848) described a new species in
Pinus from "the higher mountains of Chihuahua" recognition.
with the binomial Pinus macrophylla,a later homonym of P macrophylla Lindley (1839). Carriere 13. Pinus hartwegii Lindley, Edwards's Bot. Reg.
(1854) renamedit and repeatedEngelmann'sdescrip25: 62. Aug 1839 [& Allg. Gartenzeitung 7: 324.
tion. The type specimen (Wislizenus233, MO) was
18391.
Fig. 29
collected in 1846. Sheet MO 3377671, with a label
Pinus nontezunaeA. B. Lambertvar.hartwegii(Lindley)
written by Engelmann, was considered to be the
G. R. Shaw, [Pines Mexico] Publ. Arnold Arbor. 1: 23.
holotype (Carvajal & McVaugh, 1992). Two more
1909. Type. Mexico. Michoacan: "Campanario,in the
sheets (MO 3377672 and 3377673), bearing Engelmountainsof Angangueo," 1838, Hartweg s.n. (lectotype, P, here designated;isolectotype, W).
mann's and Wislizenus's labels, respectively, and a
Pinus
ehrenbergiiEndlicher,Syn. Conif. 151. 1847; Pinus
very fragmentedcone (MO 3534934) can be considdecandolleana Roezl var. ehrenhergii(Endlicher) Carered isolectotypes, as they are likely to belong to the
riere, Trait6 Gen. Conif., ed. 2, 2: 570. 1867. Type.
same gathering.The cone is now entirely broken up
Mexico. Hidalgo: Mineral del Monte, Dec 1837, Ehrenherg 915 (holotype. W; isotype, HAL).
but the scales are typically thick woody and have
Pinus rudis Endlicher, Syn. Conif. 151. 1847; Pinus
prominent, recurved apophyses. Parlatore (1868),
montezulnae A. B. Lambert var. rudis (Endlicher)
who had received part of the type materialon loan,
G. R. Shaw, [Pines Mexico] Publ. Arnold Arbor. 1: 22.
included neither Engelmann'sspecies nor Carriere's
1909; Pinus hartwegii Lindley var. rudis (Endlicher)
new name in the text of his prodromus.Later,Shaw
Silba, Phytologia 68: 50. 1990. Type. Mexico. Locality
not stated, 1837-1838, Ehrenberg 1083 (lectotype, W,
(1909) treated it as a variety of P ponderosa under
here designated).
the broadest possible concept of that species, rePinus northumherlandianaRoezl, Cat. Grain. Conif.
taining Engelmann'sepithet.
Mexic. 13. 1857. Type. Mexico. Mexico: Mt. PopocateSargent (1889), Lemmon (1894), and Sudworth
petl, 1857, Roezl s.n. (lectotype, Fl, here designated;
isolectotype, P).
(1897) recognized that this pine was not identical
Pinus lowii Roezl, Cat. Grain. Conif. Mexic. 18. 1857.
with P ponderosa when it was encounterednorthof
Type. Mexico. Locality not stated, 1857, Roezl s.n.
the Mexican borderin Arizona,and they described it
(lectotype, Fl, here designated).
underthree different names: respectively,P latifolia
Pinus robustaRoezl, Cat. Grain.Conif. Mexic. 19. 1857.
(later dropped in favour of a treatmentas a variety:
Type. Mexico. Mexico: Ajusco Mts., 1857, RoezI s.n.
P ponderosa var. mayriana by Sargent, 1897), P
Pinus endlichericna Roezl, Cat. Grain.Conif. Mexic. 19.
two
The
P
and
first
binomials
mayriana.
apacheca,
1857. Type. Mexico: Ajusco Mts. Original materialnot
are based on different specimens from Arizona, and
located.
the latter is a nomenclaturalsynonym for P latifolia
Pinus wilsonii Roezl, Cat. Grain.Conif. Mexic. 19. 1857;
Pinus decaisneana Roezl var. wilsonii (Roezl) Carriere,
C. S. Sargent(Little, 1944).
Trait6Gin. Conif., ed. 2, 2: 544. 1867. Type. Mexico.
Martinez (1944), who had no access to much of
Hidalgo: Pachuca, 1857, Roezl s.n. (lectotype, Fl, here
the older Europeanliterature,still used Engelmann's
designated).
epithet for the species but correctedthis later (MartiPinus resinosatRoezl, Cat. Grain.Conif. Mexic. 30. 1857,
non Aiton, 1789; Pinus roezlii Carriere, Traite Gen.
nez, 1948) and published a new variety: P macroConif., ed. 2, 2: 575. 1867, nom. superfl.Type. Mexico.
phylla var. blancoi (= P engelmannii var. blancoi).
Locality not stated, 1857, Roezl s.n. (lectotype, Fl, here
The lectotype, designated "by default" (Carvajal&
designated).
McVaugh,1992) is M. Martinez3450 (MEXU 2509),
Pinus iztacihuatlii Roezl, Cat. Grain. Conif. Mexic. 30.
labeled "El Salto" and "var.nov." by Martinez,and
1857. Type. Mexico. Mexico: VolcanIztaccihuatl,1857.
RoezI s.n. (lectotype, FI, here designated; isolectotype,
probablycollected by C. E. Blanco in August 1940.
P).
Martinezapparentlywrote the labels afterwardand,
Pinus scoparia Roezl, Cat. Grain.Conif. Mexic. 30. 1857.
because of this method (see McVaugh, 1972), ended
up with a differenttype locality (Santiago Bayacora)
(lectotype, FI, here designated).
in the protologue. The minor differences described
Pinus standishii Roezl, Cat. Grain. Conif. Mexic. 30.
117
?0
FIG. 29. Pinus hartwegii (A-C, E, McCarter.Hughes & Syles 5; D, Stead 303; F, Stead & Styles 625). A. Branch
with leaf fascicles and terminal bud. B. Leaf fascicle. C. Cross section of leaf. D. Shoot with leaf fascicles and pollen
cones. E. Shoot with ovuliferous cone. F. Ovuliferous cone. G. Seed scale (two views) and abortedseed. H. Seeds with
articulatewings. (Magnifications:A, B, D-H, x0.5; C, x 25.)
118
1857. Type. Mexico. Locality not stated, 1857, Roezl
s.n. (lectotype, Fl, here designated;isolectotype, P).
Pious aculcensis Roezl, Cat. Grain. Conif. Mexic. 30.
s.n. (lectotype, Fl, here designated; isolectotype, P).
Pinus amecaensis Roezl, Cat. Grain. Conif. Mexic. 30.
1857. Type. Mexico. Mexico: Locality not stated, 1857,
Roezl s.n. (lectotype, P, here designated).
Pinus papeleuii Roezl, Cat. Grain. Conif. Mexic. 31.
1857. Type. Mexico. "On Popocatepetlor Iztaccihuatl,
near 13000-14000 ft.," 1857, Roezl s.n. (lectotype, Fl,
here designated).
Pinus lindlevana G. Gordon & Glendinning, Pinetum
229. 1858 (nom. superfl.); Pinus montezlonae A. B.
Lambert var. lindlevana (G. Gordon & Glendinning)
Parlatorein Candolle, Prodr. 16(2): 399. 1868. Type.
Mexico. Locality not stated, n.d., Hartwvegs.n. (holotype, K).
Pinus suffruticosaRoezi ex Carriere,Trait6Gen. Conif.,
ed. 2, 2: 584. 1867. Type. Mexico: Original material
not located.
Pittus donnell-smithiiM. T. Masters, Bot. Gaz. 16: 199.
1891.Type.Guatemala.
Volcande Agua,
Sacatepequez:
near the summit, Apr 1890, Donnell-Smith2182 (holotype, US; isotype, K).
Tree, tall, erect, stunted near tree line, height to

25-30 m, dbh to 80-100 cm. Trunk monopodial,
erect, terete, bole free of branchesfor 2/ of length,
or retaining old branch stumps. Bark thick, scaly,
deeply fissured, divided into small or large, square
or irregularplates, outerbarkdarkbrown,weathering
grey, on young trees and branches scaly, brown.
Branchesof firstorderthick, often twisted, spreading
and assurging, often persistent; branches of higher
orders stout, spreading and assurging, forming a
dense, roundedcrown in maturetrees. Shoots thick,
rigid, scaly with persistent, decurrentpulvini, purplish brown, sometimes glaucous. Cataphylls large,
15-20 mm long, subulate,recurved,scarious,brown,
weatheringblackish grey, with ciliate to erose margins, usually persisting longer than leaf fascicles.
Vegetativebuds ovoid-conical, large, terminal buds
up to 30 x 15 mm, the lateralssmaller,not resinous;
the scales imbricatebut free at apex, subulate,scarious, reddish brown with ciliate margins, giving the
buds a rugose appearance.Fascicle sheaths of young
leaves long, 30-40 mm, with ca. 10 imbricate,appressed scales with ciliate margins, light brown,
persistent, reduced in older fascicles to (10-)15-20
mm, tight, resinous,weatheringblackishgrey. Leaves
in fascicles of (3-)4-5(-6), most commonly 5, in
dense tufts toward the ends of upturnedbranches,
persisting2(-3) years, straightor curved, not twisted,
rigid, (6-)10-17(-22) cm X (1-)1.2-1.5 mm, with
serrulatemargins, acute-pungent,dull light green or
glaucous-green.Stomataon all faces of leaves, in 6-
Flora Neotropica
10(-11) lines on the convex abaxial face and in 46(-7) lines on each adaxial face. Leaf anatomy:
Cross section triangularor transverse-triangular,
with
convex abaxial side; cuticula and epidermis thick;
hypodermisnearly uniform,with 2-3 layers of cells;
resin ducts (2-)4-10(-12), occasionally absent, medial, occasionally 1-2 internal; stele oval in cross
section; outercell walls of endodermisnot thickened;
vascular bundles 2, distinct but approximateor the
xylem strands connate. Pollen cones crowded near
the proximal end of a new shoot, subtended by
subulate bracts, ovoid-oblong to cylindrical, ultimately 20-25 X 6-7 mm, yellowish or pink-purplish, maturingto brown. Seed cones subterminal,in
pairs or whorls of 3-6 on short, stout, bracteate
peduncleswhich are hiddenby basal scales of mature
cones. Immaturecones ovoid-oblong, often slightly
curved, purplishbrown to purplish black, maturing
in two years. Maturecones ovoid-oblong, with acute,
curved apex when closed, (obliquely) ovoid with a
flattenedbase when opened, 8-12(-14) x 5-8 cm.
Seed scales ca. 150-200, partingto release the seeds,
spreadingwide, often recurvednearthe base, oblong,
thin woody, flexible or more rigid, purplish brown
to blackish with a light brown margin below the
apophysis abaxially, brown with lighter marks of
seed wings on the adaxial side. Apophysis more or
less flat, (weakly) transversely keeled, sometimes
gibbous, rhombicin outline with an angularor irregular upper margin, varying from light brown to
purplishbrown with a blackishcentre. Umbo dorsal,
depressed, flat or raised, up to 4 mm wide, obtuse
or with a deciduous prickle, grey or black. Seeds
broadlyovoid, slightly flattened,5-6 mm long, light
or darkbrown with black spots. Seed wings articulate
but effective, held to the seed by two oblique claws,
oblong with a straightand a curved side, 12-20 x
7-12 mm, browntinged with darkstripes.Cotyledons
(4-)5-6(-7).
Distribution and ecology (Map 10). Mexico: In
Chihuahua(Cerro Mohinora),S Coahuila, S Nuevo
Leon, Durango, SW Tamaulipas, Jalisco (Nevado
de Colima), Michoacan, Mexico, Morelos, Hidalgo,
DistritoFederal,Tlaxcala, Puebla,W Veracruz,Guerrero (CerroTeotepec and vicinity), Oaxaca, and Chiapas; in Guatemala in most of the SW highlands;
in Hondurason several isolated mountainsummits;
reported from the extreme N of El Salvador, but
not confirmed. Pinus hartwegii is the typical highaltitude pine of Mexico and Guatemala, where it
often forms extensive, monotypic pine forests up to
the treelineon high, isolated volcanos or summits of
mountain ranges. In Hondurasit is rare, of limited
119
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Map I0. Pinus h.rm'e~?uii.

Representative specimens examined. MEXICO. CHIextent and usually found with Abies guatemalensis,
HUAHUA:
Cerro Mohinora, I Sep 1898, Nelson 4916 (A.
Cupressus lusitanica, Juniperus standleyi, Quercus US). GUERRERO:
Cerro Zacatonal,3 km S of CerroTeotespp., Dendropanaxlempirianus,Dryimisgranadensis, pec, 15 Jan 1988, Aquino 328 (FHO, MEXU); 3 km W of
a ground cover of Ericaceae, Lycopodiaceae and CerroTeotepec, 27 Sep 1988, Aquino 514 (FHO, MEXU).
Locality not stated, 1837, Ehrenberg648 (MO);
epiphytic Bromeliaceae in a cool cloud forest type HIDALGO:
near Pachuca, in El Chico National Park, 5 Nov 1992,
usually between 2700-2850 m on the highest moun- Gardner
& Knees 5300
Nevado de Colima.
tain summits.Similar forests occur in Guatemalaand 15 Mar 1974, Styles 79 (U). JALISCO:
(FHO). MExico: Nevado de Toluca,
the southern states of Mexico, but here extensive May 1827, KarMinskis.n. (M); Valle de Mexico: 1800, er
pine forests predominate,in which P hartwegii in- herb. LindleY3164 (CGE); Salazar. 1900, Nelson 4 (A.
creasinglydominateswith rising altitude. Its altitudi- US); Ajusco Mts., 4 Dec 1903, Pringle 8789 (A. E. F,
nal range in Guatemala and Mexico is similar: GOET, K, M, MEXU, MICH, MO, NY,); Ajusco Mts.,
18-19 Aug 1906, Pringle 13782 (JE, MICH, S, US).
(2300-)2500-4000(-4300) m. At lower elevations it MICHOACAN:
Cerro Patamban, n.d., Nelson 6563 (US);
is often mixed with P montezumae,with which it is Cerro Tancitaro, La Bolita, n.d., Madrigal 3269 (IBUG,
Near Toro Sta., 14 Nov 1907, Pringle
closely related (Carvajal& McVaugh, 1992; Styles, MEXU). MORELOS:
1993), and with other pines depending on the geo- s.n. (US). NUEVOLE6N: Sierra Infernillo, 22 km SW of
Galeana, 16 Jun 1934, Mueller & Mueller 822 (MICH, NY,
graphical area. Soils are from both volcanic and P); Cerro Potosi, 26 Jul 1934, Mueller & Mueller 1261 (F,
granitic rock, of various depths but often poor in MICH, NY, P); La Joya, near Galeana, 13 Mar 1980,
nutrients.Climatically there are considerable differ- Stead & Stvles 625 (FHO); 1 km N of Mimbres, 6 Nov
ences congruent with latitude/altitudedines, with 1984, Styles, Hoogh & Favela 19 (FHO. MEXU); La
Escondido. Cerro Viejo Zaragosa, 7 Nov 1984, Styles,
heavy frost and snow during several months and
Hoogh & Favela 24 (FHO, MEXU). OAXACA:Sierra
often high winds near the treelines of the high volca- Juarez.
along Hwy. 175 from Oaxaca to Tuxtepec, 23 Apr
nos in Central Mexico. Phenology: Time of pollen 1994, Farjon & Mejia 3/7 (E, FHO, HEH. K, MEXU, U);
Lower
dispersal varies mainly with altitude and is from Cuajimoloyas,2 Oct 1973, Hill 1846 (NY). PUEBLA:
slopes of Pico de Orizaba W of Chlachicomula, 21 Sep
March(Critchfield, 1966b) to July.
120
1957, Beaman 1789 (US); Albergue PiedraGrande, 14 Jul
1960, Beaman 3642 (US); Mt. Iztaccihuatl,on S side of
mtn., along rd. between Paso de Cort6sand ParajeLa Joya,
23 Sep 1961, Beaman 4520 (US). VERACRUZ:
Cofre de
Perote, 28 Aug 1971, Dorantes 350 (FHO, MEXU, XAL);
Pico de Orizaba, (in Natl. Park), 25 Apr 1994. Farjon &
Mejia 321 (E, FHO, HEH, K, MEXU, U); Cofre de Perote,
28 Mar 1963, Little 18974 (K, MEXU, US); Cofre de
Perote, 1 km NW of the summit, 5 Sep 1986, Nee 32987
(FHO, MO, NY); Pico de Orizaba, Rose & Hay 5780
LOCALITY:
(A, US). UNKNOWN
Locality not stated, n.d..
Hartweg s.n. (K); locality not stated, 1857, Roezi s.n. (FI);
locality not stated, 1857, Roezl s.n. (FI).
GUATEMALA. CHIMALTENANGO:
Volcin Acatenango.
16 Feb 1906, Kellerman5840 (US), 7 Feb 1907, Kellerman
6073 (US). GUATEMALA:
Chinantla, May 1892, DonnellSierra los CuchumaSmith 2633B (K). HUEHUETENANGO:
tanes, along rd. between Huehuetenangoand Concepci6n.
27 Aug 1976, Boeke 99 (NY, US). QUEZAI.TENANGO:
Volcan Santa Maria, Sep 1876, Bernoulli & Cario 1089
(GOET); along rd. from Quetzaltenango,40 km toward
Guatemala City, I Oct 1983, McCarter & Hughes 41
(FHO); "from mountains near Hacienda Chaucol," 1900,
Nelson 3729 (US). SACATEPEQUEZ:
Volcan de Agua. 31
Jul 1959, Beaman 2960 (US), 15 Feb 1905, Kellermtan
4580 (US); Volcan de Agua, on crater rim, 16 Feb 1983,
McCarter.Hughes & Styles 5 (FHO), June 1892, Shannon
3622 (A, K, NY, S, US). SOLOLA:Mtn. ridge 1.5 km W
of Hwy. CA I, at Km 168, 18 Apr 1994, Farjon & Mejia
306 (E, FHO, HEH, K, MEXU, U); VolcanAtitlan, 16 Feb
Near Totonica1906, Kellermans.n. (US). TOTONICAPAN:
pan at Chuapechec,6 Mar 1993, Higman. Padilla & Styles
4 (EAP, FHO, HEH, MO); 18 km from Nahuali toward
San Crist6balTotonicapdn,I1 May 1979, Stead 303 (FHO).
HONDURAS. CORTES:Ridge between Cerro Cantiles
and Cerro Jilinco, 20 km W of San Pedro Sula, in Cusuco
Natl. Park, 19 Mar 1993, Mejia 343 (EAP, FHO, HEH,
MO, TEFH), 22 Mar 1993, D. Mejia 372 (EAP, FHO,
El Moj6n Peak, 14 km SW
HEH, MO, TEFH). LEMPIRA:
of Gracias, in Celaque Natl. Park, 15 Apr 1994, Farjon &
Mejia 300 (E, FHO, HEH, K, MEXU, U), 17 May 1992,
Mejia 107 (EAP, FHO, HEH, MO, TEFH); Mt. El Cascal,
7.3 km NW of San Manuel Colohete, 17 Feb 1993, Mejia
313 (EAP, FHO, HEH, MO, TEFH). OLANCHO:13 km
NW of Catacamas, 2 Jun 1992, Mejia 154 (EAP, FHO.
HEH. MO, TEFH).
Uses. Pinus hartwegii has fairly dense but resinous
wood. It is exploited as a timber tree where stands
are extensive and accessible, but due to high altitude
and consequent lack of infrastructure (access roads),
many forests remain virtually untouched today.
Based on a collection by C. T. Hartweg ("Campanario, in the mountains of Angangueo, Hartweg"),
Lindley (1839) described this species as a 4-leaved
pine with very stout branches. There is no original
material at CGE (Lindley's herbarium); a specimen
at K ("ex herb. Gordon, Mexico, Hartweg") bears
no indication that Lindley saw it. In P and W,
material was found with Lindley's original labels;
although no locality was stated (this is cited from
Flora Neotropica
the protologue),it is original materialand a lectotype
was chosen from it (P). The Paris sheet bears only
foliage, while in W only a mature cone has been
preserved.
There are evidently variations in cone colour in
several populations;a form with lighter brown (not
purplish or blackish) cones has often been distinguished as P. rudis. Endlicher(1847) described that
species without any reference to cone colour. In the
same publication, P hartwegii was described with
cones of a clear greyish brown. Martinez (1948)
emphasized the colour differences and designated
the name P hartwegii for the high-altitude pine
with blackish cones and P rudis for those with
brown cones. With Carvajal and McVaugh (1992)
we agree that such distinctions mean little. A
morphological assessment (Matos, 1995) using 25
characters of samples taken from two elevational
transects through populations in Mexico reported
to include both taxa was unable to separate two
morphological entities. We also found that these
charactersare far fromconsistent,and our conclusion
is likewise to sink P rudis into synonymy of P
hartwegii. Lower-altitude collections with brown
cones from W Mexico by J. N. Rose (Rose 2376,
2379, 3004), cited by Shaw (1909) underP. montezumae var. rudis, have been treated under P durangensis by Carvajal and McVaugh (1992) and in
this volume.
Perry(1991), who recognizes P rutdisand P donnell-smithii as full species close to P hartwegii,
presents a number of "distinguishing" character
states for all three in a table, which at close inspection appearto be overlappingor even similar. Pinus
donnell-smlithii was described by Masters (1891)
from materialcollected on Volcande Agua in Guatemala, near the summit at ca. 3750 m (Dolnnell-Smith
2182, holotype, US, isotype K). Like elsewhere,
it forms pure forest at this high altitude. Further
collections and fieldworkby McCarter,Hughes, and
Styles (FHO) revealed that the numberof leaves per
fascicle from that location is consistently 4-5, not
5-6(-7) as given by Perry (1991) for P donn1ellsmithii. There is only one species of Pinus present
around the crater rim of this stratovolcano;the correct name for it is P hartwegii. Leaf numberseems
to decrease furthernorth and may be predominantly
3 in places in CentralMexico (Carvajal& McVaugh,
1992), but 4-5 occurs throughoutthe range of P
hartwegii.
The numerous"new" species described by Roezl
(1857) have here been lectotypifiedfor the first time
by materialpreservedmainly in Fl and P. As Parla-
SystematicTreatment
121
tore (1868) and Shaw (1909) alreadyobserved, none which remain with a few cone scales on the branch
of them are even worthtaxonomicconsideration,but when the cone has fallen. Immaturecones ovoid,
theirtrueidentitycould (in most cases) be established 15-20 X 10-12 mm, with small, spreading spines,
only throughtypification. More details about Roezl purplish, soon pinkish brown, sometimes glaucous,
and his catalogue of pines are presented under P maturingin two seasons. Maturecones ovoid-oblong
to obliquely ovoid when closed, sometimes ovoidrmontezuniae.
attenuate,more or less asymmetrical,often curved
14. Pinus pseudostrobus Lindley, Edwards's Bot. at base, ovoid-oblong to broadlyovoid when opened,
X 6-13 cm. Seed scales ca. 140-190,
Reg. 25: 63. Aug 1839 [& Allg. Gartenzeitung7: then 7-16
parting to release the seeds except the proximal
325. 1839].
infertilescales, usually thick woody, oblong, straight
Tree, medium to large, height to 20-40(-45) m, or slightly curved, somewhatlargerand more curved
dbh to 80-100 cm. Trunkmonopodial,erect, terete, on one side of the cone, reddish brown to dark
the bole usually for %A-/2of the total height free of purplish brown, with light brown marks of seed
branches. Bark thick, scaly, with elongated plates wings on the adaxialside. Apophysisextremely variand deep, longitudinalfissures, dark brown to grey- able, from nearly flat to elongated, more so on one
brown, on young trees and branchessmooth, reddish side of the cone and toward the base, transversely
brown or grey-brown.Branches of first order long, keeled, taperingto an obtuse umbo (or umbo mucrospreading or ascending, the lower branches curved nate),rhombicor pentagonalin outline, uppermargin
down, often distinctly whorled; branches of higher angular,irregularlyundulate,or rounded,colour in
orders slender, spreading, curved down and then various hues of brown. Umbo dorsal, variable,from
upwards,or ascending. Shoots slender,smooth, with obtuse to prominent,3-15 mm long, 5-10 mm wide
(short)decurrentpulvini,glaucous or pruinose.Cata- at base, without a prickle or prickle deciduous, usuphylls 10-15 mm long, subulate-caudate,soon really darker than the apophysis. Seeds obliquely
curved, with ciliate margins, reddish brown to dark ovoid, slightly flattened,5-7 X 3-4.5 mm, ochrabrown, falling shortly after the fascicles. Vegetative ceous to grey-brown, with or without dark spots.
buds ovoid-conical, acute, the terminal buds 15-20 Seed wings articulate,effective, held to the seed by
X 10-15 mm, the laterals smaller, not resinous; two claws, thinly covering part of the seed on one
the scales imbricate, with free, spreading apices, side, obliquely-ovate, with a straightside, 20-25 X
subulate-linear,with ciliate margins, reddish brown. 7-10 mm, yellowish brown, translucent,with darker
Fascicle sheaths (15-)20-30(-35) mm long, persis- tinge distally. Seedlings: The hypocotyl of this
tent, consisting of ca. 10 initially distinct, imbricate, species is consistently longer (mean 31.8 mm) than
subulatescales, lustrousred-brown,weatheringgrey- that of P montezumnae (mean 17.5 mm), according
brown. Leaves in fascicles of 5, rarely4 or 6, in lax to a study by Caballero (1967). Cotyledons (7-)8tufts at the ends of upturnedbranches,persisting212(-13), but given as a mean of 7.2 by Caballero
3 years, slender, straight, spreading or drooping, (1967).
Distribution (Maps 4, 11, 12). Mexico: Sinaloararely more or less rigid, (18-)20-30(-35) cm x
0.8-1.3 mm, with serrulate margins, acute, (light) Durangoborder,Nuevo Le6n, SE Coahuila, E Guagreen or glaucous-green. Stomata on all faces of najuato(?),Jalisco, Michoacan,Mexico, DistritoFedleaves, in (2-)3-7 lines on the convex abaxial face eral, Morelos, Hidalgo, Puebla,Tlaxcala, W Central
and 2-4(-5) lines on each adaxial face. Leaf anat- Veracruz, Guerrero, Oaxaca, Chiapas; Guatemala
omly: Cross section triangular;hypodermis multi- (highlands);W Honduras;N El Salvador.A considerlayered,not or scarcely intrudinginto the mesophyll; able east-west gap exists at the Isthmus of Tehuanresin ducts (2-)3-4(-6), the 3 marginalducts usually tepec, separatingthe Mesoamericanpopulationsfrom
present,medial, rarely 1-2 internalor external;stele the Mexican populations by ca. 250 km. The NW
more or less terete;the outer cell walls of the endo- (Sinaloa-Durango)and NE (Nuevo Le6n-Coahuila)
dermis thickened; vascular bundles 2, distinct, the disjunctionsare separatedfrom the main distribution
xylem strands connate. Pollen cones crowded near in central Mexico by similar distances.
the proximalend of a new shoot, numerous,spreading, ovoid-oblong to cylindrical, 20-35 x 5-7 mm,
Key to the varieties of Pinus pseudostrobus
purplish or pinkish yellow, turning brown. Seed
cones subterminal,solitary or in pairs, more rarely I. Seed scales with slightly to prominentlyraised or
in whorls of 3-4 on very short, stout peduncles
elongated apophyses; umbo obtuse or elongated,
...
Flora Neotropica
122
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more or less continuing the line of the transverse

keel on the apophysis .................................................... 2
I. Seed scales with nearly flat apophyses;umbo acutish, with a distinct angle to the transversekeel of
the apophysis ..........................................................
........................ 14a2. var. pseudostrobusfo. protuberans
2. Seed scales with slightly to prominentlyraised
but not elongated apophyses; umbo not elon........................................
gated, obtuse.
............... 14al. var. pseudostrobusfo. pseudostrobus
2. Seed scales with prominently raised and, at
least in the lower half of one side of the cone,
elongated apophyses;umbo on these scales also
elongated .................................... 14b. var. apulcensis
14al. Pinus pseudostrobus
Lindley var. pseudostrobus fo. pseudostrobus. Type. Mexico. Michoacan: Near Angangueo, 1838, Hartweg s.n. (lectotype, P, sheet with foliage, here designated;
isolectotype, MO, cone scale and seeds).
Fig. 30-31
Pinus orizabae G. Gordon, J. Hort. Soc. London 1: 79.

1846. Type. Mexico. Puebla: [Vic. of] Pico de Orizaba,
1838?, Hartwegs.n. (original materialnot located) (lectotype, fig. in J. Hort. Soc. London 1: 238, here designated).
Pinus protuberans Roezl, Cat. Grain. Conif. Mexic. 27.
1857. Type. Mexico. Mexico: Contreras, 1857, Roezl
s.n. (lectotype, FI, here designated; isolectotype, BM).
Pinus angulata Roezl, Cat. Grain. Conif. Mexic. 27.
1857; Pinus protuberans Roezl var. angulata (Roezl)
Carriere,Traite Gen. Conif., ed. 2, 2: 523. 1867. Type.
Mexico, n.d., Roezl s.n.-n.v.
Pinus regeliana Roezl, Cat. Grain. Conif. Mexic. 29.
1857. Type. Mexico. Mexico: Volcan Iztaccihuatl, on
SW side of volcano, near Hacienda de Zavaleta, 1857,
Pinus pseudostrobusLindley var. estevezii Martinez,Anales Inst. Biol. Univ. Nac. Mexico 16: 188. 1945; Pinus
estevezii (Martinez) J. P. Perry, J. Arnold Arbor. 63:
187. 1982. Type. Mexico. Nuevo Le6n: Caion de Mieleras, Jun 1941, M. Mart'nez 3433 (holotype, MEXU;
isotypes. A. NY).
123
~???i?~
:
-
..
.........
MON.
--":t
II
I-,
Map 12. Pinus pseudostrobusfo. protuberans(circles); P.montezumiaevar. gordoniana (triangles).
PinuspseudostrobusLindley var.coateplecensisMartinez,
Anales Inst. Biol. Univ. Nac. Mexico 16: 187. 1945.
Type. Mexico. Veracruz: "Cerca de Coatepec," Jun
1940, M. Martinez3432 (holotype, MEXU).
Pinus pseudostrobus Lindley var. apulcensis (Lindley)
Martinez,Anales Inst. Biol. Univ. Nac. Mexico 16: 192.
1945, non Shaw, 1909; Pinus pseudostrobus Lindley
subsp. apulcensis (Lindley) sensu Stead, excl. basion. &
type, Bot. J. Linn. Soc. 89: 269. 1984. Type. Mexico.
Hidalgo: Apulco, Sep 1941, M. Martinez 3434 (holotype, MEXU).
Pinus pseudostrobus Lindley forma megacarpa Loock,
[pines Mexico Brit. Honduras] S. Afr. Dept. Forest.
Bull. No. 35: 153. 1950. Type. Mexico. Michoacan:
Ciudad Hidalgo, Loock 10618 (holotype, PRF).
Pinus nubicola J. P. Perry, J. Arnold Arbor. 68: 447.
1987. Type. Guatemala. Guatemala:40 km E of San
Jose Pinula along rd. to Mataquescuintla,25 Feb 1979,
Pern' GUA-32-79 (holotype, GH; isotypes, CHAP, E,
K, MEXU).
Pinus pseudostrobusLindley var. laubenfelsii Silba, Phytologia 68: 60. 1990. Type. Mexico. Sinaloa: In the
mtns. E of Mazatlan,Aug 1968, Laubenfels663 (holotype, SYR-n.v.; isotype, NY).
Apophysis variable, from nearly flat to prominently
raised, more so on one side of the cone and toward
the base, transversely keeled, tapering to an obtuse
umbo, rhombic or pentagonal in outline, upper margin angular, irregularly undulate, or rounded, colour
in various hues of brown. Umbo dorsal, obtuse, 3-6

mm long, 4-6 mm wide at base, without a prickle or
prickledeciduous, usually darkerthan the apophysis.
Distribution and ecology. Mexico: In the Sinaloa-Durango borderarea, in Nuevo Le6n, SE Coahuila, E Guanajuato(?),Jalisco, Michoacan, Mexico,
DistritoFederal,Morelos, Hidalgo, Puebla,Tlaxcala,
W CentralVeracruz,Guerrero,Oaxaca, and Chiapas.
Further south in Guatemala, W Honduras, and N
El Salvador. Of the varieties here recognized, var.
pseudostrobus is the most widely distributed, occurring throughoutthe entire range of the species.
The other two varieties are often sympatric.This is
a widely distributedand in many places common or
abundantpine of montane to high montane habitat
in cold-temperateto warm-temperatezones. Its altitudinal range is considerable but differs little along
the entire 2500 km of its range: (850-)1900-3000
(-3250) m; in Mesoamerica and in N Mexico its
main distributionfalls within a 1900-2400 m zone.
Annual precipitation is also variable, but with a
minimumof ca. 800 mm; in Guatemalaand fonduras it can be >2000 mm. Throughoutits range it is
an importantconstituent of mixed conifer pine and
pine-oak forests, occasionally associated with Liquidambaras on wet E slopes on the Gulf Coast side
124
Flora Neotropica
L I
?il
?,,
?
.
~~~~~~~~~~~~-~~~~~~~~~.~~~~~~~~~~~~~$.,,,,,...,....??I
-, K-"~/
:i:
c~~~~~~,~
?,
., . .
-.
30~~~~~.'4
FIG. 30. A-C, G, H. Pinus pseudostrobus var. apulcensis (Hignan. PcidillaS& StYles 26). D-F. P pseulostrobus var.
pseludostrobus(McCcarter572). A. Branchwith leaf fascicles and pollen cones. B. Leaf fascicle. C. Cross section of leaf.
D, G. Ovuliferous cones. E. Seed scale (three views). F, H. Seeds (two views). (Magnifications:A. B. D-H, XO.5; C.
x 40.)
125
FIG. 31. A. P pseudostrobusvar. pseudostrobus(Farjon & D. Mejit 329). Ovuliferous cone and scale (two views).
s var. upulcensis (B, Hartwegs..t. lectotype: C. Higman. Pxdilla & Styles 26; D, Slell & Styles
B-E. Pinus pseudostrobih
446; E, E. W.Nelson 985. holotype of P. (xuricana). B. Closed cone. C. Cone. lateral(top) and proximal(bottom) views.
scale (two views). D. Cone, scale (three views). E. Upperpartof scale (two views). F. P pseclostrobus var.pseulostrobus
fo. protuberans(Pringle 8788). Scale (two views). (Magnifications X 0.5.)
126
of the mountains of central and southern Mexico. In
the many disturbed forests it may survive as scattered
groves or as individual trees, often with an understorey of, e.g., Gaultheria, Cassia, or, when associated
with fire, grasses and/or Pteridium aquilinum. In its
driest habitat in central and northeastern Mexico
it occurs with P cembroides, Juniperus flaccila,
Quercus, and an understorey with, e.g., Agave, Buddleja, Opuntia, and Salvia, probably mostly in secondary forest. Phenology: Pollen dispersal occurs in
February-April (Critchfield, 1966b; and in specimens examined), dependent on location and altitude.
Flora Neotropica
Tualor29478 (MO, NY); "nearJalapa,4-7000 ft.," 3 Mar

1899. Pringle 8090 (A, F, GOET,JE, M, MEXU. MO, NY,
US, W); Municipio Perote. nearCoatepeque,22 Mar 1980.
Stead & Styles 688 (FHO. P).
(UATEMALA. CHIMALTENANGO:
Tecpan.5 Feb 1972,
Kemip138 (FHO). EL QUICHE:Chichicastenango, 7 Jul
1951, Schwerdtfeger47 (M); Cunun, 4 Apr 1979, Stead &
Styles 337 (ENCB, FHO). GUATEMALA:
Along rd. between
San Jose Pinula and Mataquescuintla,ca. 20 km from San
Jose Pinula. 17 Apr 1994, Farjonl& Mejia 302, 303 (E,
N of ChianFHO, HEH, K, MEXU, U). HUEHUETENANGO:
tla, CuchumatanesMtns., 4 Apr 1979, Stead & Styles 336
(FHO);Aguacatin, 4 Apr 1979, Stead & Styles 338 (FHO).
JALAPA:La Lagunilla, II Sep 1976. Mittak 8299 (FHO.
M). SACATEPEQUEZ:
Antigua, 3 Jul 1951, Schwerdtf,ser
43 (F). SOLOLA:Argueta, Sep 1870, Ber,noulli& Cario
Representative specimens examined. MEXICO. CHIA1081 (GOET); San Antonio Palopo, along shore of Lake
PAS:Las Rosas, 12 Mar 1993, Higman, Padilla & Styles 15
(EAP, FHO. HEH, MO); Arcotete. 15 Mar 1993. Higman. Atitlan, 4 Apr 1993, Higmlan,Padilla & Styles 66 (EAP,
Padilla & Styles 29 (EAP, FHO. HEH, MO). DURAN(;O: FHO, HEH, MO); San Andres Sementajab,along rd. from
Los Negros, 5 Apr 1991, Styles. Favela et al. 16 (FHO). Panajachelto Godinez, 19 Apr 1979, Stead & Styles 372
between Solola and Santa Lucia Utatlan ("Santa
GUERRERO:
Chilpancingo,at Soxamanatlin. 25 Mar 1993, (FHO):
Higmnan,Padilla & Styles 61 (EAP. FHO. HEH, MO); Lucia Uspantan"),20 Apr 1979, Stead & Styles 376 (FHO).
Cerro Teotepec. 12 km NE of Puerto del Gallo. 19 Oct TOTONICAPAN:Near Totonicapin, 6 Mar 1993, Higtman,
1983, Soto N. et al. 5857 (MEXU, MO); Puntecillas. 10 Padilla & Styles 2 (EAP, FHO, HEH, MO); near jet. of
Nov 1965, May Nab AM237 (ENCB, INIF). HIDALG(O: rds. to San Francisco El Alto and Momostenango. 6 Mar
Padilla & Styles 5 (EAP. FHO, HEH. MO);
Along Hwy. 85 NE of Zimapin, near hamlet of Maguey 1993. Higm,tan.
Verde, 28 Apr 1994, Farjon & Mejia 329 (E, FHO, HEH. San Crist6bal, 3 Apr 1993, Higman. Padilla & Styles 63
K. MEXU, U); (near)Apulco, 17 Mar 1980, Stead & Styles (EAP, FHO, HEH. MO).
HONDURAS. COPAN:Dulce Nombre, May 1977,
641 (FHO. MEXU). JALISCO:G6mez Farias,21 Feb 1980,
Stead & Styles 506 (EAP, FHO). MEXIco: Ajusco Mtns.. Chaplin 367 (FHO). LA PAZ:On mtn. slope near small
1905, Lemt1on 241 (UC); Valle de Mexico, n.d., e. herb. hamlet of El Pinar, 13 Apr 1994, Faijon & Mejia 295 (E.
LindIle 3178 (CGE); Vallede Mexico, Eslava, 31 Jan 1899, FHO, HEH, K, MEXU, U); 12 km S of San Pedro Tutule
towardGuajicirro,MontanaPacayal,28 Jun 1982,
Pringle 8054 (A. BM, E, F, GOET. JE. K, M, MEXU, along rd.
Mt. Celaque. 20 Nov
Ca. 29 km S of Patzcuaro,20 Nov HSglhes222 (FHO, K). LEMPIRA:
MO). MICHOACAN:
1961, King & Soderstromi5168 (MEXU, MICH, MS, NY, 1974. Hiazlett2277 (HEH); Cerro de las Delicias, 5 Mar
1988. McCarter572 (EAP, FHO); on ridge N of Arcagual
US); along rd. from Ocampo to San Felipe, I kin from San
13 km SW of Gracias, Mt. Celaque National Park,
Felipe, I Mar 1980, Stead & Styles 586 (FHO); near camp,
15 May 1992, Mejia 106 (EAP, FHO, HEH, MO. TEFH).
N
main
rd.
to
22
Feb
8
km
1974.
Uruap:in,
along
Carapin,
EL SALVADOR. SANTAANA:On the upper S-facing
Styles 3 (FHO); 10 km along rd. from Uruapanto Tancitiro.
23 Feb 1974. Styles 6 (FHO). NuEvo LE6N: Iturbide, 14 slopes of Cerro Montecristo, within the Montecristo NaNov 1991. Frankis 159 (E); 5 km W of Iturbide,30 Mar tional Park.ca. 16 km N of Metap6in,23 Feb 1989. Hulghes
1963, Little 18981 (IFGP. K, US); "Sierra Madre near 1262 (FHO. K. LAGU).
Monterrey,"9 Jul 1888, Pringle 1965 (E, MO. S. UC,
US. W); ParqueNac. Chipinque Monterrey,I) Mar 1980, 14a2. Pinus pseudostrobus
Lindley var. pseuStead & Styles 600 (FHO, MEXU); 8 km from Iturbide
dostrobus forma protuberans Martinez, Pinos
along rd. from Linares to Iturbide. 12 Mar 1980. Stead &
Mexic., ed. 2, 192, f. 154. 1948. Type. Mexico.
Styles 607 (FHO, M, P), 12 Mar 1980, Stead & Styles 607h
Distrito Federal: Eslava, Jun 1940, M. Martihez
At La Cumbre,along dirt rd. to Yurila,
(FHO. P). OAXACA:
3 km E of Hwy. 175, 24 Apr 1986, Gereau & Martin 1927
3431 (lectotype, MEXU, designated here).
(FHO, MO); Sierra Benito Juirez, 16 Nov 1984, Lepez et
Fig. 31F
al. 248 (UC); roadside between Suchistepec and Miahuatain.22 Mar 1895, Nelsonl 2521 (K, US). PUEBLA:Above
Apophysis nearly flat, or slightly raised only on
Coxcatlin between Apala and top of Cerro Chichiltepec. one side of the cone and toward the base,
transversely
17 Jul 1961, C. E. Smithet al. 3850 (F. MEXU). SINAIOA:
keeled, rhombic or pentagonal in outline, upper mar17 km W of Sinaloa borderalong Hwy. 40. 22 Jul 1975.
Dunn et al. 21800 (NY); in mtns. E of Mazatlain.Aug gin angular, irregularly undulate, or rounded, colour
in various hues of brown. Umbo dorsal, prominent,
1968, Laithlfenfls 665 (NY). TAMAULIPAS:"Between Los
Caballos and Agua de Vacas," May 1970. Gonsiles M. 4-6 mm long, 3-4 mm wide at base, without a
3058 (MEXU). VERACRUZ:
Along Hwy. 140, 5 km SE of
darker than the apophysis.
Las Vigas. 10 Mar 1983. Nee & 7Taylor25879 (F, FHO. prickle, usually
Distribution
and
ecology. Mexico. This form has
MICH, NY); Cerro de Tecomal, 5 km S of Los Altos
Veracruz. 12 Mar 1983, Nee & Tavlor 25936 (F, FHO, been first observed in the state of Mexico and in the
MEXU. MO, NY. XAL); Zongolica. 8 Feb 1984. Nee & Ajusco Mountains near Mexico City. The much
wider distribution given by Martinez (1948) and
Perry(1991) includesS Jalisco, Michoacan,Morelos,
Puebla, and one locality in Oaxaca. This taxon was
not recognizedby Stead and Styles, and the (original)
identificationsas this taxon, if present,on collections
seen by them were not recordedin the database.
specimensexamined.MEXICO.MEXRepresentative
Ico: Santa Rosa, Oct 1940, M. Martinez3505 (US). MixICO D.l.: Eslava, at base of Sierra de Ajusco at Eslava
19 Nov 1903,Pringle8788 (A, BM,F, GOET,

("Eslaba"),
K, MEXU,NY,P, S, U, UC);Eslava,n.d.,Pringle13262
(MICH.MO.US).
14b. Pinus pseudostrobus Lindley var. apulcensis
(Lindley) G. R. Shaw, [Pines Mexico] Publ.Arnold
Arbor. 1: 19, t. 12, figs. 6-8. 1909. Figs. 30-31
Pinusapulcensis Lindley,Edwards'sBot. Reg. 25: 63.
7: 325. 18391.Type.
Aug 1839[& Allg. Gartenzeitung
Mexico.Hidalgo:RavinesnearApulco,1838,Hartweg
s.n. (lectotype,W, 2 seed cones with labels:P aptplcensis Lindl.in Lindley'shandwriting
and phialwith
seedsandscales,designated
by Farjon(1995);isolectotype,GH,seedconeandsomeleaves).
Pinuls pseudostrobus Lindley var. oaxaicana Martinez.
Anales Inst. Biol. Univ. Nac. Mexico 16: 195, figs.

164-168. 1945, nom. inval.. sine lat. diagn.;Pinuis
oa.racana Mirov, Madrofo 14: 145. 1958; Pinus pselu-
cdostrohus
Lindleyvar.oaxacana(Mirov)S. G. Harrison,Taxon 14: 247. 1965. Type. Mexico.Oaxaca:
18Aug 1894,Nelson985 (holotype,
"NearLa Parada,"
US; isotype,A).
127
Padilla & Styles 28 (EAP. FHO. HEH. MO); near San
Crist6balde las Casas. 16 Oct 1960. Little 17963 (IFGP.
K, US); Mun. Tenejapa.20 Jan 1966, ShilornTon 650 (A,
NY); between Comitanand San Crist6balde las Casas. ca.
7 km from Comitan. 26 Mar 1979, Stead & Styles 293
(FHO, US). GUERRERO:Along rd. from Chilpancingo to
San Vicente, near Chilpancingo, 25 Mar 1993, Higman.
Padilla & St les 59 (EAP. FHO, HEH. MO); along rd.
from Xochipala to Filo de Caballo, 19 Apr 1985. Soto
N. & Aureoles C. 8261 (FHO); "Mpio. V. Guerrero.San
Bernardino,La Laguna sobre terraceriaAzumbella," 27
Feb 1982. Lott & Wendt133 (MEXU); between Filo de
Caballo and Colonia Eliodora de Castilla, 8 Feb 1981,
Stead et al. 734 (FHO). HIDALGO:
Apulco, n.d., (Hartweg?)
e.xherb. Gordons.r. (FI). OAXACA:
La Carbonera,18 Mar
1993, Higman, Padilla & Styles 36 (EAP. FHO, HEH.
MO); NE of Guelatao. ca. 50 km NE of Oaxaca. 13 Oct
1960, Little 17942 (IFGP,K. US); Mun. Ixtepeji, 19 km N
of Hwy. 190 along Hwy. 175. 13 Feb 1981. Martin 320
(ENCB, MEXU); Teotitlandel Valle, Rancho Tablas, Nov
1940, M. Martinez 3435 (MEXU); Lachatao, Jan 1941.
M. Martin. 4102 (US); "Miahuatlin" [= Santa Lucia
Miahuatlan?].1895, Nelson 2539 (US); San Pablo Macuiltianguis. "cerco del panteon viejo." 3 May 1980. Pdrez
C. 182, 183 (MEXU); distr. Coixtlahuaca,4 km SW of
Magdalena Jicotlan. 17 May 1968. Rzedowski 25712
(ENCB); Nochixtlin. along rd. to Jaltepec. ca. 3 km from
Hwy. 190. 29 Jul 1970, C. E. Smith & Kitchen5034 (US);
ca. 13 km S of Miahuatlan,along rd. to Puerto Angel. 2
Feb 1980. Stead & Styles 423 (FHO, US). 2 Feb 1980.
Stead & Styles 434 (FHO. P), 2 Feb 1980. Stead & Styles
444-447. 449 (FHO); 10 km along rd. from Oaxaca to
Tuxtepec. 26 Mar 1974. Styles 101 (FHO); 6 km SW of
Cuquila, along rd. from Tlaxiaco to Putla. 8 Jun 1985,
TorresC. 6693 (MEXU); 24 km NE of Ojite, nearTlaxiaco,
9 Aug 1985, TorresC. et al. 7144 (MEXU). PUEBLA:
San
Pedro Chapulco, 18 km NE of Tehuacin. 20 Mar 1989,
Hughes 1322 (FHO. K. MEXU); 6 km S of San Salvador
(El Seco). km 204. 28 Mar 1963. Little 18966 (IFGP,INIF.
K, US); Carpinteros.3 Jan 1974, Magana 900 (MEXU); I
km S of Carpinteros.3 Jan 1974, Magana 990 (MEXU):
Cofre
Zacatepec, Jul 1986, Nee 33008 (NY). VERACRUZ:
de Perote, Jul 1866, Hahn s.n. (P); Saltillo de la Fragua.
Nov 1979, Klaus s.n. (WU).
GUATEMALA. SAN MARCOS:Location given as
14?58'N.91?43'W, 7 Apr 1979, Stead & Styles 346 (FHO.
NY). 7 Apr 1979. Stead & Styles 349 (FHO). SOLOLA:
Mtn. ridge W of Hwy. CA 1. at Km 168, 1.5 km W of
hwy.. 18 Apr 1994. Farjon & Mejia 307 (E. FHO. HEH.
K. MEXU. U).
EL SALVADOR. CHALATENANGO:
Los Esesmiles, 6
May 1959, Allen & van Severe, 7324 (A. EAP, F, NY.
US); 4 km SSE of La Palma, 11 Jun 1970, Davidse & Pohl
2076A (FHO, MO).
Apophysis variable,prominentlyraised and partly

elongated, especially on one side of the cone and
toward the base, transversely keeled, rhombic or
pentagonalin outline, upper marginangular,irregularly undulate,or rounded,colour in various hues of
brown. Umbodorsal,prominentand/orelongated, 515 mm long, 5-10 mm wide at base, without a
prickle, usually darkerthan the apophysis.
Distribution and ecology. Mexico: Mexico, Hidalgo, Puebla, Tlaxcala, W central Veracruz,Guerrero, Oaxaca, and Chiapas. Also in the Guatemala
highlands and N El Salvador.Apparentlyrestricted
to the SE part of the range of the species, but most
common in Oaxaca, where it is perhapseven more
common than var.pseudostrobus.It is often sympatric and occupies much the same habitat but seems
absent from the driest locations.
Uses. Pinus pseudlcstrobus is one of the most
and important "hard pines" in the southern
common
Representative
specimensexamined.MEXICO.CHIA-
PAS:10 km E of Las Ollas, near hamlet of Cruzchen,E of

half of Mexico and the highlands of Guatemala and
SanCrist6balde las Casas,20 Apr 1994,Farjon& Mejfia
of Honduras. Its wood is light yellowish in
311 (E, FHO,HEH,K, MEXU,U); Las Rosas, 12 Mar parts
colour,
lightweight (specific gravity -0.45), strong,
1993, Higman,Padilla& Styles 14 (EAP,FHO, HEH,
with long intervals of knot-free wood that is only
MO); Jitotol, 14 Mar 1993, Higman, Padilla & Styles 26
(EAP,FHO,HEH.MO);Arcotete.14 Mar1993.Hig,ntn, slightly resinous. Exploitation for timber is wide-
128
spread and has led to regional depletion of the best
stands. In Mexico, it is also used as a source of
resin.
Pinus pseudostrobus was described by Lindley
(1839). based on materialcollected by Hartwegnear
Angangueo in Michoacan,Mexico. The only original
materialfound so far is a sheet in P without a cone,
which refers to "Benth. pi. Hartw.p. 58. No. 443*"
under P pseudostrobusLindl. and with a Paris label
identifying it as a Hartweg collection. Number 443
is not Hartweg'scollection numberfor this specimen.
The specimen was annotatedas "type collection" by
E. L. Little in 1969. In BM a single cone with a
label-"Pinus pseudostrobusLindl. Mexico, Anganguco-Lambert's Sale [XXVI.55] 68"-is annotated
as "type specimen" by E. L. Little. Apart from
the location, this specimen and its label profoundly
disagree with the protologue and belong with P
monteznumae; it is not accepted here as a type specimen for Pifills pseudostrobusvar.pseudostrobus.An
original cone has still to be found; Stead and Styles
(1984) reportedthat they did not see any type material. The species was described as a 5-leaved pine,
the leaves very slender and glaucous, "like those of
the Wymouth pine." The cone scales are described
as having a rhomboid,pyramidalapophysis("apex"),
with an elevated, transverseline (i.e., keeled transversely); the length of the cone is given as ca. 10
cm.
As was customaryand understandableat the time,

such a descriptionwas based on very scanty material,
often a single specimen. Little was known about the
diversity of species in Mexico, and the material
was often comparedwith specimens of species from
relatively high latitudes, which are much less variable. As Stead and Styles (1984) pointed out, Lindley's description,in the light of today's knowledge,
could apply to numerousspecies. However,any subsequent application of such names can be correct
only if it definitely includes its type as well as the
charactersgiven in the protologue.
There is an extensive synonymy for this species,
in part discussed in Stead & Styles, 1984. Because
the treatment of P pseudostroblusLindley heremainly due to the more recent typification of Lindley's two relevant species, P apulcensis (Farjon,
1995) and P pseudostrobus--differs considerably
from the one presented by Stead and Styles (1984),
emphasis in the discussion will be on some taxa
treateddifferently,or published since 1984.
Roezl (1857) published a numberof species that
were tentatively synonymized, but not typified, in
recent treatments of Mexican pines. As discussed
Flora Neotropica
under P imontezumnae,the great majority of his names
now have here been typified. One of them is P

protulberansRoezl, first cited in synonymy with P
pseudostrobusby Parlatore(1868), who had indeed
seen Roezl's materialat Florence (Fl).
Martinez(1948) recognized a numberof infraspecific taxa under P pseudostrobtus.One of these is P
pseudostrobusf. protuberansMartinez.Its characters
are describedabove. Stead and Styles (1984) synonymized it under P pseudostrobus(subsp. psetdostrobus) as P.pseudostrobusf. prottuberais(Roezl) Martinez, but Martinez did not refer to Roezl. In an
earlier edition (Martinez, 1945: 184) he referred
to Pringle 8788 (MEXU!) as "formaprotuberance
Roezl," but this is Pringle's determination, not
Roezl's type, which is somethingaltogetherdifferent.
We must simply consider this as a coincidence of
homonymy, but at different ranks, and not read too
much into the case.
Pinluspseudostrobus var. coatepecensis Martinez
is based on Martbiez 3432 from the vicinity of
Coatepec ("la region vecina a Coatepec") in Veracruz, Mexico. It has a rather small, broad cone,
which was taken as a distinction by Martinez,but in
all other charactersit is similar to P pseudostrobhts
var.pseuldostrobus.Stead and Styles (1984) synonymized it with their subsp. apulcensis, which was
defined by its relatively large cones, disregardingthe
type but basing this opinion on their "population
collection" gathered from 23 trees at Coatepec,
which was presumedto be the type locality (Stead.
1983).The inherentuncertaintiesof this methodhave
been discussed elsewhere (Farjon, 1995). Size and
shape of cones are variable quantitativetraits in P
pseudostrobus, and the morphometric statistics in
Stead (1983) fail to produce evidence for discrete
charactersbased on these measurements.
Pinus pseludostrobuis
var. estevezii Martinez,based
on materialfrom Canionde las Mieleras, near Santa
Catarina,Nuevo Le6n, was also distinguished from
P pseutdostrobusvar. pseudostrobus by the cones,
which were describedto be larger,asymmetrical,and
"harder,"with elevated apophyses and a persistent
spine on the umbo. Differences in the leaves pertain
to a thicker hypodermis and an endodermis with
thickeroutercell walls. Gaussen (1960) elevated this
variety to species but failed to cite the basionym,
resulting in an invalidly published combination.
Perry (1982) elevated it once more as a distinct
species, based on his furthercollections, e.g., W of
the town of Iturbide,Nuevo Le6n, where Zobel and
Cech (1957) and Stead (1983) also collected it.
Although more or less confirming Martinez's mor-
Systematic Treatment129
phological differences and adding a few from his
own field observations(mainly habitand bark),Perry
based his main argumenton the chemical composition of the turpentine, with high levels of light
hydrocarbonsin P estevezii and low levels in P
pseudostrobis. Trees with morphological character
states ascribed to this taxon can be found in many
other places, far beyond the region Perry (1991)
defined for it. None of these populations has been
found to be truly distinct.
Farjon (1995) published a full discussion of the
identity, including its typification, of P. apulcen.sis
Lindley, and he showed that Shaw (1909) correctly
cited the illustrationin Loudon (1842) as representing Hartweg'soriginal material.The original use of
the name as to type, confirmed by Shaw, has been
restored against the alternative of rejecting it in
favour of the usage by Martinez (1948) and Stead
and Styles (1984), with a conserved type, because
there is no case of consistent use in a differentsense
since its publicationand the taxon is now recognized
as a variety(subspecies)only. Pinlusoaxaccaa Mirov
is a taxonomic synonym of P apulcensis.
From Guatemala,ca. 40 km from San Jos6 Pinula
E of GuatemalaCity, Perry(1987) describedanother
new species, P nubicola. He also reported it from
other localities in Guatemala,Honduras,El Salvador,
Chiapas, and a location in Veracruz. Virtually all
collections are in Perry's personal herbarium(Hertford, North Carolina,United States, but not seen on
a visit March 1994 by A. Farjon);only type specimens were distributedto several institutionalherbaria. This makes evaluation rathercomplicated; the
only other accessible collection cited by Perry was
Mittak 8299, of which the FHO specimen consists
merely of a few seed scales. Collections from (the
vicinity of) some of Perry's locations were made in
1994. It was found that besides Perry'sform, several
others, both in regard to foliage and cones, were
easily gatheredwithin a few hundredyards of each
other, e.g., at the village of Las Trancas, Dept. La
Paz, Honduras.On a larger geographical scale, the
cone of Stead & Styles 641 (MEXU), part of a
"populationcollection" of P. pseudostrobus subsp.
apulcensis sensu Stead & Styles at Apulco, Hidalgo,
Mexico, matches the cone of Perry GUA-32-79
(MEXU!), an isotype of P. nubicola J. P. Perry
from Guatemala.Perry (1987) reportedhaving seen
intermediatesbetween P nubicola and P pseudostrobus and assumed hybridization and back-crossing
were responsible.
The species Pitnus pseudostrobus is a variable
and widely distributedtaxon (Stead & Styles, 1984;
Carvajal& McVaugh,1992), whose genetic diversity

apparently indicates complex relationships among
populations.Morphometricsandchemistryhave contributed inconsistent classifications, which in turn
have been shown to use one of Lindley's binomials
contraryto its type. The treatmentpresentedhere is
more conservative. New evidence, preferablybased
on moleculardata, could perhapsshow the existence
of distinct, taxonomicallymeaningfulcharacters.
15. Pinus montezumae A. B. Lambert,Descr.Pinus,
ed. 8?, 1: 39, t. 22. 1832.
Tree, medium to large, height to 20-30 m, dbh to
100 cm. Truik monopodial,erect, terete, massive in
solitary trees, with persistent branches. Bark thick,
scaly, breakinginto numerous,relativelysmall, irregular, slightly elongated plates divided by shallow
fissures, dark brown, weathering grey-black, on
young trees and branches rough and scaly, reddish
brown or greyish brown. Branches of first order
long, often massive, spreading wide; branches of
higher orders spreading to ascending in the upper
partof the tree, assurgenttoward the ends, forming
a dense, usually broad,roundedcrown. Shoots thick,
rigid, or more slender, very rough and scaly, with
persistent,decurrentpulvini, brown,sometimesglaucous. Cataplhlls ca. 15 mm long, subulate,recurved
or reflexed,scarious, with erose-ciliatemargins,dark
brown, weatheringgrey, persisting somewhat longer
than the leaf fascicles. Vegetativebuds (broadly)
ovoid, the terminalbuds 15-30 mm long, the lateral
buds smaller,not resinous;the scales imbricate,with
free, spreading apex, subulate-linear,with ciliate
margins, reddish brown to orange-brown,the ciliae
3-5 mm long, orange-brown.Fascicle sheaths (20-)
25-35 X 1.5-2.5 mm, persistent,consisting of ca. 10
imbricate,orange-brownscales with ciliate margins.
remainingtight, weatheringgrey to black, sometimes
resinous. Leaves in fascicles of (4-)5 (rarely3 or 6),
in dense tufts towardthe ends of upturnedbranches.
persisting 2-3 years, slender, flexible and drooping,
or lax to rigid and spreading,straight,(15-)20-35(40) cm X 0.8-1.3 mm, with serrulatemargins,acute
or pungent,(light) green to glaucous-green.Stonmata
on all faces of leaves, in 4-7 lines on the convex
abaxial face and 3-6 lines on each adaxial face,
weakly correlated with leaf width. Leaf anatoty:
Triangularin cross section, with a convex abaxial
side; hypodermis sometimes thin and uni-layered,
but mostly multi-layered,especially in the marginal
regions, with slight intrusions into the mesophyll:
resin ducts (2-)4-5(-6). medial or occasionally 1-2
130
Flora Neotropica
internal; stele terete; the outer cell walls of the

ovoid-oblong. apophysis of scales flat, especially
towardthe base of the cone, with a weak transverse
endodermis thickened; vascular bundles 2, separate
line.................................................. 15b. var. gordoniana
but close to each other, or the xylem strandssometimes connate. Pollen cones crowded near the proximal end of a new shoot, subtended by subulate to 15a. Pinus montezumae A. B. Lambertvar.montezumae. Type. Mexico. Veracruz:"InterCruz Blanca
lanceolate bracts, cylindrical, 20-40 x 6-8 mm,
et
or
reddish
brown.
Jalacingo, reg. frig. Nov. 28," Schiede & Deppe
initially pink
purplish, turning
s.n. (lectotype, BM, designated by McVaugh in
Seed cones subterminal,solitary or in whorls of 3Carvajal & McVaugh, 1992: 79; isolectotypes,
6, spreading or recurved on short, stout peduncles.
HAL, W).
Immature cones ovoid, often curved, purplish to
Fig. 32
purplish brown, maturing in two seasons. Mature
Pinulf occidenltalis Kunth in Humboldt, Bonpland &
cones variable,from (broadly)ovoid or symmetrical
Kunth, Nov. Gen. & Sp. PI. 2: 4. 1817, non Swartz.
1788. Type. Mexico. Locality not stated, 1806. Bonpovoid-oblong to ovoid-attenuate,often curved, with
la(nd & Humboldt 3913 (holotype. P-Bonpl.; isotype,
a more or less flattened base when opened, usually
P).
widest near the base, 8-20 x 5-10 cm when open.
Pinu,. russelliana Lindley, Edwards's Bot. Reg. 25: 63.
Seed scales ca. 175-250, partingto release the seeds,
Aug 1839 l& Allg. Gartenzeitung7: 325. 1839] Type.
the proximal,infertilescales remainingon the branch
Mexico. Hidalgo:Near Mineraldel Monte. "lalong] the
road from San Pedroto San Pablo," 1838, Hartwegs.n.
when the cone falls, thin or thick woody, oblong,
(lectotype, W, cones only, here designated).
straightor curved, with thin margins,reddishbrown
Piltns monzttez:umae
A. B. Lambert var. liidlevi J. C.
to purplishblack with a light bandunderthe apophyLoudon. Encycl. Trees Shrubs 1004, figs. 1882-1883.
sis abaxially,(light) brown, with faint marksof seed
1842. Type. Mexico. Locality not stated, n.d., Hatrtweg
31(?) (lectotype. W (ex Hort. Soc. London. 2 cones
wings on the adaxial side. Apophysis flat or raised
only), here designated).
especially on proximal scales, if raised with a transPinl.s teiinltgaensis Roezl. Cat. Grain. Conif. Mexic. 10.
verse keel, (transverse-)rhombicin outline, some1857; PiinuspseudostrobusG. Gordon & Glendinning,
times irregular,up to 20 mm wide, often rugose or
Pinetum 237. 1858, non Lindley, 1839. Type. Mexico.
Mexico: Mt. Ajusco, 1857, Roezl s.n. (lectotype. Fl.
radially striate, in various shades of ochraceous to
here designated;isolectotypes, CGE. P).
brown. Umbodorsal, depressed,flat or raised,obtuse
Piinus
grandis Roezl, Cat. Grain.Conif. Mexic. 11. 1857.
or terminatingin a small prickle, weathering grey.
non D. Don, 1832. Type. Mexico. Between the volcanos
Seeds obliquely ovoid, slightly flattened,5-7 x 4Popocatepetland Iztaccihuatl,at <9000 ft, 1857. Roe.l
5 mm, light brown,often with darkspots. Seed wings
s.n. (lectotype, P. here designated).
Pirius imonstro.s Roezl, Cat. Grain. Conif. Mexic. II.
articulate, effective, held to the seed by two thin
1857. Type. Mexico. Roezl s... photograph of seed
claws, covering the seed partlyon one side, obliquely
cone from Roezl's collection at P in G. R. Shaw's pine
oblong, 18-28 x 7-12 mm, light brown with dark
collection; original material not located (lectotype, A.
stripes. Seedlings: Hypocotyl consistently longer
here designated).
Pinus souliangeanaRoezl, Cat. Grain. Conif. Mexic. 11.
(29-37 mm) than in P pseudostrobus(14-28 mm);
1857. Type. Mexico. Mexico: Mt. lztaccihuatl,on SW
mean number of cotyledons 7.2 (Caballero, 1967),
slope of volcano, 1857, RoezI s.n. (lectotype, Fl, here
range 7-10 according to Mirov (1967).
designated:isolectotype, P).
Distribution (Maps 12, 13). Mexico: In Nuevo
Pini.ustroubezkoia,ltRoezl, Cat. Grain.Conif. Mexic. II.
Le6n, SW Tamaulipas,Nayarit,S Zacatecas,Jalisco,
1857. Type. Mexico. Mexico: Mt. Iztaccihuatl,on SW
Michoacan,Mexico, Distrito Federal,Quer6taro,Hiz s.n. (lectotype, Fl. here
slope of volcano, 1857, Roe
designated).
central
Morelos,
Puebla,
Tlaxcala,
Veracruz,
dalgo,
Piirl.s hoothiIna Roezl, Cat. Grain. Conif. Mexic. 12.
Guerrero,Oaxaca, and Chiapas;in Guatemalain the
highlanddepartments.
Key to the varieties of Pinus monteztumae
I. Leaves spreading or drooping, 1-1.3 mm wide,
with (4-)5-7 lines of stomata on the abaxial face
and 4-6 lines on each adaxial face; mature seed
cones variablein shape, apophysisof scales raised,
especially towards the base of the cone, with a
transversekeel ............................. 15a. var. mtonte-uuneue
1. Leaves slender, drooping, 0.8-1 mm wide, with
4-5 lines of stomata on the abaxial face and 3-4
lines on each adaxial face; mature seed cones
Pinu.sboucheilna Roezl. Cat. Grain. Conif. Mexic. 12.

1857. Type. Mexico. Puebla:Mt. Iztaccihuatl,on W side
of volcano, nearAmeca, 1857, Roezl s.. (lectotype. Fl.
here designated).
Pin,is elegans Roezl, Cat. Grain.Conif. Mexic. 12. 1857.
Type. Mexico. Locality not stated, 1857, Roezl s.I.
Pinus runielianu Roezl, Cat. Grain. Conif. Mexic. 12.
1857. Type. Mexico. Roezl s.n., original material not
located; photographof seed cone from Roezl's collection at P in G. R. Shaw's pine collection (lectotype, A,
here designated).
I"
131
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Map
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otzmevt
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io
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a'"rMap13 Pmnusmonezumne
Pinus spinosa Roezl, Cat. Grain.Conif. Mexic. 12. 1857.

Type. Mexico. Roezl s.n., original materialnot located.
Pinus keteleerii Roezl, Cat. Grain. Conif. Mexic. 13.
1857. Type. Mexico. M6xico: Toluca, 1857, Roezl s.n.
(lectotype, P, here designated; isolectotype, FI).
Pinus san-rafaeliana Roezi, Cat. Grain. Conif. Mexic.
13. 1857. Type. Mexico. Mexico: On the slope from
Acoculco to San Rafael, 1857, Roezl s.n. (lectotype, Fl,
here designated).
Pinus vangeertii RoezI, Cat. Grain. Conif. Mexic. 13.
1857. Type. Mexico. Mexico: Hacienda Tomacoco,
1857, Roezl s.n. (lectotype, Fl, here designated).
Pinus antoineana Roezl, Cat. Grain. Conif. Mexic. 14.
1857. Type. Mexico. Mexico: Near HaciendaLa Valeta,
1857, Roezl s.n. (lectotype, P, here designated;isolectotype, FI).
Pinus doelleriana Roezl, Cat. Grain. Conif. Mexic. 14.
1857; Pinus grandis Roezl var. doelleriana (Roezl)
Carritre,TraitdGen. Conif., ed. 2, 2: 532. 1867. Type.
Mexico. Mexico: Toluca, 1857, Roezl s.n. (lectotype, P,
here designated; isolectotype, Fl).
n.o..z u . .
Pinus paxtonii Roezl, Cat. Grain.Conif. Mexic. 14. 1857.

Type. Mexico. Mexico: Hacienda Tomacoco, 1857,
Pinus vanhouttei Roezl, Cat. Grain. Conif. Mexic. 14.
1857. Type. Mexico. Mexico: Mt. Ajusco. 1857, Roezl
s.n. (lectotype, FI, here designated; isolectotype, P).
Pinus haageana Roezl, Cat. Grain. Conif. Mexic. 15.
1857; Pinus grandis Roezl var. haageana (Roezl) Carriere, Traite Gin. Conif., ed. 2, 2: 535. 1867. Type.
Mexico. Mexico: San Rafael, 1857, Roezl s.n. (lectotype, P, here designated; isolectotype, Fl).
Pinus hendersonii Roezl, Cat. Grain. Conif. Mexic. 15.
1857; Pinus monstrosa Roezl var. hendersonii (Roezl)
Carribre,Trait6G6n. Conif., ed. 2, 2: 526. 1867. Type.
Mexico. Mexico: Rfo Frio, 1857, Roezl s.n. (lectotype,
P, here designated; isolectotype, Fl).
Pinus jostii Roezl, Cat. Grain. Conif. Mexic. 15. 1857.
(lectotype, CGE (ex herb. Lindley 3177), here designated; isolectotype, Fl).
Pinus thibaudiana Roezl, Cat. Grain. Conif. Mexic. 15.
132
Flora Neotropica
'"I".
f'
iiIi
'F..
f.
iii
Viit
'?I?
'.
'I'Iru1.
.&
~,. t.,
'
I'" ,i.,~
'
'
~ij~E~
"
'
':,.
A~~~~~~~~~~'.
FIG. 32. A-F, H. Pinns 01monttez

.uma1evar. monte:.-unae(A-C, Stead & StYles536f D-F, H, Stead & StYles637). G. P
itionitezitmae var. gorloniianua (M. Martinez 3438). A. Branch with leaf fascicles and flushing terminal bud. B. Leaf
tfascicle.C. Cross-section of leaf. D. Ovuliferous cone. E. Seed scale (three views). F. Branchwith remnantcone scales.
G. Ovuliferous cone. H. Seed with articulatewing. (Magnifications:A, B. D-H. x0.5; C. x35.)
SystematicTreatment
1857. Type. Mexico. Mexico: Mt. Popocatepetl,on N
side of volcano, 1857, Roezl s.n. (lectotype, Fl, here
designated;isolectotype, P).
Pinus Iullata Roezl, Cat. Grain.Conif. Mexic. 16. 1857.
Type. Mexico. Mexico: Near village of San Mateo,
1857, Roezi s.n. (lectotype, P, here designated;isolectotype, Fl).
Pinus coarctata Roezl, Cat. Grain. Conif. Mexic. 16.
1857. Type. Mexico. Mexico: Mt. Tzompoli, 1857,
Roezl s.n. (lectotype, P, here designated).
Pinus nesselrodianaRoezl, Cat. Grain.Conif. Mexic. 16.
1857; Pinus monstrosaRoezl var.nesselrodiana(Roezl)
Carriere,Trait6G6n. Conif., ed. 2, 2: 526. 1867. Type.
Mexico. Between the volcanos Popocatepetland Iztaccihuatl, 1857, Roezi s.n. (lectotype, P, here designated;
isolectotype, Fl).
Pinus valida Roezl, Cat. Grain. Conif. Mexic. 16. 1857.
Type. Mexico. Roezl s.n., original materialnot located;
photographof seed cone from Roezi's collection at P
in G. R. Shaw's pine collection (lectotype, A, here
designated).
Pinus thelemannii Roezl, Cat. Grain. Conif. Mexic. 16.
1857; Pinus grandis Roezl var. thelemannii (Roezl)
Carriere,Trait6G6n. Conif., ed. 2, 2: 532. 1867. Type.
Mexico. Mexico: Mt. Tzompoli, on N side of mtn.,
1857, RoezI s.n. (lectotype, P, here designated).
Pinus horizontalis Roezl, Cat. Grain. Conif. Mexic. 17.
1857. Type. Mexico. Roezi s.n., original material not
located; photographof seed cone from Roezl's collection at P in G. R. Shaw's pine collection (lectotype, A,
here designated).
Pinus rubescens Roezl, Cat. Grain. Conif. Mexic. 17.
1857. Type. Mexico. Mexico: San Augustin, 1857,
Roezi s.n. (lectotype, Fl, here designated; isolectotype,
P).
Pinus carrierei Roezl, Cat. Grain. Conif. Mexic. 18.
1857; Pinus grandis Roezl var. carrierei (Roezl) Carriere, Trait6 G6n. Conif., ed. 2, 2: 533. 1867. Type.
Mexico. Hidalgo: Tulancingo, 1857, Roezl s.n. (lectotype, P, here designated).
Pinus planchonii Roezl, Cat. Grain. Conif. Mexic. 18.
1857; Pinus grandis Roezl var.planchonii (Roezl) Carriere, Trait6 Gen. Conif., ed. 2, 2: 534. 1867. Type.
Mexico. Hidalgo: Tulancingo, 1857, Roezl s.n. (lectotype, FI, here designated;isolectotype, P).
Pinus richardiana Roezl, Cat. Grain. Conif. Mexic. 18.
1857; Pinus endlicheriana Roezl var. richardiana
(Roezl) Carriere,Trait6Gen. Conif., ed. 2, 2: 542. 1867.
Type. Mexico. Mexico: Mt. Ajusco, 1857, Roezl s.n.
(lectotype, Fl, here designated;isolectotype, P).
Pinus decaisneana Roezl, Cat. Grain. Conif. Mexic. 19.
1857. Type. Mexico. Hidalgo: Pachuca, 1857, Roezl
Pinus inflexa Roezl, Cat. Grain.Conif. Mexic. 19. 1857;
Pinus rohustaRoezl var. inflexa(Roezl) Carriere,Traite
Gen. Conif., ed. 2, 2: 541. 1867. Type. Mexico. Mexico:
Mt. Ajusco, 1857, Roelz s.n. (lectotype, Fl, here designated).
Pinus ortgiesiana Roezl, Cat. Grain. Conif. Mexic. 25.
1857; Pinus tenangaensisRoezl var.ortgiesiana (Roez!)
Carriere,Trait6GCn.Conif., ed. 2, 2: 520. 1867. Type.
Mexico. Mexico: San Rafael, 1857, Roezl s.n. (lectotype, P, here designated).
Pinus rinzii Roezl, Cat. Grain. Conif. Mexic. 25. 1857.
133
Type.Mexico.Michoacan:Localitynot stated,1857.
Roezls.n. (lectotype,Fl, heredesignated;
isolectotype.
A).
Pinus rohanii Roezl, Cat. Grain.Conif. Mexic. 25. 1857.
Type. Mexico. Mexico: San Rafael. 1857, Roezl s.n.
(lectotype, P, here designated).
Pinus aztecaensis Roezl, Cat. Grain. Conif. Mexic. 26.
1857. Type. Mexico. Roezl s.n., original material not
of seed cone fromRoezl'scolleclocated;photograph

tionat P in G. R. Shaw'spinecollection(lectotype.A.
heredesignated).
Pinus zacatlanae Roezl, Cat. Grain. Conif. Mexic. 26.
1857.Type.Roezls.n.. originalmaterialnot located.
Pinus exserta Roezl, Cat. Grain.Conif. Mexic. 27. 1857;

Pinus protuberansRoezl var. exserta (Roezi) Carriere.
Trait6G6n.Conif.,ed. 2, 2: 524. 1867.Type.Mexico.

Mexico:Guarda,
alongrd.fromMexicoCityto Guernavaca, 1857,Roezls.n. (lectotype,FI, heredesignated;
isolectotype,BM).
Pinus cornea Roezl ex G. Gordon & Glendinning,Pinetum, Suppl. 74. 1862. Type. Mexico. Localitynot stated.
1857-1858, Roezi s.n. (lectotype, CGE (ex herb. Lindley 3187), here designated).
Pinus tenangaensis Roezl var. longifolia Carriere,Trait6
Gen. Conif., ed. 2, 2: 520. 1867. Type. Roezl s.n..
original materialnot located.
Pinus pescatorei Roezl ex Carriere,Trait6 Gen. Conif.,
ed. 2, 2: 555. 1867. Type. Mexico. Locality not stated.
1858, Roezl s.n. (lectotype, P, here designated).
Pinus montezumnaeA. B. Lambert forma macrocarpa
Martinez, Pinos Mexic., ed. 2, 219. fig. 177. 1948.
Type. Mexico. Morelos: Along rd. from Mexico City
to Cuernavaca, near Km 58, Jun 1940. M. Martinez
3437 (lectotype, MEXU, designated by Carvajal &
McVaugh, 1992: 79; isolectotype, MICH).
Pinus ntontezumaeA. B. Lambertvar. mezamnbrana
Carvajal, Phytologia59: 138. 1986 ("'m'ezenmiranuls").
Type.
Mexico. Jalisco: La Media Luna, 25 Sep 1981. Carvajt(l & Luce 3411 (holotype, CREG).
Leaves in fascicles of (4-)5 (rarely 3 or 6), variable, slender and droopingor more rigid, 1-1.3 mm
wide; stomata in (4-)5-7 lines on the abaxial face
and in 4-6 lines on each adaxial face. Seed con)es
variable in shape and size, broadly ovoid, ovoidoblong or ovoid-attenuate,8-20 x 5-10 cm when
open. Apophysis raised, especially toward the base
of the cone, transverselykeeled, often rugose. Umbo
raised, sometimes with a small prickle.
Note. Although Carvajal and McVaugh (1992)
reporteda maximumleaf width of 1.6 mm, but when
material associated with "doubtless montezumae"
cones was measured,no width greaterthan 1.3 mm
could be found in the collections cited here. We
assume that the thicker leaves have to be referredto
P devoniana instead.
Distribution and ecology. Mexico: In Nuevo

Le6n, SW Tamaulipas,Nayarit,S Zacatecas,Jalisco,
Michoacan,Mdxico, Distrito Federal,Quer6taro,Hidalgo, Morelos, Tlaxcala, Puebla, central Veracruz,
Flora Neotropica
134
Guerrero, Oaxaca, and Chiapas. Guatemala: In the
highland provinces. Pinus montezumae occurs in
a wide range of edaphic and climatic conditions
throughout the mountainous regions of central and
southern Mexico and the Guatemalan highlands. In
Nuevo Le6n it reaches from semi-arid pinyon-juniper
woodland up to cold-temperate mixed conifer forest.
Its altitudinal range is also great-(1200-)20003200(-3500) m overall, with occurrences below 2000
m mainly in Nayarit, Nuevo Le6n, and Tamaulipas.
It is most abundant and best developed in the temperate zone at ca. 2400-2800 m, with annual precipitation exceeding 800 mm, most of it falling from June
through September. Throughout its wide geographical range, P mrontezumae occurs together with many
other tree species, mainly in pine-oak and mixed pine
forests, but also with Abies religiosa, A. guatemalensis, and Cupressus lusitanica. Many of these forests have been depleted or even turned into smallscale farms with groups of scattered trees, among
which P montezumiae is often present. Phenology:
Time of pollen dispersal appears to be March-April
(material from Oaxaca and Veracruz).
Representative specimens examined. MEXICO. CHIAPAS:10 km NE of Las Margaritas,17 Feb 1973, Breedlove
33414 (MEXU, MO, NY). DISTRITOFEDERAL:2 km S of
San Miguel Topilejo. along rd. to Guernavaca, 27 Feb
1981, Mendez G. s.n. (MEXU). GUERRERO:Los Morros,
between Xochipala and Filo de Caballos, 14 Feb 1982,
Fonseca 193 (FHO, MEXU); Chilpancingo,25 Mar 1993,
Higman, Padilla & Styles 57 (EAP, FHO, HEH, MO).
HIDALGO:
Mineraldel Monte, "on the road from San Pablo
to San Pedro, near Real del Monte," 1838, Hartweg s.n.
(W), Dec 1839, Schiede 915 (HAL); La Majada, 20 km
NE of Zimapin, 6 Nov 1979, HertcindezM. 3863 (MEXU);
Ocotillos, Mar 1939, M. Martinez3625 (US); Apulco, 17
Mar 1980, Stead & Styles 637 (FHO, MEXU). MEXIco:
DesertaVieja. 1865, Bourgeau902 (P); Las Cruces, "Cerca
de Las Cruces, Valle de M6xico," 19 Dec 1950, Matuda
20905 (MEXU); HaciendaTomacoco 1857, Roezl s.n. (FI);
Mt. Iztaccihuatl, 1857, Roezl s.n. (P); between San Felipe
and Jala, 60 km along rd. from Toluca to Zitacuaro,2 Mar
1980, Stead & Styles 592 (FHO, MEXU); 8 km S of Rio
Frio. II Dec 1978, Vega A. 576 (MEXU). MICHOACAN:
"Municipio.Zinapecuarokm 116.5 on road from Toluca to
Morelia,"21 Jun 1988, Conzales M. 17002 (MEXU); Cerro
San Miguel, Pomocuaran,9 Apr 1979, Madrigal S. 3255
(MEXU, MS), 25 Feb 1980, Stead & Styles 536 (FHO,
MEXU); along rd. from Ocampoto San Felipe, nearZitacuaro, I Mar 1980, Stead & Styles 590 (FHO); Angangueo,
1838, sin coll., ex herb.LambertXXVI55 (BM). MORELOS:
Toro, 14 Nov 1907, Pringle s.n. (MICH). NUEVOLE6N:
Locality not stated, 5 Jul 1935, Mueller 2125 (A. MEXU,
MICH, MO); 22 km SW of Pueblo Galeana,Sierra Madre
Oriental, 18 May 1934, Mueller & Mueller 439 (MICH);
10 km NE of Laguna de Sanchez, Sierra Madre Oriental,
3 May 1983, Marshall 83-26 (FHO); near Monterrey,23
Jun 1888, Pringle 1964 (A, B, E, K, M, MO, US); "Sierra
Madre above Monterrey,alt. 2500-4000 ft.," 2 Apr 1906.
Pringle 10170 (F, GOET, M, MEXU, MICH, MO, NY,

W); I km N of 18 de Marzo, 29 Oct 1984, Rushforth497
Dec 1955,Zobel 10 (UC);Chipinque,

(E): La Encantada,
Dec 1955, Zobel 23 (UC). OAXACA:
San Jer6nimoSosola,
Oct 1940, M. Martinez3436 (MEXU); 4.5 km SW of La
Uni6n, 9 Nov 1984, Manzanero M. 508 (MEXU, MO).
PUEBLA:Miguel Hidalgo, 17 Mar 1983, Nee & Tavlor
25982 (F. FHO, MEXU, MO, NY, XAL). TAMAULIPAS:
Acahuales, along Hwy. 70 between Ocampo and Tula, 29
Apr 1994, Farjont& Mejia 332 (E, FHO, HEH, K, MEXU,
2 km SE of Palo Bendito, 26 Apr 1983,
U). VERACRUZ:
Nee & Taylor26760 (F, FHO, INIF,MO, NY, XAL); "inter
Cruz Blanca et Jalacingo,"Nov 1828, Schiede s.n. (HAL).
LOCALITY:
UNKNOWN
Locality not stated, 1857, Roezl s.n.
(Fl); locality not stated, 1857, Roezl s.n. (FI).
GUATEMALA. HUEHUETENANGO:5 km SW of San
Mateo Ixlatan, Sierra de los Cuchumatanes,6 Feb 1965,
Breedlove 8643 (F). SOLOLA:Along rd. between Solola
and Santa Lucia Uspantan, 20 Apr 1979, Stead & Styles
377 (FHO). TOTONICAPAN:
Alasca, 24 May 1979, Mittak
8894, 8895 (M).
15b. Pinus montezumae A. B. Lambert var. gordoniana (K. T. Hartweg ex G. Gordon) Silba,
Phytologia 68: 55. 1990.
Fig. 32G
Pinusgordonicana
K. T. Hartwegex G. Gordon,J. Hort.
Soc. London2: 79. 1847.Type.Mexico.Cerrode San
Juan,nearTepic,n.d., Hartwegs.n., originalmaterial
not located(lectotype,J. Hort.Soc. 2: 80, fig. (1847),
designatedby Carvajal& McVaugh,1992:79).
Leaves in fascicles of (4-)5, slender and drooping,
0.8-1 mm wide; stomata in 4-5 lines on the abaxial

face and in 3-4 lines on each adaxial face. Seed
cones ovoid-oblong or ovoid-attenuate,10-15 X 57 cm when open. Apophysis flat, especially toward
the base of the cone, with a weak transverse line,

often smooth. Umhboflat or obtuse.
Distribution and ecology. Mexico: Scattered in

the Sierra Madre del Sur N to Nayarit; also found
along the "Eje Volcanico Transversal."Its distribution as given here is probably incomplete, but it
appears to be sympatric with var. montezumae in
several locations. Its ecology is generally similar,
too, but it has not been found in the drier zones to
the northeast. Neither have we seen conclusive mate-
rial from Mesoamericanlocations.

Representative specimens examined. MEXICO.
GUERRERO:
NE slope of Sierra Madredel Sur, near Chilpancingo, 24 Dec 1894, Nelson 2180 (A); Filo de Caballo,
5 km SE of Fili de los Caballos,21 Apr 1985,SotoN. &
Aureoles C. 8322 (FHO). JALISCO:

Nevado de Colima, Jan
1942, M. Martitnez2212 (NY), Jul 1941, M. Martinez.3438
On N slope of Mt. Patamban,Jan
(MEXU). MICHOACAN:
Cerro San Juan.
1903, E. W. Nelson 6599 (A). NAYARIT:
1846, Hartweg s.n. (K, fragments), 1847, Hartwegs.n. (W,
cone). OAXACA:"Mountains W of Oaxaca," Sep 1894,
Nelson 1486 (A). PUEBLA:Buenavista. Huejotzingo. Jul
1941, M. Martine:z2002 (A).
Uses. Pinus montezumaeis exploited as a timber
tree throughoutits range. The timber is heavy and
strong and in demand for construction purposes.
Locally, especially in the southernpart of its range,
this (and other) species, growing often near habitations among fields as remnantsof more contiguous
forests, has its branches cut off due to the rural
population's incessant demand for firewood. Since
pines rarelyregenerate(only one Mexican species is
known to do so) and none can be coppiced or
pollarded indefinitely like many temperate broadleaved trees, this use will be detrimentaland lead to
the demise of the species in these areas.
As Carvajal and McVaugh (1992) have pointed
out, Pinus montezumae was described as a new
species, not as an avowed substitute for the later
homonym P. occidentalis Kunth.They lectotypified
Lambert'sname with a specimen (foliage) on a sheet
at BM, which is a part of collections made by
C. J. W. Schiede and F. Deppe. At W, a few pine
cones have been preserved from the Endlicherherbarium, among which is a cone with a small label
with the inscriptionby J. Lindley: "P montezumae
Lamb."andon the reverse:"of this cone Mr.Lambert
drew his plate." It does not say who collected it, yet
it perfectlyfits Lambert'sillustration,even as to size,
and must be consideredtype material.Loudon(1842)
referredmaterialdistributedby the HorticulturalSociety of London (Lindley?) to his P. montezumae
var. lindleyi, which is a taxonomic synonym.
Roezl's catalogue of Mexican pines. Benedict
Roezl (1824-1885) was the son of a Czech gardener,
who emigratedto Mexico in 1854, where he maintained a nurseryand a plantation(Mabberley,1985).
In June 1857 he issued a "Cataloguedes graines de
coniferes mexicains,qui se trouventchez B. Roezl &
Cie. a Napoles, pres Mexico," printed in Mexico
City. In it he named and described 82 new species
of Pinus, giving brief characters,remarkson their
beauty or excellence, and often a statement as to
where he had found them. Almost all of them are
from the Valle de M6xico and its surroundingmountains, many from the same mountain or locality.
There is no attemptto compare them. Many of the
descriptionsare completely overlappingeach other.
In 1858, a second catalogue was published in Paris,
on plant seeds for sale at Roezl & Co., in which
several new names appear,in addition to the 1857
names; but these, with one exception, received a
summarydescriptionin groups, which rendersthem
invalidly published.
The obvious mercantile purpose (especially suggested in that many names are honouringEuropean
135
nurserymenand estate owners of the time who were
likely to buy "novelties") also guaranteed a wide
dispersalof the catalogues.The publicationswere of
an ephemeral nature and are now difficult to find
in libraries. This perhaps influenced Schlechtendal
(1857-1858) to reprintall of Roezl's names in Lininaea and translatetheir descriptionsinto Latin, with
only a slight doubt expressed as to the question of
whether there really could be so many more new
pines in Mexico thanthose which Schiede and Deppe
had previouslybroughtto his attention.Roezl's catalogue of pines was furtherdistributedin France by
Vilmorin-Andrieux& Cie. and also translatedinto
Germanfor distributionin CentralEurope.Although
much of his seeds and other materialcame to England, we have not come across an English version
of the catalogue.
Contemporaryhorticulturalauthors on conifers,
such as Gordon (1862) and Carriere(1867), undertook to comment on them, but neitherhad a sound
taxonomic knowledge of the Mexican pines known
at that time on which to base their judgements.
Gordonsynonymized many of Roezl's names, often
incorrectly,while Carrieretended to accept most of
them, even validating several of Roezl's nomina
nuda by providingthem with descriptions.Parlatore
(1868), the more critical botanist, made short shrift
of them by reducing the entire list (and more-see
below) of names to synonyms of seven already
known species (see also Shaw, 1909).
What happened to the material on which these
names were based and who saw it? Shaw (1909),
who checked Gordon's reference (1862: 71, footnote), found that Roezl's collection was lost, which,
for all we know, is the case. He believed that it
might be "impossible today to find a complete authenticset." But threeherbariawere probabledepositories, and these were checked in 1994 by A. Farjon:
I) Lindley's herbariumin CGE and/or herbariato
which he distributedmaterialthroughthe HorticulturalSociety of London;2) Parlatore'sherbariumin
Fl; and 3) the material Carriere had seen, which
could have gone to Paris (P). Little of it remains in
Lindley's herbarium (CGE), but a few mounted
cones are there, bearing scant information as to
provenance. In Fl and P there are incomplete but
substantialsets, the most comprehensive of sheets
and separate cones being the one in Fl. In P, the
material of branches and cones was wrapped up
in coarse paper and stored with the carpological
collections. There is nothing in W, where Lindley
had sent otherearly pine collections for the attention
of Endlicher.
136
In the register of accessions at FI (Botanical Museum) from 1841 to 1938, on page 33 underthe date
14 October 1862, an entry is made for 435 specimens
of conifers taken from E Parlatorefrom his trip to
London. Parlatore had bought Roezl's specimens
from John Standish (1814-1875), gardener of the
Duchess of Gloucester at Bagshot Park, Surrey,England, while visiting the HorticulturalSociety in
August 1862. Parlatorewrote his determinationsand
"Parl.Conif." referringto his treatmentin the Prodroinms (Parlatore, 1868) on the labels bearing
Roezl's names. There are sheets and separatecones
in glass jars, but not all of these match up: there are
sheets without cones and vice versa. The sheets bear
original labels with the names in a large hand, also
found on the wrappings in P. This is believed to be
Roezl's writing;the rest is probablyby variousother
people who added "Mexico" (in Italian or French)
and donnedepar M. Roezl" and/orsometimes a location. In the catalogue of the Museum botanique(P),
under "entrees sorties" 1833-1864, there is on page
78 an entry under 18 December 1858, No. 30: "M.
Roezl (de Mexico) Echantillonsde Pins (cones) du
Mexique, 67" [= 67 specimens]. The labels have Fnumbers(accession) and also "Roezlnumbers";these
probably refer to one of his catalogues, but not to
the one publishedin 1858 in Paris.There is no proof
that Carriereever saw this material,but it may well
be that he did not annotate it. The wrappings had
become extremely brittle and have been removed,
retaining any pieces with text. In G. R. Shaw's
collection of pines at Harvard University (A) are
kept an album and a box with photographsof the
seed cones then at P and leaf fascicles, possibly also
from P, which Shaw probably received after the
publicationof his "Pines of Mexico" in 1909.
Investigation of this material shows that Roezl
issued many more names than the ones published in
the 1857 catalogue. Parlatoredid extremely well in
synonymizing them, save for the distinctionbetween
P montezumtaeand P gordoniana, which he did not
make, and a few errors with P hartwegii which we
now define in a narrower sense. Only a few of
Roezl's describednames are now left withouta trace
of original material in institutional herbaria. For
some of these, the descriptionis unambiguousas to
where to place it. If cited by Parlatore(1868), he
could have had a cone that subsequentlywent astray.
These names have here been synonymized along
with those for which a lectotypehas been designated.
The rest, only a few names now, is listed with
inserted cedis.
Pi'mlt montezcinae and its close ally P devoniana
Flora Neotropica
are two polymorphictaxa rangingfrom CentralMexico to Guatemala.They are also closely related to
the high-altitudespecies P hartwegii, but more distinct from it. The morphologicalcharactersused to
delimit P. nmotezumae and P devoniana are largely
quantitative and some are even subjective (Carvajal & McVaugh, 1992). The extremes of both taxa
are quite distinct,but many of the charactersused are
much weaker than those preferredin a phylogenetic
species concept (e.g., Stevens, 1991; Thiele, 1993).
A more or less congruentcombination of character
states (e.g., narrowerleaves, shorterfascicle sheaths
and leaves, and smallercones) is then used to distinguish P montezumae.But not all collections combine
these properties.Shaw (1909) therefore united both
under P montezumae, but Martinez (1945, 1948)
could not accept this and went off in the other
directionby describinga rangeof varietiesand forms
underboth species. In the presenttreatment,lumping
of the two seems prematurewithout furtherdetailed
research,which should aim firstof all at findingtruly
fixed characters (Davis & Nixon, 1992), possibly
molecularones.
Pinus Jmonitezumaevar. gordonianca: Gordon
(1847) gave a validating description of P gordoniani, a name coined by Hartweg for one of his
collections from Mexico. No specimen had been
found at K or elsewhere that could be interpreted
as original material; the illustration accompanying
Gordon'sdescriptionwas designatedas the lectotype
(Carvajal& McVaugh, 1992). A single cone at W.
from Lindley to Endlicher,labeled "P gordoniana,
Hartweg," could be original material. The leaves
Gordon described as slender and very long, to 40
cm; the cone illustrated is ovoid-attenuate, 12 cm
long (natural size) and has flat to slightly raised
apophyses and obtuse umbos. Shaw's (1909) treatment of P montezumae var. lindleyi Loudon, followed by Martinez(1945, 1948), disagrees with what
Loudon had in mind (see also Carvajal& McVaugh,
1992), which had ratherthick, rigid and short leaves,
and a cone with raisedapophyses. So P montezumnae
var. lindlevi auct., non Loudon, is indeed in need of
another name. But this need not be a new one: P
gordonlianaK. T. Hartwegex G. Gordon is available
for it, as it agrees with both Shaw's and Martinez's
concepts. We, indeed, believe it to be distinctenough
to be recognized at the rank of variety, for which
Silba (1990) provided the correct nomenclature.
Carvajaland McVaugh(1992), in synonymizing P
with P mnonitezumrae,
montezumaevar. mnezacibrana
suggested a possible hybrid origin of this taxon
involving P montezumaeand P. hartvegii. Including,
as they did, MartSnez 3438 (MEXU; Fig. 32G) in
that concept would bring P. montezumnae var gordoniana as we understand it into that putative orbit
of hybridization between two closely related, though
ecologically distinct, taxa.
16. Pinus devoniana Lindley, Edwards's Bot. Reg.
25: 62. Aug 1839 [& Allg. Gartenzeitung 7: 324.
1839]. Type. Mexico. Hidalgo: Cerro Ocotillo,
"found on the Ocotillo between Real del Monte
and Regla," 1838, Hartweg s.n. (lectotype, W, here
designated).
Fig. 33
Pinlts macrophyllcLindley, Edwards'sBot. Reg. 25: 63.
Aug 1839 [& Allg. Gartenzeitung7: 325. 18391;Pinus
iontezuniae A. B. Lambertvar. ntacrophylla(Lindley)
Parlatorein Candolle, Prodr. 16(2): 399. 1868. Type.
Mexico. Hidalgo:CerroOcotillo, "foundon the Ocotillo
between Real del Monte and Regla," 1838, Hartweg
s.n. (lectotype, W, here designated;isolectotype. K).
Pinusfilifolia Lindley.Edwards'sBot. Reg. 26: 61. 1839.
Type. Guatemala."Murray,HabitatGuatemala"(CGE),
"Santiago, Guatemala" (P), locality not stated (W),
1838?, [H(artwegs.n.] ex herb. Lindlev3179 (lectotype,
CGE, here designated;isolectotypes, P, W).
Pinlls grenvilleaeG. Gordon,J. Hort.Soc. London2: 77.
1847. Type. Mexico. Nayarit:Cerro San Juan, Hartiveg
s.n. (holotype, W).
Pinus wincesterianaG. Gordon,J. Hort. Soc. London 2:
158. 1847. Type. Mexico. Nayarit:"Cerrode San Juan
(or Saddle Mountain)near Tepic," 1845, Hartweg s.n.
(holotype, W. "Pinus Winchestericna";isotype, Fl).
Pinuls michoacaensis Roezl, Cat. Grain. Conif. Mexic.
26. 1857. Type. Mexico. Roezl s.n., original material
not located; photograph of seed cone from Roezl's
collection at P in G. R. Shaw's pine collection (lectotype, A, here designated).
Pinus magnifica Roezl, Cat. Grain. Conif. Mexic. 28.
1857. Type. Mexico. Michoacan:In mtns. near Morelia,
1857, Roezl s.n. (lectotype, FI, here designated).
Pinus ocanmpiiRoezl, Cat. Grain.Conif. Mexic. 28. 1857.
Type. Mexico. Michoacan:"On hacienda of Melchior
Ocampo, near Morelia,"1857, Roez s.n. (lectotype, Fl,
here designated;isolectotypes. A, P).
Pijnus zitacuarensisRoezl, Cat. Grain. Conif. Mexic. 29.
1857.Type. Mexico. Michoacan:Zitacuaro.1857. Roezl
Pinus nec-plus-ultra Roezl, Cat. Grain. PI. Mexic. 6.
1858. Type. Mexico. No locality given, 1858, Roezl s.n.
Pinus zitccuarensis Roezl var. nitidacRoezl ex Carriere,
Trait6Gen. Conif.. ed. 2. 2: 554. 1867. Type. Mexico.
Location unknown, 1857. Roezl s.n., original material
not located.
Pinus verschaffeltiiRoezl ex Carriere.Trait6Gen. Conif.,
ed. 2, 2: 555. 1867. Type. Mexico. Location unknown,
1857, Roezl s.n. (lectotype, P, here designated;isolectotype, Fl).
Pinus ptawlowskiana Roezl ex Carriere, Trait6 Gen.
Conif., ed. 2, 2: 556. 1867. Type. Mexico. Roezl s.n.,
original materialnot located; photographof seed cone
from Roezl's collection at P in G. R. Shaw's pine
collection (lectotype, A, here designated).
137
PittusmichoacanaMartinez,Anales Inst. Biol. Univ.
Nac. Mexico 15: 1. 1944.Type.Mexico.Michoacan:
Parangaricutiro.
May 1940,M. Martinez3443 (holotype,MEXU).
Pitus michoacana Martinez var. corliuta Martinez.Anales Inst. Biol. Univ. Nac. Mexico 15: 1. 1944. Type.
Mexico. Jalisco: Tequila Iprobablyon Volcin de Tequila, S of Tequila].Oct 1941, M. Martinez3446 (holotype. MEXU).
Pinus michoacainaMartinezvar. quevedoi Martinez,Anales Inst. Biol. Univ. Nac. Mexico 15: 1. 1944. Type.
Mexico. San Luis Potosi: El Tablon[= SierraEl
Tablon?]. Oct 1940. M. Martinez 3448 (holotype.

MEXU).
Pintis michoacana Martinezvar. cornutta Martinezforma
naivaritana Martinez,Pinos Mexic., ed. 2, 266, fig. 215.
1948. Type. Mexico. Nayarit:Compostela, May 1941.
M. Martitez 3447 (holotype, MEXU).
Pinls michoacanc Martinez forma procera Martinez.
PinosMexic.,ed. 2, 257, fig. 207. 1948.Type.Mexico.
Jalisco: Tuxpan, Los Mazos. n.d.. M. Martinez 2433
(lectotype,F. heredesignated).
Pionus nichoacana Martinez forma trumi,ida

Martinez.
PinosMexic.,ed. 2, 258, fig. 208. 1948.Type.Mexico.

Oaxaca:San SebastianEtla ['Ixtlahuaca.
Etla"on the
label,but the localitycited in Los PinosMexicanos,
ed. 2 (1948), is Telixtlahuaca,
Oaxaca],Oct 1940,M.
Martinez3444 (holotype.MEXU).
Tree, medium or sometimes large, height to 2030 m, dbh to 80-100 cm. Trunkmonopodial,usually
erect, slender or massive in old solitary trees. Bark
thick, scaly, very rough, with elongated plates divided by deep longitudinal black fissures, reddish
brown to dark brown, on young trees rough, soon
fissured.Branches of first order long, spreadingand
assurging,or ascendingnearthe top, often persistent;
branchesof higher orders sparse, spreadingand ascending, the highest order branchesassurging,forming an open, broad pyramidal or domed crown.
Shlootsvery thick (15-20 mm), rigid, curved, very
rough and scaly, with persistent, decurrentpulvini.
Cataphylls large, up to 20-25 mm long, subulate,
scarious,recurvedor reflexed,darkbrownto blackish
grey, with erose-ciliate margins, persisting ca. one
season longer than the leaf fascicles. Vegetativebuds
large, terminalbuds 20-40 mm long, lateralssmaller,
ovoid-acute, not resinous; the scales imbricate but
spreadingapically, subulate-linear,orange-brownto
reddishbrown, with long-ciliate (3-5 mm) margins,
giving the buds a villose appearance. Fascicle
sheaths very long, up to 30-40 mm, with ca. 10
imbricate,orange-brownscales with whitish ciliate
margins, persistent, on mature fascicles (20-)2535(-40) x 2-3 mm, tightly enclosing the fascicle,
turningdark brown or blackish, often very resinous.
Leaves in fascicles of 5, rarely 4 or 6, in dense,
wide-spreading tufts toward the ends of upturned
Flora Neotropica
138
'
i
FIG. 33. Pinus (Ievolnia(li (A-C. Higman. Padilla & Styles 62; D. G. Stead & Styles 28IA; E, F. Hughes 1349). A.
Shoot with leaf fascicles, cataphylls. bud and immatureovuliferous cone. B. Fascicle sheath. C. Cross section of leaf. D,
E. Maturecones. F. Basal seed scales of cone attached to branch.G. Seed scale (three views). H. Seed with articulate
xO.5:
B. D-H.
C. x
x30.)
A, B.
D-H, x
wing. (Magnifications:
(Magnifications:A.
win~.
O.5:C.
30.)
branches, persisting 2-3 years, rigid, straight, or
flexible and drooping, (17-)25-40(-45) cm X 1.11.6 mm, with serrulate margins, acute-pungent,
bright,lustrous(dark)green. Stomataon all faces of
leaves, in (4-?)5-9(-10) lines on the abaxial face
and 3-6 lines on each adaxial face. Leaf anatomy:
with
Cross section triangularor transverse-triangular,
flat or convex abaxial side; cuticula and epidermis
thick; hypodermismulti-layered,with (irregular)intrusions into the mesophyll; resin ducts large, 3(-4),
medial, near the edges, sometimes 1-2 (smaller)
ducts internal;stele terete or weakly trigonous;outer
walls of endodermisnot thickened;vascularbundles
2, distinct but approximate,often partly surrounded
by sclerenchym layers. Pollen cones crowded near
the proximalend of a new shoot in short,often dense
clusters, subtended by long, subulate bracts with
ciliate margins, cylindrical, 20-40 mm long, pinkpurplish,turninglight brown. Microsporophyllspeltate-cordate,with a radially striate abaxial surface
and hyaline-erose margins.Seed cones subterminal,
solitaryor in pairsor whorls of 3-4. Immaturecones
initially often erect, after pollination recurving on
thick, bracteate peduncles, ovoid-oblong, purplish
brown,maturingin two years. Maturecones on short,
persistent peduncles, leaving a few scales on the
branch when falling, variable in size and shape,
typically ovoid-oblong, with an oblique base, or
curvate to cornute (often by insect-damage), 15-35
x 8-15 cm when open. Seed scales ca. 175-225,
partingwhen the seeds are mature,spreadingwide,
straight or slightly recurved, thick woody, rigid or
flexible with some force, margins thin, light brown
or purplishblack on the abaxial side, (light) brown
with lighter marks of seed wings on the adaxial
side. Apophysis mostly raised, transversely keeled,
rhombic in outline, with irregularsides, up to 25
mm wide, often radially striated, in various shades
of brown. Umbo dorsal, raised, flat or depressed,up
to 7 mm wide, terminatingin a small, usuallydeciduous prickle, grey-brown. Seeds obliquely broad
ovoid, flattened,8-10 X 5-7 mm, light brown,often
with dark spots. Seed wings articulatebut effective,
held to the seed in two oblique claws, obliquely
ovate to oblong, with a straightside, 25-35 X 1015 mm, light brown with darker stripes. Seedling:
Numberof cotyledons not observed;many seedlings
develop a "grass stage" with delayed apical growth
of the stem.
Nayarit,Jalisco, Zacatecas,Aguascalientes,San Luis
Potosi, Queretaro,Hidalgo,Michoacan,Mexico, Distrito Federal, Morelos, Tlaxcala, Puebla, Veracruz,
139
Guerrero,Oaxaca, and Chiapas. Guatemala:In the
southernhighlands. In both countries it is a constituent of relatively open, often secondary pine-oak
forests; secondly of secondarymixed pine forest, or
with Pinus oocarpa invading burnedmountainsides.
It is well-adaptedto withstand fires by its juvenile
"grassstage" similar to that of P palustris of the SE
United States. The pines most commonly associated
with it vary somewhat from north to south and
include P. oocarpa, P. montezumae, P. pseudostrobus,
P. maximinoi, and, less frequently, P. cembroides at
lower altitudes and P hartrwegiiat higher altitudes.

Common are also Quercus and Liquidambar and,
in the understorey,Calliandra, Leucaena, Acacia,

Dodonaea, Gaultheria, and Mimosa. Its altitudinal
rangeis (700-)900-2500(-3000) m. Pinus devoniana
grows on a variety of soils, often of volcanic origin.
The climate is warm-temperateto subtropical,with
annualprecipitation1000-1500 mm and a dry season
from November to May. Phenology:Time of pollen
dispersal not recorded.
Representative
specimensexamined.MEXICO.CHIAPAS:Comitan. 11 Mar 1993. Higman. Padilla & Styles 10
(EAP,FHO,MO.ESNAC);Bochil,14 Mar1993,Higilan,
Padilla & Styles 23 (EAP.FHO. MO, ESNAC): Motozintla.
16 May 1945. Matuda 5534 (MEXU. MO, NY); 7 km
from Comitanat FranciscoSarabia, I Apr 1974. Styles 121
(FHO); 4 km E of Amatenango del Valle, along Hwy.
190 to Comitan, 27 May 1985, Villaseior & Thomais829
Omiltemi, Plan de Potrerillos.
(MEXU, NY). GUERRERO:
24 Apr 1985, CasteloN. 446 (MEXU);along rd. to Chilpancingo, 5 km fromChilapa.24 Mar 1993, Higmatn,Padilla &
Styles 54 (EAP. FHO, MO, ESNAC); SierraAlquitran,10
Jun 1968, Kruse 1778 (MEXU); 2 km from Los Morros
along rd. to Filo de Caballos, 5 Mar 1984, Lorea 2986
(FHO, MEXU); near Taxco. Jun 1940. M. Martitnez3445
(MEXU); W of Petlacala.4 Jan 1938, Me.ria9083 (F. K.
MO, NY, S, UC. US); 1I km W of Chilpancingo, 13 Nov
1973, Quero et al. 2264 (FHO, MEXU, UC); Filo de
Caballo, 20 Apr 1985. Soto N. & Aureoles C. 8281 (FHO).
HIDALGO: 20 km NE of Pachuca along rd. to Huasca at
El Octal, Gardner& Knees 5259A (E). JALISCO:Cerro La
Bufa, 2 Feb 1927, Mex'.r 1627 (A, BM, F. MICH. NY.
UC); Talpa de Allende, on rd. to Cuale, 8 Nov 1984.
Rusliforth609 (E); Sierra del Tigre, 10 km from Ciudad
Guzmfin, 10 Mar 1974, Styles 51 (FHO). MICHOACAN:
CerroAzul, nearMorelia,28 Mar 1910,Arsene5477 (MO);
Dos Aguas, 19 Feb 1980, Klaus s.n. (WU); Los Conejos,
10 km SW of Uruapan, 1900, Little 17837 (K. MS, US);
17 km along rd. from Ciudad Hidalgo to Mil Cumbres.9
Feb 1980, Stead & Styles 466 (FHO). MORELOS:
CuernaCerro Los
vaca, Nov 1979, Klaus s.n. (WU). NAYARIT:
Lobos, NE of Santa Mariadel Oro, 19 Aug 1959, Feddema
664 (MEXU. MICH);8 km W of Chapalilla, I Nov 1984,
Marshall 84F-104 (FHO); loc. unknown, Feb 1941. M.
Matrtinez2413 (F, GH, US); El Tortuguero,30 Oct 1979,
Salazar J. I (MEXU); 6 km NW of Xalisco. 30 Mar 1987.
Dirt rd. to San Lucas.
Tellez V 10121 (MEXU). OAXACA:
near Hwy. 190, ca. 30 km from Oaxaca. 23 Apr 1994.
Flora Neotropica
140
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Piu
Farjon & Mejia 314 (E, FHO. HEH, K, MEXU, U); La

Soledad Tectitlan, 26 Apr 1986, Gereau & Martin 2000
(FHO, MO); Presa Benito Juarez,near Santiago Lachiguiri,
ca. 55 km NW of Tehuantepec,29 Mar 1989, Hughes 1349
(FHO. K. MEXU); Portillo de Nejapa, 17 km NW of El
Coyul. 8 Nov 1993, Hughes 1773 (FHO, K, MEXU);
Ayoquezco, 29 Oct 1979, Ortega C. s.n. (MEXU); 5 km E
of San PedroCoatlan, 18 Mar 1988, TorresC. et al. I 1850
(MEXU). PUEBLA:Near Calchicomula, Jul 1901, Rose &
Nochistlan, n.d., M. Martiinez
Hay 5781 (A); ZACATECAS:
LOCALITY:
2429 (A). WITHOUT
Specimen labeled "Pinus
filifolia" presentedto Parlatore,1862, ex herb. Balfour s.n.
(FI); specimen labeled "Pinus Winchesteriana,Marquisof
Winchesterspecimen of seeds 1845," ex herb. Gordon s.n.
(K, det. not certain);specimen labeled "Pinus Devoniana,"
1857, Roezl s.n. (Fl).
Cubulco, Sep 1870,
GUATEMALA. BAJAVERAPAZ:
LoBernoulli & Cario 1085 (GOET). CHIMALTENANGO:
cality not stated, 3 Jul 1951, Schwerdtfeger44 (F); Zaragoza, along Pan American Hwy. between Patizicia and
Chimaltenango,22 Apr 1979, Stead & Styles 381 (FHO).
GUATEMALA:
Along rd. from San Jose Ocafia to San Juan
Sacatepeque. 22 Mar 1979, Stead & Styles 278 (FHO).

HUEHUETENANGO:
Along Hwy. CA I between Huehuetenango and Colotenango,above the Rio Cuilco, 19 Apr 1994,
Farjon & Mejia 309 (E, FHO, K, MEXU, U, HEH);
Malacatacingo,3 Apr 1993, Higman, Padilla & Styles 62
(EAP, ESNAC, FHO, MO); near village of Gracias a Dios
W of Finca la Trinidad, ca. 25 km N of Nenton, 2 Apr
1989, Hughes 1357 (FHO, K); 13 km from Aguacatan. 5
Apr 1979, Stead & Styles 339 (FHO). QUEZALTENANGO:
Sacatepequez, Mar 1892, Donnell-Smith 2632 (K, US);
along Hwy. CA I. near Chiquibal, 18 Apr 1994, Farjon &
Mejia 308 (E, FHO, HEH, K, MEXU, U); 2 km SW of
village Alotenango, on track to Volcin de Fuego, 9 Mar
1984, McCarter& Styles 184 (FHO); Santa MariaCauque,
7 Feb 1978, Stead & Styles 142 (FHO). SOLOLA:Volcin
SantaClara.5 Jun 1942, Stevermark47038 (F). TOTONICAPAN: Casa Blanca, 19 km along rd. from Totonicapan to
Santa Cruz del Quiche, 12 May 1979. Stead 310 (FHO).
Uses. Pinus devoniana is a common tree throughout the mountainous parts of southern Mexico and
Guatemala, often encountered in open, degraded
pine-oak or pine forest. As it is not a very tall,
straight-boledtree and has brancheslow on the trunk,
it is not often selected as a timber tree. On the
other hand, it is locally used and, like other pines,
increasinglyfor firewoodby the growing ruralpopulation.
This species-in its more extreme forms easily
distinguishable from its relatives (in particularP.
,nontezuinae)by its thick, shaggy branches,its very
long and stiff, lustrous green leaves with very long,
dark, resinous fascicle sheaths, and the very large,
thick woody cones-occurs in a bewildering array
of polymorphismsespecially of cone forms. Also,
intermediatesoccur between this taxon and P. montezumae, with charactersof either species shared in
populations of one or the other. Carvajal and
McVaugh (1992) compared this situation with the
species complex which Stead and Styles (1984)
called the "pseudostrobus"group, and spoke of the
"montezumae"group, in which P hartwegii is to be
included. Shaw (1909) considered them as a single
species, P. montezumae, with four varieties: var.
montezumae,var. lindlevi, var. rudis, and var. hartwegii, but in his monograph(Shaw, 1914) he dropped
even these. He considered the wide range in size
of both leaves and cones of no taxonomic value and
believed that they probably increased with warm,
subtropicalclimatic conditions furthersouth. Numbers of leaves per fascicle increase in a similar
fashion, according to Shaw, and he believed all
these characters to be "accurately correlated with
altitude and exposure." His lumping (Shaw, 1914),
however, did not prevent him from recommending
monographicstudy of the "montezumae"group, an
undertakingwe still await today (Styles, 1993: 409).
The confusion originates at least in part also
from a profusion of names published by Lindley,
Gordon, and Roezl, and then again after Shaw's
monographic work by Martinez, who included a
number of varieties and forms under a new name:
Pinus michoacana.
Lindley (1839) described three "quite new" species from C. T. Hartweg'scollections of pines from
Mexico, all gathered from the vicinity of Real del
Monte in Hidalgo. Hartwegbroughtcones (for seeds)
and foliage (judging fromthe illustrationsmade from
it in Loudon, 1842), but the latter has rarely been
preserved. We regard P devoniana and P macrophylla, as a single taxon for which the first name
should be retained. A year later, Lindley (1840)
published P filifolia from Hartweg's collection on
the Volcan de Fuego in Guatemala,with very long
141
leaves (45 cm). Like the othermaterial,it was distributed throughthe HorticulturalSociety of London,of
which Lindley was secretary, to estate owners in
GreatBritain,and also to Endlicherin Vienna,where
good cone specimens (W) have "miraculously"survived the ravages of the Second WorldWar.
Gordon's (1847) names P grenvilleae and P.
wincesterianawere probablybased on a single gathering made by Hartwegon Cerro de San Juan near
Tepic in the state of Nayarit. Because no original
material has been found in Gordon's herbariumat
K, Carvajaland McVaugh(1992: 51) designatedthe
relevant published illustrationsof P. grenvillae and
P. wircesteriana as the lectotypes of these species.
However, as with Lindley's taxa, there is original
material of these in W and Fl (the seeds of "P.
Winchesteriana"in K cannotbe identifiedunambiguously) which must be considered the holotypes (W)
and isotype (FI), of Gordon's taxa. They are to
supersedethe lectotypes (illustrations)designatedby
Carvajaland McVaugh(Art. 9.13 in Greuteret al.,
1994).
It has, in addition, been possible to retracemany
of Roezl's "new species" to original materialin A,
CGE, and especially Fl and P (see for discussion
under P montezumae),and they have accordingly
been lectotypifiedhere. None of them deserves recognition as distinct taxa, as Parlatore (1868) had
alreadyconcluded.
Martinez (1945, 1948), reacting to Shaw's very
inclusive treatmentof P. montezumae,set up a "grupo
Michoacana"with a new species: P miichoacana.He
considered this species likely to correspond with
Lindley's species P devoniana, P russelliana, and
P. macrophylla,but probablyfelt he could not use
any of these names as his only source of information
for them seems to have been Shaw (1909, 1914). He
proceeded to describe a number of varieties and
formsbased on sizes and shapesof cones, apophyses,
and umbos of the scales. However,to his six varietal
shapes of apophyses (Martinez, 1948: 259) several
more, as well as intermediates,could be added from
the materialhere examined.A furtherproblemis that
they do not correlate with cone sizes and shapes,
such as Martinezsuggested.
Geographically,there is in our view no clear separation into subspecific taxa and in Guatemala,from
where Martinez did not study any collections, as
many taxa could be assembled. Perry (1991), who
followed Martinez'snomenclature,especially maintained var. cornuta but, apart from his findings in
turpentine compounds, gave no new evidence.
Curvedcones do not always coincide with pyramidal
142
apophyses and/orraisedumbos (e.g., Farjon& Mejia
309). This "variety"seems to be sympatricwith the
species sensu stricto. That none of these authors
had a clear notion of Lindley's concept of Pinus
devoniana is aptly illustratedby the lectotype in W.
It consists of three cones, two of them (bound with
rope to preventthem from opening) ca. 25 cm long,
only 5-6 cm wide and very "cornute,"and all three
labeled in Lindley's handwriting "P Devoniana
Lindl."Very recently,materialfrom very near to the
type locality (or at it) was collected (Gardner &
Knees 5259A, E) with cones matchingthe type collection at W. We agree with Carvajaland McVaugh
(1992) that there are, from the existing evidence, no
convincing reasons to maintain these varieties nor
to make the necessary new combinations under P
devoniana. However, the polymorphyof the "mtontezumale"group would appear to ask for more detailed research using different and perhaps more
useful approaches, of which molecular methods
should particularlybe mentioned.
17. Pinus douglasiana Martinez, Madrono 7: 4.
1943. Type. Mexico. Nayarit:Cerro Juanacata,Jul
1940, M. Martfinez3429 (holotype, MEXU).
Fig. 34
80-100 cm. Trunkmonopodial, straight, columnar,
terete and slender, usually clear of branches2/ of its
length. Barkrough,scaly, divided into large, irregular
plates and deep fissures,dark reddishbrown, weathering grey-brown; on young trees and branches
smooth or scaly, reddish brown. Branches of first
order long, slender, spreading, or ascending in the
upperpartof the crown; of second and higherorders
spreading or ascending, flexible. Crown in young
trees pyramidal, in older trees generally rounded,
dense or open. Shoots uninodal, smooth or rough
with prominent, decurrent, mostly exfoliating pulvini, dark brown, not glaucous. Cataphyllssubulatecaudate, spreading or recurved, with erose-ciliate
margins, brown. Vegetativebuds ovoid-conical, the
terminal buds 15-25 x 10-15 mm, the laterals
smaller, not resinous, brown;the scales free, spreading or recurved,subulate-linear,with ciliate margins.
Fascicle sheaths persistent,remaininglong, (15-)2035 mm, with 7-10 imbricate,bracteate,light lustrous
brownscales with ciliate margins,weatheringgreyish
brown.Leaves in fascicles of 5, rarely4 or 6, spreading and drooping in dense tufts, persisting 2-2.5
years, slender, lax or sometimes more rigid, 22-35
cm X 0.7-1.2 mm, margins serrulate,apex acute,
Flora Neotropica
light yellowish green to glaucous-green.Stomata on
all faces of leaves, in 3-4 lines on the abaxial face
and 3-4 lines on each of the adaxial faces. Leaf
anatomlv:Cross section triangular;hypodermismultilayered, with numerous intrusions into the mesophyll, some connecting with the endodermis(septal);
resin ducts (2-)3, medial, large; stele terete; outer
cell walls of endodermisthickened;vascularbundles
2, connate but distinct. Pollen cones numerous,
crowded near the proximal end of a new shoot,
subtendedby subulatebracts,forminglong, spiculate
clusters, cylindrical, 20-25 x 4-5 mm, pinkish
brown at maturity.Microsporophyllsdistally peltate,
ca. I mm diam., with an erose-denticulate, lightcoloured margin.Seed cones subterminal,solitary,in
pairsor whorls of 3-4 on stout, recurved, 10-15 mm
long peduncles which fall with the cone. Immature
cones on bracteatepeduncles, ovoid, with pungent
umbos, ca. 15 x 10 mm, purplish or glaucous.
Maturecones ovoid to ovoid-attenuatewhen closed,
often slightly curved, more broadly ovoid with a
flattenedbase when opened, then 7-10 x 5-7 cm.
Seed scales ca. 110-130, partingto release the seeds
except at the base, (thin) woody, oblong, straightor
slightly curved, light or dark brown abaxially, light
brown, with often indistinct lighter marks of seed
wings adaxially.Apophysis nearly flat or raised and
transverselykeeled, radiallystriateor grooved, rhombic or pentagonal in outline, nearly symmetrical
aroundthe cone, 11-17 mm wide, light brown, reddish brown or dark brown. Umbo dorsal, raised,
transverselyrhombic,5-7 mm wide, darkerthan the
apophysis, prickle absent. Seeds obliquely obovoid,
slightly flattened, 4-5 x 3-3.5 mm, light grey or
brown, often with dark spots. Seed wings articulate
but effective, held to the seed by two oblique claws,
oblong, with a straightand a curved side, 18-24 X
7-9 mm, light yellowish brown, translucent,sometimes with a darkerbrown tinge. Number of cotyledons not observed.
Distribution and ecology (Map 9). Mexico:
Mainly in Jalisco, Michoacan, Mexico, and N Morelos, but extending northwardinto Nayarit and the
crest of the Sierra Madre Occidental on the border
between Sinaloa and Durango; southward it occurs
locally in Guerreroand Oaxaca. The latter was collected as Pinuspseudostrobus;in recentyears several
populationsformerlyso identifiedhave turnedout to
be P douglasiana (cf. Critchfield & Little, 1966:
map 52). It is a constituentof mostly mixed pine or
pine-oak forests at (1100-)1400-2500(-2700) m in
warmto temperateclimatic zones. The annualprecipitation differs with altitude but is roughly 1000 mm
143
'd,
FIG. 34. Pinus douglasiana (A, C, D, Styles 24: B, Stiles 76; E, Higman, Padilla & Stiles 51; F-H, Farjon & D.
Mejial344). A. Branch with leaf fascicles, pollen cones, and immatureovuliferous cone. B. Shoot with flushing leaf
fascicles and young ovuliferous cones. C. Leaf fascicle. D. Cross section of leaf. E, E Ovuliferouscones. G. Seed scale
(two views). H. Seed (two views). (Magnifications:A-C, E-H, x0.5; D, x40.)
Flora Neotropica
144
in most areas. Common associated pines are Pilns
pseudostrobus,P herrerae,Pi leiophylla, P lawsonii,
P ayacahuite in the southern part of its range, and
sometimes P oocarpa and P devoniana at lower
elevation and drier sites, respectively.At the highest
and wettest sites it can occur with Abies, Picea (in
Durango). or Cupressus lusitanica. In many areas
Quercusspp. are co-dominant;a shift in forest composition toward broad-leavedtrees may occur also
by selective cutting of pines. Phenology: Pollen dispersal was observed to occur in Michoacanat 1500
m near the end of February(Styles 24) and may
occur up to a month later at high altitudes.
dermis were found to be similar in P maximinoi

where the two species are sympatric (i.e., in the
southernpart of the range of P douglasiana). Shaw
(1909) had observed the same structureof the hypo-
between Bolafos and Guadalajara,20 Sep 1897, Rose 3025
ana and P nmaximinoi versus the persistent pulvini
dermis in P mnaximinoi, including Hartweg 620, the
type collection from Guatemala.

Distinction between P doulglasiana and P pseu-
dostrobus can be similarly vague, as was also observed by Stead and Styles (1984). Few individual
characters appear to have a diagnostic value by
themselves and only in combinationcan they be used
to distinguish between these taxa. The most useful
appearto be the hypodermaldevelopment, which is
"normal"in P pseudostrobus,and the cone peduncle,
Representativespecimensexamined.MEXICO.Duwhich persists on the branchin the latterspecies but
ChavarriaNuevo. 3 Apr 1991, Styles, Favela et al.
RANGO:
7 (FHO, MEXU). GUERRERO:Carrizalde Bravo, 20 Aug falls with the cone in the other two species. Carvajal
1984, Lorea3124 (FHO, MEXU). JALISCO:
Along rd. from and McVaugh(1992) mentionthe decurrent,exfoliatTonayato CiudadGuzman,nearEl Jazmin,4 May 1994, ing bractbases (pulvini) on the shoots of P douglasiFarjon& Mejia 344 (E, FHO, HEH, K, MEXU, U);
of P pseudostrobus to distinguish them "quickly
and without much question."However, the frequent
misidentificationof these pines would indicate that
K);Tecatitlan.14 Mar1974,Styles76 (FHO.K);Cerrode such "ease" of identificationis not
experienced by
Alotica, II Mar 1974, Styles & Kemp60 (FHO). MeXIco:
Valle Bravo,NW of Lagunade Avandaro,2 Oct 1965, most collectors of pines to date.
Ca. 3.5 km S of
Sousa 2602 (MEXU).MICHOACAN:
Paracho de Verduzco, at Km 35 along Hwy. 37, 6 May 18. Pinus maximinoi H. E. Moore, Baileya 14: 8.
1994, Farjon & Mejia 350 (E, FHO, K. HEH. MEXU, U);
1966.
Fig. 35
W
(US); 10 km fromCiudadGuzmin,at G6mezFarias,21
Feb 1980, Stead & Styles 524 (FHO): Sierra el Tigre. 10

km from Ciudad Guzmin, 10 Mar 1974, Styles 50 (FHO,
of Tumbiscatio,27 Feb 1974,

Las Playitas Forest Sta.
Styles 24 (FHO. K); Dos Aguas, W of Aguililla, 2 Mar
Benito JudirezNational
1974, Styles 42 (FHO, K). OAXACA:
Park, 20 Mar 1993, Higman. Padilla & Styles 43 (EAP,
ESNAC. FHO, MO); Miahuatlan.22 Mar 1993, Higman,
Padilla & Styles 51 (EAP, ESNAC, FHO, MO). SINALOA:
Guanera.16 Feb 1980, Stead & Styles 476 (FHO, MEXU).
Uses. Pinus douglasiana is, along with other species with which it often occurs, an importanttimber
tree in most of its range. It is, however,not especially
sought out for tree-breedingpurposes, perhaps due
to the fact that it is relatively unknownto foresters.
Martinez (1943) distinguished his new species
from Pinus pseudostrobus var. tenuifolia (Benth.)
Shaw (- P maxininoi H. E. Moore) by the morphology of the leaves, which in P douglasiana are less
slender, flexible, and drooping. The hypodermis of
this species is very strongly developed; the sclerenchyma cells form several intrusions into the mesophyll, often connecting with the endodermis. The
cones are rather similar but have a thicker, more
raised apophysis of the seed scales and are smaller
than those of P pseudostrobus. These differences
were confirmedin a morphometricstudy using Principal ComponentsAnalysis (PCA) by Stead (1983),
and the taxon was recognized at species rank by
Stead and Styles (1984). The intrusionsof the hypo-
Pinus tenuifolia Bentham, PI. Hartweg. 92. 1842, non

R. A. Salisbury, 1796; Pinus pseudostrobusLindley var.
G. R. Shaw,[PinesMexico]Publ.
(Bentham)
tenuifoilia
Arnold Arbor. 1: 20. t. 13. 1909;Pinus ldouglasiana
Martinez var. mariminoi (H. E. Moore) Silba, Phyto-
NearGuatemala
logia68: 50. 1990.Type.Guatemala.
City, "CanalesMts.," 1841, Hartweg620 (holotype, K,
ex herb.G. Bentham1854;isotypes,B. BM,CGE.Fl,

K (see below), MO, NY, P, W).
Piniusescalndoniana
Roezl,Cat.Grain.Conif.Mexic.24.
1857. Type. Mexico. Mexico: Mt. Tzompoli. 1857.
Roe:lIs.n. (lectotype,FI,designatedhere).
PiinushoserianaRoezl, Cat. Grain.Conif. Mexic. 24.
1857. Type. Mexico. Mexico: Mt. Tzompoli. 1857,
RoeZs.n. (lectotype,FI,designatedhere).
Piinustiomlpoliana
Roezl,Cat.Grain.Conif.Mexic.24.
1857. Type. Mexico. Mexico: Mt. Tzompoli, 1857,
Roezls.n. (lectotype,Fl, designatedhere).
Tree,medium to tall, height to 20-40(-50) m, dbh
to 70-90(-I(X)+) cm. Trunk monopodial, straight,
slender,terete,the lower half of the bole usually free
of branches. Bark thick on the lower part of the
trunk,with relativelysmooth, longitudinalplates and
deep longitudinal fissures, grey-brown, on young
trees and branchesmore or less smooth, grey-brown.
Branches of first order slender, longest in the upper
partof the crown, often distinctly whorled,spreading
or ascending; branches of higher orders sparse,
145
'
"F
/
scale
.Seed.
(three
views).
(MagA,
X.5;C,x4
FIG. 35. Pinus maximinioi(A-C, F. Hughes 1264; D. Stead & Sn/les 135; E. Hignian, Padilla & Sn/?aes13; G-1.
McCarrter/46). A. Branchwith leaf fascicles and pollen cones. B. Leaf fascicle. C. Cross section of leaf. D. Shoot with
flushing leaves and pollen cones. E. Shoot with flushing leaves and ovuliferous cones. F, G Ovuliferous cones. H. Seed
scale (three views). I. Seed. (Magnifications:A. B. D-IA XO.5 C, X40.)
146
spreading,slender and flexible, forming a pyramidal
crown in young trees and an open or dense, rounded
crown in maturetrees. Shoots uninodal,with prominent, decurrentpulvini, green or light brown, rarely
glaucous. Cataphylls subulate-caudate, soon reflexed, 10-15 mm long, brown with ciliate, hyaline
margins, persisting longer than the leaf fascicles.
Vegetativebuds ovoid-conical, the terminalbuds 1520 mm long, the laterals smaller, not resinous; the
scales free and spreading,subulate-linear,with ciliate
margins, light brown. Fascicle sheaths 15-25(-30)
cm long, persistent,consisting of 8-10 initially distinct, imbricate, orange-brown scales with ciliatewhite margins, later (lustrous) grey-brown to grey.
Leaves in fascicles of 5, rarely 4 or 6, in dense
tufts, persisting 2-2.5 years, slender, lax, drooping,
sometimes pendulous,20-35 cm X 0.6-1(-1.1) mm,
with serrulatemargins,acute, (yellowish) light green
to glaucous-green.Stomata on all faces of leaves, in
2-4 lines on the abaxial face and in 2-3 lines on
each of the two adaxial faces. Leaf anatomy: Cross
section triangular;hypodermismulti-layered,thicker
in the marginal areas, often with some intrusions
into the mesophyll, especially on the abaxial side of
the leaf where they may contactwith the endodermis;
resin ducts (2-)3(-4), medial;stele terete;outerwalls
of endodermal cells thickened; vascular bundles 2,
with the two xylem strands often connate. Pollen
cones crowded nearthe proximalend of new leading
shoots, sometimes also on lateral shoots, densely
clustered, subtended by subulate-lanceolate,brown
bracts, cylindrical, 30-40 x 5-8 mm when fullgrown, light pinkish brown, turning darker. Seed
cones subterminal, solitary or in pairs on distinct,
stout, curved peduncles which fall with the cones.
Immaturecones ovoid or elliptic, up to 15 x 10 mm,
with small, spreadingspines, lustrouslight brown or
glaucous. maturing in two seasons. Mature cones
narrowlyovoid to ovoid-attenuatewhen closed, more
or less ovoid, slightly curved, with an obliquely
flattenedbase when opened, (4-)5-10(-12) X (3-)
4-8 cm when open. Seed scales (100-)120-160,
readily parting to release the seeds, thin woody,
flexible, oblong, straight, spreading 90? or often
(strongly) reflexed, mostly symmetrical around the
cone, dark (purplish) brown, with a distinct, light
brown band below the apophysis abaxially and two
light brown marks of seed wings on the adaxial
side. Apophysis flattenedor slightly raised and then
transversely keeled, 8-15 mm wide, irregularly
rhombic to pentagonal in outline, variably light
brown. Umbo dorsal, raised, curved, transverserhombic in outline, 4-8 mm wide, up to 4 mm long,
Flora Neotropica
darker than the apophysis, weathering grey. Seeds
obliquely ovoid, slightly flattened,4-6 x 3-4 mm,
ochraceous or dark brown, with or without dark
spots. Seed wings articulate, effective, held to the
seed by two oblique claws, oblong, with a straight
side, widest near the middle, 13-22 x 4-8 mm,
yellowish brown, translucent.Numberof cotyledons
not observed.
Distribution and ecology (Map 15). Mexico:
Mainly in the southern half, in Sinaloa, Jalisco,
Michoacan, M6xico, Hidalgo, Tlaxcala, Puebla, Veracruz, Guerrero,Oaxaca, and Chiapas; and in the
mountainousparts of Guatemala,Honduras,El Salvador,and NW Nicaragua.The occurrencein Sinaloa
appears to be a clear disjunction (see also Perry,
1991). Pinus maximinoi is a species with a wide
ecological amplitude,occurringfrom wet subtropical
forest, where it is a gap pioneer, well up into the
cooler cloud forestson high mountainsin Mesoamerica. In Mexico it also occupies drier sites as a
constituent of pine or oak-pine forest or woodland.
Its altitudinal range is great: (450-)600-2800 m,
with an optimum at 900-1800 m. In the northwest
of its range the species occurs at 1500-2800 m. It
occurs on a variety of soils under various climatic
conditions; in Mesoamerica annual precipitation is
in the range of ca. 900-2500 mm, with the wettest
conditions on the Atlantic and Pacific slopes of the
mountains. Under these conditions it occurs frequently with Pinus tecunumanii and Liquidambar
styraciflua, at lower altitudes with P oocarpa. In
secondary broad-leavedforest other pines may join

in: P devoniana, P pseudostrobus, and pine wood-
land may prevail undera regime of grazing or burning, with the undergrowthdominated by grasses or
the fern Pteridium aquilinur.
In central Mexico, it
grows at the higher sites with Abies religiosa, P

ayacahuite, P patula, P pseudostrobus,P douglasiana, and often Quercus in mixed pine or oak-pine
forest, where precipitationis more moderatebut the
seasonal temperaturerange greater,with some frosts
occurring in winter. Phenology: The time of pollen
dispersal of this species is found to be FebruaryMarchin Mesoamerica;data for centralMexico were
not available.
Representative specimens examined. MEXICO. CHIAPAS:At Toliman, 38 km W of Huixtla, 1972, Breedlove &
Thorne30968 (MEXU, MO); above Colonia Chiapasalong
rd. to Leyra,II Mar 1993,Higman,Padilla& Styles 13

(EAP,FHO,HEH,MO);Soyalo, 13 Mar 1993,Higman.
Padilla & Styles 22 (EAP, FHO, HEH, MO); San Luis, near
Siltepec, 23 Jan 1945, Matuda5317 (NY); La Grandeza,19
Rinc6n Viejo,
May 1945, Matuda5583 (NY). GUERRERO:
CerroCabezade Toro,10 km W of Aguade Obispo,26
.-i.
LP L CY
147
? I 1 _ ? 1
A
? 1 L
~ ?~
l I~a
r .
-.
r~~~~~~~~~~~~~~~~~~~~~~~~~~
a
Solid
-
~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~~~
*
Map
15 Pinus mawninn
IM
**0t4~~~~~~~~~4
Feb 1969, Kruse 2406 (MEXU); PuertoSoleares, 6 km SE

of El Carrizal. 3 Mar 1984, Lorea 2973 (FHO, MEXU).
Sierra Manantlan,16 Apr 1987, Hughes & Styles
JALISCO:
140 (FHO, K); Mascota, 14 Aug 1976. Lamas & Torres60
(UC): along trail from San Sebastianto Arroyo Seco, 8 Jan
1927, Mexia 1421 (BM, F, MICH, MO, UC). MBxico:
Temascaltepec.4 May 1934, Hinton 5775 (NY); Valle de
Mexico, at Tenango, 10 Dec 1950. Matuda 20873 (NY);
Nevado de Toluca, 15 May 1954, Matuda et al. 31020
(MEXU). MICHOACAN:
Campanario,near Morelia. 4 Jul
1911,Arsene 6041 (P, US); CerroSan Miguel, 21 Apr 1979,
MadrigalS. 3283 (ENCB, INIF,MEXU, MS); Uruapan,25
Feb 1980, Stead & Styles 533 (FHO, K); Las Playitas,
forest sta. W of Tumbiscatio, 27 Feb 1974, Styles 25
(FHO); 8 km from Uruapan,6 Mar 1974, Styles 49 (FHO).
OAXACA:
San Juan Lachao. 22 Mar 1993, Higman. Padilla & Stiles 52 (EAP, FHO, HEH, MO); Las Galeras, ca.
100 km NE of Oaxaca, 14 Oct 1960, Little 17953 (IFGP,
US); San Gabriel Mixtepec. I Feb 1980, Stead & Styles
421 (FHO, K). SINALOA:
San Ignacio, Jan 1940, M. Martinez 2392 (US); PuertoBlanco, Jun 1940, M. Martinez3428
Las Visas, 2 km E of rd. Xalara(MEXU). VERACRUZ:
Visas, 30 Aug 1989, Zamora C. 1002 (MEXU).
GUATEMALA. ALTA VERAPAZ:
Finca Chajsei, S of
Coban, 3 Feb 1978, Stead & Stiles 134 (FHO); between
Segoum and the mines of Caqupej, 3 Feb 1978, Stead &
El Rancho, 6 Jan
Styles 135 (FHO, K). BAJAVERAPAZ:
Between
1906, Kellerman5026 (UC). CHIMALTENANGO:
Antigua and Lake Atitlan. 18 Feb 1938, Schrenk 2 (MO,
as determined by A. Farjon). EL PROGRESO: W end of

Sierra de las Minas, above Morazan, 16 May 1979. Stead
332 (FHO). GUATEMALA:Chinautla."in montibuspraeruptisque 'Canales' dictis. necnon ad pagum Chinanta [Chinautla] prope Guatemala. et summis jugo Choacas prope
Salama," 1842, Harhtweg 92 (NY); Chinautla.3 Jun 1977,
Mittak 8488 (FHO, M). SACATEPEQUEZ:
Santa Maria
Cauque, along rd. from Guatemala to Chimaltenango, 7
Feb 1978, Stead & Stiles 141 (FHO. MEXU). SOLOLA:
Between San Pedro and Santiago Atitlan. 4 Apr 1993.
Higman, Padilla & Styles 65 (EAP. FHO. HEH. MO).
SUCHITEPEQUEZ:Finca Moca, 30 Oct 1934, Skutch 1566
(F, NY, US).
HONDURAS. COMAYAGUA:
Plan Cerro Maneadero.
10.5 km E of Lago Yojoa, 14 Mar 1993, Hawkins 631
(EAP, FHO, HEH, MO, TEFH); El Achote. 18 Feb 1928,
Standlev 56144 (A, F, US). COPAN:Cerro Azul National
Park, 12 km NW of Florida, 9 Feb 1992, D. Mejia 40
Cusuco National
(EAP, FHO, HEH, MO, TEFH). CORTES:
Park, near visitors ctr., 18 km W of San Pedro Sula, 21
Mar 1993, D. Mejia 359 (EAP, FHO. MO. TEFH); Cerro
San Ildefonso, between Buenos Aires and Banaderos, 27
Mar 1963, Molina R. 11600 (EAP, F, NY. US); 14 km from
Cofradiaalong rd. to Buenos Aires, 14 Mar 1978. Stecad&
MORAZAN: Between El
Styles 254 (FHO). FRANCISCO
Hatillo and Lo de Ponce, rd. into Montana La Tigra. 24
Nov 1958, Molina R. 8685 (A, EAP, F, US). INTIBUCA:
El
Achote, above the plains near Siguatepeque, 28 Jul 1936.
Yunckeret al. 6205 (A, F, K, MICH, MO. NY, S). LA PAZ:
148
Alongrd.to Marcala,13Apr1994,Farjon& Mejia297 (E,
La Planta, between
FHO, HEH, K, MEXU, U). LEMPIRA:
electricpowerplantand Don Tomascampsite,19 Nov

1991,Hawkins98 (EAP,FHO,MO,TEFH);10 km along
rd. fromErandique
SW to Candelaria.
2 km NE of Cerro
Congolon,9 Mar1991,Hughes1428(FHO);Mt.Celaque
NationalPark,8 km SW of Gracias,19 May 1992, D.
El File,
Mejia 113 (EAP, FHO, MO, TEFH). OLANCHO:
2 Jun1992,HawIkins461 (EAP,
13kmNNEof Catacamas,
FHO,MO,TEFH);MontafaCoyolito.15 km NE of San
Esteban.14 Sep 1983,McCarter146(FHO).
EL SALVADOR. SANTAANA:Cerro Montecristo, 16
km N of Metapan,23 Feb 1989,Hughes1264 (FHO,K,

LAGU).
San Rafael del Norte, 2 Mar
NICARAGUA. JINOTEGA:
1977,Chaplinc312 (FHO).19 Jan 1970,Kemp12 (FHO).

Uses. Pinus ilaximinoi is an importanttimbertree
in most of its range, where it is exploited with other
tall-growing pines. It has also been the subject of
experimentalforestryin varioussubtropicaland tropical countries underprogramssuch as those initiated
by the Central America & Mexico Coniferous ResourcesCooperative(CAMCORE).The wood is relatively soft and light and easily worked.
Bentham (1842), when describing various plants
collected by C. T. Hartweg, named and described
Hartieg 620 from Guatemala as Pinus tenuifolia.
which is a later homonym of P tenuifolia R. A.
Salisbury(1796), a synonym of P strobusLinnaeus.
Moore (1966) correctedthis nomenclaturaloversight,
but in the meantime P. tenuifolia Benthamhad been
reduced to a variety of P pseutdostrobusby Shaw
(1909). Later,Shaw (1914) synonymized it with the
latter species, but Martinez(1945, 1948) restoredit
as a species, using the illegitimate name.
Hartwegcollected this pine nearthe city of Guatemala in the "CanalesMountains."The type collection
(K) consists of two sheets, one labeled "620 Guatemala Hartweg,"the other "620 Pinus sp." and with
two locations given. It is thus not certain that both
sheets representthe same gathering,despite the same
number.The firstsheet, from W. D. Hooker's herbarium, bears the inscription"Pinus tenuifolia Benth.";
the second, from G. Bentham'sherbarium,lacks this
name. Both are sterile shoots; separate cones have
not been found. The specimen from the Bentham
herbariumis considered the holotype; the one from
Hooker's herbariumis possibly an isotype. On the
reverse of the BM sheet, three localities are mentioned, with the "CanalesMts." as the first.Annotations like this seem to serve as much to describe
where the taxon occurred as to indicate the locality
where it was collected.
Roezl (1857) describeda few "new species" from
Mt. Tzompoli near Mexico City, of which original
Flora Neotropica
material could be traced and is here designated as
lectotypes of these obsolete names. (For a discussion
of Roezl's catalogues, names, and collections, see
under P. montezumae.)
As has been discussed under P. douglasiana, the
distinctionbetween P maximinoiand P douglasiana
has proved to be difficult, leading to misidentifications in the field, in herbaria,and in the literature.
Stead and Styles (1984) concluded that P maximinoi
is essentially a Mesoamerican species that extends
into centralMexico as far as Michoacanand Jalisco,
but is there restricted to higher altitudes and the
Pacific side of the mountains. Gross morphology
shows differences in leaf width, with very thin leaves
in P mtaximinoi(not exceeding I mm), with (as
a result) fewer stomatal lines. There is, however,
considerable overlap in the length of the fascicle
sheaths, which contradictsthe figures given by Stead
and Styles (1984) and Carvajaland McVaugh(1992),
the latterauthorsonly citing specimens from western
Mexico. The ovuliferous cones are usually smaller
in this species and have thin woody scales with
flattened apophyses. Such differences are quantitative and, to some extent, even continuous; often a
certaindeterminationcan be made only by considering a combinationof characters.
19. Pinus lumholtzii B. L. Robinson & Fernald,
Proc. Amer.Acad. Arts 30: 122. 1894. Type. Mexico. Chihuahua:Near Coloradas, May 1893, Hartman 541 (holotype, GH; isotypes, A, K, US).
Fig. 36
Pinutspatltul B. C. Seemann,Bot.Voy.Herald336. 1856;
non Schlechtendal
& Chamisso.1831.Type.Mexico.
"BetweenMesquitaland SantaTeDurango/Nayarit:
1961(holotype,K;isotype,BM).
resa,"1850,Seetmaunn
Tree, medium size, height to 20 m, dbh to 50-70

cm. Trunkmonopodial,straight,terete, usually erect.
Bark thick, scaly, very rough,divided into elongated,
irregularplates and deep, wide, longitudinalfissures,
greyish brown to dark grey, on young trees and
branches scaly, flaking very soon, reddish brown,
turning grey. Branches of first order long, slender,
spreading or ascending, persistent; branches of
higherordersslender,flexible, drooping,the ultimate
branches ascending. The crown is broad, rounded
and usually open. Shoots ridged between decurrent
pulvini, initially glaucous, turning reddish brown,
then grey. Cataplhlls subulate, up to 10 mm long,
scarious, with ciliate-hyaline margins, red-brown,
early deciduous. Vegetative buds ovoid-conical,
acute, the terminalbuds ca. 15 x 8 mm, the laterals
149
Al
o,
AA
~,vty ~
~ ...
FIG. 36. Pinus lumholtzii(A, C. Wilkinson& St$les 3; B. D, E, G. H. Stead & Styles 474; F, Hughes 973). A. Shoot
with bud and leaf fascicles. B. Shoot with leaf fascicles and disintegratingfascicle sheaths. C. Fascicle sheaths. D. Cross
section of leaf. E, F. Ovuliferous cones. G. Seed scale (two views). H. Seeds with articulatewings. (Magnifications:A.
B. E, F. x0.5; C. G, H, x I; D, x 30.)
150
smaller, resinous; the scales imbricate,more or less
free at base, narrowlytriangular,acute, with ciliate
margins, reddish brown. Fascicle sheaths (20-)2535 mm long on juvenile fascicles, consisting of ca.
10 initially imbricate,ligulate-subulate,thin, reddish
brown scales, soon coming apart, forming a tuft of
curling scales at the base of the fascicles before they
fall at maturity,leaving the fascicles bare. Leaves in
fascicles of 3 (exceptionally 2 or 4), remote, usually
extremely pendulous, persisting two years, lax but
not thin, (15-)20-30(-40+) cm X (1-)1.2-1.5 mm,
with serrulatemargins,acute, light green. Stomataon
all faces of leaves, in 8-14 lines on the convex
abaxial face, in 4-5 lines on each adaxial face,
conspicuous. Leaf anatomy: Cross section transverse-triangular,with convex abaxial side; hypodermis multi-layered,especially in the marginalregions,
with minor intrusionsinto the mesophyll;resin ducts
4-10, medial, sometimes 1-4 internal; stele wide,
oval in cross section; outer cells of endodermis not
thickened; vascular bundles 2, distinct, the xylem
strandsoften connate. Pollen cones concentratednear
the proximal end of a new shoot but sparsely clustered, subtended by imbricate, thin, reddish brown
bracts,cylindrical, 20-30 X 5 mm when mature,at
first pinkish, turningyellow. Seed cones subterminal
or lateral,solitary (sometimes in whorls of 2, rarely
3) on 10-15 mm long, curved peduncles, breaking
off easily and remaining with the soon deciduous
cones. Immaturecones broadlyovoid to subglobose,
with antrorselycurved spines, maturingin two seasons. Mature cones ovoid to ovoid-attenuatewhen
closed, ovoid to ovoid-acute when opened, then (3-)
3.5-5.5(-7) X (2.5-)3-4.5 cm. Seed scales ca. 7090, parting gradually,spreading wide or the lower
scales remainingclosed, thick woody, oblong, widest
near the apex up to 10-12 mm, the smaller basal
scales remaining connate, dark red-brown on the
abaxial side, dark brown with light brown marks
of seed wings adaxially. Apophysis slightly raised,
thickened along the distal margin, obscurely transversely keeled, those on the basal scales gibbous,
rhombicto pentagonalin outline, ochraceousto reddish brown. Umbo dorsal, often tilted toward the
distal marginof the apophysis, flat or slightly raised,
up to 5 mm wide, with a minute, soon deciduous
prickle,grey-brown,often darkerthanthe apophysis.
Seeds obliquely obovoid, slightly flattened,3-5 mm
long, dark brown, often with black dots. Seed twings
articulate,effective, held to the seed by two oblique
claws, oblique, widest below the apex, (8-)10-14 X
4-6 mm, light yellowish or greyish brown. Number
of cotyledons not observed.
Flora Neotropica
the Sierra MadreOccidental,in Chihuahua,Sinaloa,
Durango,Nayarit,Jalisco, Zacatecas,Aguascalientes,
and Guanajuato.The altitudinalrangeof this species
is (1500-)1700-2600(-2900) m, which corresponds
to the lower and middle slopes of the Sierra Madre.
It grows usually mixed with several species of Quercus and other pines in pine-oak forest, or on the
wetter western slopes of the SierraMadre, in mixed
pine forest. Associated pines are, e.g., P leiophylla,
P arizonica, P douglasiana, P teocote, and P oocarpa: on more mesic sites one can expect Pinus
avacalhuite and Pseudotsuga menziesii, while in dry
habitatsP. cembroidescan occur with it. The annual

precipitation,mostly as summer rains, is a moderate
500-600 mm except on the driest and wettest sites.
Phenology:Time of pollen dispersal not observed.
Representativespecimensexamined.MEXICO.CHISep 1936,Gentry
7 Jul 1936.
2873 (A. F, K, MO,S, UC, US);Basaseachic,
LeSueur557 (ENCB,F,MO,UC,US);PuebloNuevo,Mar
HUAHUA:SierraCanelo,Rio Mayo,24
1895, Lumholtzs.n. (A); Mt. Mohinora, 1 Sep 1898. Nelson
4914 (F). DURANGO:

LasVeredas,"km20.8 LasVeredas,"
2 Oct 1960, Courbasson & Mahieux s.n. (P); Miradorlos
Balcones,S of Durangoalongmainrd.,2 Apr1991,Styles,

Favela et al. 4 (FHO). JALISCO:Chiquilistlan. 28 May
1892,M. E. Jones492 (US);Sierrade la Campafa.along
rd. to Mascota,ca. 12 km NW of Los Volcanes.23 Oct
1952,McVaugh 13733(BM);25 km E of Arandas,14/15
Nov 1970,McVaugh
24330(MICH);CerroEl Picacho,E
of Tenamaxtlan, 25 Dec 1985, Perez dle la Rosa 1024
(MEXU);12 km S of Etzatlan,on mtn.trailto Ameca,23

Oct 1904, Pringle 10014 (A, E, ENCB.GOET,K. M.
MEXU,MICH,);9 km fromTapalpatowardChiquilistlan,
22 Sep 1986, Wilkinson & Styles 3 (FHO). NAYARIT:
MesetaJuanacata
off rd.fromIxtlanto Tepic,14Feb 1980,
Stead & Styles 474 (ENCB, FHO, MO). SINALOA:
Rosario,
just W of El Palmita, 31 Jan 1962, Breedlove 1693 (DS);
5 km E of Los Loberos.along rd. from Mazatlinto

Durango.30 Apr 1987, Hughes973 (FHO,MEXU);El
Palmito, 28 Dec 1983, Sanders et al. 4449 (UC).
Uses. Pinus lumholtziiis generally known as "pino

triste"for its striking pendulous foliage. It has been
overexploited with other pines in some areas for
timber,but due to its scatteredoccurrence,especially
in pine-oak forest and in drier sites, it is not a
commercially importanttree.
Shaw (1909) cited Pinus patula sensu Seemann
(1856), non Schlechtendal& Chamisso (1831) as a
synonym of P lumholtzii. Seemann 1961 (BM, K)
indeed belongs to this species, and Seemann's interpretationof P patula is incorrect.
Robinson and Fernald (1894), in an account of
new plantscollected on an expedition led by archaeologist Carl Lumholtz,cite Hartman541 as the only
collection underthis new species and apparentlydid
151
-
1.,
~~~~~~~~~~~
I~~ ~ ~
a
''''
..
":
i~~~~~a
?I
'~
?CI.~~~~~~~~~~~I
r?'
Ma6 iuiIn/ot,
(crls
not consult other herbariato find if this strikingly

different pine had not been collected earlier.
Carvajal and McVaugh (1992) cite a collection
(McVaugh24330, MICH) with non-pendulous leaf
fascicles; a similarspecimen is representedby Styles,
Favela et al. 4 (FHO). Both have ratherrigid, short,
spreading leaves 14-15 cm long but otherwise are
entirelysimilar to typical P lumholtzii.These collections were made in Jalisco and Durango,respectively,
and, being apparently very rare, represent single
"aberrant"individuals within populations of "normal" P lumholtzii.
20. Pinus oocarpa Schiede ex Schlechtendal, Linnaea 12:491. 1838.
Tree, medium to tall, height to 10-15 or 30-35 m
(dependenton variety and site condition), dbh to 40-
at/
arpn/
tinls
60 or 100-125 cm. Trunkmonopodial,erect, straight

or tortuous,terete, often clear of branches for ?Vof
tree height. Bark thick, rough, scaly, breaking into
small or large, longitudinal plates and shallow fissures, darkbrown to grey-brown,on young trees and
branches thin, scaly, reddish brown. Branches long,
often tortuous,spreading,forminga roundedor irregular,open crown. Shoots roughand scaly, with prominent, decurrentpulvini, reddish brown. Cataphylls
10-15 mm long, subulate, recurved, scarious, with
erose-ciliate margins, brown, weathering blackish
grey. Vegetativebuds ovoid-oblong to fusiform, the
terminal buds 15-25 mm long, the laterals ovoidacute, smaller, not resinous; the scales imbricate,
with free, recurved apices, subulate, ciliate at margins, brown. Fascicle sheaths up to 25 mm long,
persistent,not reducedin length, with imbricate,red-
152
brown scales, weatheringto nearly black. Leaves in
fascicles of 3-5, in rigid tufts on thick branchlets,
persisting2-3 years, straight,rigid or less commonly
lax, (I l-)14-25(-30) cm x 0.8-1.6 mm, with serrulate margins, acute-pungent, lustrous (yellowish)
green. Stomata on all faces of leaves, in 6-8(-10)
lines on the abaxial face and in 3-5 lines on each
of the adaxial faces. Leaf anatomy: Cross section
with a convex abaxial side;
(transverse-)triangular,
hypodermis irregularly developed, with intrusions
into the mesophyll often connecting resin ducts to
the endodermis; resin ducts 4-8, usually septal,
sometimes medial or external;stele wide, triangular
in cross section; endodermis with uniform, thinwalled cells; vascular bundles 2, approximate, or
with the xylem strands connate. Pollen cones
crowded near the proximal end of a new shoot
but loosely arranged,subtendedby scarious bracts,
oblong-cylindrical, 15-20 x 5-6 mm, pink or reddish, turningyellowish brown. Seed cones subterminal, at the base of a subsequentshoot, solitary or in
whorls of 2-4 on stout, up to 35 mm long, recurved,
bracteate peduncles which eventually fall with the
cones which are persistent for several years after
seed dispersal. Immaturecones ovoid to subglobose,
often wider than long, purplish, then light brown,
maturingin two seasons. Maturecones broadlyovoid
to subglobose, semi-serotinous, when opened often
wider than long, with a flattenedbase, 3-8(-10) x
3-9(-12) cm when open. Seed scales ca. 70-130,
parting slowly except the proximal scales (semiserotinous), thick woody, symmetrical, oblong,
straight or slightly recurved, dark purplish brown,
with light brown marksof seed wings on the adaxial
face. Apoplhysisnearlyflat or slightly raised, in some
cones pyramidal, (weakly) transversely keeled,
rhombic to pentagonalin outline, the apical margin
entire or undulating, lustrous, ochraceous to light
brown, weatheringgrey. Umbo dorsal, flat or raised,
sometimes curved, obtuse or rarely with a minute
prickle, dark grey-brown. Seeds obliquely ovoid,
slightly flattened,4-8 X 3-4.5 mm, blackish grey,
often with black spots. Seed wings articulate,effective, held to the seed by two thin claws, 8-18 x 48 mm, greyish brown. Seedlings: Numberof cotyledons not observed. Juvenile leaves persist 3-4 years
and are ca. 30 x 0.5 mm; seedlings 2-4 years of
age may sprout from the root after fire.
Distribution (Maps 17, 18). Mexico: From the
Sierra Madre Occidental SE to Mesoamerica, in S
Sonora, Sinaloa, SW Durango,Nayarit,S Zacatecas,
Jalisco, Michoacan, Mexico, Distrito Federal, Hidalgo, N Puebla, Morelos, Tlaxcala, Guerrero,Oa-
Flora Neotropica
xaca, S Veracruz,and Chiapas; widespread in the
highlands of Guatemala,in Honduras,El Salvador,
and NW Nicaragua. Reports (Critchfield & Little,
1966; Perry, 1991) of P oocarpa from Belize are
here considered to refer to P tecunumanii.
Key to the varieties of Pinus oocarpa
i. Leavesin fasciclesof 5 (sometimesalso 3-4 on
treeswithmostly5), (17-)20-25(-30)cm X 0.81.4 mm;seedcones5-8(-10) x 5-9(-12) cm.....
........................................................
20a var. oocarpa
1. Leaves in fascicles of 3 (rarely 4), (11-)14-17
(-20) cm x 1.2-1.6 mm; seed cones 3-5.5 x 35(-6) cm...............................................20b var. trifolittal
20a. Pinus oocarpa Schiede ex Schlechtendal var.

oocarpa. Type. Mexico. Michoacan: "Zwischen
Ario und dem Feuerberge Jorullo." Schiede s.n.
(no original material has been found in HAL or
elsewhere); neotype: Michoacan:Las Playitas Forest Sta., along track to Cahas, Styles 36 (neotype,
FHO, here designated;isoneotype, K).
Fig. 37
Pinus oocarpa Schiede ex Schlechtendalvar.oocarpoides
Endlicher,Syn. Conif. 152. 1847. Type. Mexico. [Hartweg s.n.] ex herb.LindleY3150 (CGE, labeled as "Pinus
oocarpol)ites.H. L.," is possibly original material).
Pilnusoocarpa Schiede ex Schlechtendal var. mlcflantzoi
Martinez, Anales Inst. Biol. Univ. Nac. Mexico 11:
70. 1940. Type. Mexico. Hidalgo: San Jose Ocotillas
("habitatin Ocotillos, prope Huasca"), Mar 1939, M.
Martfinez3460 (holotype, MEXU-2 sheets).

100-125 cm. Leaves in fascicles of 5 (sometimes
also 3-4 on trees with mostly 5), straight, rigid or
less commonly lax, (17-)20-25(-30) cm x 0.8-1.4
mm. Seed cones 5-8(-10) x 5-9(-12) cm when
open. Seed scales ca. 100-130.
Distribution and ecology. Distribution as given
for the species. It has the largest range of all of the
pines in our region and is especially abundant in
Mesoamerica.It extends over a NW-SE distance of
ca. 3000 km and consequently is found under very
different ecological conditions. This is expressed in
its altitudinalrange, (200-)500-2300(-2700) m, and
in the variation of annual precipitation. 700-3000
mm. These variations are greatest in Mesoamerica
and are less pronouncedparticularlyin the far NW
part of the range. Especially precipitationlevels increase toward the southeast. Yet its optimum is at
middle elevation (1000-1500 m) with abundantrainfall. Seasonality is mainly expressed in a (long) dry
season from October to June in much of its range.
Nearly everywhere, fire is an integral part of the
ecosystem, but man-madefires, often deliberate,add

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Organization for Flora Neotropica

The paper used in this publicationmeets the requirementsof

Libraryof Congress [8508]

Organizationfor Flora Neotropica

The New YorkBotanicalGarden

Board Representativeto UNESCO

Claudio Delgadillo M. (1996-2001)

ORGANIZATION FOR FLORA NEOTROPICA

Founded and Conductedunderthe Auspices of UNESCO

The Scientific PublicationsProgramat The New York Botanical Garden

With Accounts of Wood Anatomy by IAN D. GOURLAY,

has been so intensively studied as the genus Pinus.

herbariain the United States, and those of Seemann

Early in this period, the curious efforts by B.

3. Collectors in the Caribbean, 1851-1914, with

We characterizethe time after the FirstWorldWar

cies into Baja California(P attenuata,P coulteri, P

study using morphologicalcharactersof the pinyon

MORPHOLOGY AND ANATOMY

ally, resulting in a hierarchical,symmetricalsystem

ated in the winterbuds (see below), and development

very low spiral), but they are cyclic (whorled) in

FIG. 1. Morphologyof foliage in Pinus. A, Ha. P. montezumae.B. P. lambertiana.Ga. P durangensis.Gb. P. caribaea

.........,, .-,,~ ....~.~~~~~~~~~~~~~~~~~~

FIG;. 3. Scanning electronmicrographs

taxa of subgen. Strobus and do not occur in other

in cross section under magnification of thin slices

which produces the phellem (-- periderm) outward

and a thin layer of phelloderm inward. Commonly

... ..... ....

~~crrtunucr~cr.M,I! strr,hif~rrllis seeds

wings; a = seed scale with seed cavity tone seed

a = abaxial view, b = adaxial view, I = apophysis. 2 = terminal

Morphology and Anatomy

surface is often irregularor longitudinally rugose.

posed to sun and rain are most developed (see

Comparedto other genera in the family, the seeds

folia, and to a lesser extent P nelsonii and P

seed scales. Upon dispersal they are normally still

WOOD ANATOMY (by Ian D. Gourlay)

taxodioid pits. Some genera contain xylem

and colour, and show a more gradualtransitionfrom

of the "hard"pines (Diplox'i-loit subgen. Piiiiis). A. Transverse

its. Magnficatons x100.

ray tracheids, e.g., P ponderosa, P taeda, P

CENTRAL AMERICAN PINES

Khoshoo, 1956; Saylor, 1972, 1983; Mehra, 1988).

the monotypic genus Pseudolarix (n = 22; 2i =

Morphologicaldifferences between Pinus leiophylla and P lumiholtzii,two species forming subsect.

be a geographical dine in several morphological

is a serious omission. Unfortunatelyalso, only three

cembroides, P edullis, and

P monophylla-thereby leaving out all the more

POLLENMORPHOLOGY(by Marie H. Kurmann)

to date. For each of the species, she listed the size

... : :`< .: ::::: :.:: .: : :...

which combines quantitative and morphological

considerationof preparationtechniquesand a combination of structuraland quantitativecharactersmay

THE ROLEOF MONOTERPENESIN THE TAXONOMYOF THE MEXICAN

The Role of Monoterpenesin the Taxonomyof the Mexican and MesoamericanPines

THE USE OF MONOTERPENE DATA

sition of Piins monophylla and P quadrifolia (An-

derson et al., 1970) found that B/-phellandrene,

raised to species rankfrom that of variety within the