ANNALS OF THE
MISSOURI BOTANICAL GARDEN
VOLUME 84
1997
Colophon
This volume of the ANNALS of the Missouri Botanical Garden has been set in APS Bodoni. The text
is set in 9 point type while the figure legends and literature cited sections are set in 8 point type.
This volume has been printed on 70# Vintage Gloss. This is an acid-free paper Mee to have a
shelf-life of over 100 years. Vintage Gloss is manufactured by the Potlatch Paper Compan
pane used in the ANNALS are reproduced using 300 line screen halftones. The binding used
n the production of the ANNALS is a proprietary method known as Permanent Binding.
The ANNALS is printed and distributed by Allen Press, Inc. of Lawrence, Kansas 66044, U.S.A.
© Missouri Botanical Garden 1997
ISSN 0026-6493
VOLUME 84
ALMEDA, FRANK. Chromosomal Observations on the Alzateaceae (Myrtales)
BACIGALUPO, NÉLIDA M. (See Elsa L. Cabral y Nélida M. Bacigalupo) _
BENÍTEZ DE ROJAS, CARMEN & WILLIAM С. D'ARCY. The Genus Ly-
cianthes (Solanaceae) in Venezuela
BERRY, PAUL E. Book Review. Guide to the Vascular Plants of Central French
Guiana. Part 1 by Scott Mori et al.
BLACKMORE, STEPHEN (See Sandra Knapp, Viveca Persson & Stephen
Blackmore
BROWN, K. D. (See J. F. Smith, J. C. Wolfram, K. D. Brown, C. L. Carroll &
. S. Denton)
CABRAL, ELSA L. Y NÉLIDA M BACIGALUPO. Revisión del Género Gal-
ianthe subg. Ebelia stat. nov. (Rubiaceae: Spermacoceae) __________
CARR, GERALD D. (See Harold Robinson, Gerald D. Carr, Robert M. King
& A. Michael Powell)
CARROLL, C. L. (See J. F. Smith, J. C. Wolfram, K. D. Brown, C. L. Carroll
&
enton)
CHASE, N MARK W. (See Douglas E. Soltis, Pamela S. Soltis, Daniel L. Nick-
rent, Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A.
Sweere, Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M.
Swensen, Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. Gillespie, W.
John Kress & Kenneth J. Sytsma)
CHAW SHU-MIAW (See Douglas E. Soltis, Pamela S. Soltis, Daniel L. Nick-
rent, Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A.
Sweere, Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M.
Swensen, Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. Gillespie, W.
John Kress & Kenneth J. Sytsma)
CROAT, THOMAS B. A Revision of Philodendron Puen? Philodendron (Ат-
aceae) for Mexico and Central America
DENTON, D. S. (See J. F. Smith, J. C. Wolfram, K. D. Brown, C. L. Carroll &
D. S. Denton)
D'ARCY, WILLIAM G. (See Carmen Benítez de Rojas & William G. D'Arcy)
D'ARCY, WILLIAM С. A Review of the Genus Eccremocarpus (Bignoniaceae)
FRITSCH, PETER W. A Revision of a (Styracaceae) for Western Texas,
Mexico, and Mesoamerica _______--
GILLESPIE, LYNN J. (See Douglas E. Soltis, Pamela S. Soltis, Daniel L. Nick-
rent, Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A.
Sweere, Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M.
Swensen, Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. Gillespie, W.
John Kress € Kenneth J. Sytsma) __
1997
305
857
167
907
67
50
857
103
705
GOLDBLATT, PETER € MASAHIRO TAKEI. Chromosome Cytology of Iri-
daceae—Patterns of Variation, Determination of Ancestral Base Num-
bers, and Modes of Karyotype Change __
GOLDBLATT, PETER & ANNICK LE THOMAS. Palynology, Phylogenetic Re-
construction, and Classification of the Afro-Madagascan Genus Aristea
(Iridaceae)
GU HONG-YA & PETER C. HOCH. Systematics of Kalimeris (Asteraceae:
Astereae)
HAHN, WILLIAM J. (See Douglas E. Soltis, Pamela S. Soltis, Daniel L. Nick-
rent, Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A.
Sweere, Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M.
Swensen, Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. Gillespie, W.
John Kress & Kenneth J. Sytsma)
HAUK, WARREN D. A Review of the Genus Cydista (Bignoniaceae) _____-
HILU, KHIDIR, W. & JOHN L. JOHNSON. Systematics of Eleusine Gaertn.
(Poaceae: Chloridoideae): Chloroplast DNA and Total Evidence
HOCH, PETER C. (See Gu Hong-ya & Peter C. Hoch)
HOOT, SARA B. (See Douglas E. Soltis, Pamela S. Soltis, Daniel L. Nickrent,
Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A. Sweere,
Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M. Swensen,
Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. Gillespie, W. John Kress
& Kenneth J. Sytsma)
JOHNSON, JOHN L. (See Khidir W. Hilu, & John L. Johnson)
JOHNSON, LEIGH A. (See Douglas E. Soltis, Pamela S. Soltis, Daniel L.
Nickrent, Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A.
Sweere, Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M.
Swensen, Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. Gillespie, W.
John Kress & Kenneth J. Sytsma) __
KING, ROBERT M. (See Harold Robinson, Gerald D. Carr, Robert M. King
& A. Michael Powell)
KNAPP, SANDRA, VIVECA PERSSON & STEPHEN BLACKMORE. A Phy-
logenetic Conspectus of the Tribe Juanulloeae (Solanaceae)
KRESS, W. JOHN (See Douglas E. Soltis, Pamela S. Soltis, Daniel L. Nickrent,
Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A. Sweere,
Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M. Swensen,
Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. Gillespie, W. John Kress
& Kenneth J. Sytsma)
KRON, KATHLEEN A. (See Douglas E. Soltis, Pamela S. Soltis, Daniel L.
Nickrent, Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A.
Sweere, Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M.
Swensen, Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. Gillespie, W.
John Kress & Kenneth J. Sytsma)
285
263
762
841
762
841
893
67
KUZOFF, ROBERT K. (See Douglas Е. Soltis, Pamela 5. Soltis, Daniel L.
Nickrent, Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A.
Sweere, Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M.
Swensen, Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. we og ae W.
John Kress & Kenneth J. Sytsma) ___ n
LE THOMAS, ANNICK (See Peter Goldblatt & Annick Le Thomas) ______
LI ЛЕ Gee Ei Xi-wen & Li Fo пон оо
LI XI-WEN & LI JIE. The Tanaka-Kaiyong Line—An Important Floristic Line
for the Study of the Flora of East Asia
MÉDAIL, FRÉDÉRIC & PIERRE QUÉZEL. Hot-Spots Analysis for Conser-
vation of Plant Biodiversity in the Mediterranean Basin ____
NICKRENT, DANIEL L. (See Douglas E. Soltis, Pamela S. Soltis, D Daniel L.
Nickrent, Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A.
Sweere, Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M.
Swensen, Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. Gillespie, W.
John Kress & Kenneth J. Sytsma)
PERSSON, VIVECA (See Sandra Knapp, Viveca Persson & Stephen Black-
more) .
PIESSCHAERT, FREDERIC, ELMAR ROBBRECHT & ERIK SMETS. Dialy-
petalanthus fuscescens Kuhlm. (Dialypetalanthaceae): The Problematic
Taxonomic Position of an Amazonian Endemic
PIRE, STELLA MARIS. Género Galianthe subg. Ebelia (Rubiaceae: Sperma-
coceae): Estudio Palinológico
POWELL, A. MICHAEL (See Harold Robinson, Gerald D. Carr, Robert M.
King & A. Michael Powell) ___
QUEZEL, PIERRE (See Frédéric Médail €: Pierre Quézel) _______----
ROBBRECHT, ELMAR (See Frederic Piesschaert, Elmar Robbrecht & Erik
ets)
ROBINSON, HAROLD, GERALD D. CARR, ROBERT M. KING & A. MI-
CHAEL POWELL. Chromosome Numbers in Compositae, XVII: Sene-
cioneae III
ROHWER, JENS G. The Fruits of Jasminum pon (Oleaceae), and the Dis-
tinction Between Jasminum and Meno
SCHUTTE, ANNE LISE. A Revision of the Genus a di bete Фа). А
SMETS, ERIK (See Frederic Piesschaert, Elmar Robbrecht & Erik Smets) —
SMITH, J. Е, J. С. WOLFRAM, К. D. BROWN, С. L. CARROLL & D. S.
DENTON. Tribal Relationships in the Gesneriaceae: Evidence from DNA
Sequences of the Chloroplast Gene ndhF ____
SOLTIS, DOUGLAS E., PAMELA 5. SOLTIS, DANIEL L. NICKRENT, LEIGH
A. JOHNSON, WILLIAM J. HAHN, SARA B. HOOT, JENNIFER A.
SWEERE, ROBERT K. KUZOFF, KATHLEEN A. KRON, MARK W.
CHASE, SUSAN M. SWENSEN, ELIZABETH A. ZIMMER, SHU-MIAW
CHAW, LYNN J. GILLESPIE, W. JOHN KRESS & KENNETH J. SYTSMA.
Angiosperm Phylogeny Inferred from 18S Ribosomal DNA Sequences __
112
SOLTIS, PAMELA 5. (See Douglas E. Soltis, Pamela 5. Soltis, Daniel L. Nick-
rent, Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A.
Sweere, Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M.
Swensen, Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. Gillespie, W.
John Kress & Kenneth J. Sytsma)
SWEERE, JENNIFER A. (See Douglas E. Soltis, Pamela S. Soltis, Daniel L.
Nickrent, Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A.
Sweere, Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M.
Swensen, Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. Gillespie, W.
John Kress & Kenneth J. Sytsma)
SWENSEN, SUSAN M. (See Douglas E. Soltis, Pamela S. Soltis, Daniel L.
Nickrent, Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A.
Sweere, Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M.
Swensen, Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. Gillespie, W.
John Kress & Kenneth J. Sytsma)
SYTSMA, KENNETH J. (See Douglas E. Soltis, Pamela 5. Soltis, Daniel L.
Nickrent, Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A.
Sweere, Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M.
Swensen, Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. Gillespie, W.
John Kress & Kenneth J. Sytsma)
` TAKEL MASAHIRO (See Peter Goldblatt & Masahiro Takei)
TAYLOR, CHARLOTTE M. Conspectus of the Genus Palicourea (Rubiaceae:
Psychotrieae) with the Description of Some New Species from Ecuador
and Colombia _
THOMPSON, MAXINE M. Survey of Chromosome Numbers in Rubus (Rosa-
ceae: Rosoideae) ____
WOLFRAM, Ј. C., (See J. Е Smith, Ј. С. Wolfram, K. D. Brown, С. L. Carroll
& D. 5. Denton)
Nickrent, Leigh A. Johnson, William J. Hahn, Sara B. Hoot, Jennifer A.
Sweere, Robert K. Kuzoff, Kathleen A. Kron, Mark W. Chase, Susan M.
Swensen, Elizabeth A. Zimmer, Shu-miaw Chaw, Lynn J. Gillespie, W.
John Kress & Kenneth J. Sytsma) _
128
50
Annals
of the
Missouri
Botanical
Garden |
1997 $
Number 1
Volume 84, Number 1
Winter 1997
Annals of the
Missouri Botanical Garden
The Annals, published quarterly, contains papers, primarily in systematic botany,
contributed from the Missouri Botanical Garden, St. Louis. Papers originating out-
side the Garden will also be accepted. Authors should write the Managing Editor
for information concerning arrangements for publishing in the ANNALS. Instructions
to Authors are printed in the back of the last issue of each volume.
Editorial Committee
Henk van der Werff
Editor, .
Missouri Botanical Garden
Amy Scheuler McPherson.
Managing Editor,
Missouri Botanical Garden
Diana Gunter
Editorial Assistant,
Missouri Botanical Pads
Vicki Couture
Secretary
Teresa Johnson
Publications Order Processor
Ihsan Á. Al-Shehbaz
Missouri Botanical Garden
Gerrit Davidse
Missouri Botanical Garden
Roy E. Gereau
Missouri Botanical Garden
Peter Goldblatt
Missouri Botanical Garden
Gordon McPherson =
Missouri Botanical Garden
P. Mick Richardson
Missouri Botanical Garden
For subscription information contact Depart- _
ment Eleven, РО. Box 299, St. Louis, MO _
63166-0299. Subscription price is $110 per
volume U. S., $115 Canada and Mexico, $135
all oth Four issues per volume. The
journal Novon is included in the eubscription
price of the ANNALS.
amepher@admin.mobot.org «езщ Deis.
dept] 1@mobot.org (orders 5) >
http://www.mobot.or, тв.
© Missouri Botanical bs 1997 _
The mission of the Missouri Botanical Garden i is to discos
their environment, in order to preserve i and enrich life.
The ANNALS
OF THE MISSOURI BOTANICAL -
GARDEN (ISSN 0026-6493) i is pitied quar-
terly by the Missouri Botanical 2345
Tower Grove Avenue, St. Louis, MO 631 “fi Pe-
riodicals postage paid at St. Louis, MO and ad-
ditional mailing offices. POSTMASTER: Send ad-
dress changes to ANNALS OF THE MISSOURI
BOTANICAL GARDEN, Department Hs PO.
66-0299
Box 299, St. Louis, MO 631
d share сол ве about planis a and
M © Mae man e metet UO 19 7 Pome ae
Volume 84. Annals
Number 1 of the Y
1997 Missouri
Botanical
Garden
ANGIOSPERM PHYLOGENY Douglas Е. Soltis, Pamela S. 801115,2
Daniel L. Nickrent,? Leigh А. Johnson,?
INFERRED FROM 18S William J. Hahn," Sara B. Hoot,’
RIBOSOMAL DNA Jennifer A. Sweere,* Robert К. Kuzoff,?
SEQUENCES! Kathleen А. Kron,* Mark W. Chase,”
Susan M. Swensen,? Elizabeth A.
Zimmer,* Shu-Miaw Chaw,? Lynn J.
Gillespie," W. John Kress,'! and
Kenneth J. Sytsma”
ABSTRACT
Parsimony analyses were conducted for 223 species representing all major groups of angiosperms using entire 18S
ribosomal DNA (rDNA) sequences. Although no search swapped to completion, the €— recovered are ero
concordant with those retrieved via broad analyses based on the гараа: gene rbcL. The general congruen 185
rDNA and rbcL topologies further clarifies the broad eS of angiosperm phylogeny. In all pst ا the first- агч
аге ичк, followed by the cera family о сов This same за order of e arly-branching taxa is pre-
rved with several suites of outgroups. In most searches, the remaining early-branching taxa represent Piperales and
olds orders of subclass Ma аана њим сені. With the exception of Acorus, the monocots are supported as
monophyletic and a have as their sister Ceratophyllum. In most analyses, taxa with o urate pollen form
ag at ык. aei p ihe ral gt a large eudicot clade is composed primarily of taxa having triaperturate др a,
Two e present within the eudicots, one consisting largely of Rosidae and a second corresponding
dai s AG jiu: ie sensu lato T duh IMS Dilleniidae and Hamamelidae are highly (n atone x data sets of
rDNA sequences also permit an analysis of the -— of molecular — of this blems deriving
from both the ee of indels and uncertain alignme t of 18S rDNA sequences hav: dag ide ова in previous
me With the exception of a few well-defined г ен пена апі ر are relatively uncommon in 18S
NA; sequences are therefore easily dined by eye across the ai rms. Indeed, здер indels i in highly —
regions appear to be phylogenetically informative. Initial analyses саси ts both stem and loop bases are importan
sources of phylogenetic information, although stem positions are prone to M ue sus н Of the €
changes analyzed, only 27% destroy a base-pairing couplet; 73% maintain or restore base pairing
! This n earch was supported in y grants from the National Science Foundation (DEB 9307000 to DES, DEB
9407984 to DLN, DEB 9303266 to К DEB 9407350 to KAK, DEB 9306913 to SMS and Loren H. Rieseberg), the
National Poa Council, Republic of China (2818F) to SMC, the Mellon ое grant (to EAZ, PSS, and DES), the
Betty W. Higinbotham Trust (Department of Botany, Washington State Uni о LAJ and ВКК), and the Scholarly
Studies Program of the Smithsonian Institution (to WJK and EAZ). We dink | D. У wofford for access to PAUP* 4.0. and
5. is for i imony jackknife analysis. We also thank B. Alverson, J. € M. Hershkovitz, D. Olmstead,
J. Palmer, Y.-L. Qiu, J. Rodman, and Q.-Y. Xiang for prm plant material and DNAs used in this study. We appreciate
the valuable advice a D: Swoffo wd on analyzing large data and the help of M. ны in using the UNIX version
of PAUP*. We also thank D. Olmstead and M. Sanderson ча "helpful Pen on the manuscript.
ANN. Missouni Bor. GARD. 84: 1-49. 1997.
Annals of the
Missouri Botanical Garden
Although the angiosperms are almost universally
considered to be monophyletic, many basic ques-
tions of angiosperm phylogeny remain unanswered,
including: (1) what are the first-branching angio-
rms? (2) what is the ancestor of the monocots?
(3) what are the major groups of angiosperms and
the relationships among these groups? Despite in-
tensive study, these questions have been difficult to
answer for a variety of reasons. Most notable, per-
haps, is the inadequacy of the fossil record alone
to answer these questions conclusively. In addition,
the apparent rapid radiation of the angiosperms fol-
lowing their origin resulted in few morphological
synapomorphies among lineages at the base of the
angiosperm tree, hindering attempts to resolve re-
lationships among major groups (Crane et al.,
1995). Finally, the angiosperms ры relatively
few morphological characters for comparison а!
higher levels. For example, recent Дыра апају-
ses of morphological characters for angiosperms
(Donoghue & Doyle, 1989a, b) and all seed plants
(Doyle et al., 1994) included only 54 and 82 char-
acters, respectively. As recently demonstrated by
Doyle et al. (1994), careful analysis of both mor-
phological and molecular data is required to un-
derstand angiosperm phylogeny.
During the past dede several attempts have
been made to reconstruct the phylogeny of the an-
giosperms. Morphological and molecular analyses
usually identify the Gnetales as the extant sister
group to the angiosperms, in either the shortest
trees or those slightly longer (e.g., Crane, 1985,
1988; Donoghue & Doyle, 1989a, b; Doyle €: Don-
oghue, 1986, 1992; Loconte & Stevenson, 1991;
amby & Zimmer, 1992; Chase et al., 1993; Doyle
et al., 1994; Nixon et al., 1994; but see Goremykin
et al., 1996; Chaw et al., 1997). Molecular phylo-
genetic analyses include those based on rbcL se-
quences (Chase et al., 1993), partial 185 and 26S
ribosomal RNA sequences (Hamby & Zimmer,
1992), and rbcS amino acid sequences (Martin &
Dowd, 1991). These analyses tend to identify many
of the same major groups of taxa, but they often
present different views of relationships among these
groups.
In the largest phylogenetic analysis of angio-
(1993) presented the results
of two parsimony analyses of DNA sequences from
the chloroplast gene rbcL for 475 and 499 species
of seed plants. More recently, Rice et al. (1997)
have reanalyzed the 499-taxon rbcL data matrix to
search for shorter trees. The benefits to the system-
atics community of performing these large phylo-
genetic analyses of seed plants in general, and an-
giosperms in particular, have been considerable.
These studies provide comprehensive, explicit phy-
logenetic hypotheses of higher-level relationships
in the angiosperms. Furthermore, the need for sim-
ilar studies of angiosperms based on other char-
acter sets has been recognized, and such studies
have been encouraged (e.g., Chase et al., 1 }
Particularly important is the comparison of chlo-
roplast-based phylogenetic estimates (Chase et al.,
1993) with topologies derived from analyses of nu-
clear genes.
sperms, Chase et al
For reasons reviewed elsewhere, phylogenetic
analyses based on nuclear DNA have largely in-
volved portions of the rDNA cistron (e.g., Mindell
& Honeycut, 1990; Hillis & Dixon, 1991; Hamby
& Zimmer, 1992; Sanderson & Doyle, 1993a; Nick-
rent & Soltis, 1995). Analyses of 18S rDNA and
rRNA sequences have been used for phylogenetic
inference at higher taxonomic levels in animals
(e.g., Sogin et al., 1986; Field et al., 1988; Wain-
right et al., 1993; Wada & Satoh, 1994), protozoa
(Schlegel et al., 1991), algae (Buchheim et al.,
1990; Huss & Sogin, 1990; Kantz et al., 1990;
Hendricks et al., 1991; Chapman & Buchheim,
1991; Bakker et al., 1994; Ragan et al., 1994; Ol-
sen et al., 1994), fungi (Forster et al., 1990; Swann
& Taylor, 1993; Hinkle et al., 1994), lichens (Gar-
gas et al., 1995), bryophytes (Mishler et al., 1994;
Capesius, 1995; Kranz et al., 1995), gymnosperms
(e.g., Chaw et al., 1993, 1995, 1997), and even
among the deepest branches of life (Wolters & Erd-
i, Taiwan, Republic of China.
ttawa, Ontario K1P 64P, Canada.
SA Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560,
"Бера of Botany, University of Wisconsin, Madison, Wisconsin 53706, U.S.A.
Моште 84, Митбег 1
1997
Soltis et al. 3
18S Ribosomal DNA Phylogeny
mann, 1986; Olsen, 1987; Woese, 1987; Embley et
al., 1994; Bhattacharya & Medlin, 1995).
Despite this wide usage in other major groups of
organisms, the 185 rRNA gene has received com-
paratively little attention in angiosperms. In large
part this reflects the enormous interest in, and dem-
onstrated utility of, rbcL sequences for inferring
phylogeny, particularly at the family level and
above (e.g., Chase et al., 1993; Morgan & Soltis,
1993; Olmstead et al., 1993; Kron & Chase, 1993
Qiu et al., 1993; Rodman et al., 1993). In addition,
skepticism apparently exists among many angio-
sperm systematists regarding the utility of 18S
rDNA sequences for inferring plant phylogeny. Ear-
ly analyses of 18S rDNA or rRNA sequences in
angiosperms (e.g., Nickrent & Franchina, 1990;
Hamby & Zimmer, 1992; Nickrent & Starr, 1994),
while in general pointing to the possibile phyloge-
netic utility of these data, raised concerns that 18S
rDNA may be too evolutionarily conservative to ad-
dress phylogenetic questions at the family level and
above and that insertion and deletion events (in-
dels) occur frequently in at least some portions of
18S rDNA, making alignment of sequences diffi-
cult. In addition, other basic background informa-
tion regarding the molecular evolution of the 185
rRNA gene is not available for angiosperms. For
example, given that 18S rRNA, as well as rRNAs
in general, have inherent secondary structure that
includes characteristic loop (non-paired) and stem
(paired) stretches of RNA, should changes in the
encoding stem and loop bases be considered equal-
ly informative in phylogenetic analyses? Models of
rRNA evolution suggest that paired (stem) bases
will undergo compensatory changes to maintain the
appropriate base pairing. However, empirical stud-
ies of angiosperm rRNA structure are few (e.g., Se-
necoff & Meagher, 1992), and available data sets
have not been used to evaluate art of evolution
of the 18S rRNA gene in angiosperm:
The history of the use of 18S RNA and rDNA
sequences for phylogeny reconstruction in angio-
sperms was recently reviewed (Nickrent & Soltis,
1995). To date, the largest studies of 18S sequences
are those of Hamby and Zimmer (1992) and Nick-
rent and Soltis (1995). Zimmer and collaborators
conducted phylogenetic analyses using direct se-
quencing of rRNA from approximately 60 species
of vascular plants, of which 29 were dicots and 17
monocots (Zimmer et al., 1989; Hamby & Zimmer,
1992). These investigators sequenced portions of
both 18S and 26S rRNA, yielding a total of 1701
base positions per taxon, 1097 base positions from
the 18S gene and 604 from the 26S gene. The
shortest trees obtained had a number of features in
accord with existing classifications, but sampling of
nonmagnoliid taxa was sparse may explain
some of the unusual relationships suggested among
more derived angiosperms. Furthermore, many of
the nodes were poorly supported. As a result of the
unusual relationships suggested for some taxa and
the poor resolution obtained in this study, angio-
sperm systematists remained unsure of the utility
of 18S and 26S rRNA (and rDNA) sequences for
inferring phyloge
More dr porum and Soltis (1995) com-
pared the rate of evolution and phylogenetic reso-
lution of entire 18S rDNA sequences with those for
the chloroplast gene rbcL using a taxonomically
similar suite of 59 angiosperms. Pairwise compar-
isons showed that rbcL is generally about three
times more variable than 18S rDNA. However, be-
cause of the longer length of 18S rDNA, the ratio
of the number of phylogenetically informative sites
per molecule is only about 1.4 times greater for
rbcL than for 18S rDNA. Exploratory parsimony
analyses of angiosperms showed that several clades
were strongly supported by both rbcL and 18S
rDNA data sets. Nickrent and Soltis D con-
cluded that complete 18S rDNA sequences are suf-
ficiently variable to conduct phylogenetic valid at
higher levels within the angiosperms.
Here we explore further the higher-level phylo-
genetic relationships within the angiosperms using
entire п 1 A sequences. More specif-
ically, we provide phylogenetic hypotheses for flow-
ering plants based on analyses of four 185 rDNA
data sets, differing in both the number of taxa and
the inclusion of indels as additional characters. We
also compare the phylogenetic estimates based on
18S rDNA sequences with those obtained from phy-
logenetic analysis of rbcL sequences (Chase et al.,
1993). Using the phylogenetic estimates, we ех-
amine patterns of molecular evolution of 18S rDNA
by assessing the frequency of insertions and деје-
tions, the prevalence of compensatory changes, and
the relative phylogenetic importance of stem versus
loop changes in angiosperm 18S rDNA.
MATERIALS AND METHODS
SPECIES SAMPLED AND SOURCES OF PLANT MATERIAL
The species included in this analysis are given
in Table 1, along with family membership, general
collection information, and GenBank accession
numbers for the 18S sequences. In Table 1, and
throughout the text, we generally follow the taxo-
nomic circumscriptions of Cronquist (1981) for di-
cots and Dahlgren et al. (1985) for monocots. This
Annals of the
Missouri Botanical Garden
4
ИУ Еров “MS ју VAN NIS ‘888Z Mayo 9vooemoo[orsuy "т гвигрриро. штарзу
$661
ИУ £80071 ЛОЗЕ NIS ‘2262 то әвәәвицәо[ојѕиу "suns рѕојиәшо) pnj20jorsuy
jos y spes o0Sztn SA 9971 noyunjg әвәов!үелү `1 2424 паорон
ичен oz0cen sn чуо оваовоолу joumg pijofipissos DIYINGA
ицен #20291 SIM ‘6569 чүн ovaovay "1 Suasnyod 109
$661
У BLOVE1 цо Y полу NIS “16s тог оваовлу `1 тшојод кај +
за Y CT
sos y «0$ 66tcvn NIS 8267 то оваотиХоофу отрэитир ритиошорилодој
asoy Y ‘no0
jos y jos AGI SM '9922 51105 овас ћу (ysing) vnpousanr шттошот
say y иең 9902 SN “PEPE-Z6 зәл әвәовиәціиү yeg asuajodou um4ydo10)4)
10044 090491 SIM РОБЕ 27426 ovooruouuy supaSpaf отр
o40 190481 М сре 950Y) әвәовиошиү ds puojosp 4
SHS y snpos L6vcvn Sn ФР yng овоов | |олодшу тея ородоцош орјелодшу
ичен €90ctn SIM ‘6289 шуо овоовру у Army ‘ds e...
uopur]
ицен 6902 SIM ‘8989 шуо arooep![[A1tuy Y your vopfipunsws sumymg 4
ssy A 12021 SN ‘610686 VN оваовуту uoq "9 палодиту штуу 4.
ey) 181670 ІЅУН ‘IZEI “mY? овоотешету “1 этой тиру
105 96tcvn SA “TEL mnaoyysiay овоовогту un psupdxa pruo3p4ja]
snos sovern SM «т uvo 9eooerprur ү а трпит. V
эшо toten SM ‘SISE SNOS Y NOS әвәәвләәү v] umaqna 199y
uosuqof 06261 SM “00-86 uosuyof әвәзецүивәү səəy DIM] SÁYIDISAYID A
Aq yuequesy попио anew] әәлповуләцәпод Ауто satoadg
aouanbag
‘S198 wep uoxei-66[ 941 10u Inq ‘səs vep иохез-922 әц ш asad suaodsordue әвоці возеотрш | "s — ba. пәдеҷа[е родивше are soroods ‘sorruey uryt
"(мәз цес о) #шрлоәәр sjooouour tjsmbuor) о} Випрлооов кор) seram (4 peonequrs рабина ae sotoodg o опешвл oouonbos үү $81 10} pezá[eue sereds `q APL
18S Ribosomal DNA Phylogeny
Soltis et al.
Volume 84, Number 1
1997
y Aydoyesary "ү шпзләшәр ump]4ydojo.9;)
sos LISZHN пом 22016 MÒ әвәовүү | са MUS
SHIPS EG ПЭМ ZHI #5942 әвәәв]о[ецЧәг) [qe] гмортопој зл
0661 ‘tury? a Fa ae
жем 00991Х лыд 29 зоо NIS ‘7682 Tuo221N 98990118819 aS (qn)
DIS 2263 1u22421N әвәовипепввгу т ођојповтћо ри 9
^ M OW “9 ‘4 unossry овоовошво "| оХрара vouw)
ie ~ ~ ies 0100 “u's poarsurg оваовпојаа во) эң (T) sng вод омоцашле
. ~ . а
1 oeoovi[ojude -xe (чапу) nyobou 04221001. 4
ru Ha oh "ан Ra Lor ‘22017 psouids p1u1J420204
a. Hirn “urs 2рацәцс-Јү avoovsedde’) WPJ DUDILA/SDY 2u102]7)
=П[05 Y snjos TISZA OW o равна: a AGUAS
eis гей preted 686 uasunf әвәдвүпивішеғ) "1 ишә D11290]
он ин HOIN ‘#86 uasunf oeaov[nuedure?) "пе 2војтио omundun)
sapos Y snjos MEGAN pct akin ac С sa
зчәгҹәтү BIESEN n a ig papa вена
Mb Po х bal "126 сова ovaovxng "1 suana4aduios snxng
- SM ‘180-56 uosuyof әвәәвГәүррпд "woue14 npiapp vfapppng
os Med (INI “u's га 9voovrunug "uduoag (1) »souiZnun] vijozaag
ا 90Scvn 1 ! Se па вајари
24 опаолош
ssar Y uH SL0cV(1 SN '99р2-26 ssary оваовпошола n40jfijmpuod тилоот 019
0661 ‘snis jews
-ad 3 umouyun әвәовә1ввв1{ чцәчәор 02114 921
клен ким mper umouyun 9vootorsstug он ("1) puppy sisdopiqpiay
LLOOTX 6861 T + рни A 1 ign
“1219 ТИ ‘f ово2еи! 82104 boef юуиәүпәәп$ ni 1
JuorqotN Olesen OM 1 | am | d
йоту LOS@bN SIM “us uossaazy avaororquiog g 9 ари
E 29 DA
M 162611 SM ‘00-86 uosuyof avaoetuousig шәә$ рај D121 d
1661
7866 umouyun "uj: nsouinj3 snu]y
SX *opuo[e] у рлелес̧ : as pr sce чывы es |
ee 1049 NIS ‘1682 Tuo4421 риса а:
9016" 9 Худото)
N “Sz6 100H INP CT) зартолоцоцј шті,
anang 1 Y ‘ZION may ovoovtuodag "ds muofəquíç
m corer ПЭМ ‘SZZ 2504) әвәәвтиойәң noumBups x voyou viuoSog
' DA,
911105 soszrn A 229-2261 My rica. MAY Pa DA
4q живио) uon smy] 9o1nos/19u9n0A Ayre y satoads
эчә м
"penunuo) “| ajqe,
Missouri Botanical Garden
Annals of the
му "9 BUY ejueg oqouey
UISUIMS осртерл ILII Пея (18944) 02042140]2 02500]
sujos I£Scvn NON “93016 MÒ оваов| Ху тица ве] ‘ds wnyAydiuydog
чогу v6cevn 9661 “чогу NON “ws иолу овоов||ш45 `1 D1ofiuaomı DIA)
ѕѕәгу Y шең LLOGIN SN ‘E9PE-Z6 55213 оваовлодк у лорвлцоб snzpujsoqpp тәй)
S661
JUODOIN _ LVLVET “SOS 29 UAIN AIS ‘STOE ruouso1N овоовуповпу) "PIM 2201042. отовту)
Пеш
snos оестрл SVO ‘SETZ 191194 оваовшошпу) штаојпишта. штутодоголо)
uosuoMg тострл 3 “16 2504) аваовифтопу) "трпе nzojfimua) оадоду
51106 (САТА 20 | о та guy gjuesg оцэиву овоовјешововвол“)у пу unonuofio DULOSOSSOL)
sn]os SESE SA ‘ESIZ uDRıoy aeaor[nsSe1) uasne[) шттиполфта umpos
1911194 » youl
впјоб 129% SA TSIZ UDI LO y] 9BIDP[NSSPAL) -eH риотошоттр Чыл
SOS 9csecvn 10829 “9 'g uoidunung әвәәвүпв6вл7) (uosieA $) пртовла мири
uasqooef
suos scecvn SA ZSIZ чойо 9BIDB[NSSPIL) Y лодпн риргәтиләш ортол)
ѕѕәгу Y UY? y os0zra ‘SN ‘OILE-F6 55917 989081807) ssang 57101008100 511807)
ed
впјоб 29 зпјоб расту 216 'фу ррошу әвәәвилог)у м (qunyy) poruodof m3umjəy
sn]os x SH[OS €zSzPn пар, фу Surqáns әвәәвилогу [Novy 427$0210]02 D130407)
suos ГАТА | тү “TEN "$ n әвәәви10”) ‘qund, n3ruodpf pqnony
поло oresen ON hx 11999 әвәэв|пл[олио”) Ђовј тәгәрәү тәошойј
чцен 920cvn SIM “vs suvag овоовиешшој ‘ds тю]
ичен LOGIN SIM 7999 "ng оваовопцојо "Т грриштто umnpjo)
snog таесрп ПЭМ 7881 чогу аваовл ој) "т vipofiupo Day
suog ozszrn Y£L-V9 °9 ^g "do1j pyryouey 9vooput[eqosÁ) човиза рѕојиәшо) DUDIT]
snos 6181 пе-92 9 ‘doy, pjroare 4 ovooput[eqosÁn) "1 090% snupjpqos&np")
JUIIADIN 9ecoen NIS "2202 Tu24321N әвәәвцив2о|ц”) “MS $u32$210041D umnusokpag
с661
поло ОСУП *sn]os 29 IUJN NIS ‘9682 тој avaoe1podouay;) “1 naop4ajo т1әриніс̧
‘sonz Y prog
snog 8тесрл SA ‘OPSZ 51105 әрәәрүАцйтргәләг) -ә!$ штәтиойо[ штухуатр12227)
Aq yuequasy uone олтелојг] әәолтовуләцәпод Хш» 8212946
аопопбос
‘panunuory сү QPL
+
"(peusiqndun ‘asey)) sunooeg | ann, ay) 01 pajas Ápueysip quo st iZurumyy yey) sarpur вооџопђов “1994 Яшәп oeaoerunooe]4 jo sisáeue опоџовојкца po[reiop 149091 у,
зпјоб ecran QNI “us aoud әвәэвтиюзәсу ер шпәләшә штиоо)
0821 “pur
snis Otccta “) `9 BUY murs оцоиеу amwe xa se[iino(] vondi nup)
иоѕицо[ 092611 ‘Z91098 °9 `8 unossiy »vooeuambno y "шјави а suapuajds vusmbnoy
100] 8061€1 HIIN ‘9626 y20ruzay | kanoj, оштхг Diwan]
pozn 6ESZHN SIM ‘2212 wossoa]y 9eaoerunoor| 4 завио) отдруоә птици,
`N чойшип ој
E әләәмс̧ поета -eıoqe] [eorsoporg ешүолег) avaonqey штатив штета
<86
Е £Z9Z0X apouu22 uwouxun vooeqe 4 uio (7]) wow auna
snjos LESZEN ASW “ws гр ход] AS 4 "ds prurynog
5 эпо$ 9esen 9261 год avaoeqe 4 eran шып ogy
‘song
100 18627 006 тд әвәәвәүәзйп д X PIOS олрио«јод pojaidng
E 10$ сестр ред 10 Y sujos оваоеифоц па pura md тоц
“stu
oa safes resen БӨРЕВ-О14 ssconqoqdng -nH (пем) мудрим воой
105 [35:4 21] 0€20-98 “Py Эш ses d uong soni] opion] vydany
ES 100) 990151 2016 MÒ LE | "MO заргошјо Druuoong
човицо 26261 662 чогу әвәәвоия “WY иойэоэрш шттитоэрд
човицој ZLZ6V1 SM ‘S80-P6 uosuyof әвәәвәия (71) ssan-pan ворбуйорормү
OMS Vern ‘еар әтә) , Әәвәов:рәція “SwA CS ошобашоз Dspaydy
vu zpzsed LL ‘2626-1, əvəvvapəydq Jamg ponis vspoydy
pozn 9c9ct SIA “U's uoszaa]y ЛО “WPS "N 0170/0] “yo vauvojg
IUIN 060671 IS ‘868% телу әвәэвиўвәрү GUAY y олојјәдшт =пи?оәру
uory C6ZEP NON "FOO£ чогу әвәәвиәд puptuiXina sosddsouy
106 ZECTH "Я uouresuouieq IVIIPIISON(] sisuados 11950]
os $ =1[0$ osien OIN ‘166 vasuof anaonoesdiq "ds snopsdig
USO! ese IS '966g телу әвәәешә[и] иву (90) 0200 drag
човицо 82671 NON '291 чогу овооввио дец] орујодот хрјог)
uosuqo 822671 ON ‘81068 SMH avoorrsuadei(y notuoddp] visuadpi]
E 2226 uosvidduog
човиомб [rs под ewg “9 *g enug ovaovosne(] огоуірпи ѕәјәшодәр
Ф 4q ueque?) uone am IDINOS/IHYINOA, Anu y soradg
~
58 panunuor "pomi
Missouri Botanical Garden
Annals of the
*hps E€8LZHN NSO 'Р82'11 `P 12 Ássnaig wampun] "Hd ритггририлој suojov]
9100]
он £9091 SVL "ws pug f әвәәврц iL (00H) vorumusp) siséudosy
ON період
100H 29067 ‘POS6 Ўишируу 3 појдрјо) энәгери (тоя "D zppaesonq стотројо
изд
100H бЄ8С/Л HOIW 7822 220 овоово хә "хцәтуү штаоалра штоујј
uosuyof 26267 1 SM ‘S00-Z6 uosuyof araoe[[AudoıpAH "SIEM хә мој, 4070219 DIJ290Y q
эпос с810%П SM ‘TIPE SNOS Y yog овоововивлран Чета 1181213] студгрорца
sos 1822+n SA 012 UDI LO y овоововивлран зло], оурудолоош родирархн
31H
-PH L90cvn SIM ‘Z889 чуо 9vooequroeéH хә Хәлең sipignjoa vamog
ѕѕәгу y uqeH Z80zbN SA 8111-08 ssary овоовшоопен Bue] vapur тиоотон
SHS ESSEN SA ‘9162 SOS X 5106 овознрнош ин "т отрова oquipmbr]
snis csecvn 900£6 ni) "атара ‘Ogy оваоврцешешвн ‘ds радиу
JTopqorg (Lin
snog осетрп М ‘ESP 045 овоов влојеН хә syueg) 012212 $190.10 DY
suyos 28161 ALM “us Злодојота овоовлоџипо) uopur] DIDIUDU рлоџипо
su[os óvecvn OH “u's ирраој овоовшвјпеволе) ‘J JOOH vorupuisp] nondapopajag
“sug soy "pu
sos АД 1e3n9 впциперд “шпиечлән овоовшвјпеволе) Чиш$ ^q `[ хојдилу vassnoy
SHIOS £VISCI SM ‘0222 suos X 51106 FOE EO Чета umamo ойу
791
suos Lvecvn XAL 692 zayoung әвәәвиүпвволсу -muey snyofipunos иошоколоја
"ил
snos 9р6 SM FEIT uv2iojy IBIDBLIB|NSSOLL) (‘zomn]) зпаспоцој ошоо
пос сретрћ SM 'IOF6 2404 овоовшвјпеволе) “1 отапа род
зпјос y зпјос prszpn EEET-ZS 9 ‘A JI» n овоовмејпеволе) Ашвом sisuaquimboo птиојр005
‘[Meg 199) хә злепоцј,
вос срстрћ М ‘LO6PT-2261 мом овоовџејп5воле) SISUaLDISDEDPOUL DIXALT
вос ISTEPN 907079 'q1y Surq&ug ILII) "4282$ uafoyppos D119)
9199MG ЛҮП SN ‘IZZE-16 10 19 55019 овоовђопо) ошпја ("[qny) suam штио
2199M [317471 Sn “РЕР Adsao обоовјоцо) "uduoag wumaojfipou wnjauy
929935 9ТРЕРП Sn "crar әтәји) 999091900 "1 иошгиг штиво
snos 29 snjog evecvn NIS 626% 1191] IBIDBLIQUSIL) ‘oe SUDIPPDA STYYUDUÁYISI Y
Áq yuequa«) uon олпјелој и] аолпов/лоцопол, Aue] saroadŞ
aouanbag
"ponunuo) ^q әчү,
ва ас ован وا ا ا Km соса... -_ __- _
+
Soltis et al.
Ф
о
Е
=
=
$
Ф
Е
=
o
>
18S Ribosomal DNA Phylogeny
1997
S661 "pno
JU9.DJOIN ПРРаЛ “0105 29 109121 AIS 2063 Mayan овоовло 4 (әлем) suadias рошолодој
suos (24 | SM ‘BE znaoyysiog овоовоовјо АЦА "] риромошр 02201014
58913 $ Чең vVLOcVO SN ‘SOSE-Z6 55915 IBIDRIPÁ[IA ds mzyoyuņəH
зоон 985/7 м 825 25995 әвәовләлеаіед ug 9q19qua umosadAy
snos c6LcV(1 M ‘997 99) ILBIMIUO J psonjonifns тлиогра
[ос Tl6zcvn M 42768 95095 IBIDPPIYII) [pur] штрароха wmpu)
[ос 06271 DIS ‘918Z Mayan овоов 4) qxoy рәортиәшт пар)
вше 026291 AV ‘9572 42111109 овоовлави() вле) у рирцирх MYD) 4
uosuyof 682671 SA 'Р00-96 uosuyof обоовој) "| vandosna 9216)
зпјос 29 5пјоб 68270 O8T-PL фу 2uiqKug IBIDRSSÁN "euoe(] ртоитштәр DIIYIOId UD)
su[os 9882cvn SA '09 znaoyys42H 9vootutzejoAN "т odoppf sinqpapy
S661
JUSIADIN РОРРЕЛ *sn]og 29 PUAN AIS ‘9062 1u2321N овоововцашА N әшед рзолодт nanuduiAA
snos 282000 DIS 9508 11910 оваовууподо “ds sayruaday
wo SE8SZT EILL ZIZ6 100H а ANION ‘saad (PIM) Рәт] oqunjayy
ѕѕәгу Y иең £80270 sn “u's ‘prey ‘104 ‘$: IBIDESNIA RIOD DIVDUMUNID юп
SOS 29 SNOS 982сӯП SIM 10502 SMI аваоввиџој "ure Diafiajo n2u10]y
зчәгәту 866РСТ DIS #262 11911 3890810 T AP ToN
5661
JUSIDIN 089SZ1 ‘SJOS 29 JUAN AIS 810€ Masynyy оваовдолопојј "т ?40jfiun одолоиоју |
Bomy 8евсуп SM ZE znaoyys12H дваовш ап [OW "] VID ponaa ognjopy
100H VI62£1 AJN ‘IEIZ pjoasaopf овоовшлодв шоу sion D4ffoo рлодкошј 4
100H EGET HOIN 2292 ZWN овоовшлодв шоу 7| sisuappups штилодктиоуј
ssary ицен 6L0cvn ап PELE-P6 Ssary oRooRIURIL A TIPA 4070919 DIUDID YA]
jjoznx Lc8cvn SIM “U's uos120y оваовлјејј "p ummsanj umdiXssoy
поло орорал AIS “5063 14244NN овоовт а јејј e] 042212209 птутајруј
ШЕРИМ LOPEZ DIS 006% 1u243421N әвәәвиг] “ү әииәләа штит]
зпјос Y зпјос v8LZPN HDIW “6098 y2o1uzay әвәәрцивишг] “PIEM зартоотпалокола тәҹләо] 4
MEU) SZL6cd ISVH '2960Н 9gooeI[TT wey wmupsouuof шту 4
SA '9rec sujog 29 sujog 9eoogIne'T səəN (INN) umpiqpp spafosspg
JUSIADIN C6LIET DIS 'eróc 1u24421N ovoov[eqezrpaeq 231992 (MoH) орритђ тлдгуу
uosuyof L8Z6V1 SM ‘100-S6 uosuyof 3899E IUE"] "1 ојтомлхојашро. штшју
Aq yueques попео onje1o]r] 9o1nos/19qonoA Áprure y soroadg
аопопђос
"penunuo) Tp [QL
Annals of the
10
Missouri Botanical Garden
зпјос 66Lcvn SA “сст& upnziopy әвәовишецу ysing snauimsups snyjouvay
Пе
1909 689/7 ПЭМ ‘08016 MÒ пр укыл my vunssioyduns D2141041UDX
S661
JUIN 260721 *snjog 29 1uoryotN DIS ZE6Z 1uo4491N оваовјпоппиву ZJUBI") snopips SNINIUNUDY
100H 888811 ноти Pionog 3p $504 эво "qétres (Y ойн mdo) 4
ques
AIN Y помјођ OW “us ‘12m0 IBIDROLÁ Y "ug орла vod 4
JUAN 11881 DIS TE6Z 1191] оваовошпа Ур штрираг опита |
әш d A ЭЧИ
100H LE8SLT TAN ‘OIL svjgnoq әвәәвә]од,{ ‘L `Э шпәәрмоә шпшләбзоәр]] 4,
2661 ове
JU91XOIN сетрал ‘SOS Y JUSMDIN -e[) PIUBS *eruoji[e?) Jo ‘ATU әвәәвә1од{ ла Y DS]99x2 рту
џовоцој S666V1 SM “900-S6 uosuyof SINE "ds ounid
SUS
чәшшу “919IMS 862200 DIS 2266 1u24121N] әвәәвио8 Код "1 олаћат 04070207)
snos 26101 на 2951 2/00 әвәовүе8 од "PIM 240jfionpd vjp24joq
2661
Бону s snis РӮСТ "snos 29 3ueryotN SA “SI-Z6 uosuyof РЕЗОН НОШЕН “Sung Dd? три)
uosuyof LLZ6V1 SA “U's UOS1INDA овоовшошојод лег) Suapunos Dango?)
uero y Ава °9 y (998
AR I1Z66VT SA '020-96 ‘How эр иоѕицог ЗА АРА -opue1g) 801108 pip Foyruvoy
Vo “oy
SNOS 7 opeq “оцу *seuo1e10qe']
*iouumz “олоомб 962cvn qoo1uo[) (ума эәїшопәс)) IBIDROS “| shoul 727
7861 7861
©те 19 RMIBARL, сс)00Х “Te 19 BMIBABL uMouxun a2900 qf "1 pays D210
роу c6Lcvn DIS “u's тиолуо у оваовшввашпја "шет DIDMAMD ободштја
SHES M
“ISUIUTZ "олоомб VOLGA SA ‘HISZ 81105 Y 5140S bek i. O le
2661 qd d
роп 5 SH[OS СЪ18С1 ‘SOS Y UAJN VSM “us 81әә8ә1у овоовлодвот а хә “моң шпоиодог штіойѕота
Aq ҳивдиәх) попео IINIT аолпов/лоцопој Күрше sardadg
аопопбос
грепипиођ сү e[quj,
= 1091
-IPON 29 MP STESEN NIS ‘TVO ON 112409 ovoogure[nudo1os ПИ а suoma омриту 4
"reg
100H ГА УЧА | HOIW “u's yaoruzay авоовлрива ос (zoın]) SISIUIYI DIPUDSIYIS
8005 29 знјоб er8zvn пча 1625-01 YM UPA OS ‘qun, sisuadpo D144
SHIPS EGA SA “ESZZ su]os Y 1105 әвәовелугхес̧ ооң о1уо/гЯәуш vspsfixog
> suis 09921 SM ‘ZEZZ чгравн овоов8елјтхеб 1 sepiopos wnsoyruad
5 зпјос 60821 SM “U's s1105 2р suos IBIDRÍRIJIBS syurg pDuquaf DISSDULD qd
Ej зпјос x зпјос 808ZPN OW 2282 suvuna IBIDRBRIJIXES quny, n22Djj&udoX402 vrunuoy
Е nona CION)
а. sgos Tb18Z1 (YIN. “us spwoyy IRIDPÍRIJIBS umpo9nyrods uoppjadoinda]
5 51105 6£187X SM '6F6I $UJOS Y 51105 IBIDRBRIJIXBS sejgnoq purup4onu раоџотон
a с661
© SROS 161861 “SIJOS 29 їйәл{о! SM ‘6/7 51106 Y 51105 әвәоввлугхес̧ “ABD DIOfIYIUOS розира 4
Е
В. HLYad
© 2 зпјос L0gcvn “62р ULDIY Y sjouuy әвәәв8елухес̧ "pug отрипога audsowasy
Ф = souof
2 e 5105 908cv(1 SA “8£911 2719249 OS (офа) әтрәшләрит vrurykog
E e SD[os 29
T лошшт7 “9199MG совсрл Sn '921 yrs овоовлипес тп ттиләә snanampg
©661
мәә ГАА! *sn[os Y juen[otN DIS '0P6Z 1u24491N авоовлипеб "qunqj, DIDp409 тихттој
suos v08Z9N SA “us uvo IBIMPIUIIBLIRS "1 vasndind DIUIIDLIDS
TA Y Pyy
100H 1788/7 Hd “us m) IBIHPXOPOJUIÍ LS (АЦО) отоәипо охоротиәдс̧ |
uory 080€vN 9661 ‘чогу ПЭМ ‘621 2504) әвәәв1оЧес uskoy ("1) viodoz олорируј
TUTAN Є1ЄӨЄП AIS 162 119 оваоврша ес "ue рурүпәтирЧ омотолјеоу 4
"pueis
sn[os €09cvn AIS ‘TELS 1u24321N овоовјејшес X suooH »7p]022un] sus)
1001] Opr8SZT ISVH ‘8199 42u2ng оваовијеб 190YUINS 01906
= WAIN TIESEN NIS 2263 1444NN geo q штуирто snam) 4
= SOS cogcvn ASO “u's Fumy oesoeiqny "т suada vjjouonpy
n su[os loszra SA ‘012 uvziopy IBIDRSOY 19917; houg) 1әтоуира vavndg
Е 59106 67281 Ha “us uosqoiq 4 Я | yosteg ("1) 2215194 пит]
= SHS 608cvn SA ‘TELS uow 9999990 ssea(] шәр типоца
а Aq yuequesy uonejr әлегә әолпоѕ/ләцәпод Áprue q вогоо С
Ф әәиәпһәс̧
E i
3o
5 о: "penunuo) 'T AQEL
ВИРУ ie TNT >»
Annals of the
Missouri Botanical Garden
овоов | |Ацдо8А7
UT] шт12и0ѕ UNIMON)
81105 vc8cvn HOIN “u's uossapuy
ssary Y ицеН T90cvn SN '9922-16 55213 оваовло ти wquo1oN штәитшр19 1901217
род Y WIEN LTOOZW 9961 T94 Y UNEN umouyun овоовпше7, "т оршта тшо7
ssory y uyeH po0ozrn SN ZEPE-Z6 Ssary овоовооцлошивХ "ds DIOYLIOYIUDX
SHS 27 1918
чәшш17 *91994G €Z8ZPN Sn Zr ng овоовлоји у 104 72) Y ^W Г warn skuq
зпјос Y su[os ecacvn Sn 6 yng овоовлоји у доошоо SUNA]
пос Ic8cvn DIS ‘EGEZ 1910] 9B9OBORSIA "| штдајо U4
зпјоб O0cgcvn DIS ‘26g 1u24121N әвәәвәги[] -[M9 29 “dousey 12421ppo с.
зпјоб 61867п SM ZISG 51195 Y 810S a3899BUI[(] DIDLISS 00011127
"prauyos
snos EAN NON “Z0006d "10 әвәәвшү[\ "M `2 sisuaupuum& s112)
51105 11871 NON ‘02 25992 овеовлоштј, pyofiuyn pio,
S661
snos 96L1€1 “епүо$ 29 JUSDOIN ПЭМ “ETT 2504) oeoov[ooudoa, "т зтош wnjoandosy
'0007 29 p[oq
81105 9T98cvn NON 93006 MÒ оваовлрпороцоодј, -918 зартођоло uo1puapot204],
jjozuxy 628601 SIM ‘812 vosseqqy эвде ‘пепо пииршәәѕ Dayan]
snos ЧЕ ТАД NIS 626% 1u244]91N овоовоцј, "f poruodpf решо)
snos vI9cvn NON 60006 "10 звоовциоовлој, АЦО ззигилу UOLJUIIDA/A],
Чен SLOGAN SIM “9969 ЧОН оваовов|туаооој, “Т чзигаро ојјгирр)
sn
иең £90ZPN гробрбд uepies) "jog LINOSSIJA 38908208], ‘ds 0220],
погу L6cevn 9661 “1014 NON '9002 чогу аваовоојашке ‘biy pim moruod sovojduiAg
пол 96cevn 9661 “чогу NON “ооё чолу әвәәвәвлА15 ше] риромошр XDIAIS
с661
JUSIADIN 6TvbZ1 ‘SOS Y IAJN AIS EP6Z 1u244291N aevoortursiedg Pug штагоодата шттир9100
"игд (гриојцоојцоб
uosuyof vLZ6VI SM 200-86 uosuyof овоовивјоб x “ueyo) рлоуіотра visjofunag
qual
PIN Y MOQ LIESEN NIS “144$ OW 7/20) әвәәриеүпцаоләс "хцотиј DIDJ029UD] SUDMNPƏd
quai
AIN Y [[9M[0) 9reeen AIS ‘OLGAS VO 7/2799 овоовџејпудолос "gueg smymupuo зпаоооц а)
Aq yuequesy uOHE110 олпувлој] 991n0$/19q9noA Á[rure y saradg
зопопбос
enunuo) T әү,
Volume 84, Number 1
1997
Soltis et al. 13
18S Ribosomal DNA Phylogeny
approach parallels that in nit et al. (1993) and
should facilitate compariso
As with the broad sed of rbcL sequences,
close examination of the genera included in this
study will reveal an uneven taxonomic distribution.
Some groups are relatively well represented (e.g.,
Saxifragaceae sensu stricto [Saxifragaceae s. str.]
and allies, ranunculids, Asteridae sensu lato [As-
teridae 8.1.]), whereas others are not as thoroughly
sampled (e.g., portions of Dilleniidae and Magno-
liidae). However, our selection of taxa was not ran-
dom. We attempted to include samples from all ma-
jor angiosperm orders and subclasses sensu
Cronquist (1981). Furthermore, in selecting genera
for sequencing, we tried to sample representatives
from each of the major clades recovered in the
analyses of Chase et al. (1993) (e.g., rosid I, asterid
I, asterid III, etc.), as well as from the various sub-
clades present within those major clades. We also
used, when available, the same DNAs used previ-
ously for the sequencing of rbcL (Chase et al.,
1993). If a given DNA was no longer available, we
attempted to obtain leaf material of the same spe-
cies, and if that failed, from a congeneric species.
Another factor that influenced our choice of taxa
was sequence quality. As discussed in detail below,
one outcome of this study was the discovery that
many available sequences are erroneous, some
highly so. We therefore attempted to eliminate any
dubious sequences from our data sets. In addition,
some available sequences were not included be-
cause they were incomplete or contained numerous
ambiguities or extensive gaps.
Several laboratories were involved in this proj-
ect; hence, several different protocols were used to
anual sequencing strategies were
employed, 7096 of the sequences analyzed were
generated via automated sequencing. The general
methods used for PCR amplification and subse-
quent manual sequencing of 18S rDNA are provid-
ed in Nickrent (1994), Nickrent and Starr (1994),
and Bult et al. (1992). General methods for the au-
tomated sequencing approach for 18S rDNA are
given in D. Soltis and Soltis (1997). The base com-
position of the oligonucleotide primers used for
PCR and se ng are provided in Nickrent and
Starr (1994) and Bult et al. (1992).
ALIGNMENT OF THE 185 rDNA SEQUENCES
With the exception of a few, small, well-defined
regions, alignment of 18S rDNA sequences was
easily accomplished by eye across all taxa. This
general ease of alignment is due not only to the
highly conserved nature of the 18S rRNA gene, but
also to the fact that most length mutations involve
insertions or deletions of a single base pair.
Straightforward alignment of sequences was further
facilitated by the fact that most indels in 185 rDNA
are confined to a few specific regions that are par-
ticularly prone to variation in sequence
and length, such as the termini of helices E10-1,
17, E23-1, and 43 (see also Nickrent & Soltis,
1995). Because they were difficult to align over a
broad taxonomic scale, no attempt was made to
align four small regions of 185 rDNA over all of
the taxa analyzed: positions 230-237; 496-501;
666-672; 1363-1369 (see Appendix). These base
positions correspond to the sequence of Glycine
ax (Eckenrode et al., 1985), which provides a
convenient reference sequence because of the
availability of a proposed ribosomal RNA second-
ary structure model (Nickrent & Soltis, 1995).
These four regions of ambiguous alignment were
subsequently eliminated from the um ic
analyses, following оог and Olsen (1990). I
addition, the extreme 5’ and 3’ ends of the se-
quences were not included in the analyses. Posi-
tions 1-20 were excluded because they correspond
to the forward PCR primer. Because most of the
sequences were саму readable at, or just before,
base position 41, we began analysis of our data set
at position 42. At the 3’ end of the 18S sequences,
base positions 1751-1808 (on Glycine) were often
difficult to read and also were eliminated from the
analysis. Some sequences are incomplete at the 3’
end and are approximately 1700 base pairs in
length. The total length of the aligned 185 rDNA
sequences was 1850 base pairs.
Two indels, each of a single base pair, were de-
tected in highly conserved regions of the 18S rRNA
gene not prone to insertion-deletion (Table 2). One
indel (A), an apparent deletion based on outgroup
comparison, is present in all higher dicots (i.e., the
large clade consisting of Rosidae and Asteridae
s.l). A second indel (B), an apparent insertion, oc-
curs in all members of the saxifragoid clade (also
referred to as Saxifragales; D. Soltis & Soltis,
1997). These two indels were included as charac-
ters in two of the phylogenetic analyses, as de-
scribed below.
PHYLOGENETIC ANALYSIS
We constructed four data sets for phylogenetic
analysis: (1) a data set of 223 angiosperms plus five
members of Gnetales as outgroups, without the two
indels noted above; (2) taxon sampling as in (1),
but with indels A and B (see also Table 2) included;
Annals of th
Missouri Botanical Garden
Table 2. Potentially phylogenetically informative in-
dels located in conserved regions of 18S rNDA. Indel A
is one-bp deletion characterizes all higher eudicots
clade. Base positions correspond to the last position given
in the sequence of Glycine max.
Indel A
1529
Glycine CCGGGTA ATCTTTC -
Trochodendron CCGGGTAATCTTTGA
Indel B
1406
Glycine TATGGCCGCTTA -GGC
Heuchera TATGGCGATTTAAGGC
(3) a data set of 194 angiosperms, plus five Gne-
tales as outgroups, without the two indels; (4) taxon
sampling as in (3) above, with indels A and B in-
cluded. For data sets 2 and 4, the indels A and B
were added to the data matrix, and the position
scored as either present (1) or absent (0).
Four data sets were used for several reasons. First,
the approach used permitted an assessment of the
phylogenetic informativeness of the two indels. Sec-
ond, our goal in constructing the two smaller data
sets was to improve the phylogenetic analysis by re-
moving incomplete and/or possibly erroneous se-
quences and by reducing the size of the matrix to
make the problem more tractable. The two smaller
data sets (3 and 4) differ from the larger data sets
(1 and 2) in the removal of several taxa having long
branch lengths (e.g., Dillenia, Acorus) and several
taxa for which the sequences were incomplete (e.g.,
several of the ranunculids). In addition, representa-
tives from some of the larger clades (e.g., monocots,
Asteridae s.l.) and from some families for which mul-
tiple sequences were available (e.g., Annonaceae,
Aristolochiaceae, Proteaceae) were removed to con-
struct data sets 3 and 4
The outgroups were five members of Gnetales:
Ephedra sinica, E. torreyana, Gnetum nodiflorum,
G. gnemon, and G. urens. Gnetales were the logical
choice of outgroup because they appear as the ex-
tant sister to the angiosperms in most recent phy-
logenetic analyses (e.g., Crane, 1985, 1988; Doyle
& Donoghue, 1986, 1992; Donoghue & Doyle,
1989a, b; Loconte & Stevenson, 1991; Chase et al.,
1993; Doyle et al., 1994; Nixon et al., 1994). In
addition, to ascertain the topological impact of oth-
er outgroups, particularly with regard to the first-
branching angiosperms, we conducted several other
searches. Using the smaller 18S data sets (194 an-
giosperms), both with and without indels, we used
as outgroups: (1) the five Gnetales and Zamia pum-
ila; (2) the five Gnetales, Zamia pumila and Cycas
revoluta. Similarly, for the large data sets (223 an-
giosperms) we used as outgroups: (1) the five Gne-
tales and Zamia pumila; (2) the five Gnetales and
a recently acquired sequence of Welwitschia mirab-
ilis.
Because of the large number of taxa involved, we
used two basic search strategies. The first method
was a heuristic search sisse using PAUP* 4.0
(Swofford, pers. comm.) and t dap extent
PAUP 3.1.1 (Swofford, 1993) gea
RANDOM taxon addition, and TBR e hue.
ping. These searches were permitted to run for a
week or more using either a Macintosh Quadra 650
or Sun Sparc Server 600P. These searches did not
produce trees as short as those produced by the
method — and will not be discussed further.
The ary search strategy was inspired by
Maddie et aL (1992) and suggested by D. Swof-
ford (pers. comm.). For each of the four data sets,
we used 50-100 consecutive searches without
MULPARS using RANDOM taxon addition and
NNI branch-swapping. We then performed multiple
searches (300-500 replicates; a Sun Sparc Server
600P typically required 19-25 hours to complete
five replicates) using RANDOM addition, MUL-
PARS, and TBR Беју swapping, where only two
trees (NCHUCK = 2) of a specified length
(CHUCKLEN) or longer were saved per replicate.
The length of the shortest trees obtained in the NNI
searches was used as the initial CHUCKLEN value.
As shorter trees were found, additional searches
were conducted with lower CHUCKLEN values.
This approach prevented the searches from being
overwhelmed with trees.
The final portion of this search strategy involved
use of the shortest trees obtained above as starting
points for subsequent searches, again with MUL-
жаныр and TBR branch swapping. These searches
ere permitted to run for weeks or months using
Macintosh Quadra/Centris 650 or PowerMac 6100
100 computers. Typically no more than 2000-
5000 trees were saved in any of these searches. We
used starting trees of several different lengths when
implementing this final portion of the search strat-
egy to explore tree space from multiple perspec-
tives and to prevent the analysis from stalling while
swapping on suboptimal trees id Soltis & Soltis,
7). For data set 1, 94 starting trees of lengths
3929, 3930, 3934, 3936, Pe 3938, 3939, 3940,
and 3941 were used (shortest tree ultimately ob-
tained had a length of 3923 steps). For data set 2,
78 starting trees of lengths 3938, 3939, 3940,
—
Volume 84, Number 1
1997
Soltis et al. 15
18S Ribosomal DNA Phylogeny
3941, 3942, 3944, 3946, and 3947 were used
(shortest tree ultimately obtained had a length of
3930 steps). For data set 3, 192 starting trees of
lengths 3506, 3508, 3509, 3512, 3514, 3515, and
3517 were used (shortest tree ultimately obtained
had a length of 3501 steps). For data set 4, 96
starting trees of lengths 3513, 3514, 3515, 3516,
3517, 3520, and 3521 (shortest tree ultimately ob-
tained had a length of 3507 steps). Many of these
searches resulted in trees one to several steps lon-
ger than the shortest trees ultimately obtained;
these longer trees were also examined to help as-
certain the general support for some branches. Im-
plementing the above search strategy for the data
sets described ultimately entailed well over two
years of computer time.
Implementing decay analyses (Bremer, 1988;
Donoghue et al., 1992) is impractical with data sets
of this size. To obtain an estimate of support for the
18S rDNA topologies, we applied the parsimony
jackknife approach (Farris et al., 1997) to data set
1 (this analysis was kindly conducted by S. Farris).
The jackknife is a resampling approach, similar to
the bootstrap, in which the characters of a data set
are resampled to generate replicate data sets. Each
replicate is analyzed, and the proportion of repli-
cates supporting a given conclusion (in this case a
clade) is considered a measure of support. Jack-
knife percentages can therefore be interpreted sim-
ilarly to bootstrap percentages. In this analysis,
1000 replicates were conducted, and a minimum
jackknife value of 50 (CUT = 50) was used (i.e.,
only clades supported by jackknife values of 50%
or greater were retained). If a node is supported by
one uncontradicted character, the jackknife value
is 63% (Farris et al., 1997). Thus, clades with val-
ues of 63% or more are strongly supported; nodes
with values of 51–62% are less well supported, and
those with values of 50% or less receive no support.
RESULTS AND DISCUSSION
I. THE EVOLUTION OF THE 185 rRNA GENE
The accumulation of a large data set of entire
18S rDNA sequences has permitted a more thor-
ough assessment of the general evolution of the 18S
rRNA gene. Unlike protein-coding genes, such as
the widely sequenced rbcL, matK, and ndhF, there
is no clear frame of reference for aligning sequenc-
es or revealing errors. For example, with protein-
coding genes, translation of a sequence to amino
acids will potentially reveal some errors such as
frameshifts and internal stop codons. No such in-
ternal check is available, however, for rDNA. As a
result, general features concerning the evolution of
18S rDNA have, in large part, been greatly mis-
understood. In particular, insertion and deletion
events have long been considered common in 18S
rDNA; concomitantly, alignment was Ponce
highly problematic. Until now, the exi
of angiosperm 18S rDNA sequences was insuffi-
cient to assess these views.
INSERTION-DELETION AND ALIGNMENT
This study reveals that indels are not widespread
in the 185 rDNA sequences of angiosperms, but
instead are confined to a few, small regions of the
gene. Furthermore, with the exception of these
same small regions, alignment of 18S rDNA se-
quences is straightforward. Several possibilities
may explain the misconception that the 185 rRNA
gene is highly prone to insertion and deletion. First,
the literature contains a number of erroneous 18S
rDNA sequences. We have resequenced the 18S
rDNA of over 20 taxa, and have found that some
published sequences are in error by as many as 33
bases, which corresponds to 1.896 of the total gene.
In several instances, we discovered numerous er-
rors in the 18S rDNA sequence for a taxon using
the same DNA originally used to produce the re-
ported sequence. These errors in previously gen-
erated sequences are not confined to base compo-
sition, but also involve the presence of what we
refer to here as "false" insertions and deletions. For
cluded a large number of “false” insertions relative
to all other angiosperms. Integrating our new se-
quence for Zea into the angiosperm data matrix and
removal of the previously published sequence re-
sulted in the elimination of 14 indels from our 18S
rDNA data set, four of which were alignment gaps
that had to be added to all other angiosperms. We
were able to remove additional alignment gaps
through the resequencing of other taxa for which
18S rDNA sequences were reported previously. As
a point of comparison, the total length of the
igned sequences in the data matrix of Nickrent
and Soltis (1995) for 64 taxa was 1853 bp. In con-
trast, the length of the aligned sequences in our
228-taxon data matrix is only 1850 bp. The rese-
quencing of additional taxa for which published
sequences cause numerous alignment gaps would
likely decrease further the total length of the
aligned sequences.
The numerous erroneous 18S rDNA sequences
in the literature ү result from inherent dif-
ficulties in sequencing rDNA. Secondary structure
in the rRNA for which this gene codes is also pres-
Annals of the
Missouri Botanical Garden
Table 3. Area initially thought to be prone to insertion
and deletion; however apparent gaps result from sequenc-
ing difficulties. The underlined portion of the Hydrangea
sequence shows the actual base composition that we have
determined to be present for many taxa for this area.
215
*
Hydrangea AAAGGTTGACGCGGGCTTTGCCC
Glycine AAAGGTCAACACAGGCTCTGCCT
AAAGGCCAAC----GCTTTGCCC
AAAGGTCAACGCTTGCTTCGGCT
AAAGGCCAAC----GCTCTGCCC
Francoa AAAGGTCGAC----GCTTTGCCC
Podophyllum _ AAAGGTCAACG- - -GCTTTGCCC
AAAGGCCGAT--CGGCTCTGCCC
AAAGGTCAAC???????
AAAGGTCGA
Heuchera
Lepuropetalon
Prunus
Austrobaileya
f". I. Lidl
р
Вихиз
ent in the gene itself and may lead to compressions
and associated sequencing problems. More than
one molecular systematist with substantial experi-
ence in the sequencing of chloroplast genes such
as rbcL has referred to the sequencing of 18S rDNA
as “tricky.” We have found that preparation of sam-
ples via cycle sequencing followed by automated
sequencing yields reliable 185 rDNA sequences
that appear more accurate than most manually gen-
erated sequences. The critical procedural step is
likely the cycle sequencing reactions, in which sec-
ondary structure is reduced or eliminated by high
temperature. Several specific regions of 18S rDNA
are particularly difficult to sequence. These include
base positions 215-230, 1355-1365, and 1705-
1715 (all positions mentioned in this paper corre-
spond to those of Glycine max; Eckenrode et al.,
1985), as well as several of those small regions not-
ed earlier that are prone to insertion and deletion
(positions 230-237; 496-501; 666-672;
pe (see Appendix).
e will use the first of these regions (base po-
ear 215-230) to illustrate the errors that can
result in 18S rDNA sequencing. Based on manual
— (generated by D. Soltis & R. Kuzoff), the
base composition of this region in Saxifragaceae
“aa several other rosid families initially appeared
to involve a large deletion relative to some other
available sequences (see Table 3). Similarly, the
18S rDNA sequences generated manually by other
investigators, representing a diversity of —
sperms, typically were lacking one or more bas
pairs in this region. Alternatively, побио
scored this region as uncertain, using either “?” ог
“N.” Thus, sequences available prior to this study
suggested that this region was highly prone to in-
sertion and deletion. As a result, alignment of this
region was initially difficult. Alignment problems
were exacerbated by the apparent occurrence of
base substitution in the region. Further compound-
ing the difficulty of alignment is the fact that the
region just 3’ of this area actually is prone to in-
sertion-deletion, as well as to considerable varia-
tion in primary sequence. Base positions 230-237
correspond to one of the variable helix termini not-
ed above. Cycle and automated sequencing of over
100 taxa, however, revealed no indels in the area
of posan 215-230. In fact, after resequencing
this region in many taxa for which manual sequenc-
es initially suggested the presence of numerous in-
dels, we have concluded that indels in this region
are either extremely rare or nonexistent. This region
is G-C rich; as a result, sequencing “stops” often
occurred, yielding only a portion of the base pairs
actually present in the region. Alignment of these
incomplete sequences suggested numerous indels
in this region, leading to the misconception that
indels were frequent. Similar sequencing problems
were encountered in other portions of the 185
rRNA gene. Taken together, these regions had con-
tributed to the view that insertion and deletion are
common in 185 rDNA
Although the frequency of indels has been over-
stated for the 185 gene, several regions of 185
rDNA are, in fact, prone to variation in primary
sequence and length. However, these regions are
small, easily located, and, as noted by Nickrent and
Soltis (1995), typically confined to the termini of
helices on the proposed secondary structure —
for 18S rRNA (e.g., Nickrent & Soltis, 1995). Е
such regions, represented by base positions 230-
237, 496—501, 666—672, and 1363-1369, corre-
spond to the termini of helices E10-1, 17, E23-1,
and 43, respectively (see Appendix). These regions
are difficult to align over a broad taxonomic scale,
such as all angiosperms, and were therefore not
used in our phylogenetic analyses (see Materials
and Methods above). On a lower taxonomic scale
(e.g., closely related families), however, even these
highly variable regions are easily aligned, permit-
ting the use of these regions in more focused stud-
ies (e.g., Polemoniaceae and related Asteridae s.l.,
Johnson et al., unpublished; portions of Saxifrag
ceae s.l, D. Soltis & Soltis, 1997; Orchidaceae,
Cameron & Chase, unpublished).
Because indels in 18S rDNA are neither as prev-
alent nor as problematic as previously thought,
alignment of clean 18S rDNA sequences is straight-
forward. With the exclusion of the few small regions
noted above, alignment of over 200 angiosperm se-
quences was straightforward and easily accom-
Volume 84, Number 1
1997
Soltis et al. 17
18S Ribosomal DNA Phylogeny
plished by eye. This is also true for the alignment
of 18S rDNA sequences on a broader scale across
vascular plants (P. Soltis et al., unpublished). As
noted above, the resequencing of some taxa greatly
facilitated the alignment process in angiosperms.
PHYLOGENETICALLY INFORMATIVE INDELS
Not only do indels in angiosperm 185 rDNA se-
quences not cause alignment problems, but some
indels may be phylogenetically informative at the
level of our investigation. Here we do not consider
those indels located within the variable regions not-
ed above, but only those indels located in highly
conserved regions not normally prone to changes in
length. Two such indels in particular (Table 2) ap-
pear to be a informative across the
angiosperm
One indel involves an apparent deletion of one
base pair that unites all higher eudicots (Table 2,
indel A). This base pair is present in сше,
monocots, paleoherbs, Magnoliales, Laurales, ran-
unculids, Trochodendraceae, Tetracentraceae, Pro-
teaceae, Sabiaceae, Platanaceae, and Nelumbona-
ceae and is absent from all higher dicots (i.e., the
large Rosidae clade and Asteridae s.l.). Thus, the
distribution of this indel agrees with the results of
phylogenetic analyses based only on base substi-
tutions (Figs. 1, 2, 4; all figures, plus Appendix,
follow Lit. Cit.). In addition, the distribution of this
indel also agrees with topologies based on analyses
of rbcL sequences. It appears, however, that this
base pair may also have been lost independently in
two of the monocots analyzed here (i.e., Calla and
Chlorophytum).
The second phylogenetically informative indel
involves an apparent insertion (Table 2, indel B)
that unites all iê of a ebr clade
(Saxifragaceae s. str. and close allies; this is the
Saxifragales of D. Soltis & Soltis, 1997). The sax-
ifragoid clade also is united by base substitutions
and represents one of the most strongly supported
clades resulting from the phylogenetic analyses.
Additional examples of potentially informative
indels can be found at lower taxonomic levels. For
example, in Zingiberales, two insertions, each of a
single base pair, are found at positions 117 and 260
in all members of the Zingiberaceae (Kress et al.,
1995). None of the other approximately 70 mono-
cots for which 18S rDNA has been sequenced ex-
hibits these insertions. Similarly, an insertion of one
at NARE 655 appears to unite members of
Vise:
ом additional 185 rDNA sequencing
will reveal more examples of phylogenetically in-
formative indels. However, the discovery of such
indels depends on the availability of a large data-
base of accurate sequences. Previously published
sequences containing errors and ambiguities,
“false” indels, and incompletely sequenced regions
have made assessment of the phylogenetic potential
of indels in 185 rDNA impossible.
STEM VERSUS LOOP CHANGES/COMPENSATORY CHANGES
The secondary structure of the 185 rRNA tran-
script may have significant implications for phylog-
eny reconstruction using rRNA or rDNA sequences
(e.g., Mishler et al., 1988; Dixon & Hillis, 1993).
The question remains as to whether both loop bases
(non-pairing bases) and stem bases (pairing bases)
should be used in phylogeny reconstruction and, if
so, whether bases from stems and loops should be
considered equally informative and independent.
Assuming near-perfect compensatory mutation
(substitutions that maintain or restore stem base
omplementarity—e.g., Noller, 1984; Curtiss &
Vournakis, 1984; Wheeler & Honeycutt, 1988) in
stem regions to maintain secondary structure of the
18S or 26S (28S) rRNA, Wheeler and Honeycutt
(1988) recommended that stem bases either be
eliminated from phylogenetic analysis or weighted
by one-half relative to loop bases. However, in their
analyses of 28S rRNA sequences from vertebrates,
Dixon and Hillis (1993) found that characters from
both stems and loops contain phylogenetic infor-
mation. They also found that the number of com-
pensatory mutations in stem bases was less than
40% of that expected under a hypothesis of perfect
compensation to maintain secondary structure. Dix-
on and Hillis therefore suggested that the weighting
of stem characters be reduced by no more than
20% relative to loop characters in phylogenetic
analyses.
The large database of 18S rDNA sequences re-
ported here affords the opportunity to address these
and other issues regarding the impact of the sec-
ondary structure of the 18S rRNA transcript on
phylogeny reconstruction in angiosperms. Although
it is not our goal to examine such issues in detail
here, we will provide some initial observations re-
garding the relative importance of both stem and
loop mutations and the prevalence of compensatory
mutations.
We followed the definitions of stem and loop bas-
es used elsewhere (e.g., Dixon & Hillis, 1993):
stem bases are those that participate in base-pair-
ing interactions; loop bases do not engage in base
pairing in the mature rRNA. Mapping base substi-
tutions on the proposed 18S rRNA secondary struc-
Annals of the
Missouri Botanical Garden
ture for Glycine max (see Appendix), we examined
120 positions (in 60 taxa) at which өч
informative base substitutions had осс ased
on the results of the searches; dada ым.
Emphasis was placed on those substitutions that
provided synapomorphies for those clades that ap-
pear in all shortest trees and that represent several
different taxonomic levels (e.g., Asteridae s.l., Car-
yophyllidae s.l., monocots, glucosinolates, santa-
loids, Caryophyllales, saxifragoids, celastroids,
were stem bases, and 50 (42%) were loop
bases. Although this initial survey considers only a
subset of synapomorphous base substitutions, it
suggests that both stem and loop regions appear to
contain phylogenetic information, with perhaps a
omewhat greater proportion of informative sites
found in stem, rather than loop, regions. This topic
certainly requires a more rigorous examination. The
relative information content of stem versus loop
bases may, in fact, vary at different taxonomic lev-
els. For example, some of the more variable loop
regions (several of which were removed from these
hylogenetic analyses because the sequences were
difficult to align) may hold relatively more infor-
mation at lower levels (among and within closely
related families) than at higher taxonomic levels
(ordinal and above), at which the sequences be-
come too divergent for confident alignment.
The frequency of compensatory changes was ex-
amined in 21 stem regions located throughout the
18S rRNA gene. Following the general approach of
others (e.g., Dixon & Hillis, 1993), we considered
two classes of substitutions within stem regions.
The first class involves substitutions that change
one pair of complementary bases to another pair of
complementary bases. This includes “double com-
pensatory” substitutions in which one pair of com-
plementary bases is converted to another (e.g., C-G
to A-U). This class also includes changes that re-
quire only a single substitution event. That is, be-
cause uracil can pair with guanine as well as with
adenine, it is possible to have a single change from
one base-pairing couplet to another (e.g., U-G to
C-G; U-A to U-G). The latter represent one type of
“single compensatory” substitutions (sensu Dixon
& Hillis, 1993). The second class of stem substi-
tutions involves those that change one pair of com-
plementary bases to a pair of noncomplementary
bases, or vice versa (e.g., C-G to G-G; or C-C to
C-G). For example, a change of C-G to G-G de-
stroys a base-pairing couplet. Conversely, a change
from C-C to C-G creates a base-pairing couplet and
represents another example of a “single compen-
satory” substitution. Of the 216 stem changes we
analyzed, 19% were “double compensatory”; 46%
were single base substitutions involving uracil that
changed one base-pairing couplet to another (“sin-
gle compensatory”); 8% changed a pair of noncom-
stroyed a base-pairing couplet. Of these changes
that result in mispairing of nucleotides, over one-
third are adjacent to loop regions. Hence, the loop
regions may simply be expanded in these instances.
Nearly three quarters (73%) of the stem mutations
we analyzed maintain or restore base pairing and
would be considered compensatory. In their com-
parable analysis of 28 rRNA sequences, Dixon and
Hillis (1993) observed that only 47% of the muta-
tions maintained or restored base pairing. Our re-
sults are more similar to observations for 5S rRNA
(Curtiss & Vournakis, where approximately
88% of the base siltation analyzed from stem
regions were compensatory.
A similar pattern of molecular evolution is seen
for 18S rDNA within a single family, Polemoni-
aceae, and its closest relatives (Johnson et al., un-
published), where 228 variable nucleotide positions
were examined. Although most of these 228 posi-
tions are located on stems (133 compared to 95
loop characters), the average number of substitu-
tions per site over the potentially informative char-
acters is greater for the loop characters (5.0) than
for the stem characters (3.1). Using one of the most
arsimonious trees as a framework, Johnson et al.
(unpublished) also considered in more detail 67
substitutions that either unite or appear within Po-
lemoniaceae. Of these substitutions, 36 (53.7%) oc-
cur in loops. Considering just the 31 stem substi-
оно ee (74.2%) either maintain or restore
base- - The remaining eight stem substitu-
tions (es 8%) result in mispairing of nucleotides,
with four sites located adjacent to loops
These initial studies of the relative information
content of stem and loop regions and the frequency
of compensatory changes have implications for the
use and relative weighting of stem and loop bases
in phylogeny reconstruction. These data reinforce
the findings of others (e.g., Dixon & Hillis, 1993;
Smith, 1989) that both stem and loop regions pro-
vide important information for phylogeny recon-
rDNA suggests that perhaps stem characters should
receive less weight than loop characters in future
analyses. However, weighting of stem versus loop
е is more complex than it might seem ini-
tially. Recent work with 18S rDNA sequences in
оннан (Johnson еї al., unpublished) dem-
Volume 84, Number 1
1997
Soltis et al. 19
18S Ribosomal DNA Phylogeny
onstrates that loop regions evolve more rapidly than
do stem regions. Thus, in more focused studies in
which it is possible to align and use the entire 18S
rDNA sequence, stem and loop regions should per-
haps be given equal weight. In broader studies in
which the rapidly evolving loops are removed due
to alignment difficulties and only the more con-
served loop regions are included in the analysis,
stems should be downweighted; however, more de-
tailed analyses are required to estimate appropriate
weights.
II. PHYLOGENETIC RELATIONSHIPS
Each broad phylogenetic analysis yielded
thousands of most parsimonious trees; it is likely
that shorter trees exist for all four data sets and
that additional classes of most-parsimonious trees
were not recovered. Nonetheless, we feel that it is
significant that analyses of three of the four data
sets suggest the same general topology. The shortest
trees obtained from searches of data sets 1 and 2
are essentially identical, and differences between
the shortest trees from analysis of these two data
sets and data set 4 are minor and weakly supported.
Although phylogenetic analysis of data set 3 re-
vealed many of the same major clades recovered
by searches of the other data sets, relationships
among some of these clades differ; most notable are
the weakly supported, unusual positions of mono-
cots and saxifragoids (see below)
All searches revealed the same major clades
(e.g., Rosidae, Asteridae s.l., Caryophyllidae s.l.,
monocots, saxifragoids), as well as the same suite
of taxa as sister to all remaining angiosperms. In
general, the trees obtained in these exploratory,
broad analyses of 18S rDNA sequences depict re-
lationships very similar to those obtained in broad
analyses of rbcL sequences (Chase et al., 1993).
The general features observed in the shortest trees
obtained from the four searches are discussed be-
low. The several unusual relationships among major
clades suggested by analyses of data set 3 are dis-
cussed in more detail below under “Differences
Among the Shortest Trees.”
FIRST-BRANCHING FAMILIES
Phylogenetic analyses of three of four data sets
(data sets 1, 2, and 4) suggest that those taxa having
uniaperturate pollen (monosulcate and monosul-
cate-derived) ји ethereal oils appear as early-
branching ап rms, forming a grade (labeled
monosulcate Hera in Figs. 1, 2, and 4). Those
plants having triaperturate pollen (tricolpate and
tricolpate-derived), with a few exceptions (see be-
low), and tannins and alkaloids as secondary com-
pounds form a clade (labeled eudicot clade). The
latter group has been referred to as the eudicots
(Donoghue & Doyle, 1989b; Doyle & Hotton, 1991;
Chase et al., 1993). Although the term eudicot has
been variously defined, we will use the Chase et al.
(1993) definition to facilitate comparison between
the two studies. A eudicot clade was also recovered
in analyses of rbcL sequences (Chase et al., 1993),
but, instead of forming a grade, as they do here,
those plants with uniaperturate pollen form a weak-
ly supported clade in the rbcL trees
There are two major exceptions to the general
correspondence between the eudicot clade and the
distribution of the triaperturate pollen types (other
an the obvious departures observed in the trees
derived from analysis of data set 3). First, the Win-
teraceae and several families of paleoherbs (Chlo-
ranthaceae, Lactoridaceae, Aristolochiaceae) all
possess uniaperturate pollen, yet appear within the
eudicot (triaperturate) clade in the shortest trees
obtained in analyses of data sets 1 and 2 (Figs. 1,
2). These exceptions may reflect low taxon density
and/or the low resolving power of 18S rDNA se-
quence data (see below); these taxa seem to be un-
stable in position in the various searches. In broad
analyses of rbcL sequences, in contrast, these taxa
are clearly members of the uniaperturate clade.
second exception involves Illiciaceae and
Schisandraceae. Unlike the examples above, how-
ever, which we suspect represent spurious phylo-
genetic placements, Illiciaceae and Schisandraceae
appear to be true early-branching angiosperms (see
below), yet possess triaperturate pollen. These fam-
ilies similarly appear as early-branching angio-
sperms in analyses based on rbcL sequences (Chase
et al, 1993; Qiu et aL, 1993). As reviewed by
Doyle et al. (1990), however, the tricolpate condi-
tion in Illiciaceae and Schisandraceae is different
from that which characterizes eudicots. Hence, the
18S rDNA analyses further support the rbcL-based
inferences of Qiu et al. (1993) that Illiciaceae and
Schisandraceae represent an independent evolution
of tricolpate pollen
Four families of ecd Magnoliidae consistently
appear as sister taxa to all remaining angios
analyzed: Amborellaceae and a clade of Austro-
baileyaceae, Illiciaceae, and Schisandraceae. The
latter three families form one of the most strongly
supported clades in this study (jackknife value of
94%). In searches of data sets 1 and 2, a clade of
Austrobaileyaceae, Illiciaceae, and Schisandraceae
is the sister group to all other angiosperms, fol-
lowed subsequently by Amborellaceae; in analyses
of data sets 3 and 4, the positions of these two
20
Annals of the
Missouri Botanical Garden
lineages are reversed. Surprisingly, given the close
relationship suggested between Illiciaceae and
Schisandraceae by others (e.g., Cronquist, 1981;
Qiu et al., 1993), /llicium is sister to Austrobaileya-
Schisandra in all four analyses. These four genera,
with Nymphaeales, form a clade in the rbcL anal-
yses of Chase et al. (1993) and Qiu et al. (1993).
In the shortest trees resulting from searches of
all four data sets, from one to several families of
paleoherbs (sensu Donoghue & Doyle, 1989a) sub-
sequently follow Austrobaileyaceae, Illiciaceae,
Schisandraceae, and Amborellaceae. Nymphae-
aceae immediately follow these four families in all
shortest trees. In searches of data sets 1 and 2,
Nymphaeaceae form a clade with Piperaceae and
Saururaceae (represented by Peperomia and Hout-
tuynia and Saururus, respectively; jackknife value
of 85%), whereas in searches of data set 4, these
same three families form a grade with Nymphae-
aceae as sister to all remaining angiosperms, fol-
lowed by a clade of Piperaceae and Saururaceae.
In searches of data set 3, Nymphaeaceae also follow
Austrobaileyaceae, Illiciaceae, Schisandraceae,
orellaceae, but Nymphaeaceae are then
followed by saxifragoids, an unusual placement dis-
cussed in more detail below
Amborellaceae, followed by (1) a clade of Aus-
trobaileyaceae, Illiciaceae, and Schisandraceae, (2)
Nymphaeaceae, (3) a clade or grade of Piperaceae,
Saururaceae, Aristolochiaceae, and Lactoridaceae
(similar to Fig. 4), appear as the first-branching an-
giosperms when Zamia and Cycas are used as ad-
ditional outgroups (see Materials and Methods). In
preliminary analyses of a larger data set of 271
angiosperms using species of Welwitschia, Gnetum,
and Ephedra as reet о ое а clade
of A
and Nymphaeaceae again е as successive sis-
ters to all remaining angiosperm
The position of woody анлау as first-branch-
ing taxa in these 18S rDNA trees is in general
agreement with traditional views of angiosperm re-
lationships (e.g., Cronquist, 1968, 1981; Stebbins,
1974; Takhtajan, 1969, 1980) that suggest that
woody Magnoliidae are the most primitive extant
angiosperms. The morphological analyses of Don-
oghue and Doyle (1989a) and Loconte and Steven-
son (1991) also support the woody Magnoliidae as
the most ancestral living group of angiosperms.
Other data also point to the antiquity of at least
some of these genera. For example, Endress and
Honegger (1980) determined that the pollen of Aus-
trobaileya resembles Clavatipollenites, one of the
oldest probable angiosperm fossils, and concluded
that Austrobaileya may be “especially archaic
among the angiosperms.” If the 18S rDNA infer-
ence is correct in suggesting that Amborellaceae, a
family lacking vessel elements, are among the first-
branching angiosperms, this амын may support
the hypothesis that the angiosperm re primi-
tively vesselless (Bailey, 1957; eui 1981;
Young,
In contrast = this study, analyses based on par-
tial 18S and 2 sequences suggest sted that
chiales, Piperales,
Nymphaeales) is the sister taxon to all other flow-
ering plants (Hamby & Zimmer, 1992). However,
of the four woody families of Magnoliidae appearing
as first-branching taxa in our 18S rDNA trees (Am-
borellaceae, Schisandraceae, Illiciaceae, and Aus-
trobaileyaceae), only Illiciaceae were sampled by
amby and Zimmer (1992). Other phylogenetic
analyses similarly support the position of some pa-
leoherbs as first-branching taxa among the angio-
sperms (e.g., Doyle et al., 1994; Nixon et al., 1994).
Paleoherbs are sister to other angiosperms in trees
based on a combination of morphology and rRNA
sequence data and in those derived independently
from morphological and rRNA data (Doyle et al.,
1994). However, this topology is only weakly sup-
ported by morphological data, with trees rooted
next to Magnoliales only one step longer. Further-
more, the rRNA data set employed by Doyle et al.
(1994) is that of Hamby and Zimmer (1992), which,
as noted above, lacked several critical woody mag-
noliids.
Broad phylogenetic analyses of rbcL sequences
(Chase et al., 1993; Rice et al., 1997) place the
aquatic genus Ceratophyllum as sister to all re-
maining angiosperms. This placement of Cerato-
phyllum also has been suggested on morphological
grounds (Les, 1988; Les et al., 1991; Nixon et al.,
1994), although alternative trees in the latter study
place the paleoherb family Chloranthaceae as sister
to the remaining flowering plants. A number of flo-
ral features of Ceratophyllum also conform to the
view that the genus represents a primitive angio-
sperm (Endress, 1994). However, Ceratophyllum
does not appear as first-branching in any of our
phylogenetic analyses. Searches involving three of
the four data sets (1, 2, and 4) place Ceratophyllum
as sister to the monocots, a finding in general agree-
ment with earlier rRNA sequence analyses (Hamby
& Zimmer, 1992).
Subsequent to the Amborellaceae, Austrobailey-
aceae, Illiciaceae, Schisandraceae, Nymphaeaceae,
Piperales, in analyses of three of four data sets
(1, 2, and 4) are additional families and orders of
Magnoliidae: Annonaceae, Calycanthaceae, and
Volume 84, Number 1
Soltis 21
18S ce DNA Phylogeny
Lauraceae, all woody families traditionally consid-
ered among the most primitive extant angiosperms.
With the exception of the shortest trees resulting
from analysis of data set 3, the monocots also ap-
pear as an early lineage of angiosperms. The mono-
cots are monophyletic, with the exception of Acorus,
which does not appear closely related to the other
member of Araceae included (Calla). In analyses
of the two data sets (1 and 2) that included Acorus.
Acorus follows Nymphaeaceae—Piperales as the
subsequent sister to all remaining angiosperms. In
analyses of rbcL sequences, Acorus was considered
“phylogenetically isolated" as sister to the remain-
ing monocots (Duvall et al., 1993). Phylogenetic re-
sults based on 185 rDNA sequences also suggest
that Acorus is anomalous among monocots. Given
its long branch length and unexpected position, the
18S rDNA of Acorus should be resequenced, and
additional monocots should be added to the data
set before the affinities of this enigmatic genus are
addressed further.
Because our sampling of monocots was limited,
to permit more thorough treatment elsewhere, re-
lationships within the monocots will not be dis-
cussed here in any detail. Nonetheless, several tra-
ditionally recognized groups of monocots appear to
be monophyletic, including Zingiberales, Liliales,
and higher commelinoids. Furthermore, the bro-
meliads are grouped with the grasses and allies, as
expected (Duvall et al., 1993). The two best sup-
ported clades within the monocots are Zingiberales
(Maranta, Zingiber, Costus, Canna, Heliconia, and
Musa; jackknife value of 5896) and a clade com-
posed of Sparganiaceae, Cyperaceae, Poaceae, and
Bromeliaceae (Sparganium, Cyperus, О! Zea,
and Glomeropitcairnia; jackknife value of 59%).
Surprising results, given rbcL topologies (Duvall et
al., 1993) and morphological features, include the
placement of Orchidaceae (Oncidium) as the sister,
or one of the sisters, to the remaining monocots and
the placement. of Arecaceae (Veitchia) and Alis-
ia) within the Asparagales (Figs.
1, 2). These unusual placements should not be con-
sidered seriously, however, due to the low repre-
sentation of the monocots.
EUDICOT CLADE
Analyses of three of four data sets (1, 2, and 4)
clearly reveal a eudicot (or triaperturate) clade
(Figs. 1, 2, 4), with the following successive sister
groups at its base (Figs. 1, 2): Proteaceae, Nelum-
eae, Р! lade of ranunculids,
Trochodendraceae/Tetracentraceae, and a clade
composed of Winteraceae (Drimys), Aristolochi-
aceae (Aristolochia, Asarum, Saruma), Lactorida-
ceae, Sabiaceae, and Chloranthaceae (Hedyosmum).
The latter clade is an unexpected grouping (see be-
low) of paleoherbs (Aristolochiaceae, Lactorida-
ceae, Chloranthaceae), woody Magnoliales (Winter-
aceae), and eudicots (Sabiaceae). With the
exception of Aristolochiaceae, Lactoridaceae, Chlo-
ranthaceae, and Winteraceae, the presence of the
remaining taxa on branches at the base of the eu-
dicots closely parallels results retrieved from the
phylogenetic analyses of rbcL sequences (Chase et
In the shortest trees obtained in analyses of data
set 4, the distinction between the monosulcate
grade and the eudicot clade is less clear than in
the shortest trees obtained from the analyses of data
sets 1 and 2 (see “Lower Eudicots/Monosulcates,"
Fig. 4). Platanaceae, Trochodendraceae/Tetracen-
traceae, ranunculids (which are paraphyletic), Pro-
teaceae (Knightia), Buxaceae, Sabiaceae, and a
clade of Chloranthaceae (Hedyosmum)/Winteraceae
(Drimys) again appear as sister groups to the re-
mainder of the eudicot clade. Also in this same
lower eudicot/monosulcate grade, however, are Cal-
ycanthaceae, Annonaceae (Mkilua), and Lauraceae
(Sassafras), uniaperturate families pe appear in a
some ranunculids (Fig
In the shortest trees Ny in petam of all
data sets, the remainder of the eudicot clade is es-
sentially composed of two large subclades, one con-
sisting largely of Rosidae plus some Dilleniidae
and the other corresponding to the Asteridae s.l.
(labeled Rosidae and Asteridae s.l., respectively, in
Figs. 1, 2, 4). With a few exceptions, most notably
the placement of the monocots within the Rosidae
clade, these two large clades also are present in the
trees derived from searches of data set 3. The Ros-
idae and Asteridae s.l. clades were also recovered
in broad analyses of rbcL sequences (Chase et al.,
1993; Olmstead et al., 1992, 1993; Rice et al., sub-
mitted), although the placement of Caryophyllidae
s.l. is very different in the 185 rDNA and rbcL
topologies (see below). These two large clades, Ros-
idae and Asteridae s.l., reflect the basic division of
higher dicots into two major groups (Young & Wat-
son, 1970), with (1) polypetalous corollas and non-
tenuinucellate ovules and (2) sympetalous corollas
and tenuinucellate ovules, respectively. Below we
discuss in more detail the major clades of eudicots
based on phylogenetic analyses of 18S rDNA se-
quences. We use informal names in most instances
to refer to strongly supported clades (e.g., celas-
troids, saxifragoids, ranunculids), some of which
differ dramatically from traditional views of rela-
Annals of the
Missouri Botanical Garden
tionship, but formal taxonomic change may be war-
ranted for many of these.
Ranunculids. The searches of the eS data
sets (Figs. 1, 2) recovered a clade (labeled “Ran-
unculids” " containing Lardizabalaceae, ee
ceae, Ranunculaceae, Menispermaceae, Euptele-
aceae, Fumariaceae, Sargentodoxaceae, and
Papaveraceae. This same clade was found in the
broad analyses of rbcL sequences (Chase et al.,
1993); it represents the core of the Ranunculales
(sensu Cronquist, 1981) and corresponds closely to
the Berberidales of Thorne (1992) and the Ran-
unculiflorae of Dahlgren (1980). Analyses of not
only rbcL and 18S rDNA sequences, but also atpB
sequences, place Eupteleaceae (Hamamelidae)
within this clade (Hoot & Crane, 1995). Also part
of this clade in the 18S rDNA analyses is Sargen-
todoxaceae, a family typically placed in Ranuncu-
lales and allied with Lardizabalaceae (e.g., Cron-
quist, 1981). In contrast, analyses of rbcL
sequences placed Sargentodoxaceae with Fabaceae
(Chase et al., 1993). This result is due to the mis-
identification of leaf material in the rbcL analysis
(Qiu, pers. comm.). Reanalysis of Sargentodoxa for
rbcL places it as sister to the Lardizabalaceae (Hoot
& Crane, 1995; Hoot et al., 1995).
Searches involving the two small data sets (3 and
4) employed fewer representatives of Ranunculales.
In trees resulting from searches of data set 4, these
taxa form a grade as some of the early-branching
eudicots. In trees from searches of data set 3, in
contrast, the ranunculids appear polyphyletic. The
placements of the ranunculids in analyses of data
sets 3 and 4 may well reflect their decreased rep-
resentation (lower taxon density) in these searches.
In preliminary analyses of a 271-taxon 185 rDNA
data set including more ranunculids, the ranuncu-
lids again form a monophyletic group.
Saxifragoids. All analyses of 185 rDNA se-
quences (Figs. 1-4) reveal a clade composed of
Heuchera, Boykinia, Saxifraga (Saxifragaceae s.
str.), Crassula, Sedum, Dudleya, and Kalanchoe
(Crassulaceae), Pterostemon, Tetracarpaea, Ribes,
and ltea (Grossulariaceae), Penthorum (placed in
Saxifragaceae by Cronquist, 1981), Altingia and
Liquidambar (Hamamelidaceae), н акни
Cercidiphyllaceae, Daphniphyllaceae, and Раеоп
aceae; this clade is referred to here as йар
This clade is one of the most strongly supported
findings of this investigation (jackknife value of
68%). The same saxifragoid clade (also referred to
as Saxifragales) was identified in an analysis of 130
18S rDNA sequences aimed at elucidating the af-
finities of the morphologically diverse members of
Saxifragaceae s.l. (D. Soltis & Soltis, 1997). The
monophyly of this clade is supported not only by
base substitutions, but also by the presence of an
insertion (see Table 2) located in a portion of the
18S rRNA gene that is highly conserved in length.
An identical clade (referred to as rosid III) is re-
vealed in the 499-taxon analysis of rbcL sequences
(Chase et al., 1993) and is also retrieved in prelim-
inary analyses of a 271-taxon 18S rDNA data set
including more Hamamelidaceae, as well as in phy-
logenetic analyses involving matK sequences
ri Rl р & Soltis, unpublished) and prelimi-
па with atpB sequences (Hoot,
и As reviewed in more detail by D.
Soltis and Soltis (1997), this small clade is note-
worthy in that it contains taxa traditionally placed
in three subclasses: Paeoniaceae (Dilleniidae); Ha-
mamelidaceae, Daphniphyllaceae, Cercidiphylla-
ceae (Hamamelidae); the remaining taxa are all
members of Rosidae.
Although this saxifragoid clade is recovered by
analyses of both 18S rDNA and rbcL sequences,
this same group of taxa has never been recognized
in any classification. Whereas Saxifragaceae s. str.,
Ribes, Itea, Tetracarpaea, Pterostemon, Penthorw
and Crassulaceae are considered closely related
members of Rosidae in virtually all recent treat-
ments (e.g., Cronquist, 1981; Thorne, 1992; Takh-
tajan, 1987; Dahlgren, 1980, 1983), the affinities
of the rosid family Haloragaceae and the dilleniid
family Paeoniaceae have been considered enigmat-
ic (e.g., Cronquist, 1981). The hamamelid families
found in this clade (Hamamelidaceae, Cercidiphyl-
laceae, and Daphniphyllaceae) typically have not
been considered close relatives of Saxifragaceae s.
str. and allied rosids. The relationships of these
more anomalous members of this clade are dis-
cussed in more detail by D. Soltis and Soltis (1977).
Glucosinolate clade. Another clade revealed by
all analyses (Figs. 1—4) comprises glucosinolate-
producing taxa. The families that compose this
clade in Figures 14 are 7 of the 15 families known
to produce glucosinolates (mustard oil glucosides):
Limnanthaceae, Brassicaceae, Cap eae, Mor-
ingaceae, Сапсасеае, Bataceae, and Tropaeola-
ceae. Whereas Brassicaceae and Capparaceae have
long been as closely related, the re-
maining families included in this study (Limnan-
thaceae, Moringaceae, Caricaceae, Bataceae, and
Tropaeolaceae) are morphologically diverse and
have been placed in distinct orders (e.g., Cronquist,
1981; see review by Rodman et al., 1993). The ge-
nus Drypetes (Euphorbiaceae) also produces glu-
cosinolates, but it does not appear to be closely
Volume 84, Number 1
1997
Soltis et al. 23
18S Ribosomal DNA Phylogeny
related to the glucosinolate clade in any of the four
searches. Phylogenetic analyses of 18S rDNA se-
quences involving additional glucosinolate taxa fur-
ther demonstrate the monophyly of the glucosino-
late-producers, with the exception of Drypetes, and
also clarify relationships among the members of
this clade (Rodman et al., submitted). These results
closely parallel findings based on the phylogenetic
analysis of rbcL sequences (Rodman et al., 1993;
Chase et al., 1993) and morphology (Rodman,
1991). Thus, both rbcL and 18S rDNA sequence
data indicate that there were two independent ori-
gins of the mustard oil syndrome (see Rodman et
al., 1993, submitted).
Nitrogen-fixing clade. Species of only 10 fam-
ilies of angiosperms are known to form symbiotic
associations with nitrogen-fixing bacteria in root
nodules (Fabaceae, Betulaceae, Casuarinaceae,
Coriariaceae, Datiscaceae, Elaeagnaceae, Myrica-
ceae, Rhamnaceae, Rosaceae, and uU
These families are distributed among four of Cron-
quist's (1981) six subclasses of dicotyledons, im-
plying that many of these families are only distantly
related. Recent phylogenetic analyses of rbcL se-
quences reveal, е: that representatives of all
ten of these families occur together in a single
clade (“nitrogen- -fixing clade": Soltis et al., 1995).
ciations with nitrogen-fixing bacteria, including
Moraceae, Urticaceae, Polygalaceae,
Fagaceae, and С i i
Analyses of three of four 18S rDNA data sets
(Figs. 1, 2, 4) suggest an alliance of taxa similar to
that revealed by rbcL sequences. This clade in
large represents a subset of the taxa present
in the rbcL-based nitrogen-fixing clade. The fami-
lies in the 18S rDNA-based nitrogen-fixing clade
include Betulaceae, Cas
to be part of this alliance based on analyses of rbcL
sequences. However, neither Rosaceae nor Faba-
ceae, two families involved in nitrogen-fixing sym-
bioses, are included within the 18S rDNA nitrogen-
fixing clade, although both families are part of this
alliance in the rbcL-based trees (Soltis et al., 1995).
Searches involving the two larger data sets (1
and 2) also place three families of Malvales (Mal-
vaceae, Bombacaceae, and Tiliaceae) within the ni-
trogen-fixing clade; these taxa were not part of the
nitrogen-fixing clade in the rbcL-based trees. In
analyses of data set 4, however, these three families
of Malvales are not part of the nitrogen-fixing clade
(Fig. 4). No clear nitrogen-fixing clade emerged in
analyses of data set 3; instead, these taxa are
of a grade that represents the first branches of a
primarily rosid-dilleniid clade (Fig. 3).
Asteridae sensu lato. Analyses of all four 185
rDNA data sets also reveal an expanded Asteridae
clade (Asteridae s.l.) that agrees closely with that
recovered by analyses of rbcL sequences (Olmstead
et al., 1992, 1993; Chase et al., 1993). In addition
to the conventionally cireumscribed Asteridae, this
clade also includes a number of families placed in
Dilleniidae, such as Ericaceae, Clethraceae, Pyro-
laceae, Styracaceae, Ebenaceae, Actinidiaceae,
Sarraceniaceae, Fouquieriaceae, Theaceae, and
imulaceae. Also present in Asteridae s.l. are Nys-
saceae, Pittosporaceae, Apiaceae, Araliaceae, and
Hydrangeaceae, all members of Rosidae. In addi-
tion, Eucommiaceae, a member of Hamamelidae,
and Byblis, a genus of carnivorous plants usually
placed in Rosidae, also appear within Asteridae s.l.
All analyses also place an expanded Caryophylli-
dae (Caryophyllidae s.l.) within the Asteridae s.l.
clade, an unexpected result that is discussed in
more detail below.
Within Asteridae s.l, several subclades or
grades can be identified agree, in > part,
with some of the groups identified in analyses of
rbcL sequences (Chase et al., 1993; Olmstead et
al., 1993). Perhaps most noteworthy of these is the
ericalean grade (the asterid III clade of Chase et
al., 1993) observed in all of the shortest 185 rDNA
trees (Figs. 1—4). Other clades of Olmstead et al.
observed to be monophyletic, including
reducta Boraginales, Gentianales, Asterales
s.l., and Lamiidae. Additional asterid taxa should
be sequenced for 18S rDNA to assess more rigor-
ously the monophyly of these groups and their in-
terrelationships.
Caryophyllidae sensu lato. All analyses of 18S
rDNA sequences reveal a clade composed of Мус-
taginaceae (Mirabilis), Chenopodiaceae (Spinacia),
Phytolaccaceae (Phytolacca
nia), and Molluginaceae (Mollugo).
ilies represent Caryophyllales (e.g., Cronquist,
1981), the monophyly of which is supported in this
study by a jackknife value of 58%, as well as by
numerous lines of and molecular
Annals of the
Missouri Botanical Garden
data (e.g., Rodman et al., 1984; Rettig et al., 1992).
Sister to this clade of Caryophyllales is another
strongly supported clade comprising Plumbagina-
ceae and Polygonaceae (jackknife value of 77%):
this group collectively represents Caryophyllidae
(sensu Cronquist, 1981). The monophyly of Cary-
ophyllidae is only weakly supported by cladistic
analysis of morphological, chemical, anatomical,
and palynological features (Rodman et al., 1984).
Analyses of 18S rDNA sequences also suggest that
two families of carnivorous plants, Droseraceae and
Nepenthaceae, are sister to Caryophyllidae, and we
refer to this entire assemblage as Caryophyllidae
s.l. (Figs. 14).
Phylogenetic analyses of rbcL sequences simi-
larly recovered a Caryophyllidae s.l. clade com-
posed of Caryophyllales, Polygonaceae, Plumbagi-
naceae, Droseraceae, and Nepenthaceae (Chase et
al., 1993). One of the broad analyses of rbcL se-
quences (search A, Chase et al., 1993) placed Vi-
taceae and Dilleniaceae with this expanded Cary-
ophyllidae clade. In the analyses of 18S rDNA
sequences, Vitaceae were not sampled, and Dille-
niaceae are well removed from Caryophyllidae s.l.
The anomalous placement of Dilleniaceae near the
monocots (Figs. 1, 2) is discussed below.
Santaloids. Analyses of all four data sets reveal
a monophyletic santaloid clade or Santalales, which
are represented here by only three families (Opi-
liaceae, Santalaceae, and Viscaceae). However, in
preliminary analyses in which Santalales are rep-
resented by seven families (Opiliaceae, Santala-
ceae, Viscaceae, Eremolepidaceae, Misodendra-
ceae, Loranthaceae, and Olacaceae), santaloids
again form a clade. These seven families are widely
considered to form a natural group based on mor-
phology (e.g., Cronquist, 1981) and have been
shown to form a clade in previous, smaller analyses
of 18S rDNA sequences (Nickrent & Franchina,
1990; Nickrent & Soltis, 1995).
Although santaloids appear monophyletic, the
position of this clade varies among the analyses. In
analyses of data sets 1 and 2, santaloids are sister
to Polygala and closely related to the legumes.
Analysis of data set 4 again places santaloids with
Polygala and a legume (Pisum), as well as with
Gunnera. Analysis of data set 3 results in an un-
usual placement of santaloids with several paleo-
herbs. These findings parallel those of Chase et al.
(1993) based on rbcL sequences in which the po-
sition of santaloids differed greatly between the
476- and 499-taxon searches. In the former, san-
taloids and Gunnera form the asterid V clade; in
the latter, santaloids are sister to the caryophyllids,
but again appear near Gunnera. Thus, whereas both
rbcL and 18S rDNA searehes occasionally place
santaloids near Gunnera, analyses of three of the
four 18S rDNA data sets place santaloids close to
Fabaceae and Polygalaceae.
Celastroids. Another small clade revealed in all
analyses consists of Lepuropetalon and Parnassia
(Saxifragaceae s.l.), Brexia (Grossulariaceae), and
uonymus (Celastraceae). This clade, labeled ce-
lastroids (Figs. 1—4), was also recovered in analyses
of rbcL sequences (Chase et al., 1993; Morgan &
Soltis, 1993). Although this initially appears to be
an eclectic assemblage (Brexia is a genus of small
trees; Lepuropetalon spathulatum is the smallest
terrestrial angiosperm), embryological and morpho-
logical data also unite these taxa (reviewed in Mor-
gan & Soltis, 1993). The celastroid clade consists
of two pairs of genera, each of which is strongly
supported: Lepuropetalon—Parnassia (jackknife =
100%) and Brexia-Euonymus (jackknife = 67%).
These same two pairs of genera also were revealed
in analyses of rbcL sequences (Chase et al., 1993;
Morgan & Soltis, 1993).
Cunonioids. Bauera and Ceratopetalum (Cu-
noniaceae) and Eucryphia (Eucryphiaceae) form a
clade with a jackknife value of 5396. A close re-
lationship among these genera also was revealed by
a cladistic analysis of morphological features (Huf-
ford & Dickison, 1992). Bauera, Ceratopetalum,
and Eucryphia constitute the core of a very well
supported clade (jackknife value of 8996) labeled
cunonioids (Figs. 1—4) that also contains Cephalo-
taceae, a family of carnivorous plants, and Sloanea
(Elaeocarpaceae). A close relationship of Cephal-
otaceae to these same representatives of Cunoni-
aceae and Eucryphiaceae also is suggested by anal-
yses of rbcL sequences (Chase et al., 1993; Morgan
& Soltis, 1993). Sloanea was not represented in the
broad analyses of rbcL sequences. Other taxa that
appear closely allied with Cunoniaceae, Eucryphi-
aceae, and Cephalotaceae in rbcL analyses include
Tremandraceae and Oxalidaceae; these families
were not included, however, in the 18S rDNA anal-
yses.
Other noteworthy relationships. As recently re-
viewed (Qiu et al., 1993), the placement of Lacto-
ridaceae has been controversial, with relationships
to Magnoliales, Laurales, and Piperales all pro-
posed. Analyses of rbcL sequences suggested a
close relationship of Lactoridaceae to Aristolochi-
aceae (Chase et al., 1993), and analyses of 185
rDNA sequences similarly suggest that these two
Volume 84, Number 1
1997
Soltis et al. 25
18S Ribosomal DNA Phylogeny
families are sisters (Figs. 1—4), an inference strong-
ly supported by a jackknife value of
Additional, small monophyletic groups also merit
brief discussion. Bombacaceae, Tiliaceae, and Mal-
vaceae (represented by Bombax, Luhea, and Gos-
sypium, respectively) form a strongly supported
clade (jackknife value of 78%) in all 18S rDNA
analyses, in agreement with both traditional treat-
ments (all are members of Malvales) and topologies
based on rbcL sequences. However, Sloanea
(Elaeocarpaceae—Malvales) does not appear with
Bombacaceae-Tiliaceae-Malvaceae in any of the
18S rDNA trees (Figs. 1—4). As noted above, this
malvoid clade sometimes is embedded within the
nitrogen-fixing clade (Figs. 1, 2), a placement at
odds with analyses based on rbcL sequences. This
unusual placement could be the result of insuffi-
cient taxon density in that many of the closest pu-
tative relatives of Malvales were not included here
(e.g., Anacardiaceae, ME Leitneri-
aceae, Sterculiaceae, Dipterocarpacea:
On a broader scale, all 185 rDNA relie sug-
gest that Hamamelidae comprise a number of phy-
logenetically distinct lineages. For example, Troch-
odendraceae, Tetracentraceae, and Platanaceae
appear near the base of the eudicots in trees de-
rived from searches of data sets 1, 2, and 4 (Figs.
1, 2, 4). Eupteleaceae also appear near the base of
the eudicots, but as part of the ranunculid clade.
Three traditional families of Hamamelidae, Hama-
melidaceae, Cercidiphyllaceae, and Daphniphylla-
ceae, are part of a well supported saxifragoid clade
(Figs. 1—4). Still other families of pcd
(1.е., Betulaceae, Urticaceae, Moraceae, and U
maceae) are part of the nitrogen-fixing clade, s
Eucommiaceae are d within the Asteridae s.l.
The pronounced polyphyly of Hamamelidae was
similarly revealed by analyses of rbcL sequences.
Both 185 rDNA and rbcL sequence data suggest
similar placements for representatives of this sub-
class.
Topologies based on 18S rDNA sequences also
reveal the polyphyly of subclass Dilleniidae. Taxa
attributed to Dilleniidae appear in several phylo-
genetically well separated clades. Paeoniaceae ap-
pear in the saxifragoid clade, Nepenthaceae and
Droseraceae appear in Caryophyllidae s.l., Cappar-
ales, Batales, and Violales appear in the glucosi-
nolate clade, and several orders (e.g., Violales,
Ebenales, Ericales, Diapensiales, Primulales, and
Theales) appear in Asteridae s.l. Other represen-
tatives of Dilleniidae (e.g., Turneraceae, Elaeocar-
paceae) are scattered throughout the large Rosidae
clade
DIFFERENCES AMONG THE SHORTEST TREES
The shortest trees resulting from analyses of data
sets 1 and 2 are essentially identical (Figs. 1, 2)
and in turn are very similar to those derived from
searches of data set 4 (Fig. 4). The most unusual
topology results from searches of data set 3 (Fig.
3). For example, the distinction between the mono-
sulcate grade and the eudicot clade does not occur
in the shortest trees from this analysis, with the
monocots part of a predominantly rosid assemblage
and saxifragoids appearing as one of the early-
branching lineages of angiosperms. The ranuncu-
lids are not monophyletic in trees from searches of
data set 3, with two genera (Hypecoum and Dicen-
tra) appearing as sister to the monocots and the
remaining ranunculids appearing as part of a clade
that occupies the unusual position of sister to As-
teridae s.l. (see Asteridae s.l. Plus, Fig. 3). How-
ever, some of our numerous searches of data set 3
found trees only one step longer than the shortest
trees that have a topology essentially identical to
that resulting from analysis of the other small data
et (4).
Although searches of data sets 1, 2, and 4 yield-
ed similar topologies, several weakly supported dif-
ferences also exist among the shortest trees found.
For example, in trees derived from analyses of data
sets 1 and 2, one group of paleoherbs (Aristolochi-
aceae, Lactoridaceae) appears within the eudicot
clade, rather than within the monosulcate grade, as
would be expected. In contrast, in trees derived
from the smaller data sets (3 and 4), Aristolochi-
aceae and Lactoridaceae appear within the mono-
sulcate grade, close to other families of paleoherbs
(e.g., Piperaceae, Saururaceae). In addition, the
аро trees obtained from analyses of data sets 1
and 2 show a more well defined break between the
monosulcate grade and lower eudicots than do trees
from data set 4 (compare Figs. 1, 2, and 4). For
example, trees resulting from analyses of data set
4 place the monosulcate families Calycanthaceae,
Annonaceae, and Lauraceae with Proteaceae (see
Fig. 4, Lower eudicots/monosulcates). These and
other differences may be the result of insufficient
taxon density in certain portions of the tree, incom-
plete analysis, or lack of signal (see Caveats below).
Ф
ANOMALOUS PLACEMENTS
Perhaps the most unusual consistent feature of
the 18S rDNA trees involves the placement of Car-
yophyllidae s.l. within Asteridae s.l. Although Car-
yophyllidae s.l. form a well supported clade, the
position of this clade within Asteridae s.l. is not
strongly supported. Some of the many searches con-
26
Annals of the
Missouri Botanical Garden
ducted retrieved trees only two steps longer than
the shortest trees obtained in which Caryophyllidae
s.l. are not part of Asteridae s.l., but appear instead
within the Rosidae clade. Furthermore, analyses of
130 dicot 18S rDNA sequences aimed at resolving
the relationships of Saxifragaceae s.l. did not place
the Caryophyllidae s.l. within Asteridae s.l., but in-
stead showed the caryophyllids to be embedded
within a rosid clade (D. Soltis & Soltis, 1997). Al-
though the placement of Caryophyllidae s.l. varies
in the broad analyses of rbcL sequences (Chase et
al., 1993), this clade does not appear closely relat-
ed to the asterids in any of the shortest trees ob-
tained. The 476-taxon analysis places Caryophyl-
lidae s.l. within a clade of rosids, whereas the
499-taxon analysis places them near the split be-
tween the clades of higher eudicots (i.e., Rosidae
and Asteridae s.1.).
Other anomalous placements include the posi-
tion in some analyses (Figs. 1, 2) of one group of
paleoherbs (Chloranthaceae, Aristolochiaceae, Lac-
toridaceae) plus Winteraceae of Magnoliales near
the base of the eudicot clade. These taxa often are
considered to represent early-branching or primi-
tive angiosperms in both analyses of rbcL sequenc-
es (Chase et al., 1993; Qiu et al., 1993) and recent
classification schemes (e.g., Cronquist, 1981;
Thorne, 1992; Takhtajan, 1987). Based on phylo-
genetic analyses of rbcL sequences (e.g., Chase et
al., 1993; Qiu et al., 1993), for example, Chloran-
thaceae, Aristolochiaceae, and Lactoridaceae are
part of the monosulcate clade. As noted above,
however, in some of our searches (see Figs. 3 and
4), Chloranthaceae, Aristolochiaceae, and Lactori-
daceae do appear closer to the base of the angio-
sperms with other monosulcate taxa.
Analyses of data set 4 recovered a clade con-
sisting of Sagittaria (Alismataceae) and Cuscuta
(Cuscutaceae), placed in the Rosidae clade. In
idae s.l., respectively, in agreement with traditional
views and with trees based on rbcL sequences
(Chase et al., 1993). The unusual relationship sug-
gested by searches of data set 4 likely results from
the more limited taxon sampling of this data set
(fewer monocots are included, for example, com-
pared to data sets 1 and 2) and long-branch attrac-
tion. Sagittaria and Cuscuta have very long branch-
es (e.g., 39 and 65 steps, respectively, in Fig. 2) in
all of the shortest trees obtained. In analyses of data
sets 1 and 2, the long branch of Cuscuta also seems
to affect the placement of Ipomoea (Convolvula-
ceae), with both appearing in Lamiales instead of
Solanales.
The three subfamilies of Fabaceae, Papilionoi-
deae, Mimosoideae, and Caesalpinioideae (repre-
sented by Pisum and Glycine; Albizia; Bauhinia,
respectively), although present in the same small
clade with several other families in Figures 1 and
2, do not form a monophyletic group in any of our
searches. Rather than representing a true case of
discordance between 18S rDNA and rbcL trees, this
likely represents either the lower limits of resolu-
tion of 18S rDNA sequences (see below) or retrieval
of only a small sample of all equally most parsi-
monious trees (i.e., the strict consensus of all short-
est trees, had they been found, would have led to
the collapse of this part of the tree). In support of
the former conclusion is the observation that more
focused phylogenetic studies of 18S sequences rep-
resenting only Rosidae, some of which swapped to
completion, similarly suggest a polyphyletic Faba-
ceae; bootstrap analyses indicate that these rela-
tionships are poorly supported, however (D. Soltis
& Soltis, 1997, unpublished).
The position of the monocot genus Acorus (Ara-
ceae) (Figs. 1, 2) also is unusual. Rather than ap-
pearing with the monocots, Acorus appears as an
early-branching angiosperm, as it did in a previous
analysis of 64 18S rDNA and rRNA sequences
(Nickrent & Soltis, 1995). Other anomalous place-
ments include the position of Dilleniaceae near the
monocots (Figs. 1 and 2) and the unexpected po-
sition of Oncidium (Orchidaceae) as a first-branch-
ing monocot.
Several taxa are noteworthy not only because
their phylogenetic positions are unusual, but also
because their phylogenetic position varies from
search to search. For example, the close relation-
ship of Gunnera to the monocots (Е igs. 1 and 2) is
unexpected, but it is not seen in the trees resulting
from analysis of data sets 3 and 4 where Gunnera
appears in a clade with Santalales, Polygalaceae,
and Pisum (Fabaceae). The relationship of Gunnera
also is uncertain in rbcL topologies, in which its
placement varies from being embedded within As-
teridae s.l. (the 476-taxon search) to sister group of
the higher dicots (the 499-taxon search).
COMPARISON WITH HAMBY AND ZIMMER (1992)
Hamby and Zimmer (1992) used partial 18S and
26S rRNA sequences to examine relationships
among land plants. Because their analyses involved
only 46 angiosperms, taxon sampling clearly differs
between that and the present study. Nonetheless,
brief comparison of the topologies resulting from
the two studies is instructive.
In most of the shortest trees obtained here (Figs.
Volume 84, Number 1
1997
Soltis et al. 27
18S Ribosomal DNA Phylogeny
1, 2, 4), as well as in the study of Hamby and
Zimmer (1992), Ceratophyllum is allied with the
monocots. Both studies also concur in suggesting
that Nymphaeaceae appear near the base of the an-
giosperm radiation. Nymphaeaceae are the sister
up to all other angiosperms in Hamby and Zim-
тегз (1992) shortest trees; however, Amborella-
ceae, Austrobaileyaceae, and Schisandraceae were
not included in that study. In all of our shortest
trees, Nymphaeaceae follow the latter three families
and Illiciaceae as the sister group to all remaining
flowering plants.
Another similarity between the shortest trees in
both studies is the placement of Drimys (Wintera-
ceae). Drimys occupies an unusual phylogenetic
position in trees presented by both Hamby and
Zimmer (1992) and Nickrent and Soltis (1995), ap-
pearing as sister to Glycine and Pisum (Fabaceae),
rather than as an early-branching angiosperm. Dri-
mys occupies an unusual position in trees derived
from the current analyses, as well, appearing among
the lower eudicots. In trees resulting from the anal-
ysis of data set 3, Drimys again appears with Pisum.
The 18S rDNA sequence of Drimys exhibits a num-
ber of substitutions not found in other magnoliids.
In an attempt to ascertain the relationships of Win-
teraceae, we sequenced two species of Drimys, D.
winteri and D. aromatica, and they have identical
sequences. More recently, another member of Win-
teraceae (Pseudowintera) has been sequenced for
18S rDNA (Hoot, unpublished); this sequence is
nearly identical to the sequences for Drimys. Add-
ing Pseudowintera to the analysis does not alter the
unusual position of Winteraceae (trees not shown).
The unusual phylogenetic relationships that exist
among the eudicots in the shortest trees of Hamby
and Zimmer (1992) probably derive from insuffi-
cient sampling in that study. The present analysis
with its greater representation of eudicots reveals
relationships much more in accord with recent clas-
sifications (e.g., Cronquist, 1981; Takhtajan, 1987)
and/or the rbcL topologies of Chase et al. (1993).
Thus, the present study suggests that many of the
highly unusual relationships seen in Hamby and
Zimmer are likely to reflect low taxon density rather
than an inherent inability of 185 rDNA sequences
to resolve relationships
CAVEATS
A number of limitations are inherent in any large
taxa, including uncertainty regarding maximum
parsimony, insufficient taxon sampling and/or den-
sity, the presence of “older,” erroneous 18S rDNA
sequences in the data matrix, and the overall lower
rate of evolution of 185 rDNA compared to rbcL.
We discuss these potential factors in more detail
below.
An analysis of this magnitude cannot be expect-
ed to achieve maximum parsimony in a reasonable
amount of time. It is likely that we did not find all
classes of most-parsimonious trees, despite a
search strategy (cf. Maddison et al., 1992) designed
to identify multiple islands (Maddison, 1991) of
shortest trees, and that even shorter trees exist that
were not recovered. Furthermore, although our
search strategy involved well over two years of com-
puter time, no search swapped to completion; there
is no assurance, therefore, that these trees repre-
sent even a local parsimony optimum. Although it
is, of course, impossible to know how far from com-
pletion any search is when it is truncated, the
search design used here offers an insightful basis
for comparison. Data sets 1 and 2, and 3 and 4 are
identical except for the inclusion of two gap char-
acters (indels) in data sets 2 and 4, each of which
apparently accounts for only four steps on the
shortest trees obtained. Thus, the fact that the
shortest trees obtained in searches of data set 2 are
seven steps longer than those obtained in searches
of data set 1 indicates that the shortest trees ob-
tained in our searches of data set 2 are three steps
less parsimonious than trees derived from searches
of data set 1. A similar comparison of the searches
of data sets 3 and 4 reveals that the shortest trees
from searches of data set 4 are two steps less par-
simonious than those obtained from data set 3.
We also sampled among the large set of equally
parsimonious trees following Sanderson and Doyle
(1993b). Using trees obtained in searches of data
set 1, we examined the number of distinct compo-
nents (clades) as a function of the size of the sample
of trees (number of trees). We wanted to determine
whether increasing the set of trees uncovers new
components that bear on the relationships of par-
ticular taxa or, in contrast, includes different sub-
sets of the i
tions on the same theme (Sanderson
improved methods of phylogenetic analysis of large
data sets will ultimately be one of the central issues
of phylogeny reconstruction during the next several
years (see discussions in Chase et al., 1993; Doyle
28
Annals of the
Missouri Botanical Garden
et al., 1994; Mishler, 1994; P. Soltis & Soltis,
1997).
Although the anomalous relationships described
for some taxa may be unsettling, extremely short
branches characterize most of the major clades in
the 18S rDNA trees. The internal support for many
branches is very low, as indicated by the parsimony
jackknife analysis (Farris et al., 1997). Although
the monophyly of the angiosperms is well supported
(jackknife value of 100%), few major clades within
the angiosperms have high jackknife values. For
example, large clades such as eudicots and Rosidae
do not have jackknife values above 50%; the sax-
ifragoids represent the largest clade having a high
jackknife value (jackknife value of 68%). The other
monophyletic groups with high jackknife values are
relatively small, such as cunonioids, Zingiberales,
Malvales, Caryophyllales, Lactoridaceae—Aristolo-
chiaceae, and Schisandraceae-Illiciaceae—Austro-
baileyaceae. Significantly, a number of major clades
seen in all shortest trees, as well as in trees many
steps longer than the most parsimonious trees, do
not have jackknife values above 50%, including
monocots, glucosinolates, Caryophyllidae s.l., and
Asteridae s.l. The majority of high jackknife values
correspond to pairs of sister taxa representing ter-
minal nodes (e.g., Calycanthus—Sassafras, Brexia—
Euonymus, Lepuropetalon—Parnassia, Plumbago—
Cocoloba, Helwingia—Phyllonoma, Tragopogon—
Tagetes, Francoa—Greyia, Trochodendron—Tetracen-
tron, Menispermum-Tinospora).
Examination of trees obtained from searches that
found trees one or a few steps longer than the short-
est trees also suggests low internal support for some
branches. The phylogenetic position of the mono-
cots appears weakly supported. In some searches
of data set 2, for example, trees only one step longer
than the shortest trees place the monocots within
the eudicots, as part of Rosidae, a position also
observed in the shortest trees obtained from search-
es of data set 3 (Fig. 3). Although all of the starting
trees and shortest trees showed Amborellaceae, Il-
liciaceae, Schisandraceae, and Austrobaileyaceae
to be at the base of the angiosperms, one search of
data set 2 resulted in trees two steps longer than
the shortest trees and placed these four families
near the monocots, with Acorus and Oncidium as
the first-branching angiosperms. Trees two ste
longer than the shortest trees show the Asteridae
s.l. embedded within Rosidae, rather than sister to
this large clade. In trees two steps longer than the
shortest trees found for data set 3, Caryophyllidae
s.l. are not part of Asteridae s.l. but instead are
part of the large Rosidae clade.
These few examples illustrate well the uncertain-
ty that surrounds some angiosperm relationships in-
ferred from analyses of 18S rDNA sequences. Fur-
thermore, because relatively few character-state
changes occur on many of the branches, a small
amount of homoplasy or error in the data set may
be sufficient to distort some relationships.
Additionally, some of the anomalous placements
could reflect insufficient and/or uneven taxon sam-
pling. The somewhat uneven taxonomic distribution
of the sequences presently available means that
some groups, such as Asteridae, and much of Ros-
Dilleniidae, Caryophyllidae, and several orders of
Rosidae are under-represented.
The importance of sufficient taxon density is re-
vealed here by some of the differences in topology
observed between trees resulting from analyses of
the smaller and larger data sets. Many of the taxa
not present in the two smaller data sets (3 and 4)
represent monosulcates and lower eudicots. It is
this portion of the overall topology that shows the
most spurious relationships in trees derived from
analyses of these two small data sets (the distinc-
tion between the monosulcate grade and eudicots
largely breaks down in Fig. 3, for example). In con-
trast, the much more thoroughly represented Aster-
idae s.l. and Rosidae clades are little affected by
slightly decreased representation in data sets 3 and
4. These findings lend further support to the im-
portance of sufficient and equal taxon density in
attempts to infer angiosperm phylogeny (e.g., Syts-
ma & Baum, 1996).
One of the major lessons of this study is that the
18S rRNA gene is difficult to sequence, apparently
due in large part to the secondary structure inher-
ent in the rRNA. As a result, many published se-
quences are erroneous, some highly so, and the ex-
tent of insertion and deletion events has been
greatly overestimated. We reiterate that whereas the
total length of the aligned 18S rDNA data matrix
of 64 taxa used by Nickrent and Soltis (1995) was
1853 bp, the length of our 228-taxon data matrix
actually is shorter, 1850 bp. After resequencing
over 20 dubious 18S rDNA sequences, we were
able to remove numerous "false" indels and reduce
the length of the aligned sequences. The great ma-
jority (70%) of the 18S rDNA sequences used here
were generated via cycle sequencing followed by
automated sequencing, an approach that provides
more reliable rDNA sequences. Additional “older”
18S rDNA sequences should be replaced with se-
quences generated via this approach.
The overall slower rate of evolution of 188 rDNA
compared to rbcL (see Nickrent & Soltis, 1995)
Volume 84, Number 1
1997
Soltis et al. 29
18S Ribosomal DNA Phylogeny
contributed, in part, to the widespread belief that
18S rDNA sequences would not contribute greatly
to phylogenetic inference in angiosperms. Although
this study and other recent papers employing entire
18S rDNA sequences (e.g., Nickrent & Soltis,
1995; Kron, 1996; D. Soltis & Soltis, 1997; Rod-
man et al., submitted; Johnson et al., unpublished)
have dispelled this notion, 188 rDNA sequences
will, in most cases, not elucidate relationships to
the degree possible with the more rapidly evolving
rbcL. In some groups such as Orchidaceae, how-
ever, 185 rDNA has been found to evolve faster
than rbcL (Cameron and Chase, unpublished).
FUTURE CONSIDERATIONS
These exploratory analyses clearly illustrate the
phylogenetic potential of 18S rDNA sequences for
elucidating angiosperm relationships at higher tax-
onomic levels. Future attempts to conduct broad
phylogenetic analyses of 18S rDNA sequences
should not only add more taxa, but should also in-
volve the resequencing of the 18S rRNA gene for
some of those taxa for which erroneous sequences
are suspected.
This study suggests that a broad, nuclear-based
phylogenetic hypothesis for the angiosperms is
achievable via sequence analysis of the 18S rRNA
gene. One of the strengths of 185 sequence data
appears to be the ability to recognize a suite of
groups that appear in all shortest trees (e.g., glu-
cosinolate clade, saxifragoids, Caryophyllidae s.l.,
Asteridae s.l., celastroids). This may reflect substi-
tutions that occurred in highly conserved portions
of the 188 rRNA gene during the early diversifi-
cation of a lineage, resulting in a well-supported
clade. Such substitutions are rare, however, and the
result is limited resolution in some areas of the 185
rDNA topologies. Thus, our results also clearly re-
support possible with rbcL
. Increased
sampling of angiosperms for 18S rDNA sequence
analysis is desirable. However, to achieve a nucle-
be necessary to include all, or portions of, the 26S
RNA gone es. well, The willy of portions of the
been demonstrated for angiosperms (Hamby &
ps ite ae: bo dos ade eal pe the
ganisms (e.g., & Chapman, 1991; Chap-
man & Buchheim, 1991; Chapela et al., 1994; Wa-
ters et al., 1992).
CONCLUSIONS
This study provides general insights into the
structure and evolution of the 185 rRNA gene in
angiosperms and dispels certain “myths” about its
evolution. Indels are neither as common nor as
problematic for alignment as previously believed.
Instead, they are largely confined to a few, small,
specific regions that correspond to the termini of
in helices present in the p secondary
structure for 185 rRNA. When these few, short ar-
eas are eliminated from consideration, alignment of
18S rDNA sequences is straightforward and easily
accomplished by eye across all angiosperms. Con-
versely, indels are rare throughout most of the 185
rRNA ve^ M present, they panne involve a
single base urthermore, indels present in
highly regions of the gene in Mh
be phylogenetically informative, such as the inse:
tion that unites saxifragoids and the deletion ^i
unites higher eudicots.
Initial attempts to evaluate the impact of sec-
ondary structure of the 18S rRNA transcript on
phylogeny reconstruction in angiosperms suggest
that both stem and loop regions appear to be
sources of phylogenetic information, with a slightly
greater proportion (58% vs. 42%) of informative
sites found in stem rather than loop regions. Of the
stem changes we analyzed, only 2796 destroyed a
base-pairing couplet; 73% restored or maintained
stem base pairing and hence are considered com-
pensatory. The most frequent type of stem change
observed involved single base substitutions that
changed one base-pairing couplet to another (e.g.,
U-G to C-G; U-A to U-G). The high frequency of
compensatory change indicates that some down-
weighting of stem characters relative to loop bases
future broad analyses of 18S
dance include the presence of a tricolpate or eu-
dicot clade, which in turn includes two large clades
corresponding mostly to Rosidae and Asteridae s.l.,
respectively. However, the latter clade also includes
Caryophyllidae s.l. in 18S rDNA trees, but not in
trees retrieved from analyses of rbcL sequences. In
addition, the monocotyledons are monophyletic
(with the possible exception of Acorus) and gener-
ally appear with other taxa having monosulcate pol-
len. One of the most noteworthy differences be-
tween this study and that of Chase et al. (1993)
concerns the first-branching angiosperms. The
Annals of the
Missouri Botanical Garden
woody magnoliids Amborellaceae, Illiciaceae,
ae and AC il consistent-
ly appear as first-branching angiospe е
always followe d by the paleoherb ~ ветру
Ceratophyllum is closely allied with the monocots
and does not appear as sister to all other angio-
sperms, as in analyses of rbcL sequences (Chase et
al., 1993). Monophyletic groups apparent in all
analyses include Caryophyllidae s.l., Asteridae s.l.,
saxifragoids, glucosinolate-producing taxa, santa-
loids, and cunonioids. Other clades apparent in
most analyses include ranunculids and nitrogen-fix-
ing taxa. Thus, this analysis identifies major clades
of angiosperms that are largely consistent with
those inferred from rbcL analyses
This study further pre RE that 18S rDNA
quences contain sufficient information to conduct
пет ч studies at higher taxonomic levels in
the s. Additional phylogenetic analyses
of Sarath по bi be conducted using a larger
18S rDNA data set that improves taxon sampling
for Magnoliidae and Dilleniidae in particular. In
constructing this larger data set, some taxa
which published sequences are available mere
first be resequenced.
Although comparative sequencing of the entire
18S rRNA gene holds great promise for retrieving
phylogeny at the family level and above in the an-
giosperms, this nuclear gene will rarely elucidate
familial and generic relationships to the extent pos-
sible with rbcL (see also Nickrent & Soltis, 1995).
Due to the slower rate of evolution of 185 rDNA
compared to rbcL, it likely will be necessary to se-
quence the 26S rDNA as well to obtain a nuclear-
based estimate of phylogeny comparable to that
achieved with rbcL. Lastly, because of the general
congruence of 18S rDNA and rbcL topologies for
angiosperms, this study concomitantly suggests that
18S rDNA and rbcL sequences should be combined
to provide a more accurate estimate of angiosperm
phylogeny. One can anticipate that other sequences
(e.g., atpB and 26S rDNA) will ultimately also be
combined with rbcL and 18S rDNA sequences to
provide a larger data set from which to infer a more
complete picture of angiosperm phylogeny.
Literature Cited
Bailey, 1. W. 1957. The ponti, and limitations of
wood anatomy in the study of the phylogeny and AE
sification of angiosperms. J. Arnold Arbor. 38:
Bakker, F. T., J. L. Olsen, W. T. Stam & C. Van
1994. The Cladophora complex ү шеп. ig
views based on 18S rRNA gene sequences. Molec . Phy-
logenetics and Evol. 3: 365-382.
Bhattacharya, D. & L. Medlin. 1995. The phylogeny of
plastids: А review based on comparisons of small-sub-
unit ribosomal RNA coding regions. J. Phycol. 31: 489-
498.
Bremer, K. 1988. The limits of amino acid sequence data
«e hee ag phylogenetic reconstruction. Evolution
8;
rii M. A. & R. L. Chapman. 1991. Phylogeny of
the colonial green flagellates: A study of 18S and 26S
rRNA sequence data. BioSystems 25: 85-100
, M. Turmel, Zimmer & R Chapin
1990. Phylogeny of uen mydomonas gt Sas
based on cladistic an + of р? чиш 185 rRNA 5
quence data. J. Phyco L2 6: 689—6
Bult, C., M. Kallersjó & Y. Sub. Ue о аи Dion and
ea ga: of 16/188 rDNA T ол а total
plant DNA. Pl. Molec. Biol. Re : 273-284.
анк I. 1995. A molecular и a bryophytes
based on the nuclear encoded 18S rRNA genes. J. Pl.
Physiol. 146: 59-63.
Chapela, I. H., S. A. Rehner, T. ut Schultz s: U. G. Muel-
ler. 1994. Evolutionary "hist of the symbiosis be-
tween fungus-growing ants ut iuit imet: [yen 266:
Buchheim. 1991. Ribosomal RNA
logeny and em of green algae. C. R. C. Crit Ra
Pl. Sci. 10: 343-368.
ho M. W., D. E. Soltis, R. G. mico D. Morgan,
. Les
< arn, Jr.,
ett, S, [луд а & ү. А. Рћујо
кане of seed plants: An analysis of mucleotide se-
quences from the D gene rbcL. Ann. Missouri Bot.
Gard. 80: 528—
Chaw, S.-M., H. Lon . Wang, A. Zharkikh & W.-H.
Li. 1993. "The phylogenetic position o of Taxaceae based
on 18S rRNA sequences. J. y olec. Evol. 37: e.
, Н.-М. Sung, Н. Long, A. Zharhikh & W.-H
1995. The phylogenetic positions of the conifer pe
Amentotaxus, Phyllocladus, and Nageia inferred from
S
‚А.
1997. Маны: Ен of gymnosperms and 5
plant evolution: Analysis of 185 rRNA sequences. e
Molec. Evol.: in press.
Crane, P. R. 1985. Phylogenetic analysis of seed plants
and the origin of the angiosperms. Ann. Missouri Bot.
Gard. 72: 71
988. Major clades €— oe in the
“higher” gymnosperms. Pp. 2 п С. B. Beck (ed-
itors), Dion and Evolution е dose Nee a, Columbia
Univ. s, New York.
. Pedersen. 1995. The
and early diversification of angiosperms. Nature 374:
27-
Cronquist, A. 1968. The Evolution in ¡peones of
Higher Tor, Houghton,
Mifflin, Bos
An Integrated System E Classification of
EE Plants. Columbia Univ. Press, N
м Yor
Curtiss, W. C. & J. N. Vournakis. 1984. Ginis of
Volume 84, Number 1
1997
Soltis e 31
185 d DNA Phylogeny
ies substitutions in eukaryotic 5S rRNA: Selection а
or of RNA secondary structure. J. Mole:
Evol. 20: 351-361.
Dahlgren, R. T. 1980. A revised system of classification
of the angiosperms. Bot. J. Linn. Soc. 80: 91-124.
—————. 1983. General aspects of angiosperm evolution
and a Te о Nordic J. Bot. 3: 119-149.
ive Clifford & P. F. Yeo. 1985. The
onocotyledons: kein Evolution,
ig ine er- Meron N York.
D M. T B D. M. Hillis ieee d RNA sec-
ary structure—Com арене tory mutations and més
= for ai вв dris Molec. Biol. Evol. 1
ГА
Donoghue, М. J. & J. А. Doyle. 1989а. Phylogenetic
analysis of и and the тер of Ham
amelidae. Pp. 17-45 in P. R. Crane & S. Bla aces
(editors), ткн 5 po el History of
the Hamamelidae. ia arendon Press, Oxford.
& 89b. Phylo; genetic "sed of seed
rphol
plants and angiosperms و on m al char-
acters. Pp. 181-193 in B. Fernholm, K. Bremer & H.
Jornvall (editor d jm Hierarchy of Life: тармақ апа
hology in Phylogenetic Analysis. Elsevier Science
Publisher Amste: As
Im: ead, J. Е Smith & J. mer.
1992. тыы та of dk [vas
on rbcL sequences. Ann. Missouri Bot. Gard. 79: 333—
345.
Doyle, J. A. & M. J. Donoghue. 1986. Seed plant phy-
logeny and the origin of the ig mro A е uc
oen cladistic approach. Bot.
1992. Fossils mi seed стра TN
eny analy Brittonia 44: 89-
& . Hotton. 1991. РИУ of early
angiosperm polled 1 ina ee ca context. Pp. 169-195
in S. Blackmore & S. H. es (editors), Pollen and
Spores. ure Press, O: эчи
‚ М. J. Donoghue & E. A. Zimmer. 1994. Inte-
gration of pera and ribosomal RNA data on
the origin of angiosperms. Ann. Missouri Bot. Gard. 81:
419-450.
DOE Мама Lotes 1990. Early Creta-
s tetrads, zonasulcula e pollen and Winteraceae.
IL I Cla distic sini and таб визе у Amer. J. Bot. 77:
568.
Duvall, M. R., M. T. и М. W. Chase, W. D. Clark, W.
J. Kress, H. С. Hills, L. E. rri а cene Bs
Gaut, E. A. Zimme га C H. Lee Phylo-
genetic hypotheses for the kel ape e ош
Pea sequence data. Ann. Missouri Bot. Gard.
к V. K, J. Arnold & В. B. Meagher. 1985.
Comparison of the nucleotide sequence of soybean 18S
rDNA with the sequences of = small-subunit
59-
D. M. Kil. 1994. Bio-
diversity at the еле level: The domains, kingdoms
and api of ries gras eke. = io sigo 21-31.
Endress, P. K of the floral
structure in ee PL. $n. Evol. 8: 175-183.
———— & R. Honegger. 1980. The pollen of the Austro-
baileyaceae and its phylogenetic significance. Grana
19: 177-182.
Farris, J. S., * А. Ађеп, М. врте D. signed
A. G. Kluge. 1997. Parsim
жешн и Cladistics “1. ‘in press.
me K Gy G: J-Olsen; D: J; D" Бој чађи
M. T. Сй, Е. С. Raff, N. R Pace & R. A. Raff.
1988. epos phylogeny of the animal kingdom. Sci-
ence 239:
Forster, H., + р. јави H. Elwood & M. L. Sogin. 1990.
Sequence analysis of the small subunit ribosomal RNAs
of three zoosporic fungi and Av ian for fungal evo-
lution. үн ser 82: 306—
Gargas, A., P. T. De Priest "fi w Taylor. 1995. Positions
of multiple insertions in SSU e of lichen-forming
fungi. Molec. Biol. Evol. 12: 208—
Goremykin, V М; ш, Bobrova, J з А. Troitsky, А. Ап-
ton 1996. Noncoding remount sie
the ede. keen chloroplast inverted re
dition to rbcL data do not support Gnetalean “affinities
of angiosperms. Molec. Biol. Evol. 13: 383-396.
Hamby, K. R. & E immer. 1992. Ribosomal cvm
asa phylogenetic tool i in plant systematics. Pp. 50-
in P. S. Soltis, D. E. Soltis & J. J. Doyle (editors), de
се Systematics of Plants. Chapman and Hall, New
Hendricks, LR De m Bi 2n de bns Neefs, A
s& hte The evolutionary posi
tion ol the rhodophyte "frin umbilicalis and the Ms
ote уе о scottii among other eukary-
otes as from complete sequences of small
ribos ru сан RNA. J. Molec. Evol. 32: 167-177.
Hillis, D. M. T. Dixon ibosomal DNA:
Mar evolution and phylogenetic inference. Quart.
Rev. Biol. 66: 411-453
Hinkle, G., J. K. Wetterer, T. R. Schultz & M. L. Sogin.
1994. Phylo ogeny of the attine ant fungi based on anal-
ysis of small subunit н mal RNA gene sequences.
Science 266: 1695—
Hoot, S. B. & P. R E
ships in the Ranuncu ie b e on
Evol. 9: 119-131.
` 1995. Inter-familial relation-
molecular system-
m & Р. i ioi 1995. Phylogenetic
A. Culha
relationships of ~ zabalaceae and жашыны ox-
асе roplas
: Chloroplast and die DNA sequence evi-
Rc do 2 Evol. 9: 195-1 €
Hufford, & W. C. Dickison I geo pun
rond of Cunoniaceae. € ва. 1 17: 181-2
. 1990. Роан posi-
ella 8 species within the Chlorococ-
n complete small-subunit en
RNA sequences. J. е Evol. 31: 43 2
пе Е. С. Theriot, Е. A. Zimmer & К. L. Сћар-
1990. Тһе Козя and Mieromona-
puertos -Cladistic =» of nuclear rRNA ве-
quence "a Phycol. 26: 711-721.
ranz, H. D., ce = v gese Capesi sius, Ch. W.
Sensen & у. А. Низ 5. The origin of land plants:
Phylogenetic cune eit ioe К harophytes, Bryo-
phytes, and vascular plants inferred from complete
small subunit ribosomal RNA gene sequences. J. Mo-
lec. E
Kress, W. Е ahn, Т. М. Evans & Е. A. Zimmer.
1995. Using the evolutionary radiation of the fam-
ilies of the Zingiberanae ч morphological, molecu-
lar, and fossil ie er. J. Bot. 82 (suppl.): 142.
Kron, 1996. a relationships nt Empe-
traceae, Epacridaceae, and Ericaceae: Evidence from
nuclear ribosomal 18S sequence data. Ann. Bot. (Lon-
don) 77: qa 03.
Chase. 1993. Systematics of Ericaceae,
E Epacridaceae and related taxa based
32
Annals of the
Missouri Botanical Garden
upon пер sequence data. Ann. Missouri Bot. Gard. 80:
735-74
Les, D. H bos The origins and affinities of the Cera-
tonhyllaceae. Taxon 37: 326-345
, D. K. Garvin & С. Е Wimpee. 1991. Molecular
evolutionary history of ancient aquatic angiosperms.
Proc. Natl. Acad. USA. 88: 10119-10123.
. Stevenson. 1991. Cladistics of the
96.
iscovery and importance ^ mul-
tiple islands of most- -parsimonious trees. Syst. Zool. 40:
5-328.
— ———, M. Ruvolo & D. L. Swofford. Meta Geographic
origins of айна mitochondrial DNA: Phylogenetic ev-
idence from control region sequences. Syst. Biol. 41:
111-124.
Martin, P. G. & J. M. Dowd. 1991. Studies of angiosperm
phylogeny por protein sequences. Ann. Missouri Bot.
Gard. 78: 296-337.
— D. P. & R. L. Honeycutt. 1990. Ribosomal RNA
n vertebrate es: a ia and Шоо applications.
n. Rev. Ес “mer 121:
Mishler, B. D. и ескі of molecular "i
morphological ntes Amer. J. Phys. Anthropol. 94: 1
156
— ——, К. Bremer, С. J. Humphries & S. P. Churchill.
1988. The use of nucleic acid sequence data in phy-
logenetic Piatt ep yrsa 37: 391-395
Lo As s, M. m, K. а Renzaglia,
Tos.
chhei
D.3. ‘Garbary C. F Delwiche, F. X.
Kantz & R. L. Chapman. 1994. Phylogenetic relation
ships of the * "green cni and “bryophytes.” Ann.
souri Bot. Gard. 81:
Morgan, D. R. & D. A Sulis 1993. Phyl
tionships among m
based o
ene rela-
mbers of Бахйтарасане sensu lato
on rbcL toe data. Ann. Missouri Bot. em)
80: 631—660.
Nairn, C. J. & R. J. Ferl. 1988. The — nucleotide
sequence of the small- мац ribosomal RNA coding
mia ere: co im-
plications. J. Molec. Evol. 27: 133-141.
масай D. 1. 1994. From fl ^ film: Rapid se "—
ing methods for bes m plant species. Bio
Tec — 16: 4
C. R. Fra нын 1990. Phylogenetic Mim
ships of the Santalales and relatives. J. Molec. Evol. 31:
301
& D . Soltis. 1995. A comparison of angio-
sperm deii Кош nuclear 18S ape and rbcL
sequences. ore hae Bot. Gard. 82: 208-234
& E. M. 1994. High cc of nucleotide
substitution in iari r small- subunit (18S) rDNA from
vemm rasitic rans: plants. J. Molec. Evol. 39: 62—
iiie. К. C., W. L: Crepet, D. Stevenson & E. M. Ег
1994. A "d of seed plant phylogeny. i
Missouri Bot. Gard. 81: 484—5:
"НЕ
Noller, 1984. Structure of ribóéorel RNA. Annual
Rev. ries 53: 119-
Olmstead, R. , K. M. Scott & J. D.
Michae
Palmer. Ty Monophyly of y Asteridae and iden-
tification of their — lineages inferred from DNA se-
quences of rbcL. - Missouri Bot. Gard. 79: 249—
265.
— ——^ B. Bremer, К. M. Scott & J. D. Palmer, 1993. A
parsimony analysis of the Asteridae sensu lato based on
rbcL sequenc n. Missouri Bot. Gard. 80: 700-722.
Olsen, G. J. 1987. Earliest phylogenetic _branchings:
Comparing rRNA-based evolutionary trees inferred with
various ee. Cold Spring Harbor Symp. Quant.
Biol. 52: 825-83
Olsen, J. L., W. T. з ат, 5. Berge r & D. Menzel. 1994.
185 rDNA and evolution in the Dasycladales es
rophyta): Modern living fossils. J. Phycol. 30: 729-744.
Qiu, ан M. Y. Chase, D. H. Les & С. R. Parks. 1993.
siste of the Magnoliidae: Cladistic
nalyses of nucleo sequences of the plastid gene
vin L. Ann. Midi: Bot. ard. 80: 587-606.
Ragan, M. A., C. J. Bird, E. L. Rice, R. R. Gutell, C. A.
Murphy & R. K. Singh. 1994. A molecular phylogeny
of the marine red algae (Rhodophyta) based on nuclear
small-subunit rRNA gene. Proc. Natl. Acad. U.S.A. 91:
7276-7280.
Rathgeber, Ј. € 1. Capesius. 1990. Nucleotide Minor.
of the intergenic spacer and the 18S ribosomal RNA
gene from mustard (Sinapis alba). Nucl. Acids Res. 18:
12
Rettig, J. H., H. D. Wilson & J. R. Manhart. 1992. Phy-
logeny of the Cason lle elie sequence data. Taxon
41: 201-209.
Rice, K. A., M. J. Donoghue & R. G. Olmstead. 1997. A
reanalysis of the large rbcL data set: Implications for
future phylogenetic studies. Syst. Biol. in dris
Rodman, J. E. 1991. A taxonomic analysis o diy.
id Veni: plants, Part 2: Cladistics. Syst. Bot
16: 6
8% us ‚ D. E. Soltis & К. J. Sytsma. Dual
origin of mustard oll biosynthesis inferred from congru-
ent nuclear and plastid gene phylogenies. Proc. Nat
Aca dug 4 S.A., submitted.
К Olives: R. | Nakamura, J. U. McClammer,
Jr. & + H.B Bledsoe. 1984. A taxonomic analysis and
revised classification of Centrospermae. Syst. Bot. 9:
23.
: e, K. Karol, E. Conti, K. J. Sytsma &
Ј D. Раван. 1993. Nucleotide sequences of the rbcL
e ке the monophyly of mustard oil plants. Ann.
Missouri Bot. Gard. 80: 686-699.
ышын М. J. & J. J. Doyle. 1993a. Reconstruction of
organismal gene phylogenies from data on multigene
families: E in homoplasy, and сопћ-
dence. DUM Biol. 4
E c ogenetic relationships in
North a Азтара, Jus us (Fabaceae) based on chlo-
roplast DNA restriction site variation. Syst. Bot. 18:
395408.
&M onoghue. 1994. Shifts in diversification
rate with Kog ои of angiosperms. Science 264: 1
1593.
Savard, L. & M. Lalonde. 1. Sequence of 185 rDNA
of actinorhizal t€ не (Betulaceae). Pl. Molec.
Biol. 16: 725-72
Schlegel, M., H. J. = М. L. Sogin. 1991. Molec-
ular evolution i in hypotrichous. ciliates: Sequence of the
ag prem ribosomal RNA g — from Onychodro-
quadri xytricha granulifera (Oxytri-
chida, мга Ciliophora). J. Molec. Evol. 32:
Senet J. F. & R. B. Meagher. 1992. In vivo analysis
plant RNA structure: Soybean 18S ribosomal an
d Фе ныр лао carboxylase small subunit
RNAs. Pl. Molec. Biol. 1 234.
Smith, A. B. 1989. RNA d data in phylogenetic
|
|
|
1
|
|
|
Volume 84, Number 1
1997
Soltis et al. 33
18S и DNA Phylogeny
reconstruction: з the limits of its resolution. Cla-
distics 5: ee
Sogin, M. L., У uu & J. H. Gunderson. 1986.
сва ‘diversity of е: small subunit
rRNA genes. Proc. Natl. Acad. U.S.A 1383-1387.
Soltis, D. ан $ P. S. Soltis. 1997. Phylogenet
ships a Saxifragaceae sensu lato: A compariso
topologies oed in 188 rDNA and ++ његова
Amer. J. Bot. in pao
n gan, S. M. Swensen, B. C.
Mullin, J. M. Dowd & К С Martin. 1995. Chloroplast
gene sequence data suggest a single origin of the pre-
oe for gems nitrogen fixation in angio-
. Proc. Natl. Acad. U.S.A. 92: 2647-2651.
Soltis, P p s. &D E. Soltis. 1997. lee nere —
: I bids data sets. Bol. Soc. Bot. México. In
Stebbins, б. L. 1974. Flowering е раја cam Above
the Species Level. Harvard Uni York.
Swann, E c. rs W. Taylor. 1993. Hikes: чөй ч basid-
— it Big 18S rRNA gene perspective. Mycologia
Swf, D T 1993. uH Былыр ig Analysis Us-
g Pars — version 3.1. or Biodiv к Il-
linois Natural H ar Survey, ipd
& G. J. Olsen. 1990. Phylogeny acude.
Pp. 411—501 in D. "Hillis € C. Moritz (editors), Molec-
ular era Sinauer, Sunderland, Massachusetts.
Sytsma, K. J. € 1996. ко nd
nies and the т. of the р.
314-340 in D. W. Taylor & L. J. Hickey (dio, Tor
bin. Plant Origin, Evolution, and Phylogeny. Chapman
and Hall, New York.
Takaiwa, F., K. Oono & M. Sugiura. 1984. The complete
nucleotide sequence of a rice 17S rRNA gene. Nucl.
1–5448.
Flowering Plants: Origin and Dis-
persal. Oliver & Boyd, Edinburgh.
Outline of ex pre = flowering س
plants Magli Bo
> = op odi со. of
bere 0.55. R, pê grad.
Thorne, R. F. 1992. An п updated classification of the flow-
ering plants. Aliso 1
Unfried, 1., U. Stocker à P. Gruendler, 1989. мо»
EY of the 18S rRNA gene from Arabidops
iana Соло. Reged Herein Res. 17: 7513.
Wada, H. & N. Sa Зона relationships
among extant clase s. echi noderms, as inferred from
sequences o NA, coincide with ag angen a nid
= ed from the fossil record. J. Molec. Evol. 38: 4
„сш Р. О., О. Hinkle, М. L. Sogin € 5. K. Stickel.
1993. Monophyletic "y sm К the Metazoa: An evolu-
tionary link with fungi. Science 260; {тине
Buchi
Waters, D. A., Ки. A y & К.
ا 1992. Preliminary Bred د f the Poe
of bryophytes
pea жая, ere Bot.
459—466.
Wheeler, W. С. LL Honea 1988.
quence „лыы ein lomas E Evolutionary and
phylogenetic — Melo: ¿vol.
he mS. pe ү” . Bacterial Solution, Microbiol. Rev.
Ст Pu x А. Erdmann. 1986. Cladistic noe jog of
had rRNA and 16S rRNA secondary and primary stru
re—The evolution of eukaryotes - their мин о to
bd teria. J. Molec. Evol. 24: 152-166.
Young, D. A. 1981. Are the angiosperms primitively ves-
selless? Syst. Bot. 6: sedeo
Young, D. J. & 970. classification of
dicotyledons: A € of the upper Lovell of hierarchy.
Austral. а Bot. 18: 387-433.
sR x Hamby, M. L. Arnold, D. A. Leblanc
E. iot. . Ribosomal RNA ронун
and flowering | plan evolution. Pp. 205-214 in B. Fern
holm, K. Bre & J. Jórnvall (editors), The 1 Histerchy
of Life. падну Science Publishers, Amsterdam.
34 Annals of the
Missouri Botanical Garden
1A
EUDICOTS
100 r- Calycanthus
| — Sassafras
Tacca
Bowiea
Chlorophytum
Allium
Sagittaria
95 — Hippeastrum
Eucharis
Gladiolus
Isophysis
Xanthorrhoea
E Veitchia
— Cyanella z
Helmholizia o
as Maranta 2 =
2 Zingiber O =
E Costus О б
Саппа me [72]
Heliconia =
E M Eb n O
== 61 Sepa >
21 а Е
59
e e
Sparganium ~
баб йолт m
m Call.
Onc
dmm ape
Dillenia
Gunnera
100 r- Mkilua
— Isolona
Acor S
100 [ .—— Nymphaea E
Houttuynia [e]
Ls] = Peperomia m
Saururus ё
Amborella o
ы sr ва |
ustrobaileya
— Шїсїи т
г— Gnetum gnemon e
e Gnetum i e a
Gnetum urens 3
Ephedra f torreyana ©
Ерћеага зтса o
Figure l. Strict consensus of 5294 shortest trees resulting from the exploratory dices analysis of 223 species
of angios rms. Eac shortest trees has de f 3923 steps, 0.535. Parsimony Jackin
values (Farris et al., 1997) of 50 or above (based on 1000 replicates) are given ates nodes (run time = 949 sec.).
Because of its size, the tree has been broken he foir parts (1A, 1B, 1C, and 1D).
Volume 84, Number 1
1997
Soltis et al.
18S Ribosomal DNA Phylogeny
35
П
Figure 1B.
ASTERIDAE S. L.
ROSIDAE
Hedyosmum
Aristolochia
Asarum
Saruma
S
Lactori.
Drimys
ja
Trochodendron
Tetracentron
Sargentodoxa
oo
ptis
anthorhiza
lenispermum
позрога
aulophyllum
одорћућит
иргејеа
5 0
Ото С
ејит
Knightia
Placospermum
S8U3HO3T1Vd
SGMNONNNVY
SLOSIGNA H3M01
MONOSULCATE GRADE
36 Annals of the
Missouri Botanical Garden
ASTERIDAE S.L. TC
Co
S
SGIODVUAIXVS
54
1
Daph iphyll
niphyllum
Cercidiphyllum
ч Liquidambar
Altingia
ао
ao
is
Ceanothus
Morus
00 Pilea à
Symbegonia
onia
1
2 Bombax
ossypium
Luhea
fo Ејавадпиз
Tetrameles
—[Lt Bes
Abobra
98 س Francoa
Greyia
Alnus
Casuarii
Bauera
89 Ceratopetalum
Eucryphia
Cephalotus
Koelreuteria
Guaiacum
SNIXIJ-N39O.LIN
3VGdlSOMN
SGIOINONND
Tm
5 E
SGIO1VLNVS
100 Punica
67 rexia
Saloy LSY139
SALVIONISODNTD
LOWER EUDICOTS
Figure 1C.
Volume 84, Number 1
1997
Soltis et al.
18S Ribosomal DNA Phylogeny
37
Figure 1D.
Drosera
зауно NVAIVOIA
locos
Philadelphus
Hydrangea
Helwingia
Phyllonoma
3vanalsv
ROSIDAE
LOWER EUDICOTS
AVGITIAHdOANVD
1D
4 9 IVUIUILSV
38 Annals of the
Missouri Botanical Garden
4 2A
La
Veitchia
E па Ye anella
к,
пїа
|
o
SLODONOW
Glomeropitcairnia
Ze
la
Oryza
erus
Sparganium
Elasis
3Qvu5 31VWITASONON
lu
Colchicum
Ca ми
Onc
^ Ceratophyllum
| œ
3]
©
=
®
3
5
"i
“TR
tm
Sau3Hod Wd
9 Ł Schisandra
4— Austrobaileya
icium
Gnetum то
40 [
Gnetum кт
Ып Gnetum
pe Ephedra s nba
| — Ephedra torreyana
5апомо 1 по
Figure 2. Опе of 2508 shortest trees resulting from the па phylogenetic analysis of 223 species of angio-
sperms; two T were included in the analyses. Each of the shortest trees has a length of 3930 steps, СТ = 0.235,
and RI — 0.540. Arrows indicate nodes not present in the strict consensus of all shortest trees. The letters A and B
indicate the perpe of the indels described in Table 2. Because of its size, the tree has been broken into four parts
(2A, 2B, 2C, an
Volume 84, Number 1 Soltis et al. 39
1997 18S Ribosomal DNA Phylogeny
2B
А з ASTERIDAE S. L.
ROSIDAE
Hedyosmum
19 Aristolochia
2 5 4
S8U3HOd'T1Vd
5 = Trochodendron
s Tetracentron
bia
6
2 3 Sargentodoxa
Ranunculus
Coptis
Xanthorhiza
Menispermum
ll
~ A
о а
[^]
CIA | ој >
SLODICNA чэмоп
]
inospora
5 0 Caulophyllum
; Ё Podophyllum
SOMMINANVEY
MONOSULCATE GRADE
Figure 2B.
Annals of
Missouri an Garden
Figure 2C.
ASTERIDAE S. L.
SGdIO9 VilHIXVS
ONIXI4-NIDOY.LIN
SGIOINONND
SGIOTV.LNVS
Saiouisv 132
S31V IONISOO(119
LOWER EUDICOTS
2C
AVdISOY
Моште 84, Митбег 1
1997
Soltis et al.
18S Ribosomal DNA Phylogeny
41
7
ЗУОГПАНЗОАУ О
3avu5 NVATVOIS
3vanalsv
Camptotheca
ROSIDAE
Figure 2D.
LOWER EUDICOTS
2D
715 3avani1sv
42 Annals of the
Missouri Botanical Garden
3A
ASTERIDAE S. L.
PLUS
ROSIDAE PLUS
Hedyosmum |
imys
59
238
Е
la
SGIO TV INVS
à
Sd83HO31Vd
3
SCIODVUsIXYS
N
3
с
| |
58
0n
5
5апоно по
Ephedra torreyana
e 3. Опе of 8897 shortest trees resulting from the exploratory РО pip Би of 194 species of angio-
Bil Each of the shortest trees has a length of 3501 steps, CI = 0.249, and RI = 0.531. Arrows indicate nodes
not present in the strict consensus of all shortest trees. Becis of its Sec ш tree w beet broken into three parts
(3A, 3B, and 3C).
Volume 84, Number 1
Soltis et al.
18S Ribosomal DNA Phylogeny
3B
=
С
=
8
|
n
>
=
=
2
Ё
e
=
=
о
a
2
8
2
5
2
8
5
a
ASTERIDAE S. L. PLUS
EARLY-BRANCHING TAXA
Figure 3B.
3V0ISOY
44 Annals of the
Missouri Botanical Garden
ROSIDAE PLUS
Ceratophyllum
Gunnera
SOMNMINANVA
4
`1 `$ IVAITIAHAOAYVO
3avu5 NV31V2I3
7175 зУаМа15У
3vanialsv
EARLY-BRANCHING
TAXA
Figure 3C. | |
Volume 84, Number 1 Soltis et al. 45
1997 18S Ribosomal DNA Phylogeny
4A
EUDICOTS
Tacca
Bowiea
Chlorophytum
Hippeastrum
Xanthorrhoea
| Helmholtzia
Maranta
ES Zingiber
Costus
Canna
Heliconia
E Musa
UE
S.LODONON
| 8 рта
en
Me -
Colchicum
Calla
3QVvu5 ALVIINSONOW
Oncidium
z Ceratophyllum
Nelumbo
A Aristolochia
| Азагит
i Lactoris
S83HO31Vd
Amborella
[ — Gnetum gnemon
Gnetum nodiflorum
Gnetum €
Ephedra
yg 27. Ephedra rana
SANOYILNO
Figure 4. Strict consensus of 2582 shortest trees resulting from the exploratory ph f 194 species
of angiosperms; m Eg were nis rege in the analyses. Each of the shortest trees im a in of 3507 Moe. "C =
0.249, and RI — 0.536. The letters A and B indicate the occurrence of the indels described in Table 2. Because of
its size, the tree ба been broken = ded parts (4A, 4B, and 4C).
46 Annals of the
Missouri Botanical Garden
ASTERIDAE S. L.
ROSIDAE
Calycanthus
Sassafras
Mkilua
Knightia
Akebia
Ranunculus
Xanthorhiza
lenispermum
Сашорћућит
ر Euptelea
E Dicentra
Hypecoum
г— Trochodendron
7 Tetracentron
Platanus
MONOSCULCATE
GRADE
SGMNONANVY
SALVIINSONOW/SLODICNA YIMOT
Figure 4B.
туар —— Jg
Volume 84, Number 1
1997
Soltis et al.
18S Ribosomal DNA Phylogeny
47
Figure 4C.
в
E
[7]
ONIXIJ N3903.LIN
аиега
Ceratopetalum|
Eu ja
potus
bizi
SGIOINONND
ou
2
||
$ t
$
Зао 15 77132
SALVIONISOSNID
ños
Cercidiphyllum
ا
вешт $
Polygala
lia
Viscum
Gunnera
LOWER EUDICOTS/
MONOSULCATES
SQIO9 viralxvs
SQIO 1V.LN
4C
ЗУО5ОМ
48 Annals of the
Missouri Botanical Garden
4D
Drosera
>
‘
7175 AVGITIAHdOANVD
Acanthogilia
Philadelphus
Hydrangea
TS AVGMALSV
wingia
fos по inal
3vanaisv
Olea
Pach ystachys
Parmentiera
Buadleja
Lamium
Мала)
Pri
Bourr
Pd AA
Mitchell ese
[pomo
Г] Bruntelsia
Montinia
Vahlia
Aeschynanthus
Berzelia
Camptotheca
ROSIDAE
LOWER EUDICOTS/
MONOSULCATES
Figure 4D.
Volume 84, Number 1
Soltis
18S eat DNA Phylogeny
PPENDIX. Proposed se 0 eia for the 18S и. шч of Glycine max er from Nickrent
ed on the rimary
sa 1995). This structural m
h
ach ac 39 42
АЧ Legh ^ СА Ue
Čuauaasagica А
E ANNA A “de ^26 A" "ccá aca ^ ^ccucaa ©си©си
254 чада, GAUACUUUAUdC A сад“ Aa IC cA Bonn SAE dêk el
AcU, "UY y u y PUC UG VUGA 9GAGUU GGA G
З A ay g OVA E ph as
uUa, с-а 9% ura ОТА ЖУА
у ч a-c J^ P Meo)
a-c Es
y 37 ac Я аи,
у аса 0-С, СА
ч e E] ФА oe A9 ^a 40 e-d Sanu
HH oem с 9 ЗЕН 8:9, 5-8
a FONS asa А с- а 926-00
ова UA A s. о, в
AA Any 24 с-а AU cA; ë
e^, yr E23-9 0-с А-0 V u
ARO u.a а
Е23-8 5,7 FER a.u
zu ACA ОТА а.о а Fg
ш 76 ura sau
A € ,qaA-U ae
с aca me АНА
^ u^ v
А ^
Qu ^
uc асисџа ro
e Epi Tf Y
са СаА АС a
AA ^ y Au
E23-7 E ^ м,
а се; 23a Y
G^ ‘Aru v
yt uut 799^ а ^4
AU. Uca.y © E23-1
AD 06 Zu
A as le
"G D
aTe- а
E23-6 рен aS
823-2 СА $6
^
с-а d c
к^ ut А u
v.a 18, Ui
G-c A^^Uc v -c Tu
c-a ш c Au А A-U cac, Uns, QUUAA
v cê e ч n) * ACCU A v iC
a.u Y 2 џа-сУ -A cA, 33 u^
"and Wiek 21 r 19 32 | олег и) 4
с 46-6, | ¡AQUE-G= o x AGGAA "ud Aca A AGG ear А9 New [дА Р
c-a y Us ا tenes etait 4
Е23-5 9: u » ê E дсџаи UUCCCQU o x 4
$-c u u `u? 00000 yg^ C @ А cee
u e €; ARO о 4 uM u- ac-%
б М C оо لا لا A ^ ANA
Aan T u 2 с v 9, А Mop
„ш Ч-А Su 3 5 E Ag ^ Auc AG DA СУ
V чах ` -© и Y LA
17u délalacéló u 8 reste > A, 1000 De
rog
A فر e ٠ لاي c-a” Gaacacau ФА 50 ^ AU
u ¿EE ц 4-ио ve ur^ ey
са с-а, A a у a
^ 1650-G v c a с
^cácaa ^ aau А А A
a dum $-$ H 48 vy4^
AGUCCU d
: £P od d
A, 98-1807
c-a ®©
с-а
Gea
yea 1 тА. с
[^o 2% Д De
ме басе , g oM eU. а y-imo
А Ao аА? а“ ^ А-у
даба КА ^u Y Y
А A u Ави A Ora
14 acuAcc hee Н ас
еа са UCA E 9:0
Лао e ^ae. a’ А * =
5 a
a ar Go
А А А
а, لار o
¿eu UNA GAA^ e.
y $
9. с ЕЮ 10
“е
мее
ѕ Баѕ
1 icula: rly pr e to variation in prim
1369); fii а are difficult to align over a broad өл mi scale and were no
text).
72,
included i in our Ерни антен (ѕее
ЕсКепгоде
ary sequence and
al.,
f Glycine (
ry ноев ions are indicated by hick lin nes. The posi
>
осе
А“
&
om
1995) an
ength (positions, 230-
43
vauaaAc? 4,
Abb, A
СЕРА bau ола
Zoco
e^
TRIBAL RELATIONSHIPS IN
THE GESNERIACEAE:
EVIDENCE FROM DNA
SEQUENCES OF THE
CHLOROPLAST GENE ndhF'
J. Е Smith?, J. C. Wolfram, К. D.
Brown?, С. L. Carroll?, and D. S.
Denton?
ABSTRACT
ecies from the Scro
the closest a -species outgroup fo:
d Paulowni order to be
e tribal relationships of the Gesneriaceae are investigated using ndhF sequences. A e analysis Р 70 tax
grou
tter assess relationships within the family. The smaller analysis resulted in a
, resulted in two ыен
ние Eu of о steps. The Корее. are identified as the sister to the remainder of the way and cou uld
P
potentially i iis ed a
ei place
rioideae is su
e
its Epson upported.
the remaindér of the
oe de aer lineages within
with the a data tribe Trichospo
ement of [pee e у in Cyrtandroideae does not have support from this analysis, whereas
Alternatively, Coronanthereae could be segre,
but in order. to korp a paraphyletic Gesnerioideae would ei
rieae, EN and Beslerieae are identified as mono-
inningia sensu lato отой Within саа анде.
ge, heterogeneous tribe Didymocarpeae are identified, and
reae poa o he pipa ic. The a of chromosome numbers
nodal anatomy, dabis Bod and stem modification are examined based on these molecular trees.
Investigations of higher level cladistic relation-
ships (generic, familial, and above) have recently
drawn a great deal of attention (Annals of the Mis-
souri Botanical Garden Vol. 80(3); Olmstead et al.,
1992, 1993; Donoghue et al., 1992; Cantino, 1992;
Judd et al., 1994). These. analyses have provided
tremendous insights toward our classification sys-
em and process of classification, frequently draw-
ing attention to families that have been separated
on the basis of primarily woody versus herbaceous
taxa (Cantino, 1992; Judd et al., 1994) or tropical
versus temperate (Judd et al., 1994). More recently
an investigation of the Lamiales sensu lato has in-
dicated that the largest family in this order, Scroph-
ulariaceae, is unlikely to be a monophyletic group
усыны et al., 1992, 1993; Olmstead & Reeves,
A thorough investigation of the Scrophular-
laceae ‚шап DNA sequences from both the rbcL
F genes has indicated that the family is
comprised of at least two monophyletic groups with
several genera not having any strict affinity to
Scrophulariaceae or other related families nl
in the analysis (Olmstead & Reeves, 1995).
wise, Olmstead and Reeves (1995) found nn sev-
eral families traditionally segregated from ge
Scrophulariaceae are best included as members
one of the two major lineages (e.g., a repete de
Although most members of the Lamiales s.l. ar
temperate, there are some primarily tropical groups
(Gesneriaceae, Acanthaceae, Bignoniaceae). In or-
der to better assess whether the division between
ese families represents another artificial segre-
gation based on distribution (tropical vs. temperate)
or woody versus herbaceous (e.g., Bignoniaceae vs.
Gesneriaceae), a thorough investigation of the Ges-
neriaceae was deemed necessary to complement
the investigations that have already demonstrated
monophyly of Acanthaceae (Scotland et al., 1995)
and Bignoniaceae (R. Olmstead, pers. comm.), but
have not sampled widely in the Gesneriaceae.
1 We are indebted to the following for sharing plant material: L
n, D. Turley, wart, M.
(AGGS) seed Pie We us ‘thank Richard Olmstead and Michael Kiehn for "ep
Arboretum, M. Killer:
Gesneriad and Gl
ó, B. Nordenstam, R. Dun
хад Somer
us;
E. Skog, W. L. Wagner, J. K. Boggan, Eta
„Је Кини B. Stewart, M. Evans, and the A
comments on the manuscript. Funding for this project was provided by NSF grant DEB-9317775 and a grant fro
AGGS to JFS
2 Depart нем of Biology, Boise State University, 1910 University Drive, уйы Idaho, 83725, U.S.A.
83404, U.S
Current address: 1263 Londonderry, Idaho Falls, Idaho,
ANN. MISSOURI BOT. GARD. 84: 50-66. 1997.
e
Volume 84, Number 1
1997
Smith et al. 51
Tribal e in Gesneriaceae
The Gesneriaceae are a mid-sized to large plant
family comprising approximately 2 500 s
cies in 120-135 genera, distributed primarily in
the tropics with a few temperate species in Europe,
China, and Japan (Heywood, 1978; A. Weber, pers.
comm.). The majority of species in the Gesneri-
aceae are herbaceous perennials, but can be an-
nuals, shrubs, lianas, and trees. Many species
(20%) are epiphytic, and the Gesneriaceae rank
among the top ten plant families in terms of abso-
lute numbers of epiphytic taxa (Madison, 1977;
Kress, 1986). Given the diverse habits of the Ges-
neriaceae, it is not surprising that there is a wide
array of morphological variation within the family.
Corolla tubes may be long and prominent as in Col-
umnea L., or short as in Saintpaulia Wendl. Leaves
are opposite in the majority of the family, but ani-
sophylly, leading to an alternate arrangement with
abscission of the smaller leaf, is common. Many of
these morphologically diverse features of the Ges-
neriaceae are hypothesized as adaptations to the
epiphytic habit (Ackerman, 19
The Gesneriaceae are a Рао, of the Lamiales
s.l. and are distinguished from other families in the
order by the combination of five-lobed corollas, pa-
rietal placentation, and presence of endosperm in
most taxa (Cronquist, 1981). However, because
many of these characters vary within some mem-
bers of the Gesneriaceae (including variation within
individuals of some species), there has been con-
siderable confusion regarding the placement of
some genera. For example, members with axile pla-
centation can be classified incorrectly with the
Scrophulariaceae, and those genera lacking endo-
sperm potentially may be classified with the Acan-
thaceae and Bignoniaceae.
There have been relatively few cladistic analyses
performed within the Gesneriaceae (Kvist, 1990;
Crisci et al., 1991; Boggan, 1991; Smith & Sytsma,
1994a, b, c; Smith, 1996), and only one (Smith,
1996) performed at the tribal level. A cladistic
analysis is desirable to help resolve relationships,
to determine if the family is monophyletic, and to
improve classification within the family by rear-
ranging tribes and subfamilies to reflect phyloge-
netic relationships.
Classifications of the Gesneriaceae traditionally
recognize two subfamilies (Gesnerioideae and Cyr-
tandroideae) (Bentham, 1876; Burtt, 1962, 1977;
Fritsch, 1893, 1894), but others have included an
additional subfamily (Coronantheroideae: Wiehler,
1983; Episcioideae: Ivanina, 1965). The division of
the family is largely based on the uniform (Ges-
nerioideae), or uneven (Cyrtandroideae) enlarge-
ment of the cotyledons after germination (Burtt,
1962). Another character that has been useful in
separating the subfamilies is the presence (Gesner-
ioideae) or absence (Cyrtandroideae) of endosperm
in the seed. In addition, the Gesnerioideae have a
neotropical distribution and most species have in-
ferior or semi-inferior ovaries, whereas the Cyrtan-
droideae are primarily paleotropical with superior
ovaries. However, the geographic distribution and
ovary position are not consistent within the subfam-
ilies. Therefore, although the Cyrtandroideae can
be defined by a synapomorphic character (uneven
cotyledon development), the Gesnerioideae have
been characterized by a symplesiomorphic char-
acter common to dicotyledons in gener
The two subfamilies have been divided further
into 9-17 tribes (Bentham, 1876; Burtt, 1962,
1977; Fritsch, 1893, 1894; Ivanina, 1965; Wiehler,
1983; Burtt & Wiehler, 1995). The classification
schemes differ due to the characters emphasized.
For example, Fritsch (1893, 1894) placed the Col-
umneae in the Cyrtandroideae based on their su-
perior ovary. Later, the Columneae were moved to
the Gesnerioideae due to the presence of uniform
cotyledons (Burtt, 1962, 1977) and combined into
the Episcieae based on nodal anatomy (Wiehler,
1983). This paper presents a cladistic analysis of
DNA sequences in order that phylogenetic relation-
ships among taxa may be more clearly resolved,
and a more stable classification scheme proposed.
The gene ndhF is a chloroplast gene that in to-
bacco encodes a protein of 740 amino acids pre-
sumed to be a subunit of an NADH dehydrogenase
(Sugiura, 1992). The use of ndhF sequences for
systematic studies has provided a far greater num-
ber of characters to resolve relationships than stud-
ies using rbcL. The reasons for the increased num-
ber of characters are that the gene is approximately
50% longer than rbcL (2103 vs. 1431 bp in tobacco
[Wolfe, 1991]) and has a nucleotide substitution
rate that is approximately two times higher than
L based on comparisons of rice and tobacco
(Sugiura, 1989). In recent studies using this gene
in the Acanthaceae, Scotland et al. (1995) found
three times the number of characters compared to
rbcL, and Olmstead and Sweere (1994) discovered
60% more variable characters with ndhF in the So-
lanaceae. Likewise, Clark et al. (1995) have found
that ndhF sequences are informative for resolving
relationships within the Poaceae, and Olmstead and
Reeves (1995) have resolved several clades in a
polyphyletic Serophulariaceae. The larger number
of variable characters makes ndhF sequences ideal
for taxonomic groups that have not been resolved
well using rbcL data, such as members of the As-
~
~
52 Annals of the
Md Botanical Garden
Table 1. i СНА in this study with Genbank submission numbers and voucher specimens. JFS - James
F. Smith, WLW - n L. Wagner, DEB - Dennis E. Breedlove, SI - Smithsonian Institution, LG - Longwood Gardens.
Letters in папа dica Mick where vouchers are deposited.
Genbank
Species Voucher number
]
Achimenes skinneri Lindl. SI 94-606 U62177
pares micranthus C. B. Clarke JFS 643 (WIS) U62169
myla parasitica (Lam.) Kuntze SI 94-570 U62171 !
jonas. meridensis Klotzsch JFS 1182 (WIS) U62158 1
Anna mollifolia (W. T. Wang) W. T. Wang & K. Y. Pan Skog 94-498 U62188 [
Asteranthera ovata (Cav.) Напз!. Stewart 12234 (SRP) U62 3
Besleria affini. rton LG870575 U62162 | Г
оеа hygroscopica Е. Muell. 51 89-04 062205 h
Chirita sinens SI 94-111 U62189
Codonanthe elegans Wiehler SI 82-45 U62178 ,
Colw schiedeana Schlecht. JFS 288 (WIS U62164
Cyrtandra hawaiensis C. B. Clarke WLW 6753 (BISH) U62172 ’
Cyrtandra umbellifera WLW 6701 (BISH) U62165
yrtandromoea acuminata Benth. & Hook JFS 3539 (SRP) U62173
гамета racemiferum Benth I 85-98 U62156 y
Didissandra frutescens Clarke SI 94-512 U62190
Didym us albomarginata H 4-5 U62207 oO
Drymonia зена фину (J. D. Smith) E. Moore JFS 2248 (WIS) U62159 |
Fieldia australis Cunn. Stewart s.n. (SRP) U62196 3
Gasteranthus corallinus (Fritsch) Wiehler SI 94-243 U62163 5
Gesneria pedicellaris Alain SI 94-567 U62192
Gesneria christii Urban 5 U62191 l
Gloxinia sylvatica (HBK) Kunth Dunn 9012051 (SRP) U62157
Hemiboea henryi C. B. Clarke SI 85-157 U62180 j
Kohleria spicata (Kunth) Oerst. SI 94-552 U62181
Lysionotus —€— Maxim SI 94-158 U62182
itraria coc Stewart s.n. (SRP) U62193
Mola A Hat Franch. о U62168
Monopyle macrocarpa Benth. no voucher U62197 |
Napeanthus costaricensis Wiehler U62198 |
n macrostoma Leeuwenberg Feuillet (U U62161
Negria rhabdothamnoides F. Muell. Nordenstam 8608 (S U62195
Nematanthus hirsutus (Mart.) Wiehler Olmstead & Reeves, 1995 L36404
Niphaea oblonga Lindl. SI 78-354 U62160 i
Opithandra primuloides (Miq.) B. L. Burtt SI 93-073 U62183 |
Ornithoboea wildeana Craib. SI 93-075 U62166
Paliavana prasinata (Ker-Gawl.) Fritsch SI 78-368 U62174
Paraboea rufescens (Franch.) Burtt Skog s.n. (US) :
Petrocosmea flaccida Craib SI 85-196 U62184 |
Primulina tabacum Hance SI 93-040 U62167 |
Ramonda myconi (L.) Rehb. Katzenstein s.n. (SRP) U62185
hynchoglossum notonianum (Wall.) B. L. Burtt SI 94-378 U62179 |
Rytidophyllum tomentosum (L.) Mart. SI 77-235 U62200
Rytidophyllum чол Ноок. SI 94-524 U62199
aintpaulia rupi B. L. Burtt SI 94-49 U62176
Sarmienta repens Ruiz & Pavén Stewart s.n. (SRP) U62194
Sinningia (Lietzia) brasiliensis = & Schmidt) Wiehler Dunn 9104014 (SRP) U62175
Sinningia cooperi (Paxt.) W SI 94-340 U62201 |
Sinningia пећи Cla I 94-554 U62186
Solenophora obliqua D. L. Denham & D. N. Gibson DEB 71542 (CAS) U62202 :
Remi holstii Engl. Olmstead & Reeves, 1995 L36415 |
Streptocarpus saxorum Engl. JFS s.n. (WIS) U62170
na oldhamii (Hemsl.) Soler. SI 86-106 U62187
Volume 84, Number 1
1997
Smith et al. 53
Tribal Relationships in Gesneriaceae
Table 1. Continued.
Genbank
Species Voucher number
Vanhouttea lanata Fritsch SI 94-516 U62203
Outgroups
Antirrhinum Olmstead & Reeves, 1995 L36392
Brillantaisia ЕДЫ Benth. Scotland et al., 1995 012654.
Catalpa sp. Olmstead & Reeves, 1995 136397
Celsia arcturus Jacq
Crabbea reticulata C. B. Clarke
Crossandra nilotica Oliv.
Digitalis grandiflora Mill.
Hygrophila corymbosa Lindau
Martinella obovata (HBK) Bureau & K. Schum.
Paulownia tomentosa Steu
Selago thunbergii Choisy
Schlegelia telo (Oerst.) Monachino
Scrophular
Tabebuia Verl (A. de Candolle) Britton
Verbascum tha,
Veronica catenata Pennell
Olmstead & Reeves, 1995 L36398
Scotland et al., 1995 U12655
Scotland et al., 1995 U12656
Olmstead & Reeves, 1995 L36399
еме et al., U12661
stead & Reeves, 1995 L36402
nis: & Reeves, 1995 L36406
Olmstead & Reeves, 1995 L36412
Olmstead & Reeves, 1995 L36410
Olmstead & Reeves, 1995 L36411
Olmstead & Reeves, 1995 L36416
Olmstead & Reeves, 1995 L36417
Olmstead & Reeves, 1995 L36419
teridae and Lamiales s.l. (Olmstead et al., 1992,
1993; Chase et al., 1993).
MATERIALS AND METHODS
The gene sequences used in this analysis were
generated by thermal cycle sequencing (Innis et al.,
88) of previously amplified ndhF regions. The
ndhF gene was amplified in two overlapping sec-
tions (positions 1-1350, and 972-2044) from ge-
nomic DNA isolated from fresh, frozen, or silica gel
dried material (Smith et al., 1992). Once amplifi-
cation products were obtained, the sample was pu-
rified using PCR wizard purification preps (Pro-
mega) according to the manufacturer's instructions.
The purified DNA then was subjected to cycle se-
quencing using the Silver Sequence method (Pro-
mega). This sequencing method resulted in se-
quences that could be read within six to eight hours
after amplification was completed. The products of
one round of amplification provided sufficient ma-
terial for completing the gene sequences described
here. The advantages of the silver staining proce-
dure over radioactive methods are safety, minimal
waste disposal, and speed.
CHOICE OF TAXA
The focus of this analysis was on the tribal re-
lationships of the Gesneriaceae and comparison of
the results of this analysis with one based on mor-
phological data (Smith, 1996). Genera were select-
ed to represent current and previous tribal classi-
fications within the family and, whenever possible,
to match genera used in the morphological analysis.
In some instances, a genus that had been used with
the morphological analysis (Smith, 1996) was not
readily available for the molecular analysis. There-
fore this molecular analysis contains many taxa that
have not been included in the morphological anal-
ysis, and direct comparisons will be made with a
reduced data set at a future date. The species used
in the analysis, voucher information, and Genbank
accession numbers are included in Table 1.
Genera have been selected to represent the most
recent tribal classifications with two to ten genera
from each tribe (Tables 2 and 3). In order to rep-
resent current classification systems along with ear-
lier systems, 48 genera were selected (Tables 2 and
OUTGROUP SELECTION
Outgroups were selected to root the tree repre-
senting tribal relationships within Gesneriaceae.
The best method for doing this is by outgroup com-
parison (Donoghue & Cantino, 1984; Maddison et
al., 1984). The most appropriate outgroup for the
tribes of the Gesneriaceae should be the most
closely related plant family or clade. The Gesner-
iaceae have been placed in the order Lamiales s.l.
in the subclass Asteridae (or equivalent groups of
families) in numerous taxonomic treatments (Dahl-
gren, 1975; Thome, 1976, 1983, 1992; Heywood,
1978; Takhtajan, 1980; Cronquist, 1981). However,
Annals of the
Missouri Botanical Garden
e 2. Genera of Gesnerioideae (Burtt & Wiehler, 1995) used and their classification status. NT = not treated.
Subfamilial names are underlined to readily distinguish them from tribal names. Subtribal names are abbreviated as
Colum. - Columneineae, Codon. - Codonanthinae.
Genus Wiehler, 1983 Ivanina, 1965 Fritsch, 1893-94
Gesnerioideae Gesnerioideae Gesnerioideae
Achimenes Gloxinieae Gloxinieae Gloxinieae
Gloxinia Gloxinieae loxinieae Gloxinieae
Monopyle Gloxinieae Bellonieae Bellonieae
Niphaea Gloxinieae Bellonieae ellonieae
Kohleria Gloxin le Kohlerieae
Diastema Gloxinieae Kohlerieae Kohlerieae
Sinningi Gloxinieae Kohlerieae Sinningieae
Vanhouttea Gloxinieae Kohlerieae Kohlerieae
aliavana Gloxinieae Reichsteinerieae па а
легла Gloxinieae Reichsteinerieae Sinningi
Solenophora Gloxinieae Solenophoreae apes hie
sneria Gloxinieae Gesne: Gesnerieae
Rytidophyllum =Gesneria Gesnerieae Gesnerieae
Episcioideae Cyrtandroideae
Columnea Episcieae Columneae Columneae-Colum
Codonanthe Episcieae Columneae Columneae-Codon
Nematanthus Episcieae Columneae Columneae-Colum
Alloplectus Episcieae Episcieae Columneae-Colum
rymonia Episcieae Episcieae Columneae-Colum
esleria Beslerieae Episcieae Beslerieae
Gasteranthus Beslerieae
Napeantheae Episcieae Klugieae
Coronantheroideae Cyrtandroideae
Asteranthera oronanther: Mitrarieae Coronanthereae
Sarmienta Coronanthereae Mitrarieae Coronanthereae
Mitraria Coronanthereae Mitrarieae Coronanthereae
ieldia Coronanthereae itrarieae Coronanthereae
Negria Coronanthereae Coronanthereae Coronanthereae
the relationships among these families are some- | PHYLOGENETIC ANALYSIS
what i
ous. A recent cladistic analysis of
these families based on DNA sequencing of the
chloroplast encoded rbcL gene resulted in poor res-
olution of the relationships of these families (
stead et al., 1993), ough these relationships
have been more resolved with the addition of ndhF
sequences (Olmstead & Reeves, 1995).
ee families from the Lamiales s.l. were used
as outgroups for this analysis. These were the
Acanthaceae, Bignoniaceae, and Scrophulariaceae.
Olm-
& Reeves, 1995; Scotland et al., 1995) and includ-
ed representatives from three lineages identified
within the Scrophulariaceae (Olmstead & Reeves,
1995). Initial analyses used all 16 species as the
outgroup. Subsequent analyses used only Gesneri-
aceae with Paulownia Sieb. & Zucc. as the out-
group.
Phylogenetic divergence was reconstructed using
PAUP version 3.1.1 ш 1993) to implement
Wagner parsimony (Farri ; Farris et al.,
1970; Swofford & e 1987). This program
allows parallelisms and reversals (homoplasy), and
provides an option for missing data. In this analy-
sis, trees were generated using the general heuristic
option, saving minimal trees only, with the collapse
zero-length branches, and ignore uninformative
characters options in effect. Because of the large
number of taxa in this analysis, the branch and
bound and exhaustive search options would have
consumed an excessive amount of time. Therefore,
the trees presented here are best approximations
and not exact solutions. The manner in which the
program reconstructs phylogenetic sequences is
sensitive to the of taxa presentation in the
data matrix, frequently finding islands of equally
parsimonious trees depending on the order (Mad-
Volume 84, Number 1
97
Smith et a
Tribal Relationships in Gesneriaceae
we 3. Genera of Cyrtandroideae (Burtt & Wiehler, 1995) used and their classification status. NT = not treated.
The tribe, Didymocarpeae, is abbreviated Didy. in order to show the subtribal classification system of Ivanina (1965)
).
and Е ка: (1893, 1894
Genus Burtt, 1962,77 Ivanina, 1965 Fritsch, 1893-94
Didymocarpeae Ramondeae Ramondeae
Saintpaulia Didymocarpeae Saintpaulieae Ramondeae
Opithandra Рлдутос Didy.-Roettlerineae NT
Didymocarpus Didymocarpeae Didy.-Roettlerineae Didy.-Roettlerineae
Didissandra Didymocarpeae Didy.-Roettlerineae Didy.-Oreacharineae
Anna Didymoc. Didy.-Roettlerineae NT
Chirita Didymocarpeae Didy.-Roettlerineae Didy.-Roettlerineae
Petrocosmea Didymocarpeae Didy.-Roettlerineae Ramondeae
Titanotrichum lous NT NT
Cyrtandromoea Loxonieae/Scroph Klugieae Beslerieae
Paraboea ymocarpeae Didy.-Roettlerineae Didy.-Roettlerineae
oea Didymocarpeae NT Streptocarpeae
Hemiboea Didymocarpeae Didy.-Roettlerineae NT
Primulina Didymocarpeae Didy.-Roettlerineae jeae
Streptocarpus Didymocarpeae Didy.-Streptocarp treptocarpeae
Ornithoboea Didymocarpeae Didy.-Streptocarp Streptocarpeae
Aeschynanthus Trichosporeae Trichosporeae chosporeae
Agalmyla Trichosporeae Trichospor: Trichosporeae
Lysionotus Trichosporeae Trichosporeae Trichosporeae
Cyrtandra Cyrtandreae Cyrtandreae Cyrtandreae
Rhynchoglossum Klugieae Klugieae Klugieae
Monophyllaea Klugieae lugieae Beslerieae
dison, 1991). Therefore, it is important to repeat
е analysis several times. To do this, the search
strategy of Olmstead and Palmer (1994) was imple-
mented: searching for 1000 trees each in five sub-
sequent analyses with the nearest — inter-
chan "tis (NNI) search option in effect an ars
"off." Each of the results from the five NNI d:
es was ud: as the starting tree(s) for a -— with
tree bisection reconnection (TBR) and mulpars
“on.” This strategy was used in the full analysis
with all 16 iom бини taxa designated as
outgroups. Likewise, the same strategy was used
with only the members of the Gesneriaceae and
aulownia as the outgroup, and with constraints
options.
Branch support analysis was performed to ex-
amine trees that were six or fewer steps longer than
the most-parsimonious tree (Bremer, 1988; Dono-
ghue et al., 1992; Bremer, 1994). This type of anal-
ysis provides an indication of the robustness of the
data by determining which clades persist in a con-
sensus tree as parsimony is relaxed. This analysis
was performed by saving all trees six steps longer
than the most-parsimonious trees and then exam-
ining subsets of trees one to six steps longer with
the filter option of PAUP.
ndhF sequences used here had several six
to twelve base pair insertions or deletions (indels)
inferred from gaps in the sequence alignments,
which in previous analyses had been re-scored as
binary characters and used as either an indepen-
dent data set or combined with the sequence data
phylogenetic importance (Scotland et al.,
995); therefore indels found in the Gesneriaceae
were removed and examined independently of se-
quence data for their oar. utility.
The monophyly of various tribal relationships not
obtained in the fubinde trees was ex-
amined by using the constraints option of PAUP.
These included the Trichosporeae, the Didymocar-
peae, the inclusion of Klugieae in Cyrtandroideae,
inningieae in Gloxinieae. Also, since the
analysis with all 16 outgroup taxa resulted in the
placement of Nematanthus Schrader and Klugieae
in discrepant positions from traditional classifica-
tions, an analysis with all 16 outgroups constrained
Nematanthus to the Gesnerioideae, and the Klu-
gieae from the Gesnerioideae. The position of Klu-
gieae and Nematanthus was also examined by con-
structing a user-defined tree with a topology of one
of the two most-parsimonious trees except that Ме-
matanthus was placed in the Episcieae, and Klu-
gieae was placed as sister to the remainder of the
Gesneriaceae. This user-defined tree was then the
Annals of the
Missouri Botanical Garden
starting tree for a search using TBR and mulpars
"onim
RESULTS
Complete sequences for the ndhF gene were ob-
tained for 52 species of Gesneriaceae (Table 1).
These sequences were supplemented with sequenc-
es from an additional 18 species (2 within Gesner-
iaceae and 16 from related families) from Genbank
(Table 1). The complete sequences resulted in 849
phylogenetically informative characters among all
70 species in the full analysis. А smaller analysis
focused on only the Gesneriaceae species with
Paulownia as the outgroup. Within this smaller
analysis 690 nucleotide positions were found to be
phylogenetically informative. Indels were found at
several positions in the Gesneriaceae from the se-
quences used in this analysis. Two widespread in-
sertions were a 12 bp insertion at position 1440
and a 6 bp insertion at 1548. Other insertions were
autapomorphic for species or genera used in the
analysis (unpublished results). No insertions were
used in the analysis. The 6 bp insertion was sym-
plesiomorphic for the Gesneriaceae. The 12 bp in-
sertion was also symplesiomorphic for the Gesner-
iaceae; however, sequence divergence within this
insertion provides an additional Aas aig for
the clade comprised of Columnea, Dı Mart
and Alloplectus Mart. (Fig. 4), where a siia base
pair transition characterizes these three genera.
Other base pair substitutions and insertions were
found within this 12 bp insertion but, with the cur-
rent level of sampling, were autapomorphic.
Cladistic analysis was performed initially with all
70 taxa of the four families (Gesneriaceae, Scroph-
ulariaceae, Acanthaceae, and Bignoniaceae) and all
taxa in the three outgroup families designated as
the outgroup. This analysis resulted in two trees of
0 steps each ie index (CI) = 0.30,
retention index (RI) = 0.48), all of which indicated
esneriaceae were a monophyletic family and
that the genus Paulownia (Scrophulariaceae) was
the closest outgroup (Figs. 1, 2).
Subsequent analyses were performed to minimize
computer analysis time that utilized only the Ges-
neriaceae and Paulownia as a designated outgroup.
This reduced analysis resulted in a single most-
а
thought to be monophyletic, or comprised tribes,
were examined using the constraints option of
PAUP to determine the impact of the monophyletic
grouping on the remainder of the data and to de-
termine the number of additional steps required to
construct these trees. The analysis required four
additional steps to create a monophyletic Trichos-
poreae, five for a monophyletic Didymocarpeae, two
to include the Klugieae in the Cyrtandroideae, and
four to include the Sinningieae in the Gloxinieae.
Constraining the analysis of all 70 taxa to place
Klugieae as the sister to the Gesneriaceae and Ne-
matanthus within the Episcieae resulted in four
trees 58 steps longer than the most-parsimonious
trees regardless of whether the constraint option of
PAUP, or user defined trees were implemented.
DISCUSSION
The cladistic analysis of 54 species of Gesneri-
aceae with 16 species of Scrophulariaceae, Bigno-
niaceae, and Acanthaceae as outgroups resulted in
a monophyletic Gesneriaceae with the single genus
Paulownia (Scrophulariaceae) indicated as the
closest outgroup (Figs. 1, 2). These results verified
that the Gesneriaceae are distinct from other mem-
bers of the Lamiales s.l. and not an artificial unit
based on their largely tropical distribution and her-
baceous habit as has been seen for some family
pairs (Judd et al., 1994). The full analysis is largely
in agreement with the position of the taxa in the
reduced analysis with the exception of the positions
of Nematanthus and the tribe Klugieae. The place-
ment of Nematanthus as the sister to the remainder
of the family is very far removed from its traditional
classification within the Episcieae (Fig. 1). Like-
wise the Klugieae are placed unusually in the sub-
family Gesnerioideae (Fig. 2). The most likely ex-
planation for the anomalous placement of these taxa
is the high level of homoplasy between the Ges-
neriaceae and the outgroups. This is exemplified
when 15 of the 16 outgroup species are removed
from the analysis. In the reduced analysis both Ne-
matanthus and Klugieae are in more expected po-
sitions regarding relationships to the remainder of
the family. An alternative explanation is that be-
cause of the size of the data set, PAUP did not find
the shortest tree and that a shorter tree with all 70
species exists that places Nematanthus and the
Klugieae in their more expected relationships. This
latter explanation is unlikely since searches con-
straining these taxa to their more traditional posi-
tions, or a user-defined tree that placed them there,
resulted in four trees that were 58 steps longer.
The reduced analysis resulted in a single most-
parsimonious tree (Figs. 3, 4). Three major mono-
phyletic divisions within the family correspond to
subfamilies Gesnerioideae and Cyrtandroideae (mi-
nus tribe Klugieae) and tribe Klugieae in a separate
position as a potential third subfamily. Traditional
Volume 84, Number 1
1997
Smith et a
Tribal Relationships in Gesneriaceae
57
Figure 1.
the оноор taxa, BI—
aceae,
erioidea:
Bi nece AC—
Acanthaceae, SC—Scro
Gesnerioideae
Cyrtandra hawaiensis
Cyrtandra umbellifera
Hemiboea
Lysionotus
Aeschynanthus
Petrocosmea
Opithandra
Anna
Didissandra
Didymocarpus
Ornithoboea
Paraboea
Streptocarpus saxorum
Saintpaulia
Streptocarpus holstii
Ramonda
Chirita
Titanotrichum
Nematanthus
Paulownia
GE
Crabbea
Crossandra
Schlegelia
subfamilies of the
am
EE
"је
АС
BI
Gesneri
Strict consensus of two most-parsimonious trees of 5610 steps each (CI = ig 30, RI = 0.48) ar a dag
phulariaceae, and th
n
and CY—Cyrtandroideae. The Ead ie of the Ска are displayed in Figure 2.
GE—Gesn
See text for explanation of position of Nematanthus in this cladogram
58 Annals of the
Missouri Botanical Garden
Diastema
Monopyle
Achimenes
Kohleria
Sinningia brasiliensis (Lietzia)
Sinningia ri
Gesneria christii
Gesneria pedicellaris
Rytidophyllum tomentosum
Rytidophyllum auriculatum
= Em Codonanthe
Solenophora
г Gloxinia
л Niphaea
Alloplectus
neti 2) Columnea
Drymonia
Paliavana
Sinningia cooperi
LEE vanhouttea
Napeanthus macrostoma
p ae Napeanthus costaricensis
Besleria
Gasteranthus
GE
Monophyllaea
Rhynchoglossum
Cyrtandromoea | —— SC/CY
| Asteranthera ————— GE
Mitraria |
2: Sarmienta
| СЕ
Lp
Fieldia
Figu e 2. Strict consensus of two most-parsimonious trees of 5610 steps each (CI = 0.30, RI = 0. 48) displaying
Goon iinet and some CY—Cyrta niaide. eos has been placed in either the SC—Scrophu-
e
нео г the Cyrtandroideae and is eee uch on this figure. The remainder of the Cyrtandroideae are
displayed in Figure 1. See text for the explanation of ists of the tribe Klugieae (Kl) in this cladogram.
Fig. 1 ——
коме о Om
Volume 84, Number 1 Smith et al. 59
1997 Tribal Relationships in Gesneriaceae
Gesnerioideae
4
Cyrtandra hawaiensis
Су
Cyrtandra umbellifera
Hemiboea —————— pj
Lysionotus ———— Tr
ai Aeschynanthus Tr
(14)
Petrocosmea Di
58(55 j
(18) 2403) | Opithandra b
а 1 $68 Anna
52046) +...
24(23) 288 Didissandra Di
109) »6 la Didymocarpus
Ornithoboea
Di
Primulina
15(11) Agalmyla —————— Tr
2 Boea
Di
Paraboea
35(31) ara Streptocarpus saxorum
же arad NE cana; |
6 | Saintpaulia Di
24(21
48(38) 3 Streptocarpus holstii
2500) Ramonda ———— Di
22
mus Chirita ————— Di
Titanotrichum —— — Т;
Monophyllaea
41(40) Rhynchoglossum Kl
Cyrtandromoea
US руна ——-———— "SC
Figure Single most- -parsimonious tree of 4613 steps (CI — 0.29, RI — 0.38) from the analysis of the species in
the на ia with only Paulownia (SC—Scrophu unen d — ted as the outgroup. Displayed in this figure are
the tribes of the Cyrtandroideae, KI—Klugieae, Ti—Titan eae, Di—D idy ymocarpeae, Tr—Trichosporeae, and Cy—
Cyrtandreae. The Fins pese are displayed in Figure 4. гаг
those clades. Numbers ntheses indicate those synapomorphies es are cenae i in this tree. Numbers below
branches are decay ux тош with no value indicated have a decay value
60 Annals of the
Missouri Botanical Garden
Diastema
Solenophora
Monopyle
Gloxinia Gl
Niphaea
Achimenes
Kohleria
Gesneria christii
Gesneria pedicellaris
Rytidophyllum auriculatum
Rytidophyllum tomentosum |
Alloplectus
Drymonia
Columnea Ep
Codonanthe
Nematanthus
Paliavana
Vanhouttea
Sinningia cooperi Si
Sinningia brasiliensis (Lietzia)
Sinningia richii
Napeanthus macrostoma || Ма
Napeanthus costaricensis
Besleria | Ве
Gasteranthus
Mitraria
Sarmienta
Negria Co
Fieldia
Asteranthera
Figure Single most-parsimonious tree of 4613 steps (CI = 0.29, RI = 0. 38) from the ае of the species in
the Coca with only dió pe as the outgroup. Displayed in this figure are the tribes of the
, Co—C eslerieae, Na—Napeantheae, Si—Sinningieae, ке honda Ge—Ges-
nerieae, and C E The Crtadnidea: are displayed in deae 3. Numbers along branches are the syna-
pomorphies that support those clades. Numbers i
this tree. Numbers below branches are decay ilie. Branches with no Supr indicated have a decay value of 1.
Volume 84, Number 1
1997
Smith et al. 61
Tribal Relationships in Gesneriaceae
classification schemes have placed tribe Klugieae
in the Cyrtandroideae (Table 3); however, the in-
clusion of tribe Klugieae within subfamily Cyrtan-
droideae would result in a paraphyletic Cyrtandroi-
deae. The removal of this tribe to a third subfamily
would result in a monophyletic Cyrtandroideae. The
monophyletic groups within the subfamily Gesner-
ioideae correspond highly with traditional classifi-
cation systems for this subfamily (Wiehler, 1983)
and a previous cladistic analysis based on morpho-
logical data (Smith, 1996). The relationships within
the Cyrtandroideae are less congruent with previ-
ous taxonomic treatments, mainly due to the limited
understanding and sampling of the large, hetero-
geneous tribe Didymocarpeae (Burtt, 1962).
SUBFAMILIAL GROUPINGS
The separation of the Gesneriaceae into two sub-
families (including Coronanthereae in the Gesner-
ioideae) has become well accepted during the past
30 years since the discovery of unequal cotyledon
enlargement in the Cyrtandroideae (including mem-
bers of the tribe Klugieae) and equal cotyledon en-
largement in the Gesnerioideae (Burtt, 1962). How-
ever, from a cladistic viewpoint the Cyrtandroideae
are defined by a synapomorphy whereas the Ges-
nerioideae are defined by a symplesiomorphy. One
problem with this character is that it has not been
examined thoroughly for all members of the differ-
ent subfamilies, including many of the taxa used in
this analysis.
Although an analysis of morphological data that
included cotyledon expansion did not support the
monophyly of the Cyrtandroideae (Smith, 1996), the
cladistic analysis of ndhF sequences presented
here demonstrates both a well-supported monophy-
letic Cyrtandroideae (Klugieae excluded) and Ges-
nerioideae (Figs. 3, 4). The monophyly of the Ges-
nerioideae is supported in both a morphological
analysis (Smith, 1996) and this molecular analysis
(Fig. 4). The Cyrtandroideae were paraphyletic in
a cladistic analysis of morphological data (Smith,
1996) but are well supported with ndhF sequences
(Klugieae excluded), although the position of Titan-
otrichum Solereder as sister to the remainder of the
Cyrtandroideae is supported with only 22 homo-
plastic character state changes (Fig. 3).
The placement of Coronanthereae within the
Gesnerioideae is well supported with ndhF se-
quences (Fig. 4) as it is with morphological data
(Smith, 1996). This tribe does not belong in the
subfamily Cyrtandroideae as had been proposed
earlier (Fritsch, 1894). Wiehler (1983) in his treat-
ment of the neotropical Gesneriaceae suggested a
separate subfamilial status for Coronanthereae due
the numerous autapomorphic characters pos-
sessed by members of this group, such as fusion of
the nectary to the ovary wall and high chromosome
numbers (Wiehler, 1983). The morphological data
would allow the Coronanthereae to be either a
monophyletic tribe within the Gesnerioideae or a
separate monophyletic subfamily without disrupting
the taxonomy of an r group (Smith, 1996).
However, based on the molecular data presented
here, if the Coronanthereae were raised to subfam-
ily level, it would either include the tribes Napean-
theae and Beslerieae from the Gesnerioideae or ne-
cessitate elevating these two tribes as an additional
subfamily (Fig. 4). Therefore it is recommended
that the Coronanthereae be treated as a tribe of the
Gesnerioideae rather than a separate subfamily.
TRIBAL RELATIONSHIPS
GESNERIOIDEAE
Among the relationships within the Gesnerioi-
deae, the primary lack of congruence between this
analysis and the most recent classification scheme
by Burtt and Wiehler (1995) is the polyphyly of the
Gloxinieae. However, the removal of Sinningia
Nees (including Lietzia Regel, but not including
Paliavana Vandelli or Vanhouttea Lem.) has been
proposed previously Pics 1893, 1894) as the
tribe. Sinningieae. The monophyly of Paliavana,
Lietzia, and Sinningia has ym proposed by Bog-
gan (1991), where all three genera were proposed
to be members of Sinningia as the result of a mor-
phologically based cladistic analysis of Sinningia
distic analysis (Smith, 1996), most likely due to
limited sampling among these taxa (Sinningia sen-
su stricto was represented only by Sinningia sect.
Corytholoma and Vanhouttea was not included).
The results presented here indicate that Sinningia
(including the recently combined Lietzia), Palia-
vana, and Vanhouttea should be removed from
Gloxinieae and placed in a separate monophyletic
tribe Sinningieae. Although Sinningia is paraphy-
letic in this analysis (Fig. 4), limited sampling from
this large genus leads only to a tentative conclusion
th Paliavana and Vanhouttea should be
combined into Sinningia to create a monophyletic
genus.
The sister relationship of the Beslerieae and Na-
peantheae has been hinted at based on the overlap
of several diagnostic characters between these
tribes (Skog, 1995; Skog & de Jesus, 1996). How-
ever, the sister relationship of these two tribes to
Annals of the
Missouri Botanical Garden
the Coronanthereae (Fig. 4) has not been proposed
previously. Although the morphological data did not
indicate sister group status, the data did indicate a
close affinity among these three tribes (Smith,
nóng the recent classification schemes pro-
posed for the Gesnerioideae, Burtt and Wiehler's
(1995) is the closest approximation to the results
obtained in this study. The subdivision of Wiehler's
(1983) Gloxinieae into the Bellonieae, Kohlerieae,
— Án and Solenophoreae (Ivanina,
able 2) is not supported by this cladistic
аный Likewise ме бин y Wiehler’s (1983)
Gloxinieae into Bellonieae, Kohlerieae, and Solen-
ophoreae (Table 2; Fritsch, 1893, 1894) is not sup-
ported except for the removal of the Sinningieae
(Fig. 4), which would also necessarily include Pal-
iavana and Vanhouttea (included in Fritsch’s Koh-
The placement of Napeanthus G.
gieae (Cyrtandroideae) (Table 2)
as proposed by Fritsch (1893, 1894) is inappropri-
ate.
CYRTANDROIDEAE
Burtt’s (1962, 1977) classification system for the
Cyrtandroideae is closer in agreement to this cla-
distic analysis than previous classification schemes
(Ivanina, 1965; Fritsch, 1893, 1894). However, the
monophyly of the largest tribe, the Didymocarpeae,
is not supported by this analysis (Fig. 3). Likewise
none of the subtribes created by Ivanina (1965) or
Fritsch (1893, 1894) are supported as monophyletic
ied (Fig. 3, Table 3). The Trichosporeae are not
upported as a monophyletic clade (Fig. 3). Al-
dough this tribe was well supported in the mor-
phological analysis (Smith, 1996), four а ва
steps beyond the most-parsimonious tree аге ге-
quired to make this clade monophyletic with ndhF
ta.
The position of Titanotrichum has been problem-
atic, although this genus has consistently remained
in the Gesneriaceae (Burtt, 1962, 1977; Wang et
al., 1992; Burtt & Wiehler, 1995). Titanotrichum is
a member of the Cyrtandroideae based on these
data, and perhaps may be viewed best as a mono-
typic tribe (Titanotricheae; Wang et al., 1992), sis-
ter to the remainder of the subfamily. However, the
position of Titanotrichum as the sister to the re-
mainder of the Cyrtandroideae is only weakly sup-
ported with 22 homoplastic character state changes,
and the resolution of its placement is lost in the
strict consensus of all trees only one step longer
than the most-parsimonious tree. Therefore, it is
likely that Titanotrichum, or the lineage leading to
this species, diverged early in the evolution of the
family. The placement of Titanotrichum within the
Gesneriaceae is discussed elsewhere (Smith et al.,
997).
—
The Didymocarpeae are a large heterogeneous
tribe that includes the majority of genera in the
Cyrtandroideae (Burtt, 1962, 1977; Wang et al.,
1992). In this analysis it is a paraphyletic assem-
blage that includes the Cyrtandreae and Trichos-
poreae (Figs. 1, 3). Because of the large size of the
Didymocarpeae, and the limited sampling of the
tribe in this analysis, no conclusions regarding its
monophyly, or potential division into other tribes,
are recommended at this time. Further morpholog-
ical investigations in this tribe are under way (B.
L. Burtt and A. Weber, pers. comm.), and a cladis-
tic analysis that focuses on this group will be valu-
able toward understanding its relationships. Several
well-supported nip dg groups within the Di-
dymocarpeae can be identified (Boea Commerson
ex Lamarck/Paraboea (C. B. Clarke) Ridley, Hem-
iboea C. B. Clarke/Lysionotus D. Don, Didissandra
C. B. Clarke/Didymocarpus Wallich, and Strepto-
carpus Lindl. les у It should be noted that
Didissandra an carpus, although forming a
monophyletic ذا in this analysis, are both large
heterogeneous genera and that sampling different
species may have resulted in different placement.
By focusing on morphological characters of these
groups it may be possible to identify more inclusive
monophyletic tribes out of the paraphyletic Didy-
mocarpeae. Much greater sampling within this large
group will be necessary before any major realign-
ment can begi
An ceci result of this analysis is the pa-
raphyly of Streptocarpus. The most likely explana-
tion for this paraphyly is limited sampling, with
only two species of Streptocarpus and one of Saint-
paulia. However, it is interesting to note that Saint-
рашћа is one of the few genera within the Gesner-
iaceae to have a chromosome number of n = 15
(Skog, 1984). The only other genera that share this
number are some species of Streptocarpus, includ-
ing both 5. saxorum and S. holstii, and some spe-
cies of Aeschynanthus Jack (Skog, 1984). The pos-
sibility that Saintpaulia is derived from within
Streptocarpus, as indicated by ndhF sequences and
chromosome numbers, currently is being investi-
gated with greater sampling.
The Trichosporeae traditionally have been
viewed as a monophyletic tribe defined by the pres-
ence of seed appendages not present elsewhere
within the family (Burtt, 1962, 1977; Wang et al.,
1992). Based on morphological data, the Trichos-
poreae were one of the most strongly supported
Volume 84, Number 1
1997
Smith et al. 63
Tribal Relationships in Gesneriaceae
tribes in a morphology-based cladistic analysis
(Smith, 1996). However, it is apparent from this
analysis of ndhF sequences that the selection of
characters that define the Trichosporeae is inappro-
priate (e.g., seed appendages are common in the
closely related Bignoniaceae). Alternatively, it is
possible that inadequate sampling from the Tri-
chosporeae or the large tribe Didymocarpeae may
be causing the separate placement of the three gen-
era sampled from the Trichosporeae. This latter hy-
pothesis is unlikely, because one of the more
strongly supported clades in the analysis placed Ly-
sionotus (Trichosporeae) with Hemiboea (Didymo-
carpeae) and us from the other genera of the Tri-
wende (Fig. 3
KLUGIEAE
The Klugieae are monophyletic and are the sister
group to the remainder of the Gesneriaceae (Fig.
3). The placement of this tribe in the Gesnerioideae
(Fig. 2) in the full data analysis most likely is due
to homoplasy or the result of an incomplete search
for the shortest tree. The monophyly and sister
group status of this tribe also was supported with a
cladistic analysis of morphological data (Smith,
The Klugieae possess numerous autapo-
morphic characters relative to other Gesneriaceae
such as narrow medullary rays, and verrucate edges
of the cells of the seed coat (Smith, 1996). The
placement of Cyrtandromoea Zoll. in the Klugieae
Cyrtandroideae was proposed previously by
Ivanina (1965), although other investigations indi-
cated that this genus should be excluded from the
Gesneriaceae on the basis of floral anatomy (Burtt,
1965; Singh & Jain, 1978). The placement of Cyr-
tandromoea in the Gesneriaceae is discussed else-
where (Smith et al., 1997).
EVOLUTION OF NON-MOLECULAR CHARACTER STATES
CHROMOSOME NUMBERS
v chromosome counts are synapomorphic
and non-homoplastic based on this cladistic anal-
ysis. Large numbers of chromosomes (n = 30+) are
unique to the Coronanthereae and would serve as
an additional character to separate this tribe from
the remainder of the family (Skog, 1984). A chro-
mosome base number, x, of 14 characterizes the
Gesnerieae (Wiehler, 1983; Skog, 1984). The cla-
distic analysis of morphological data was unable to
separate the Gesnerieae from the tribe Gloxinieae
though it represented a monophyletic group with-
in it (Smith, 1996). The inclusion of chromosome
numbers (which were excluded due the large num-
ber of character states) might have removed Ges-
nerieae from Gloxinieae as seen here with sequence
ta.
Most Gloxinieae sampled here (excluding Solen-
ophora Benth., Niphaea Lindl., chimenes
Pers.) have x — 13 (Wiehler, 1983; Skog, 1984).
In addition, the members of the Sinningieae that
have been examined also have x — 13 (Skog,
1984). This similarity in chromosome base number,
along with other character states, has led previous
researchers to include the members of the Sinnin-
gieae within the Gloxinieae (Wiehler, 1983). How-
ever, based on the analysis presented here, the Sin-
ningieae are best viewed as a tribe separated from
the Gloxinieae, and x — 13 is homoplastic.
Other homoplastic chromosome numbers are x —
11 (Niphaea and Achimenes), and x — 9 (Alloplec-
tus, nia, Columnea, and some Didymocarpus
species). Although most of these homoplastic
counts serve little irr utility, the count of
X — 9 serves to characterize a portion of the Ep-
iscieae. Most members s “the Episcieae have x =
9, but taxa with x = 8 (Codonanthe (Mart.) Hanst.
and Nematanthus) may represent another clade
(Fig. 4). Further sampling within the Episcieae may
reveal if this clade (Fig. 4) continues to be sup-
ported or is the result of sampling in this analysis.
Other chromosome counts in the Cyrtandroideae
are highly variable even within genera, and no pat-
tern emerges from the counts of the species that
have been included in the analysis, with the ex-
ception of the Streptocarpus/Saintpaulia counts dis-
cussed above.
NODAL ANATOMY
Another useful character for the Gesneriaceae is
nodal anatomy (Wiehler, 1983). Unfortunately only
the subfamily Gesnerioideae has been sampled
thoroughly for this character, and the lack of data
for the Cyrtandroideae necessitated the exclusion
of this character from the morphological analysis
(Smith, 1996). However, if nodal anatomy is
mapped onto the trees from this molecular analysis,
this character can provide useful phylogenetic in-
formation. The tribe Episcieae (Fig. 4) is defined
by a three-trace trilacunar node that is unique
among the Gesnerioideae, although this character
state is known from the Cyrtandroideae. The unique
presence of this character state within the Gesner-
ioideae adds further support to the monophyly of
the Episcieae. The three-trace trilacunar node may
be symplesiomorphic for the Cyrtandroideae, as all
taxa with available data for this character (Saint-
paulia, Streptocarpus, and Cyrtandra Forster & For-
ster) possess a three-trace trilacunar node except
Annals of the
oh Botanical Garden
Aeschynanthus, which has a one-trace trilacunar
node common to the Gesnerioideae.
PLACENTA
The placenta in the Gesneriaceae is either intact
or divided to the base (Ivanina, 1965). This char-
acter was included in a cladistic analysis of mor-
phological data and served as a DARS state that
brought the Episcieae, Besle and Napean-
Чет together in a single 5 (Smith 1996) as
e only taxa sampled that had divided placentae.
Ah this character state is consistent with the
relationship between the Napeantheae and Besler-
ieae, the character state is homoplastic between the
Episcieae and Napeantheae/Beslerieae based on
the data presented here (Fig. 4)
STEM MODIFICATION
Several members of the Gesneriaceae possess
modifications of the stems (rhizomes and tubers),
presumably as adaptations to periodic dry seasons
(Wiehler, 1983). The presence of scaly rhizomes is
und almost exclusively, and is widespread, within
the Gloxinieae (Wiehler, 1983). Among the taxa
sampled here, the presence of scaly rhizomes
serves as a synapomorphy for the tribe Gloxinieae,
Y
rhizomes also are known from
the Cyrtandroideae, uh Titanotrichum (Kao
& DeVol, 1972; Wan
Tubers are Tanen among species of Sinnin-
gia including Lietzia,which has recently been com-
bined into Sinningia (Wiehler & Chautems, 1995).
Although tubers serve to unite these genera, and to
separate them from the Gloxinieae, tubers are not
known from Paliavana or Vanhouttea. However, not
all species of Sinningia are tuberous, and the lack
of tubers in these species can be regarded as intra-
tribal or intra-generic variation. Tubers also are
known from several species in the Episcieae pta
page Chrysothemis Dene., Nautilocalyx Lind. e
, Paradrymonia Hanst., and корага
геена as well as one member of the Glox-
inieae (Lembocarpus Leeuwenberg). Further studies
that include these taxa will hopefully resolve the
number of times tubers have originated within the
Gesneriaceae.
BIOGEOGRAPHY
The traditional division of the Gesneriaceae into
two subfamilies (excluding the Klugieae, which
may stand best as a third subfamily) is well sup-
ported in this analysis and is in agreement with the
biogeographic distribution of these taxa. The Cyr-
tandroideae (excluding the Klugieae) are distrib-
uted almost exclusively in the eae with a
few temperate European and A species. Two
African genera were included in pr analysis, both
of which are in a single clade (Saintpaulia and
Streptocarpus). Apis one of the European taxa (Ra-
monda L. C. Richard) has been included in this
analysis; манета nothing сап be aoe regard-
ing the origin of these taxa at this tim
Members of the tribe Klugieae range from India
to south China, Taiwan, the Philippines through
Malaysia, Indonesia, and into New Guinea. Dis-
crepancies from this distribution include a single
species of Rhynchoglossum Blume found in Central
and South America. The presence of Rhynchoglos-
sum azureum (Schlecht.) B. L. Burtt in the Neo-
tropics represents a secondary dispersal event in
the family, because all other members of the Klu-
gieae are found in the Old World
The Gesnerioideae are almost aceite neo-
tropical, but with the inclusion of the Coronanthe-
reae within this subfamily the les now
encompass several Australian and South Pacific is-
land species.
Literature Cited
Ackerman, J. D. 1986. Coping A Uy ре exis-
tence: Pollination strategies. Selby.
ен С. Gesneriaceae. й 990-10 in G.
Bentham & J. D. Hooker (editors), Селин "tels
xx Reeve, London.
1991. A oe study and cladistic
of Sinningia and as ted genera with par-
cerda руна се to Tibet Шеше, Paliava ana,
and Vanhouttea (Gesneriaceae: = M.S. The-
sis, Pig University, Ithaca, Yor
Bremer, K. 1988. The limits of amino ase sequence data
in angiosperm phylogenetic reconstruction. Evolution
95-803.
== 1994. Branch support and tree stability. Cladis-
tics 10: etus.
genera.
neriaceae t s on id classi-
fication ы that fam ull. Bot. Surv. India 7
пијете above the genus; as exe
plified = ef Perec with parallels from n
groups. Pl. Syst. proh. d 1: 97-1
& H. Wiehler. 1995. Meses of the family
atr
ма И jin a ed also origin
of the Labiatae. A eo Missouri Bot. Gard. 7' 1-379.
Dele:
Chase, ltis, R. С. Olmstead, eu санан
р ird в. D. Mishler, М Duv. A. Price,
С. Hill Y.-L. Qiu, К ron. i
Hedrén, B. S. Сай sra у K.-J. Kim, С. Е
а ава а واش ن و وا کن ت a ава а са с
Volume 84, Number 1
1997
Smith et al. 65
Tribal Relationships in Gesneriaceae
Wimpee, J. F. Smith, G. R. Furnier, 5. H. Str vica E
Xiang, G. M doe = 8. Soltis, S. M. Swenson
E. Williams, P A.
Е Мегаре G.H. Шат лез. Ж
tS: s & V. A. Albert. 1993. Phylo-
pere of seed plants: An analysis of nucleotide se-
кузнец» в from the plastid gene rbcL. Ann. Missouri Bot.
580.
Gard. 80: 528—
Clark, L. G., W. Zhang & J. F. Wendel. 1995. pel id
eny of the grass family (Poaceae) based on ndhF s
436-460.
Crisci, J. V., M. M. Cigliano, J. J. Morrone & S.
Јићет. 1991. Histo]. bioge: ography of solos
America. ee ool. 40: 152-171
1981. An Integrated System of EN
of Flo: owering Plants Columbia Univ. Pres
Dahlgren, R. 1975. A system of rho “of a an
giosperms to be used to — the distribution of
асса Bot. Not. 128: 119-147.
Donoghue, M. J. & P. D. Cantino. 1984. The logic and
limitations of the or mara Peake to cla-
distic analysis. Syst. eg енн
Olm ae в de h € J. D. Palmer.
1992. Равене ‘relationships of е Баѕеа
on rbcL sequences. Ann. Missouri Bot. Gard. 79: 333—
345.
Farris, S. J. 1970. — for computing Wagner trees.
Syst. Zool. 19: 83
, A. С. Kluge e J. Eckardt. 1970. A wie
approach to phylogenetic systematics. Syst. Zoo!
172-1
dew 3 1893-1894. Gesneriaceae. Pp. 133-185 in
er & K. Prantl (editors), Die Natürlichen Pflan-
nice зе Vol. 4(3b). Wilhelm Engelmann, Leipzig.
Heywood, V. H. 1978. Flowering Plants of the World.
Prentice ger pic eet New Jersey.
Innis, M. A., H. Gelfand & M. A. D.
Br w. 1988. У ПИК e hh and direct sequencing
of polymerase chain reaction-amplified DNA. Proc.
Natl. Acad. Sci. U.S.A. 85: 9436-9440.
— Mesum of the carpological meth-
уш Gesneriaceae. Notes Roy. Bot
pii
& M. J. Donoghue. 1994.
osperm family MEN. is пад phylogenetic
pod Harvard Pap. Bot. 5: 1-5
Као, М.-Т. & С. E. Devol. *ion. T Gesneriaceae of
Taiwan. Taiwania А — en
Kress, W. J. 1986. Ti distribution of vascular
epiphytes: a inde не 9: 2-22.
Kvist, L. P. . Revision of Heppiella (Gesneriaceae).
Syst. Bot. ee 720-735.
Maddison, D. R. 1991. The discovery and importance of
multiple islands of nd -parsimonious trees. Syst. Zool.
40: 315-328.
Maddison, W. P., Donoghue & D. Maddison.
1984. Outgroup pronta and parsimony. oe Zool. 33:
pps M. 1977. Vascular epiphytes: IT systematic
and ага features. Selbyana 2: 1-13.
Омин. ЕЧ С. € J. D. Palmer. 1994. petal DNA
— tics: А review of methods and data analysis.
es E 81: 1205-1224.
A. Re eeves. 1995, Evidence for the poly-
с
phyly е th
and ndhF sequences. Ann. = Bot. Gard. 82:
176-193.
Вечен nd Mri ruis data in phy-
SS АЕ
logenetic systematics: An cal approach using
three molecular data бан іп He эа Syst. Biol.
Sie
———, B. Bremer, К. M. Scott & J. D. Palmer. 1993.
А parsimony егин sis of the Asteridae sensu lato
based on rbcL sequences. Ann. Missouri Bot. Gard.
80: 700-722
, H. J. Michaels, K. M. Scott € J. D. Palmer.
1992. Monophyly of the Asteridae and identification of
се поре lineages inferred from DNA има of
Ann. Missouri Bot. Gard. 79: се
хо, R. W., J. A. Sweere, P. A. Ree ix d
ead. 1995. Higher level МЕ тоном pr Alias
determined Ji chloroplast DNA sequences. Amer. J.
Sight Y ks D k Jain. 1978. Floral anatomy and sys-
tematic position of Cyrtandromoea. Proc. Natl. Acad.
Sci. Sage 71-74.
Skog, 1984. A review of M numbers in
the ин Selbyana 7: 252-2
1995. A possible ин E Hmmm H. E.
Moore Apes eeu Amer. J. hoa 82:
F. de Jesus. 1996. view d Basin H. E.
тр (Gesneriaceae) ышы (in p ress),
Smith, J. F. Tribal sateen within the Ges-
netiaceae: i cladistic analysis of morphological data.
Syst. Bot. чч ка
€ К tsma. 1994а. Evolution in Andean
epiphytic = Columnea (Gesneriaceae): Part I. Mor-
phological variation. Syst. bs 19: 220-235.
ifi Evolution in the Saar epi-
phytic genus Co ил а (Gesneriaceae): Part II. Chlo-
roplast DNA maritis site variation. Syst. Be 19;
317-336.
Pis 1994c. Molecules and morpholo,
ence of dat m Columnea (Gesneriaceae). PL.
eg Evol. ats
———. wn, re t Carroll & D. S. Denton. 1997.
Familia Diente of Cyrtandromoea, ор lr E
and Sanango: Three problematic genera of the Lam
ales. ey (in press
a Siam T S. Shoemaker € R. L. Smith.
1992. A qu vali tative comparison of total cellular DNA
extraction Sage Phytochem. Bull. 23: 2-9.
Sugiura, M. 1989. The chloroplast chromosomes in land
"en Am. Rev. Cell Biol. 5: 51-70
ir de chloroplast genome. Pl. Molec. Biol.
ба ч ү `1993. PAUP: Phylogenetic ede Using
Parsimony, version 3.1.1 Computer program distributed by
the Illinois Natural History Survey, comb Illinois.
& W. P. Maddison. 1987. Reconstructing ances-
tral character states under Wagner parsimony. Math.
Biosci. 87: 199-229.
Takhtajan, А. L. 1980. Outline of a classification of flow-
= иу mr emer mU Bot. Rev. 46: 225-359.
Thorn Uy gam classification of the
Ang cid. T3 n M. K. Hec
Steere & B. Wallace cor, ii Biology. Vol.
9. Plenum din New York
1983. Proposed new г feslisuments i in the angio-
sperms: Nordic J. Bot. 3: 85-117.
1992. An updat э hylogenetic classification of
flowering ver айкол Bot. Rev. (Lancaster)
225-359,
66 Annals of the
sp indy Botanical Garden
Wang, W. T., K. Pan & Z. Li. 1992. ys » pe Gesner- Wolfe, K. 1991. Protein-coding genes in chloroplast DNA:
iaceae of China. Edinburgh J. Bot. 49: 5— Compilation of nucleotide sequences, database entries and
Wiehler, H. 1983. A synopsis of the ceste Gesner- rates of molecular evolution. Pp. 467-482 in L. Bogorad &
jaceae. и 6: 1—249. І. К. Vasil (editors), The Photosynthetic Apparatus: Molec-
& A.C
hautems. 1995, A reduction of Lietzia to ular Biology and as vol. 7B, Cell Culture and So-
Sinningia. Gesneriana 1: 5-7. matic Cell Genetics in Plants. Academic Press, New York.
А РНУГОСЕМЕТТС
CONSPECTUS OF THE
TRIBE JUANULLOEAE
(SOLANACEAE)!
Sandra Knapp’, Viveca Persson”, and
Stephen Blackmore?
ABSTRACT
The tribe Juanulloeae has Ваве“ consisted of nine se of rarely collected, Acum ds and small trees:
ianaea, Dyssoch
Juanulloa, Markea, Schultesianthus
јуззосћтота,
siophyton, Ка tozoma, and Merintho-
Hee we present ie results * a cladistic study of the relationships of the species of na genera and provide
podium.
a «ре of the genera as we define
uan
oa, Markea, Meri nthopodium , Schultesianthus, а!
Solandra. which at pent is id as a separate tube Кыке Ж е ајво
in this group. key to the genera is provided and for each genus a li
hese genera have previously been
point out areas for future research in
taxa and their hon: is also given. Many
them. The nu wee ~ ae in our treatment is reduced to six: Dyssochroma,
a. Included in the key and conspectus is the genus
iscuss the groupings on the tree and
st of component
illustrated, de eras are
g t
provided for some of the previously ан taxa or for taxa where аи are difficult t
The Solanaceae are an economically important,
cosmopolitan family with over 2500 species that
have traditionally been divided into two subfami-
lies. The Cestroideae, including petunias, the ces-
trums and their relatives, have non-compressed, of-
ten prismatic seeds and tropane alkaloids. The
Solanoideae, which contain the large majority of the
species in the family, include Solanum and its rel-
atives that have compressed seeds and steroidal al-
kaloids. This traditional classification has recently
been challenged by cladistic analyses using chlo-
roplast and nuclear DNA data sets, and the family
can now be divided into approximately seven mono-
ры pan (see Olmstead et al., in press).
be Juanulloeae, first described by Hun-
ziker уен is placed in the subfamily Solanoide-
ae in both the traditional and phylogenetic systems:
its members share flat, discoidal seeds and curved
embryos with others in that subfamily. The tribe as
defined by Hunziker (1977, 1979) is delimited by
a combination of habit, anther, and seed characters
and consists of nine genera: Dyssochroma Miers,
Ectozoma Miers, Hawkesiophyton Hunz., Juanulloa
Ruiz & Pavón, Merinthopodium Donn. Sm., Markea
Rich., Rahowardiana D'Arcy, Schultesianthus
Hunz., and Trianaea Planch. & Linden. The Juan-
ulloeae are thought to be closely related to the ge-
nus Solandra Swartz, the only member of the So-
landreae. Trianaea was previously considered a
member of the Solandreae (Hunziker, 1979; Ber-
nardello & Hunziker, 1987), but was transferred to
the Juanulloeae (Hunziker & Bernardello, 1989)
owing to its ex-endospermous seeds and almost
straight embryos with oblique, accumbent cotyle-
dons. Solandra differs from the members of the
eset in its incumbent rather than accum-
bent cotyledons and its partially inferior ovary
(D'Arcy, 1973 ye Miers (1857) allied Solan-
dra, Jua arkea, Sarcophysa Miers (Juan-
idis speciosa (dh Dunal), and Dyssochroma as
the Solandreae, which he considered to be closely
related to the shrubby neotropical genus Brunfelsia
L. Brunfelsia is now considered to be related either
to Salpiglossis L. and its relatives (Hunziker, 1979)
or to Petunia L. (Olmstead et al., in press). In the
recent molecular phylogeny of the family Juanul-
loinae and Solandrinae are united in the tribe Juan-
ulloeae (see Olmstead et al., in press). The general
habit and morphological similarities of the Juan-
ulloeae and Solandra have long been recognized
and we have thus included Solandra in this anal-
ysis.
The genera of the Juanulloeae (here referred to
in the broad sense, including Solandra) are all neo-
tropical, epiphytic trees and shrubs. Many of the
species are myrmecophilous, especially in the ge-
1 We thank M. E i G. MAs R. G. Olmstead, and L. Freire de Carvalho for helpful demos about the
for
panes m. taxono
рт" US for loans and permission to samp sé она
at the Natural depend Museum for бака ná eas
of Solanaceae; М. С. Bovini in Rio de
istance; D. M. Williams for helo vi with С° ша weighting; the curators of BM, F,
nvaluable field
K, мо. NY, ОСМЕ,
Janeiro and T. Nuñez in Quito
— specimens in their care; and the SEM and Photographic Units
? Department of Botany, The Natural History red Cromwell Road, London SW7 5BD, United Kingdom.
ANN. MISSOURI Bor. GARD. 84: 67-89. 1997.
Annals
ансо? س Garden
Det. Sandra Knapp
Figure 1.
NY)
nus Markea, where two Amazonian species are as-
sociated with particular species of ants (Ducke,
191 camponoti Ducke — with
Camponotus femoratus (F) and M. formicarum
Damm. with Azteca sp.). Species = Hawkesiophyton
(here treated as congeneric with Markea) are often
associated with various species of ants a
1922), and seedlings of M. panamense Standl. p
duce enlarged basal stems even in the daie of
ants (Garwood, pers. comm.). The swollen tuber-
like stems and roots seen in these species (Fig. 1)
are often hollow and are probably used by ants for
Systematic Studies in Solanaceae purple like "carrion'
Markea formicarum Dammer
The New York Botanical Ga:
VENEZUELA, Amazonas, Dees: мо ¿Jero
Cerro de la Neblina Expe:
ess uma) just upstream
side of ue de la Neblina,
00749" 0. СА 09 50" 40 m.
ve
om gro
ubers at еге ыы MA Stems olendi
spreading, to 2 m. long; flowers hanging, odor-
les: » at m ‘Irs. y Open,
pa no odor at night; calyx light green;
lowish white on outside, the lobes
" " flrs., one lobe cupped
her zon
M. Nee 30923 17 Feb. 1985
supported i Fundacion. ا See Ре.
Markea formicarum Dammer. Specimen illustrating the swollen INE ett (Nee 30923,
nests and storage. In some ant plants, a mutualistic
relationship exists between the insect and the plant,
with high percentages of both carbon and nitrogen
being derived from ant respiration and debris de-
position (Treseder et al., 1995).
ome members of the Juanulloeae are fed upon
by the larvae of the Ithomiinae (Lepidoptera: Nym-
phalidae), specialist herbivores of the family Sola-
nacaeae. Solandra, Schultesianthus, Markea, Juan-
ulloa, and Merinthopodium are fed upon by larvae
of the genera Olyras, Eutresis, and Melinaea (tribe
Melinaeini), which are all high flying canopy dwell-
Volume 84, Number 1
1997
Knapp et al. 69
Tribe Juanulloeae
Figu A. Juanulloa учини M mi Miers,
аг =
cult. yid Beni: Gardere К 5 cm.—
B. Solandra maxima Sessé & Mocin 100, пенен аы ег-
sity of Texas, Austin, TX, originally collected at Las Tux-
tlas, Veracruz, MOS (photo J. Mallet), scale bar — 4 cm.
ers (for a complete listing of the host plant rela-
tionships of these butterflies see Drummond &
Brown, 1987). The young larvae make character-
istic *C"-shaped damage in the leaves, but hav
rarely been e and are difficult to collect а
the forest сапо
In common ah many other tropical epiphytes
the leaves of species in the group tend to be thick
and prs and the branches ime with pliable
bark. Flower shape in the Juanulloeae vari
een dim the long, red or wem orange, pre-
sumably hummingbird-pollinated, flowers of Mar-
kea coccinea Rich. and most of the species of
Juanulloa (Fig. 2A) to the greenish, open campan-
ulate flowers of = species of Trianaea (Fig. 3B),
Merinthopodium (Fig. 3A), and Dyssochroma, which
are bat-pollinated Z et al jery case,
apart from Markea panamensis, however, the flow-
ers are sympetalous and have relatively long corolla
tubes. Fruits of members of the Juanulloeae are
generally fleshy to leathery berries, with some vari-
ation in the thickness of the berry wall. Genera of
es con-
Figure 3. —A. Merinthopodium neuranthum (Hemsl.)
Donn "ei: fallen flower post-anthesis; note the broadly
nteverde, Costa scale bar —
c urere shape (Mo
26 . Trianaea speciosa (Drake) Зиме Knapp а et al.
9121. ii irn road, Bind. scale bar = 2c
the tribe have been traditionally delimited (Hun-
ziker, 1977, 1979) using combinations of the fol-
lowing characters: insertion of the anthers on the
filaments, filament insertion on the corolla tube,
and corolla aestivation. Characters and their states
are discussed more fully in Materials and Methods.
ew collections are known for most of the species
and this makes the assessment of characters diffi-
cult since the extent of variability is not known.
Woody tropical epiphytes are difficult to collect as
they often grow high in the canopy and flower only
md Many of the species of the Juanulloeae are
known only from flowering material, and thus de-
cisions whip n fruit characters tend to be rather
ad hoc at best. With few specimens available it is
nearly impossible to assess variability in charac-
ters, and a tendency to overdivide at the generic
level is apparent and understandable.
Recent си ations of molecular systematics
ave extremely useful in providing broad
a cate for directing future study. However, at
present, the limited range of taxa that has been
Annals
ра SEL Garden
able Taxa used in the cladistic analysis. Names
in parentheses are those used in Persson et al. (1994).
Nicandra Mecca (L.) Gaertner
Atropa belladonn
Lycium crum AN dl
Dyssochroma longipes (Sendtner) Miers
Dyssochroma viridiflora (Sims) Miers
Juanulloa ferruginea Cuatrecasas
Juanulloa globifera (8. Knapp €: D'Arcy) S. Knapp (Ra-
howardiana globifera)
Juanulloa membranacea Rusby
Juanulloa mexicana (Schltdl.) Miers
Juanulloa paras
Juanulloa parviflora (Ducke) Cuatrecasas
Juanulloa pavonii (Miers) Benth. & Hook. (Ectozoma pa-
Juanulloa speciosa (Miers) Dunal
Juanulloa verrucosa (Rusby) H
Juanulloa wardiana (D'Arcy) s. Kane (Rahowardiana
rdiana)
Markea camponoti Ducke
Rich.
k
arkea is Standl. (Hawkesiophyton panamense)
Markea dins Ducke
Markea ulei ныр Cuatr. (Hawkesiophyton шеї)
Markea lopezii
к aff. lo
nt обид. ыо (Hemsl.) Donn. Sm.
аката crosbianus iE 5, po (Markea
na)
Pieris: dudleyi Bernardello & Hunz.
Schultesi
Eee s uniflorus (Lundell) S. Knäpp (Markea
Sika venosus Fijos & C. V. Morton) S.
Trianaea brevipes (Cuatr.) s. Knapp (Trianaea spectabilis
var. brevipes
Teens naeka S. Knapp (Trianaea sp. 1)
naea neovisae Romero-Castañeda
Hanae nobilis Planch. & Linden
Trianaea speciosa (Drake) Soler.
Trianaea sp. nov. (Trianaea sp. 4)
sampled makes direct comparison of these results
to those from morphological studies difficult. The
sampling of morphological characters is often more
complete in any given group, especially at the fam-
ily level, and thus relationships are more resolved.
This is certainly true for this analysis of the Juan-
ulloeae. The molecular analyses of Solanaceae un-
dertaken to date have indicated a close relationship
in press). d in those analyses only four spe-
cies were , thus somewhat limiting their
iiie at i is relationships among genera
or in determining whether or not the genera as pres-
ently defined are monophyletic.
This paper presents a cladistic analysis of the
5
ological characters (Persson et
., 1994) and provides a conspectus of the genera
and species.
MATERIALS AND METHODS
Taxa used in the cladistic analysis are listed in
Table 1. By using species as terminal taxa we
hoped to test the monophyly of genera in the group
as presently defined. Nomenclatural changes ne-
cessitated by the results of this study were made in
Knapp (1995), and the new combinations are used
here. The names used in Persson et al. (1994) did
not reflect these nomenclatural changes and thus
are slightly different. Names in parentheses in Ta-
ble 1 are those used in Persson et al. (1994) when
a new combination is used here. Morphological
characters were assessed using herbarium speci-
mens from BM, CUVC, F, JAUM, K, MO, NY, and
QCNE, and voucher specimens are cited in the ap-
pendix. Palynological characters were assessed as
parsimonious trees, then bb* for finding equally
parsimonious trees. These opti
iteration can be prohibitively time-consuming for
finding shortest trees (Farris, 1988), and have been
shown to be as good as the ie option in many test
data sets (Platnick, 1989). The ensemble consisten-
cy index (CI) is a measure of homoplasy in the data
set with respect to the fit of characters to the tree.
When the fit of a character is Bh. (with no par-
ree or reversal) then the consistency index
als 1. The ensemble — index (RI) is the
Mons of apparent synapomorphy in a character
that is retained as synapomorphy on the tree (Far-
Volume 84, Number 1
1997
Knapp et al. 71
Tribe Juanulloeae
1989). Trees were also constructed using
NONA (Goloboff, 1993) to confirm the actual ver-
sus potential groupings on the tree, as NONA and
Hennig86 treat zero length branches (potential
groupings) in a different way. The following com-
mands, as recommended by еру TA were
used: hold 100, hold/20, and mult*
Successive approximation ae n w option in
Hennig86) was used to assess the reliability of fit of
characters to the most parsimonious tree. The fitting
function in the MS-DOS program PeeWee (Goloboff,
1993) was also used to assess character reliability.
Character weighting, when applied to the fit of char-
acters, emphasizes those characters that best fit the
initial tree topology. Successive weighting allows the
characters to judge themselves in terms of their reli-
ability: i.e., their best fit to the solution supported by
all the data (Carpenter, 1994). Best fit is judged by
the shortest tree (Farris et al., 1970), the shortest tree
for the weighted data in terms of tree length (Farris,
1969), or the “heaviest” tree when calculated as a
function of character weights (Goloboff, 1993). Char-
ters that are more homoplasious are less reliable
and are thus downweighted in these analyses. Suc-
cessive weighting (Hennig86, xs w option) uses the
rescaled is BN index (rc) of Farris (1989) as the
weighting fun each character: it is calculated
as the ужа ss ложи RI (retention index) times
ensemble CI (consistency index) and sc: between
0 and 10. Goloboff (1993) calculates weights as the
extra number of steps per character such that the
weight = K/K+ESi, where ESi is the extra steps per
character and K is the concavity constant (in our anal-
yses set at K = 3).
The characters were coded to be unordered, thus
. Three were selected as out-
ps (see Tables 1 and 3), Nicandra physalodes,
уны belladonna, and Lycium cestroides, represent-
ing a range of putative sister taxa for the Juanulloeae.
Choosing a range of outgroups (Watrous & Wheeler,
1981) has been thought to increase the likelihood of
obtaining an accurately rooted tree. Recent work,
however (Nixon & Carpenter, 1993), has shown that
multiple outgroups perform no better at “polarizing”
ingroup nodes, but that multiple outgroups might im-
prove inference. One difference in this data set from
used in Persson et al. (1994) is the omission of
Mandragora as one of the outgroups for the analysis.
Olmstead and Palmer (1992) had originally identified
Mandragora and Solandra as sister taxa using chlo-
roplast DNA restriction site mapping. More recent
work has revealed that Mandragora is an isolated tax-
on of uncertain affinity, possessing many autoapomor-
Table 2. Characters used in the cladistic analysis of
the Juanulloeae.
0. Habit shrubs 0, herbs 1, epiphyte
Peltate conte trichomes on leaf Med absent 0,
presen
2; amice of solitary flowers 1, few flowers 0, many
flowers 2.
3. Inflorescence axis condensed 0, elongate 1.
4. Calyx lobes shorter than the corolla tube 0, equal to
the corolla tube 1
5. Calyx texture at anthesis membranous 0, coriaceous 1
6. Calyx lobes acute 0, acuminate 1, long-acuminate 2,
rounded 3
7. Flowers radially symmetric 0, zygomorphic 1.
8. Corolla aestivation iieri oia 0, valvate 1.
9. Flower —— w tube 1, salverform 2, open 0,
campanulate
10. Corolla color seii 0, purple-purplish green 1,
red, orange, or yellow
11. Filaments straight 0, маски r
12. Filament tube absent 0, present 1.
13. Filament base glabrous 0, pubescent 1.
14. Filament base pubescence dense 0, sp: Г.
15. Filaments inserted in anther basally o, dorsally 1,
ventrall
16. Anthers included in the corolla tube 1, at mouth of
corolla tube 0, exserte
17. Anthers dehiscing пораки 0, о» h
18. Ovary superior 0, partially inferi
19. tke + conical 0, narrowly үй" in E
20. Fruit pericarp membranous 0, coriaceous 1.
21. Seeds reniform 0, rectangular 1.
22. Cells in center of testa square 0, кте E
23. Lateral cell walls straight 0, sinuate 1.
24. Dried seed color pale brown сана. огапре 2.
25. Undigested seed surface pitted 0, smooth 1.
26. Spiny supratectal processes absent 0, present 1.
27. Colpi extending nearly to the poles 0, relatively short 1.
28. Pollen without Ubisch bodies 0, covered with Ubisch
ies 1
m
29. Colpi with continuous margins 0, lalongate apertures 1.
30. Exine around apertures thickened 0, not thickened 1.
31. Tectum continuous 0, not continuous 1.
32. Beak-like margo apertures absent 0, present 1.
33. Colpus border not thickened 0, thickened 1.
34. Outline in equatorial view spherical 0, oblate
35. Outline in polar view obtuse, convex 0, acute, ou m
1.
36. Мехте same thickness as the ѕехіпе 0, пехіпе thicker
than the sexine 1.
phies, both molecular (Olmstead & Sweere, 1994;
Olmstead, pers. comm.) and morphological.
The data matrix is presented in Table 3. Most of
the characters are self-explanatory; those peculiar
to the Juanulloeae are described in detail here and
the palynological characters were discussed in
72 Annals of the
Missouri Botanical Garden
Table 3. Data matrix used for the analysis of the Juanulloeae.
Nicandra 1000100000000020200000001001001000100
Atropa 1000000000110101000000000000000000100
Lycium cestroides 0000000001100101200000000000001100100
Solandra зно 2010010000210020001010001000001000000
Solandra bolivia 2010010000110020001012222201001000000
Markea sessili; 2000012000000100100001102001101100100
Markea Scand 2000012000000100100002222201101100110 |
Markea coccinea 2000012002200100000001102001010100110 ;
Markea formicarum 2000012000000100100001102001010100110 |
Markea camponoti 2000012000000100100001102001010100110
Markea lopezii 2011001000200100100022222211001000000
Markea aff. lopezii 2010001000100100100022222222222222222
Markea panamensis 20000000000000?1110011102001000010110
Markea ulei 2000000002000101010011102001000010110
Merinthopodium neuranthum 2001000013000111200010010100000001100
1 i m 2001010013000102200022222201010100110
chultesianthus uniflorus 2110010100000100200012222201101100100
Schultesianthus ve 2110010100000100100012222201101100100 |
chultesianthus crosbianus 2110010100100110000010001001101100100
Schultesianthus megalandrus 2100010100010110000010010101100000000
Schultesianthus leucanthus 2100010100010110000010010101100000000
Sch — us riferus 2100010100010110000012222201100000000
ianthus мее 2100010100010110000010010101100000000
паннин udleyi 2100010100010110000012222201100000000
uanulloa phar 2000110000001101100011000001001000001
Juanulloa wardiana o OA ас и
Juanulloa globifera 2020110001100021110102222222222222222
Juanulloa speciosa 301001000120 T01 1000100000000 ПОНТ
Juanulloa ochracea 2000010001000101100000000000000001110
Juanulloa parasitica 2001010001200101100000001001001000001
Juanulloa ferruginea 2001010001200101000000001001000000000
Juanulloa membranacea 2001000001200101100022222201001000101
Juanulloa verrucosa 2001000001000101100020001001001000101
Juanulloa parviflora 20010100010001011000222222222222227227
uanulloa mexicana wipe ML T
Dyssochroma longipes 20101110130001001000222222 00000100100
Dyssochroma viridiflora 201011101300010020002222 ВН ЕВО
Tianaea speciosa 2001110013000112200010000000001100100
Trianaea brevipes 2010110000000112200010000001001000110
Trianaea neovisae 2000110013000112200012222222222222227
Trianaea nobilis 2001110013000112200010000000001000110
rianaea naeka 20111100130001022000222222 00000000100
Trianaea sp4 201011 2220002272222222222227222
Persson et al. (1994). Minute, peltate, glandular small pits in the mesophyll so that the ca. 24 cell
chomes (character 1) on the leaves of Shute head is at about the same level as the foliar surface.
thus were first described by Bernardello and Hun- The trichomes appear as minute reddish dots to the
ziker (1991) and are either present or vod on naked eye and are usually, but not always, more
both leaf surfaces. These trichomes are sunken into abundant on the abaxial leaf surface. Corolla aes-
—
Figure 4. One of the two equally parsimonious cladograms from the Hennig86 analysis. The tree shown here is
Ha to that eo у МОМА, and to the consensus tree. The characters are discussed in the text, and character
states are shown in Table 2. Fir Феб marked on the branches of the meri stippled marks indue reversals
and paralela pit and solid marks ноне synapomorphies
Volume 84, Number 1
1997
Knapp et al.
Tribe Juanulloeae
AR "
4vicanara
4
“uropa
3r
Lay CEUM
Juanulloa verrucosa
24 30 35
membranacea
Juanulloa parasitica
Juanulloa ferruginea
І, 1]
техісапа
Juanulloa pavonii
Juanulloa wardiana
Juanulloa 8 lobife era
Markea lopezii
Markea aff lopezii
QAI
brachycalyx
Solandra boliviana
(22. L
venosus
л: L
uniflorus
Di AS L
crosbianus
Schultesianthus megalandrus
Schultesianthus leucanthus
34
? Sch hus odoriferus
Schultesianthus coriaceus
Schultesianthus dudleyi
Markea panamensis
‘Markea ulei
Mark
sessiliflora
34 E
costanensis
(y pueri
coccinea
M зу],
formicarum
Markea camponoti
Juanulloa speciosa
ochracea
Juanulloa parviflora
M. h "m
pendulum
I4 1 1;
neuranthum
Trianaea neovisae
Trianaea brevipes
Trianaea sp. nov.
74
Annals of th
Missouri E Garden
tivation (character 8) has been coded as either
overlapping or valvate. The exact arrangement of
overlapping corolla lobes in bud is sometimes dif-
ficult to determine from herbarium specimens and
occasionally varies in a species or genus. Corolla
ontogeny has not shed any light on the relative ple-
siomorphy or apomorphy of such differing aestiva-
tion patterns in the Acanthaceae (Scotland et al.,
4). Thus, should this character be used in sub-
sequent analyses of the Juanulloeae, it should be
unordered and unweighted.
RESULTS AND DISCUSSION
In the analysis of the genera of the Juanulloeae,
two trees of length = 129, CI = 35, and RI = 71
were found by Hennig86 (see Fig. 4), and one of
these two trees was also found by NONA. If only
Nicandra is used as the outgroup, Hennig86 finds
three trees, again of length 129, one of which is
identical to the strict consensus tree. The two trees
found by Hennig86 using all three outgroup taxa
differ only in the resolution of Schultesianthus, with
tree 2 being over-resolved owing to character state
assignment from missing data. Tree 1 is identical
to that found by NONA, which does а attempt to
resolve nodes with missing data. e NONA
analysis, 15 of 50 replications were т чей 129,
indicating this is probably the shortest tree.
sing successive approximation weighting in
Hennig86 character weights stabilized after four it-
erations, giving 444 trees of L = 280, CI = 67,
and RI = 88. Reading all 444 trees back onto the
unweighted data set gave trees of length 135 or
136, which is longer than the most parsimonious
trees by 6 or 7 steps. Using the alternative weight-
ing system of Goloboff (1993) we obtained one tree
of length = 130 (not shown), one step longer than
our most parsimonious tree generated using un-
weighted data. This tree differs in the positions of
the three anomalous species of Juanulloa (see be-
low) and in the order of branching in the Trianaea
clade.
The stabilized weights from the successive ap-
proximation weighting analysis are presented in Ta-
ble 4. Weights of 10 imply perfect fit to the tree,
while weights of 0 (and lower numbers in the
PeeWee analysis) imply the character is of poorer
fit. Of the characters with weights of 10 in the Hen-
nig86 analysis, two (character 12, presence of a
filament tube, and character 26, presence on pollen
spiny supratectal processes) are uninformative
(autapomorphic) and do not contribute to group for-
mation on the tree. Only nine other characters had
weights of 10, while the rest had low weights of 5
Table 4. Stabilized character weights from Hennig86
successive approximation analysis and weights (character
fit) from PeeWee analysis (see text for explanation).
Character Character
Character weight weig
number Hennig86 PeeWee
0 10 10
1 10 10
2 8 4.2
3 0 42
4 1 4.2
5 0 4.2
6 5 15
T 10 10
8 10 10
9 5 4.2
10 0 25
11 2 6
12 10
13 1 5
14 2 5
15 2 4.2
16 1 В
17 3 15
18 10 10
19 10 10
20 1 5
21 4 fd
22 10 10
23 + 6
24 2 5
25 3 T
26 10
27 1 5
28 4 4.9
29 1 6
30 0 33
31 0 3.3
32 10 10
33 2 7.5
34 1 5
35 0 6
36 10 10
to 0, indicating the data set has a great deal of
homoplasy and very low support. Of the characters
traditionally used to define genera in the Juanul-
loeae (see Hunziker, 1977, 1979), corolla aestiva-
tion (character 8 in our analysis) and floral sym-
metry (character 7) perform better, while other
characters, such as the insertion of the filament on
the anther (character 15), are less consistent on our
tree. Micromorphological characters such as the
minute glandular trichomes characteristic of Schul-
tesianthus (character 1, see below), the relative
thicknesses of sexine and nexine layers in pollen
Volume 84, Number 1
1997
Knapp et al. 75
Tribe Juanulloeae
(character 36), and some pollen aperture characters
(character 32) are a good fit to the tree and are
useful in resolving relationships in this group.
Four main clades are identified in the Juanul-
loeae. The tribe is defined relative to the outgroups
by its epiphytic habit (character 0), and the pres-
ence of anthers included in the mouth of the corolla
tube (character 16, reversed several times in the
group). The first clade, comprising species of Juan-
Mp is sister group to the rest of the tribe (see
Fig. 4). The members of this clade share synapo-
morphies in characters 3 (elongate inflorescence—
reversed in J. wardiana, J. globifera, J. pavonii,
and J. mexicana), 9 (narrow tubular flowers), and
36 (pollen with the nexine thicker than the sexine).
Juanulloa pavonii + (J. wardiana + J. globifera)
share a large suite of characters, many of which are
shared with other taxa in the tribe (i.e., homopla-
sious): these include calyx lobes equal in length to
the corolla tube (character 4) and pale brown, reni-
form seeds (characters 21 and 24). Juanulloa war-
diana and J. globifera have been segregated as
genus Rahowardiana (D'Arcy, 1973 [1974]; ae
& D'Arcy, 1993), but the results of this analysis
indicate their relationships lie with Juanulloa, and
perhaps they are deserving of subgeneric recogni-
tion. Three species of Juanulloa are in an anoma-
lous position on the tree: J. speciosa, J. ochracea,
and J. parviflora. Juanulloa parviflora is known
only from the type, which bears a single flower, and
thus many of the characters used in this analysis
had to be scored as missing. Missing data can affect
placement of taxa and resolution of trees and
should therefore be viewed with caution (Platnick
et al., 1991). Juanulloa speciosa and J. ochracea
share a distinctive pollen type (see Persson et al.,
1994) and pollen characters contribute largely to
their position in the Trianaea (see below) clade. We
do not have confidence in this placement, and thus
have elected to keep these three species in Juan-
ulloa pending further investigation. Juanulloa spe-
ciosa was originally described as the genus Sarco-
physa by Miers (1857), and is morphologically
somewhat isolated in the genus. The relationship of
these taxa to the rest of Juanulloa is a subject that
would bear investigation using molecular tech-
niques.
The second main clade consists of Markea lopezii
Schultesianthus. This clade is united by the pos-
session of solitary flowers (character 2, reversed in
part of Schultesianthus) and basifixed anthers (char-
acter 15, in parallel with several other taxa in the
tribe). Markea lopezii and its undescribed relative
are both known only from flowering specimens, and
thus the matrix has man
prominent spiny supratectal processes (Persson et
al., 1994), and its relationships need to be inves-
tigated further. Not all species of Solandra were
included in this analysis, but these results indicate
that it along with M. lopezii and M. aff. lopezii are
the sister group to Schultesianthus and P should
be included in the tribe. However, cpDNA results
(Olmstead et al., in press), using only four taxa of
the group (Solandra, Markea, Dyssochroma, and
Juanulloa), indicate that Solandra is the sister
group of the rest of the Juanulloeae. Since no spe-
cies of Schultesianthus were sampled in the cpDNA
studies, this potential conflict remains to be solved.
Many of the species of Schultesianthus were first
described as Solandra (see Conspectus) owing to
their close morphological similarity, and the species
now recognized as Schultesianthus were thought to
be intermediate between Markea and Solandra by
D'Arcy (1973 [1974].
l the species of Schultesianthus share a num-
ber of synapomorphies, two of which (character 1,
peltate glandular trichomes and character 7, zygo-
morphic flowers) are found nowhere else on the
tree. The genus also has two pollen synapomor-
phies: character 28, copious Ubisch bodies, and
character 31, a discontinuous tectum (Persson et
al., 1994). Schultesianthus venosus, S. uniflorus, and
S. crosbianus differ from the rest of the genus as
traditionally defined (see Bernardello & Hunziker,
1991) in their flowers with non-declinate stamens
and style, but share all other synapomorphies of the
group (see also Conspectus and D'Arcy, 1973
[1974]. These species are known from very few
collections.
The genus Markea comprises the third main
clade, united by the possession of pale tan (char-
acter 24) and rectangular seeds (character 21) with
elongate testal cells (character 22) (see Fig. 4). The
species M. panamensis and M. ulei have been sep-
arated as the genus Hawkesiophyton (Hunziker,
1977) and are probably worthy of subgeneric status.
А species not treated in this analysis (it is known
only from the type, which consists of a single poorly
preserved flower), M. sturmii Cuatr., is superficially
similar to M. ulei and should be examined more
closely. The peculiar seed type seen in Markea is
perhaps related to ant dispersal, as most of the spe-
cies in this group are associated with ants in vari-
ous ways (see above and Ducke, 1915). Many ant-
dispersed seeds bear large elaiosomes (oil or fat
bodies) that make them attractive to ants (see ref-
erences in Willson, 1983). This is apparently not
Annals
Missouri о Garden
Figure 5. M of seeds A dee coccinea Rich:
ул огоз ВМ). — hole seed, scale
—B. Close-up of tec al Nue scale bar = 34.5 5
the case in Markea (see Fig. 5), but fresh material
must be examined to determine this for certain. In
the genus Datura, the elaiosome often drops off
when plants are dried and mounted on herbarium
sheets (see Persson et al., in press). The seeds of
Markea seem to have a fine reticulum of sticky
threads on top of the testa (see Fig. 5B), which may
make them stick to ants, thus causing them to be
deposited in ant garden
The fourth clade in d tribe is that containing
the genera Merinthopodium, Dyssochroma, and Tri-
anaea. Three species of Juanulloa are also present
in this clade, but as discussed above, are probably
not correctly placed here. The clade (minus the
species of Juanulloa) is united by synapomorphies
in characters 8 (valvate aestivation, reversed in
some species of Trianaea), 9 (campanulate flowers),
and 16 (filaments exserted from the corolla tube).
In general members of this clade all possess large,
campanulate, greenish or greenish purple flowers
(see Fig. 3) that are pollinated by bats (Vogel, 1958;
Cuatrecasas, 1959; Baker, 1973; Voss et al., 1980).
Voss et al. (1980), studying Merinthopodium neu-
ranthum (Hemsl.) Donn. Sm. at Finca La Selva,
Costa Rica, postulated that it was adapted for pol-
lination by traplining bats, as the flowers fell early
in the night, were borne singly, and were odorless.
Like many of the other species of the Juanulloeae
these plants are only rarely collected, but in addi-
tion to this, each sheet often consists only of a sin-
of Solandra; see Bernardello & Hunziker, 1987).
The distinctive filament base morphology men-
tioned in the generic description is not visible in
pressed specimens, but is likely to occur in all spe-
cies of Trianaea. Ovary structure, a character that
would separate Trianaea (with a 4—5-carpellate, 8—
10-loculate ovary) from Merinthopodium and Dys-
sochroma (with bicarpellate, 4-loculate ovaries),
can dry rarely be assessed even with fruiting spec-
"We d (Persson et al., 1994) that the twelve
pollen types in the tribe did not cut across the
boundaries of the main clades. However, although
they serve effectively for the purposes of identifi-
cation, palynological characters alone provided lit-
tle resolution of relationships among the taxa. The
inclusion of palynological characters in the present
analysis increases the number of character states
that can be sampled from the relatively few speci-
mens that are available for many of the species. It
contributes to the resolution of relationships and
may also serve to identify functionally correlated
syndromes of characters, such as those found in
bat-pollinated species.
his analysis clearly highlights some of the prob-
lems with the characters used to identify and rec-
ognize the genera of the Juanulloeae. As previou usly
the monophyletic groups we have recognized in this
study have not been fully resolved. The weak sup-
port for the basal clades in our tree indicates that
more work is needed at this level within the tribe.
Many of the genera have never been adequately
monographed, and field-based studies will prove
essential for the ultimate resolution of phylogenetic
relationships in the group as additional characters
that may prove useful are likely to be found in liv-
ing plants. The extension of molecular systematics
to a more comprehensive sample of the species may
Volume 84, Number 1
1997
Knapp et al. 77
Tribe Juanulloeae
also help to resolve many of the remaining uncer-
tainties.
CONSPECTUS OF THE JUANULLOEAE
(INCLUDING SOLANDRA)
s generic PU args is Dec to allow the
E. to identify her specimens and, to
some extent, ine LR For each genus a de-
1 ist of the species, and анат to
published е ог Шеш аге рго ;
The species not included in the cladistic Meis
are indicated with asterisks. pun are listed
after each accepted name. Directions for further in-
vestigation and work in progress are discussed for
each genus.
la. Corolla limb broadly campanulate, greenish or
ow, more than 5 cm long.
2a. Corolla yellow sometimes with purplish
streaks in the throat (bluish in S. boliviana),
us eate! m e stamens
declinate; ovary somewhat inferior 6. Solandra
bre ate (occasion-
b s alvate xp an ibas ventrifixed; fila-
ments conspicu ously geniculate at the
base, covering Ф nectary; ovary 8–10-
ОАВ oos у n 7. Trianaea
- Corolla vernation valvate; anthers either
asifixed or dorsifixed; filaments not
d geniculate; ovary bilocu-
w
c
pus Anthers basifixed; inflorescences
consis sting of a sin nm" ower; SE
Brazil in Atlantic forest _____
- Dyssochroma
. Anthers dorsifixed; inflorescences
of I-many flowers, with lon “
уиме e D ans =: he
n
1b. bees limb RUE to a compara.
roadly ym тн and bell-shaped, vari-
pel pentes cream to bright orange, usually
less than 5 em lon
5a. Flowers sygtnorphic, at paña ок и
ficult to see in pressed 5 ens
Si e cream fading y
fragrant; calyx thick and с
somewhat
>
or
yellovish, poe
ЖЕ
<
po
= 5
=
<
5.
ES
8
E
=
Б
2
8
y; leaves lacking minute peltate glan-
duni ићи stems glabrous ог pubes-
cent.
6a. A rs basifixed; corolla funnelform or
рел, usually yellowish ог
с, at е. 4 and nearly
salve nea), the tube and
lobes ови in ein i fruit thin-
Figure 6. Dyssochroma viridiflora 9 ут TO
J. 1818. Curtis's Botanical Magazine 45: t.
а the seeds elongate, pale or x
Markea
d
j Anthess dorsifixed; corolla ше, tu-
tube a
lobes thick and usage in texture; fruit
not yas walled as above, often some-
у, the seeds reniform, usually
at адаы АА 2. Juanulloa
1. Dyssochroma Miers, Апп. Mag. Nat. Hist. ser.
2, 4: 250. 1849. TYPE: Dyssochroma viridiflo-
ra (Sims) Miers ore Solandra viridiflora
Sims, 1818). Figure
Epiphytic shrubs or small trees, occasionally ter-
restrial; stems pendulous, the bark thin, flexible
and exfoliating, drying a dark, reddish brown.
Leaves elliptic, 5-12 X 2.5-5 cm, coriaceous, gla-
rous. Inflorescence terminal, a highly modified
cyme, congested, often consisting of a single flower,
Annals of the
Missouri Botanical Garden
glabrous. Buds elliptic, the corolla aestivation val-
vate. Pedicels glabrous. Flowers with the calyx di-
indi nearly to the base, the lobes 3-4 ст long,
gular, acuminate at the tips, coriaceous, gla-
ind corolla greenish or greenish purple, broadly
flaring to campanulate, the tube 5-6 cm long, the
lobes 1–2.5 cm long, at anthesis strongly reflexed;
filaments adnate to the corolla tube in the lower !^,
exserted or included, 7-9 cm long, glabrous along
their whole length; anthers basifixed, 1–1.5 ст
long; pollen 3-colporate with long colpi and uneven
reticulate ornamentation; ovary superior, glabrous;
style glabrous, 8-10 cm —- the stigma clavate.
Fruit and seeds not known. 2 spp., SE Brazilian
rainforests.
List of species. Dyssochroma longipes (Sendtn.)
Miers, Brazil (Markea parue (Sendtn.) Cuatr., So-
landra iiu Sen D. a (Sims)
Miers, Brazil (D i ui albidoflavum Lemaire
Markea peckoliorum Gilg, M. viridiflora (Sims)
Ducke, Solandra viridiflora Sims).
sochroma may be congeneric with Trianaea,
but since so few specimens of these plants are
known and nothing is known about the fruits or
seeds, we have maintained it as distinct pending
further "n The pollen is identical to most
species of Trianaea but differs from that of T. spe-
ciosa in that shite есы» ornamentation is discon-
tinuous, so that За. close to the poles lack muri
(see кет et al.,
The rhe species of Dyssochroma are
in fact es different stages of floral development
(L. F. de Carvalho, pers. comm.), with the included
anthers of D. longipes occurring at earlier stages of
flowering, while the exserted anthers of D. viridiflo-
ra are often seen on flowers found on the forest floor
post-flowering. Future studies being carried out at
the Jardim Bótanico de Rio de Janeiro will clarify
this situation.
2. — паа & Раубп, Prodr. Fl. Реги
t. de TYPE: Juanulloa parasitica ps
& i Feus 2A T.
Ectozoma OE m Mag. Nat. Hist. ser. 2, 3: 166.
1849. zoma pavonii Mie
Laureria leida Linnaea 8: 513. 1833. TYPE:
Laureria mexicana Schltdl.
Portaea Tenore, Atti 7a. Adum. "€ S 902, t. 1846.
TYPE: Portaea aurantiaca Ten
Rahowardiana D'Arcy, Ann. Missouri Bo. Gard. 60: 670.
pla —— 1974]. TYPE: Rahowardiana wardiana
D'A
ны ве Miers, Ann. Mag. Hist. ser. 2, 4: 190. 1849.
a Ruis & ~
Epiphytic shrubs or small trees, the stems often
hanging and pendulous from the canopy, 1-20 m;
bark of stems loose and exfoliating when dry, often
reddish or reddish brown. Leaves elliptic, ovate or
obovate, 6-30 X 5-9 cm, usually m— (sessile
in J. verrucosa), membranous or coriaceous, gla-
brous or pubescent, the trichomes grotte, sim-
ple or more often branched. Inflorescence terminal
or lateral, a variously modified cyme, occasionally
somewhat raceme-like, often many times branched,
the peduncle often very long and pendulous, usu-
ally many flowered (10-50) but occasionally re-
duced to 1-2 flowers (J. speciosa), glabrous or pu-
bescent, the trichomes uniseriate, simple or
branched, buds elongate, usually exserted from the
calyx tube, corolla aestivation quincuncial or im-
bricate. Flowers with the calyx lobed nearly to the
base or unlobed and inflated (J. speciosa), usually
somewhat fleshy and bright colored, % to equal to
the length of the corolla tube; corolla fleshy, fun-
nelform to narrowly tubular, 1-9 cm long, red, or-
ange, or cream with reddish or purple markings,
the tube either flaring at the apex (J. pavonii, J.
ferruginea) or more commonly slightly constricted,
the lobes minute and rounded or equal in length to
the tube (J. pavonii, J. ferruginea); filaments in-
serted near the base or ca. % way along the tube,
densely pubescent with uniseriate trichomes at the
point of insertion; anthers included in the corolla
tube at anthesis, very variable in size, dorsifixed;
pollen 3-colporate, either with long, narrow colpi
and scabrate-rugulate ornamentation or with short,
broad colpi and a scabrate-perforate ornamentation;
ovary ovoid to conical or beaked (J. wardiana, J.
globifera), glabrous; style glabrous, + equal to the
filaments, usually somewhat longer at anthesis, the
stigma capitate to somewhat clavate, bilobed. Fruit
green berry, ovoid to conical, usually within the
accrescent calyx, 1-3 cm long (in J. wardiana and
J. globifera seeds are borne in the ovary beak, but
mature fruits are unknown); seeds reniform, quite
large, 2-7 X 1.5-3 mm, dark brown or reddish
brown. 11 spp., Mexico to Bolivia.
Hunziker & Subils (1991).
List of species. Juanulloa ferruginea Cuatr., Co-
lombia to Peru; J. globifera (S. Knapp & D'Arcy)
S. Knapp, NW Colombia (Rahowardiana globifera
S. Knapp & D'Arcy); J. membranacea Rusby, Bo-
livia (J. pedunculata Rusby); J. mexicana (Schle-
chtendahl) Miers, Mexico and Central America to
N Colombia (J. aurantiaca Otto & Dietr., J. bicolor
Gleason, J. elliptica Dunal, J. hookeriana Miers, J.
panamensis Miers, J. sargii Donn. Sm., Laureria
mexicana Schlechtendahl, Markea tomentosa Lun-
Literature.
Volume 84, Number 1
1997
Knapp et al. 79
Tribe Juanulloeae
Fi ^ CN 1n
( ) h. & Hook. as Ectozoma Pavonni (plate 48 from Miers, J. 1857. Illustrations
gure 7
of South American bn
dell, Portaea aurantiaca Tenore); J. ochracea
uatr., Colombia to Peru; J. parasitica Ruiz & Pa-
vón, Ecuador, Peru, and Brazil
(Ruiz & Гао Pers.); J. parviflora (Ducke) Cuatr.,
own only the type)
за gi parviflora риске); Ј. a (Miers)
Benth. & Hook., W Ecuador and NW Peru (Ecto-
zoma pavonii Miers, Markea pavonii (Miers) D'Ar-
су); Ј. speciosa (Miers) Dunal, Colombia and Ес-
uador Miers); J.
verrucosa
(Rusby) Hunz., Bolivia (Markea verrucosa Rusby);
J. wardiana (D’Arcy) S. Knapp, Panama (Raho-
wardiana wardiana D'Arcy).
The flowers of most de species of Juanulloa
bes to the classic hummingbird pollination
e: they are tubular, brightly colored and
very thick and fleshy (Fig. 2A). However, J. pavonii
(Fig. 7) and J. ferruginea ( perhaps to some
extent J. parviflora) have quite different flowers,
which are greenish, with shorter tubes and some-
what reflexed lobes. These perhaps represent a dif-
ferent pollination syndrome.
Annals of the
Missouri Botanical Garden
Marekea coccinea.
Plate 45
Printed by Hullmandel & Walton.
Figure 8. Markea coccinea Rich. (plate 45 from Miers, J. 1857. Illustrations of South American Plants).
The very condensed globular inflorescence of
Juanulloa wardiana and J. globifera is unique in
the tribe, and in the family Solanaceae (illustrations
can be seen in Knapp & D’Arcy, 1993). In these
species the calyx as well as the corolla is brightly
colored and quite showy. These flowers are appar-
ently visited by hummingbirds, and produce copi-
ous nectar.
There are two types of pollen found in this genus
(Persson et al., 1994). In Juanulloa speciosa and J.
ochracea the pollen is 3-colporate with long, narrow
colpi and scabrate/rugulate exine ornamentation. In
the rest of the genus the pollen is 3-colporate with
short, broad colpi and a scabrate/perforate exine
ornamentation
3. Markea Rich., Actes Soc. Nat. Hist. Paris 1:
107. 1792. TYPE: Markea coccinea Rich. Fig-
ure 8.
Hawkesiophyton Hunz., Kurtziana 10: 39. 1979. TYPE:
Hawkesiophyton panamense (Standl.) Hunz. (basio-
nym Markea panamensis Standl.).
Lamarkea Pers., Synopsis 1: 218. 1805. TYPE: Lamarkea
|
3
|
|
|
5
i
|
ПРИШТИНУ
Volume 84, Number 1
1997
al. 81
Tribe Juanulloeae
coccinea (Rich.) Pers. (basionym Markea coccinea
Rich.).
Epiphytic shrubs or small trees, occasionally
growing on the forest floor in cloud forest. Leaves
elliptic to ovate, 5-25(-40) х 2.5-10(-20) cm, co-
riaceous or membranous, glabrous or pubescent,
the trichomes simple and uniseriate, multicellular.
Inflorescence a variously modified cyme, sometimes
superficially raceme-like, branched or simple, gla-
brous to minutely pubescent, + sessile (M. sessili-
flora), to borne on an elongate peduncle (M. coc-
cinea), 1-25 cm long, bearing 1-10 flowers at a
time, but with many scars, data on specimen labels
indicating that only one flower opens at a time.
Buds elongate, corolla aestivation imbricate or
cochlear. Pedicels glabrous or pubescent. Flowers
with the calyx lobed nearly to the base, the lobes
long-triangular, 0.8-3 cm long, often with distinc-
tively colored venation and long-acuminate tips,
glabrous or pubescent; corolla bright orange (M.
coccinea) or more often greenish purple or cream
colored, salverform (M. coccinea) to funnelform, the
tube 1-7 cm long, the lobes 0.5-1.2 cm long,
rounded and the margins occasionally somewhat la-
ciniate, not reflexed at anthesis, glabrous or pubes-
cent with simple, uniseriate trichomes on the ex-
terior surfaces; filaments adnate to and included in
the corolla tube, + equal in length to the corolla
tube or ca. % its length, pubescent at the point of
insertion; anthers basifixed, elongate, ca. 2 mm (M.
ulei, M. panamensis) to ca. 1 cm (M. sessiliflora, M.
camponoti) long, longitudinally dehiscent; pollen
3-colporate, ornamentation and apertures varying;
ovary superior, glabrous; style usually equal i
length with the filaments, or longer and held at the
mouth of the corolla tube, the stigma clavate. Fruit
an ellipsoid berry, 1-2 Х са. 1 cm, green when
ripe, the apex somewhat beaked from the persistent
style base, the pericarp thin and papery when dry,
in herbarium specimens often almost translucent,
held within the accrescent calyx; seeds many, long
rectangular, 2-5 X 0.5 mm, usually bright or-
ange. 9 spp., Panama to S Peru.
List of species. Markea camponoti Ducke, upper
Amazon basin; M. coccinea Rich., upper Amazon
basin to the Guianas (Lamarckea coccinea Pers.);
M. costanensis Steyerm., Venezuela, Cordillera de
la Costa; M. formicarum Dammer, Amazon basin,
on white sand; M. longiflora Miers, Trinidad (this
may in fact be an older name for the taxon herein
referred to as M. camponoti, but the species has not
been recollected in Trinidad since the type speci-
men, which has a single, badly damaged flower);
M. panamensis Standl., Panama and adjacent Co-
lombia нне у. panamense (Standl.)
unz., M. dimorpha C. V. Morton); M. sessiliflora
Ducke, N Amazon basin, бшшш, Roraima plateau
(M. porphyrobaphes Sandwith, M. reticulata Stey-
ire); *M. sturmii Cuatr., Colombia;
M. ulei (Dammer) Cuatr., upper Amazon basin (Ec-
tozoma ulei Dammer, Hawkesiophyton klugii Hunz.,
H. ulei (Dammer) Hunz.).
Markea as here defined is composed of two main
clades, one consisting of the small greenish-flow-
ered taxa previously treated as the genus Hawke-
siophyton, M. ulei and M. panamense, which per-
haps deserve sectional or subgeneric rank. Markea
sturmii, not treated in the cladistic analysis, is
somewhat intermediate between the small-flowered
clade and the following. The other clade consists
of five taxa, again divided into two clear groups:
one with short inflorescences and flowers drying
dark on the herbarium sheet (M. sessiliflora, M. cos-
tanensis) and the other consisting of mainly Ama-
zonian species with long-acuminate calyx lobes
with distinctive venation and usually elongate in-
florescences
The three types of pollen found in this genus
(Persson et al., 1994) correlate closely with these
oups. All are 3-colporate but they differ in exine
sculpturing and aperture type. In M. ulei and M.
panamensis the colpi are long and narrow and the
apertures distinctly beaked. Short colpi occur in
the M. camponoti type, where the ornamentation is
baculate and in the M. oo type, where it is
microreticulate to scabra
e genus Markea is in dire need of a field-
based monograph.
4. Merinthopodium Donn. Sm., Bot. Gaz. 23: 11.
1897. TYPE: Merinthopodium neuranthum
(Hemsl.) Donn. Sm. (basionym: Markea neu-
rantha Hemsl.). Figures 3A, 9.
Epiphytic shrubs or small trees, 1-10 m, often
clambering over branches and hanging from the
canopy; bark loose and exfoliating, bright silvery-
gray. Leaves elliptic to obovate, occasionally
strongly three-veined from the base, 10-15 X 5-
9 cm, glabrous to minutely puberulent along the
main veins beneath, the trichomes simple and uni-
seriate. Inflorescence an elongate, pendulous ra-
ceme, the peduncle 15-50 cm long, minutely pu-
bescent with tuberculate trichomes with
multiseriate bases bearing one or two uniseriate
simple branches, the branches breaking off leav-
ing the tuberculate bases which cause a warty
roughened texture on the peduncle, flowers borne
only at the tip or distal М, usually only 2-7 at a
time, but the inflorescence with up to 100 scars.
Buds ellipsoid, rapidly elongating to exceed the
yx tube, corolla aestiv alvate. Flowers
ngth of the corolla
tube at anthesis, membranous to coriaceous, lobe
nearly to the base, the lobes with blunt tips; co-
rolla greenish to greenish with various purple
markings, the tube 3-10 cm long, flaring early in
anthesis but at full anthesis broadly campanulate,
the lobes ca. 2 cm long, strongly reflexed at an-
thesis; filaments adnate to the base of the corolla
tube, exserted from it at anthesis, glabrous; an-
82 Annals of the
Missouri Botanical Garden
BOTANICAL GAZETTE, XXII.
MERINTHOPODIUM | NEURANTHUM, Donnell Smith.
9. Merinthopodium neuranthum (Hemsl.) Donn. Sm. (plate from Donnell Smith, J. 1897. Bot. Gaz. (Craw-
Fan 23).
thers bright yellow, dorsifixed and somewhat ver-
satile; pollen generally large, Ae залок with sca-
rate ornamentation ап ng aperture:
ovary аена glabrous; ide ion = aa
to the length of the filaments at anthesis, the stig-
ma capitate, bright green, strongly bi-lobed. Fruit
a green berry, ovoid, 1–2.5 cm long, the pericarp
bı
many, 1.5-2. х 0.
brown. 3 spp., S Mexico to N Venezuela and Co-
lombia.
List of ои „мдын pada Retiranthum
Volume 84, Number 1
1997
Knapp et al. 83
Tribe Juanulloeae
(Hemsl.) Donn. Sm., 5 Mexico and Central America
(Markea campanulata (Donn. Smith) Lundell, Mar-
kea gentlei Lundell, Markea internexa (Blake) Lun-
dell, Markea leptesthema (Blake) Cuatr., Markea
neurantha Hemsl., Me. campanulatum Donn.
mith, Me. шеиит Blake, Ме. leptesthemum
Blake) Me. pendulum (Cuatr.) Hunz., Colombia and
Venezuela (Markea pendula Cuatr.); *Me. vogelii
(Cuatr.) Castillo & R. E. Schultes, Colombia and
Venezuela (Markea vogelii Cuatr.).
The shape of the flowers of Merinthopodium de-
pends greatly on their state when collected. These
plants bloom at night and are probably pollinated
by bats (see Voss et al., 1980). Thus flowers col-
lected before full anthesis are shaped like a flaring
tube, but when fully in bloom, the limb becomes
broadly campanulate with strongly reflexed lobes.
This same situation occurs in Trianaea and Dys-
sochroma. The peculiar trichomes found on the pe-
duncle of Merinthopodium are also found in 7. nae-
ka S. Knapp of eastern Ecuador. One of the reasons
why there are many synonyms in this genus is that
specimens collected at different stages of flowering
have been regarded as distinct species. There is
also geographical variation in flower size and leaf
texture.
5. Schultesianthus Hunz., Kurtziana 10: 35.
1977. TYPE: Schultesianthus leucanthus
(Donn. Sm.) Hunz. (basionym: Markea leucan-
tha Donn. Sm.). Figure 10.
Epiphytic shrubs or small trees, occasionally ter-
restrial in wet cloud forests; bark usually somewhat
reddish, glabrous or pubescent with simple unise-
riate trichomes on the new growth; leaves borne in
congested groups separated by long naked inter-
nodes. Leaves elliptic, 6-15 X 2.5-9 ст, thick and
coriaceous, drying pale green or brownish, often
rounded at the tip, glabrous or pubescent with sim-
ple uniseriate trichomes, in addition all species
bearing minute glandular trichomes each with a
unicellular base and multicellular head sunk in a
pit on the upper and lower surfaces, giving the ap-
pearance of small red dots on dry specimens. In-
florescence a terminal cyme, variously branched,
occasionally reduced to a single flower (Sch. cros-
bianus), glabrous or variously pubescent with sim-
ple uniseriate trichomes. Buds elongate, the apex
swollen, corolla aestivation quincuncial or cochlear.
Pedicels glabrous or pubescent, usually quite short.
Flowers with the calyx very coriaceous, lobed Y
way or nearly to the base, the lobes strongly over-
lapping, glabrous or minutely glandular pubescent,
becoming woody and accrescent in fruit; corolla
funnelform to strongly infundibuliform, 4-15 cm
long, occasionally somewhat zygomorphic, strongly
and sweetly fragrant, pale green to cream (black in
S. crosbianus), yellowing with age, the lobes round-
ed and often somewhat laciniate; filaments adnate
with the corolla tube in the lower %, densely pu-
bescent at point of insertion and ca. % way to the
anthers above, regular or declinate, exserted from
the corolla tube at anthesis; anthers small and el-
liptic, or slightly larger and more elongate (in Sch.
venosus, Sch. uniflorus, and Sch. crosbianus with
non-declinate filaments); pollen 3-colporate, rugu-
late and covered in Ubisch bodies; ovary glabrous,
somewhat pointed, bilocular; style declinate or
straight, glabrous, the stigma broadly capitate, dis-
tinctly 2-lobed. Fruit a green to purplish green ber-
ry, 1-3 ст long, ovoid or somewhat pointed, woody
when dry, very leathery and fleshy when fresh, in-
vested in the accrescent woody calyx; seeds many,
reniform, large, 5-6 X 3—4 mm, pale yellowish tan
or dark brown. 8 spp., Mexico to Bolivia.
Literature. Bernardello € Hunziker (1991).
List of species. Schultesianthus coriaceus (Kun-
tze) Hunz., Antioquia, Colombia (Solandra coriacea
Kuntze); Sch. crosbianus (D’Arcy) S. Knapp, Costa
Rica and Panama (Markea crosbiana D'Arcy); Sch.
dudleyi Bernardello & Hunz., S Ecuador, S Peru;
Sch. leucanthus (Donn. Sm.) Hunz., 5 Mexico to 5
Peru (Markea leucantha Donn. Sm., Merinthopo-
dium leucanthum (Donn. Sm.) Blake, Metternichia
werklei Schumann ex Bois); Sch. megalandrus
(Dunal) Hunz., N Colombia and N Venezuela (Mar-
Кеа megalandra (Dunal) D'Arcy, M. suaveolens
Standl., Metternichia ? megalandra Dunal, Sch.
suaveolens (Standl.) Hunz., Solandra megalandra
(Dunal) Killip & Pittier); Sch. odoriferus (Cuatr.)
Hunz., Cordillera Oriental, Colombia (Solandra
odorifera Cuatr.); Sch. uniflorus (Lundell) 5. Knapp,
Chiapas, Mexico (Markea uniflora Lundell, Merin-
thopodium uniflorum (Lundell) Hunz.); Sch. venosus
(Standl. & C. V. Morton) S. Knapp, Costa Rica and
ama (Markea venosa Standl. & C. V. Morton).
D’Arcy (1973 [1974]), in his treatment of Markea
for Flora of Panama, clearly and correctly identified
the close relationships between the species of the
present genus Schultesianthus. He recognized the
three radially symmetrical flowered species, Sch.
crosbianus, Sch. uniflorus, and Sch. venosus,
members of the “perhaps generically distinct”
group including Sch. leucanthus (as Markea те-
galandra in his treatment).
Many of the species of Schultesianthus were orig-
inally described as Solandra, and S. boliviana and
Sch. coriaceus have been considered “linking” spe-
84 Annals
of the
Missouri Botanical Garden
BRITISH MUSEUM (NATURAL HISTORY)
Sample зр ane tion
PANAMA: Chiriquí; road from Gualaca
Chiriquí Grande. Roadside, north of 10s
Planes de Hornito, 8°40'N 82°13'w
1200 в.
Epiphytic shrub. Flowers white,
becoming yellow, strongly scented.
11 March 1985.
R.J. Hampshire & C. Whitefoord 469.
Figure 10. Schultesianthus leucanthus (Donn. Sm.) Hunz. (Hampshire & Whitefoord 469, BM)
cies. For discussion see comments with the generic 6. Solandra Swartz, nom. cons. 5 pa Vetensk.
description of Soland. Acad. Nya Handl. 8: 300-306. 1787. TYPE:
e flowers of EERE are among the Solandra grandiflora Swartz. Figures 2B, 11.
most showy in the Juanulloeae (see Fig. 10). They ا e
hi шагіѕа Gmelin, Systema naturae 2: 296, у -
are sweetly таван irs “+ ^t s С о Solandera Kuntze, Rev. Gen. Plantarum. 452. 1891 (or-
a creamy yellow with age. Large bees ave been thographic variant).
observed visiting the flowers, but little is known
about their biology. Woody lianas or high climbing epiphytic shrubs
Volume 84, Number 1
1997
Knapp et al. 85
Tribe Juanulloeae
id by $ буы Walworth Јан эз.
Figure 11. Solandra grandiflora (Sims, J. 1817. Curtis's Botanical Magazine 44: t. 1874).
or small trees; stems glabrous or minutely pubes-
cent, the trichomes glandular, simple or branched;
bark shiny and loose, exfoliating when dry. Leaves
elliptic, ovate or obovate, large and coriaceous,
glabrous or pubescent with trichomes like those of
the stems. Inflorescence terminal, a highly re-
duced cyme, usually with a single flower. Buds
elongate, strongly exserted from the calyx tube,
corolla aestivation quincuncial or cochlear. Pedi-
cels at anthesis stout and woody. Flowers with the
calyx 3-10 cm long, tubular, usually irregularly 2-
to 5-lobed and zygomorphic, the lobes equal in
length to the tube, occasionally somewhat pur-
plish, glabrous or pubescent; corolla very large
and showy, 13-40 cm long, zygomorphic, infun-
dibuliform to cyathiform, white to cream or yellow,
often with longitudinal purplish markings in the
throat, darkening with age, the lobes rounded, en-
tire to laciniate or fimbriate; filaments adnate to
the corolla tube, exserted or included, declinate,
densely pubescent at the point of insertion with
simple, uniseriate trichomes; anthers basifixed, 6–
13 mm long, longitudinally dehiscent; pollen
3-colporate, coarsely reticulate; ovary 2-carpel-
late, 4-locular, glabrous, partially sunken into the
receptacle and thus partly inferior; style equal in
length to the filaments, glabrous, declinate, the
stigma minute and capitate, bilobed. Fruit a leath-
ery berry, conical, green or yellow when ripe, the
calyx persistent and accrescent but splitting in
fruit; seeds round or reniform, 4-6 X 2.5-4 mm,
the cotyledons incumbent (fide Bernardello &
Hunziker, 1987). 10 spp., West Indies, Mexico to
the Amazon basin.
Bernardello & Hunziker (1987).
Literature.
List of species. Solandra brachycalyx Kuntze,
Costa Rica and Panama; *S. brevicalyx Standl., NE
Mexico; S. boliviana Britton, Bolivia; *S. grandiflo-
ra Swartz, West Indies, Central and South America
(Datura sarmentosa Lam., D. scandens Vellozo, S.
hirsuta Dunal, 5. nitida Zucc., 8. minor Griseb., 8.
scandens (Vell.) Toledo, Swartzia grandiflora
(Swartz) Gmelin); *S. guerrerensis Martínez, Duran-
Annals of the
Missouri Botanical Garden
go, Mexico; *S. guttata D. Don, Mexico; *S. lon-
giflora Tussac, West Indies, N South America (S.
laevis Hooker, S. macrantha Dunal, S. grandiflora
var. macrantha (Dunal) Voss); %5. maxima (Sessé
сто) Green, Mexico, Central and N
South America (Datura maxima Sessé & Мос̧ійо,
S. hartwegii N. E. Brown, S. selerae Dammer); *5.
nizandensis Matuda, Mexico, Guatemala, Honduras;
*S. paraensis Ducke, Amazon basin in Colombia,
French Guiana, Brazil.
The species of Solandra are widely cultivated for
their showy flowers (Fig. 2B) in the tropics and sub-
tropics pe. the world. These large yellowish
owers open in the evening, €x и bat
pollination. Species | in the genus used by the
Huichol and other groups in Meee as hallucino-
gens (see references in Bernardello & Hunziker,
1987). The partially inferior ovary of Solandra is
unique in Solanaceae. It is deeply sunk into the
receptacle at anthesis, but specimens in fruit ap-
pe to be totally superior.
andra boliviana has often been considered to
e a "linking" species between Solandra and
Schultesianthus as it is superficially similar to the
large-flowered Sch. coriaceus. Our results, however,
demonstrate that S. boliviana shares not only gross
morphological but also pollen synapomorphies with
the rest of Solandra but not Schultesianthus (see
Persson et al., 1994). The two genera are closely
related (in our eei: sister groups, but in the
analysis of Olmstead et al., in press, somewhat sep-
arate, see бейш of this article).
7. Trianaea Planch. & Linden, Prix-Courant 8: 4.
1853. TYPE: Trianaea nobilis Planch. & Lin-
den. Figures 3B, 12.
Poortmannia Drake, Bull. Soc. Philom. Paris, ser. 8, 4:
128. 1892. TYPE: Poortmannia speciosa Drake.
Epiphytic or terrestrial shrubs or small trees, of-
ten clambering over other vegetation; stems flexu-
ous, often dark purple when young; bark smooth or
verrucose, glabrous to densely pubescent, gray or
reddish brown. Leaves usually very large, 10-30 x
1-10 ст, narrowly linear to broadly elliptic, shiny
and coriaceous, glabrous to minutely pubescent,
densely hirsute in 7. naeka, the trichomes simple
and uniseriate. Buds elliptic, + halfway exserted
from the calyx tube, corolla aestivation quincuncial
or cochlear. Pedicels at anthesis short and stout to
somewhat inflated and strongly 5-angled, lobed ca.
% way to the base, coriaceous, usually drying dark
reddish brown or black, often somewhat purplish in
. Trianaea speciosa (Drake) Soler. aru et
al. 9121. TOP ess road, Ecuador), scale bar
live plants, glabrous to minutely pubescent with
simple uniseriate trichomes; corolla very large and
fleshy, 5-15 cm long, broadly campanulate, green
or greenish with purplish red markings on the
throat, Mes or densely pubescent without, the
trichomes simple and uniseriate, the lobes trian-
gular in outline, strongly reflexed at anthesis, the
margins revolute; filaments adnate to the corolla
tube in the lower М, usually strongly geniculate at
the base, connivent around the straight style, gla-
brous along their entire length; anthers elongate,
1–1.5 cm long, usually white, ventrifixed, somewhat
versatile; pollen 3-colporate, reticulate, with long,
narrow colpi; ovary glabrous or minutely pubescent,
style straight, glabrous, the stigma large, capitate
to clavate. Fruit a berry, 3-6 cm long, 8-10-locular,
green or purplish, the pericarp Pru the pulp
fleshy; seeds very small, 1-2 х 1 mm, reni-
form, but very thin, pale Riu tan. About 6
spp., Colombia to N Peru
List of species. Trianaea brevipes (Cuatr.) S
Knapp, Colombia and Ecuador (T. spectabilis Cuatr.
var. brevipes Cuatr.); T. naeka S. Knapp, SE Ecua-
dor, Cordillera del Condor; 7. пеотзае Romero-Cas-
Volume 84, Number 1
1997
Knapp et al. 87
Tribe Juanulloeae
tafieda, Sierra Nevada de Santa Marta, Colombia;
T. nobilis Planch. & Linden, Colombia and Ecua-
Peru (Poortmannia speciosa Drake, T. spectabilis
Cuatr.); T. sp. nov., NC Peru, Cajamarca.
Trianaea is one of the most poorly collected
genera in the Solanaceae. Its green flowers and
position high in the canopy make it difficult to see
and to access. The flowers of Trianaea are, like
Merinthopodium and Dyssochroma, typical bat-
pollinated flowers, and Vogel (1958) observed bats
visiting the flowers of T. brevipes in Colombia.
Persson et al. (1994) recognized two pollen types
in the genus. However, although these show dif-
ferences in the reticulum that enable them to be
distinguished, they are very similar in overall pol-
len morphology
The ovary of Trianaea is peculiar in Solanaceae
in being 4—5-carpellate rather than the more typical
2-carpellate ovary found in the rest of the family
(see Solereder, 1898, for a discussion). This char-
acter is difficult to see in flowering material, and
as most specimens have a single flower, dissection
is not desirable. Preserved collections are essential
for the future study of this genus and its relation-
ships to the rest of the green-flowered clade to
which it belongs
TAXA INCERTAE SEDIS
Markea lopezii Hunz., Cordillera Occidental,
Colombia.—In the cladistic analysis this and а re-
lated species recognized but not named by Hun-
ziker (1985) are the sister group of Solandra. The
paucity of specimens for analysis and the fact that
no fruiting specimens of Markea lopezii are known
means this ы needs further research.
The pollen of M. lopezii is anomalous in the Juan-
ulloeae (see Persson et al., 1994, and discussion
ve).
Several recent collections of epiphytic Solana-
ceae from Ecuador and adjacent Peru may certainly
represent new taxa, and as more intensive collect-
ing is done more specimens are likely to come to
light.
Literature Cited
w^ H: G. 1973; Evolutionary relationships between
owering plants and pm s in American — African
rainforests. Pp. 145- E. S. Ay-
ensu & W. p. sapiate (editors), “Tropical Fist
Ecosystems in Africa and South America: A Compar-
ative Review. Smithsonian Institution Prese, Washing-
to;
Bernadello, L. M. & A. T. Hunziker. 1987. A сенки
revision of Solandra (Solanaceae). Nordic J. В
639-652.
— ي ag The genus Schultesianthus e
lanaceae): за
. Lester, M. Nee, R. N.
Estrada peste Solanaceae MI: lg frs Chemistry,
—— Royal Botanic банни
г, J. М. 994, Successive sure reliability
vids turn dll rey 10: 5-220.
Cuatrecasas, J. New chiropterous Solanaceae from
Colombia. J. wash, Acad. Sci. 49: 269-272
D'Arcy, W. С. 1973 [1974]. Solanaceae. In: R. E. Wood-
son & R. W. Schery, Jr. — Flora of Panama. Ann.
Missouri Bot. Gard. 60:
Drummond III, B Brown, ig 1987. Ithomiinae
тамом Nymphalidae): Summary Ы oa M
MEE ~ Missouri Bot. Gard. 7.
Ded A . Plantes nouvelles ou stig connues “re la
région amazonie cage anaceae. Arch. Jard. Bot. Rio
de psi 1(1): 54-
1922. dim pis ou peu connues de la
région amazonienne ч partie). Solanaceae. Arch. Jard.
Bot. Rio be eiro 3: 250-252.
Farris, J. S. . A successive approximations approach
to быв ie Syst. Zool. 18: 374-385.
———. 1988. Hennig86: Version 1.5. Published by the
. 1989. The retention index and the rescaled con-
sistency index. ud 5: 41 md
, A. G. Kluge € M. J. Eckhardt. 1970. À numer-
ical пати to Фу im RA Syst. Zool.
2-189.
Goloboff, 2 1993. PeeWee. MS-DOS program. Published
y the author.
Hunziker, gs T 1977. Estudios sobre Solanaceae VIII.
Novedades varias € tribus, géneros, secciones y es-
pecies - Fs érica. Kurtziana 10: 7—50.
South лешип Solanaceae: A synoptic س
. С. Hawkes, В. N. Lester & A.
D. Waist (editor), bi Aes and Taxonomy of the
Solanaceae. Academic London
1985. Estudios Rid Solna XX. Markea
lopezii, nueva mee x rem imer 5; 9-12.
& L. M. Bernan 1989. Sobre la posición
ie de а ас Kurtziana 20:
215.
—— & К. Subils. 1991. Estudios sobre Solanaceae
XXXIL gad taxonómica de Juanulloa. Kurtziana
21:2
Knapp, $ ved New taxa and combinations in the Juan-
ulloeae (Solanaceae). tm 5: 281-283.
W. G. D'Arcy. 1993. Rahowardiana жыш
(Solanaceae), а new species from Colombia. Novon 3
429-430.
Miers, J. 1857. Illustrations of South American Plants.
bic 2. H. Bailliére, London.
Nix . Carpenter. 1993. On outgroups.
Cladistics 9; 413-426.
Olmstead, R. G. & J. D. Palmer. 1992. oplast
DNA phylogeny of the Solanaceae: a lation -
ships and character evolution. Ann. Missouri Bot.
79: i
& . Sweere. 1994. Combining data in phy-
logenetic omda: An empirical approach using
three molecular data sets in = Solanaceae. Syst. Biol.
— Spengler, | vi Bohs ај р. Palmer,
In press. ic and prov
88
Annals of the
Missouri Botanical Garden
Solanaceae based on chloroplast DNA. In Solanaceae
IV. Royal Botanic Gardens, Kew.
NE Mis S. Knapp & S. Blackmore. 1994.
rpho and systematics of tribe Juanulloeae A.T.
ila 0 Бошан, Rev. Palaeobot. Palynol. 83:
130.
Pollen
— ——— & ——— In press. Pollen paid rey
and the phylogenetic Hl ag of Datura L. rug-
mansia Pers. In: Solanaceae IV. Royal Bolas Кыс,
Кем.
Platnick, N. I. 1989. An empirical comparison of micro-
puter parsimony programs, II. Cladistics 5: 113-
C. E. Griswold & J. A. Coddington. 1991. On
missing entries in cladistic analysis. Cladistics 7: 337—
343.
Scotland, R. W., P. K. Endress & T. J. Lawrence. 1994.
ara ontogeny and aestivation in the Acanthaceae.
t. J. Linn. Soc. 114
Zwei Beiträge = ee der So-
lanaceen. Ber Deutsch; B 242-260.
Treseder, K., K. Davidson & D. W. ызыны 1995. АЬ-
sorption of ant- -provided carbon vo "o nitrogen by
a tropical epiphyte. Nature 375: 137—
Vogel, S. 1958. Fledermausblumen in са Oes-
terr. Bot. Z. 104: 491-530
oss, R uye, M. Fisher & R. Cort
. D. Wheeler. The outgroup
comparison method of character „о Syst. Zool. 30:
-11.
Willson. M. F. 1983. Plant Reproductive Ecology. John
Wiley & Sons, New York.
Appendix. Selected Specimens Examined.
Atropa belladonna L.—Morocco: Mohamed 146 (BM)
Lycium cestroides Schltdl.—Bolivia: Fiebrig 2213 c
Nicandra physaloides (L.) Gaertner—Cultivated: Higgins
149 (BM).
7769 (BM), LAUNE & De Haas 2543
Dyssochroma viridiflora (Sims) Miers— Brazil: Anon s.n.
(BM), Mello pues 1822 | (К); Риз 1894 (К).
Juanulloa ferruginea Cuatrecasas—Peru: Vásquez, Jar-
eg Arévalo & Lépez 4305 (NY), Encarnacion 1285
Dyssochroma longipes (Sendtner) Miers—Brazil: Sellow
M).
Nas globifera (S. Knapp & D'Arcy) S. Knapp—Co-
lombia: Cogollo, Ramírez & Alvarez 2895 (JAUM).
Juanulloa membranacea Rusby— Bolivia: Britton & Rusby
s.n
Juanulloa mexicana (Schltdl.) Miers—Colombia: Law-
rance s.n. (BM); Costa Rica: Anon s.n. (BM); Mexico:
Cedillo & er 133 (BM).
Juanulloa ochracea Cuatrecasas—Colombia: Schultes &
Cabrerg s.n. BM); Soie: Baker & Trusher 6118
N Ayala, Vásquez, Torres & Calderón s.n.
(NY), Klug . 2360 (K).
Juanulloa parasitica Ruiz € Pavón—Bolivia: Nee &
Coimbra 40085 (NY); Peru: Foster 9924 (NY), Plow-
& Schunke V. 11765 (K).
Juanulloa p (Miers) Benth. & Hook.—Ecuador: Pa-
BM), dei & Alcorn 14351 (NY); Peru:
peris & Schunke 427 (NY).
Meet speciosa (Miers) Dunal—Colombia: Luteyn, Du-
nt & Buritica 4884 (NY), Anon s.n. (BM).
Juanulloa verrucosa (Rusby) Hunz.—Bolivia: Steinbach
8466 (BM).
Juanulloa уље (D'Arcy) 5. Кпарр—Рапата: Mc-
Donagh, umpel & Plumptre 251 (BM), Luteyn
4096 (NY
Markea camponoti Ducke—Brazil: Daly, Campbell, ње
da Silva, Bahia & dos Santos D879 (NY), Guedes s.
(BM); Guyana: Hay 5414 (K).
Markea coccinea L. C. Richard—Brazil: Egler 47679
(NY), Mori, Mora Cardoso, deSilva & Sothers 20215
(NY); Colombia: Schultes & Cabrera s.n. (BM); Guyana:
Myers 5953 (K); Peru: Foster, Fernández & Vivar 10766
NY).
Markea costanensis rm.—Venezuela: Steyermark &
Davidse 116312 ШО Steyermark & Davidse 116951
(NY).
Markea formicarum Ducke—Peru: Ayala, Vasquez, Torres
& Calderón 2563 (NY); Venezuela: Nee 30923 (NY).
Markea lopezii Hunz.—Colombia: Croat 50164 (MO), Gen-
try, Juncosa € Gomez 40820 (NY).
Markea РЕН is Standl.—Colombia: Forero, Jaramillo
NY); Panama: de Nevers, Herrera &
Charnley эү (К), Gentry 8761 (NY), Kirkbride & Bris-
ton 1454 (NY), McDonagh, Lewis, Gumpel & Plumptre
101 (BM).
Markea poe ora Ducke—Guyana: Anon 3974 (K), i
man 633 (K), Pipoly 7556 (NY) Brazil: Ducke s
(
Майга. ulei (Dammer) Cuatr.—Peru: a & Jaramillo
9647 (NY), Ramos & Lima 17170 (NY)
Merinthopodium neuranthum . Sm.—Be-
ne Hazlett & Coe 2748 (NY), Peck 508 (NY); pro.
bia:
6096 (NY), McPherson & ^m 10175 (ВМ); Vene-
Pop ee Lebron-Luteyn, Ruíz-Terán & Пират
TT coriaceus (Kuntze) Hunz.—Colombia: Ne
& Callejas 32545 (NY), Tracey s.n. (K), Zarucchi, pass
& Betancour 6034 (NY
d саан (D’Arcy) S. B ee
ica: Davidse, Herrera & Grayum 28688 (NY); Panama:
Folsom, Channell, Dressler & er 7117 (MO), de
Nevers & Charnley 6058 (BM).
Schultesianthus dudleyi Bernardello & Hunz.—Ecuador:
Madsen 86180 ma Madsen & ти 75160 (ВМ).
Schultesianthus leuc Sm.) Hunz.—Panama:
ampshire $e Whiteford ies өзде
бейта А nal) Hunz.—Costa R
Haber & Crue 84 8402 (BM), петта, 502 (ВМ); rane
Steyermark & Steyermark 95;
Schultesianthus uniflorus (Lundell) E Knapp—Mexico:
Breedlove & Smith 31710 (MO).
лн venosus (Standley € C. V. Morton) 5.
& Solomon
048 (NY), Croat 51550 (MO).
iden brachycalyx Kuntze—Costa Rica: Lankester s.n.
(K); Panama: Hammel 7360 (BM).
Trianaea brevipes С же. S. Knapp—Ecuador: Palacios &
van der Werff 36 73 (NY
Trianaea naeka S. Knapp—Ecuador: Neill & Cerón 7449
N
Volume 84, Number 1 Knapp et al. 89
| 1997 Tribe Juanulloeae
Trianaea neovisae Romero-Castañeda—Peru: Díaz & (NY), Gentry, Bonifaz & Loor 30950 (MO), Dodson &
| б: 2944 (МО). Dodson 18592 (NY); Spruce 5527 (К); Peru: Barbour
Trianaea nobilis Planch. & Linden—Colombia: Silver- 4113 (MO).
| stone-Sopkin 1 188 (MO); Ecuador: Zak 1028 (К). Trianaea sp. nov.—Peru: Díaz, Osores & Bustamente 3935
| Trianaea speciosa (Drake) Soler.—Ecuador: Boeke 547 (MO).
|
3
1
3
4
1
A REVISION OF THE GENUS
XIPHOTHECA (FABACEAE)!
Anne Lise Schutte?
ABSTRACT
The genus Xiphotheca Eckl. & Zeyh. consists of nine species, all. endemic to the Flora Capensis region of South
morphologica
that there are two distinct groi
sects. Seans and Xiphotheca. Full descriptions, diagnostic os illustrations, and distribution maps of the
species are presented.
When De Candolle (1825a, b, 1826) described
the genus Priestleya, he established two sections:
P. sect. Aneisothea (with the calyx base attenuate)
and P. sect. Priestleya (with the calyx base intrusive
or “thrust in”). In 1836, Ecklon and Zeyher con-
stituted the genus Xiphotheca by according generic
status to P. sect. Aneisothea. This concept was not
кеу by en successors Ји a 1836; Wal-
, 1839; Ben reek 5: у, 1862;
Be Ss vito chose to follow ‘De Can-
dolle’s classification. Recently, however, Schutte
and Van Wyk (1993) died. sional evidence in
support of Ecklon and Zeyher’s (1836) concept and
subsequently reinstated the genus Xiphotheca. The
name Xiphotheca is a compound word, derived from
the Greek words “xipho-” meaning sword-like and
*-theca" meaning case or container, which refers to
the shape of the pod.
Xiphotheca is a genus of papilionoid legumes
comprising nine species, which are all endemic to
the Cape fynbos region of South Africa. It is a mem-
ber of the tribe Liparieae, which differs from the
closely related tribe Podalyrieae in the fusion of the
stamens into an open sheath or a closed tube. The
Podalyrieae have the stamens free almost to the
base (Polhill, 1976, 1981a, b). In a recent phylo-
genetic analysis of dod ae the genera
of the Podalyrieae and Liparieae, Van Wyk and
Schutte (1995) showed that ен Мов Атрћића-
lea Eckl. & Zeyh., and Coelidium Vogel ex Walp.
are undoubtedly monophyletic, but that more re-
search is needed to clarify the positions of some of
the other genera. Data, gained from chemical in-
vestigations, have brought new insights into the re-
lationships between the two tribes. The results of
this oe will be published elsewhere.
ostic characters of Xiphotheca are the
ae unspecialized bright yellow flowers; the
decussate, 2-flowered inflorescences; the non-intru-
sive calyx base (except for one species); the fusion
of the bract with the base of the pedicel; the pres-
ence of bracteoles (albeit sometimes strongly re-
duced); the laterally compressed pods, which are
constricted between the seeds; and the presence of
anabasine as a major alkaloid. From its closest rel-
atives, Amphithalea and Coelidium, it differs in the
ћаре of the seed aril, which lacks the extension
toward the lens; the generally higher ovule number;
and the absence of ammodendrine as a major al-
kaloid.
Apart from a brief synopsis, in which the no-
menclature, synonymy, and typification of the spe-
cies are discussed (Schutte & Van Wyk, 1993), no
taxonomic treatment of the group has been pub-
lished since Harvey’s in 1862. The aim of this pa-
per is to present a revision of Xiphotheca, in which
a phylogeny is proposed for the genus, followed by
an identification key, full agi illustrations,
and distribution maps of the speci
MATERIALS AND METHODS
Data on the morphological variation of the taxa
OL, C, G, G-DC, JRAU, K, LD, LINN, MO, NBG,
P, PRE, S, SAM, SBT, TCD, UPS, W, WU, and Z
' This study formed part of a Ph.D. thesis in Botany at the Rand Afrikaans University. J am indebted to B.-E. van
Wyk for кылы gee the study. The directors and s
My husband, J. H. J. V
Financial support from the Foundat
ы Departm ent of Botany, Rand jt
Republic of South Afri
taff of the mentioned herbaria are thanked for loa:
ok (Cape iugi Conservation kindly offered advice and assistance during collecting trips.
are a и is acknowledged
ns niveis РО. , 2006
South Africa. Present — Compton Habent National. Bolo Institute, Private Bag X7,
ans of specimens.
Auckland Park, Johannesburg, Republic
of
7735 Claremont,
ANN. Missourt Bor. GARD. 84: 90-102. 1997.
Volume 84, Number 1
1997
Schutte
Revision of Xiphotheca
(acronyms as in Holmgren et al., 1981), as well as
from fresh or preserved material collected during
field trips.
The methods applied in the oer a study are
described by Van Wyk et al. ). Voucher
specimens of the material used sa dà extraction
of alkaloids are also listed.
Cladograms were generated by using the com-
puter software package Hennig 86 (Farris, 1988).
Character states were polarized using the method
of outgroup comparison. It is important to note that
autapomorphies for the different species have been
omitted from the analysis, since they serve no pur-
pose as grouping characters. The “mhennig*,”
“bb*,” and “ie” algorithms were applied to produce
trees of minimal length.
MORPHOLOGICAL CHARACTERS
HABIT
The variation in habit found in Xiphotheca in-
cludes single-stemmed, tree-like shrubs up to 2.5
m tall, many-stemmed, virgate shrubs up to 1.2 m
tall, and prostrate or straggling shrublets of up to
0.5 m tall. Adaptations to survive recurrent fires
have had a major influence on the life forms and
habit of the taxa, since all are restricted to the fire-
prone fynbos vegetation of the Cape (Le Maitre &
Midgley, 1992; Schutte et al., 1995). There are two
main fire survival strategies: sprouters and non-
sprouters. Sprouters have a lignotuber from which
new growth takes place after fire, resulting in a
many-stemmed appearance at ground level. Non-
sprouters, on the other hand, can only reproduce
from seed after fire and are easily recognized by
the presence of a single main stem, at least at
ground level (Schutte et al., 1995).
t the specific level, the ability to resprout or
reseed after fire is a taxonomically important and
very useful character in Xiphotheca. For example,
X. canescens (Thunb.) A. L. Schutte & B.-E. van
Wyk and X. elliptica (DC.) A. L. Schutte & B.-E.
van Wyk look morphologically very similar, es-
pecially on herbarium sheets, but have different
fire-survival strategies.
Fire survival strategy is not included as a char-
acter in the cladistic analyses because it is a poly-
morphic character. In. X. fruticosa, some popula-
tions are sprouters, while other populations consist
of non-sprouting individuals.
LEAVES
All the species of Xiphotheca have simple, pet-
iolate, and distinctly pinnately veined leaves. Stip-
ules are invariably present, but reduced in size.
Leaves are generally flat, except in X. phylicoides
A. L. Schutte & B.-E. van Wyk, where the margins
are strongly revolute. The vestiture of the leaves
varies among the species from pubescent to seri-
ceous to tomentose or velutinous, and some of the
species can be identified by their leaf indumentum.
INFLORESCENCES
Xiphotheca has axillary, simple racemose inflo-
rescences, with geminate flowers. At the infrage-
neric level, inflorescences are particularly useful in
distinguishing between some of the species. In X.
canescens, X. cordifolia A. L. Schutte & B.-E. van
Wyk, X. elliptica, and X. phylicoides, the inflores-
cences are distinctly pedunculate. They are either
borne on lateral twigs, as in X. canescens, X. cor-
difolia, and X. elliptica, or borne on the main stem
as in the remainder of the species. In X. fruticosa
(L.) A. L. Schutte & B.-E. van Wyk, X. guthriei (L.
Bolus) A. L. Schutte & B.-E. van Wyk, X. Јапсео-
ata (E. Mey) Eckl. & Zeyh., and X. reflexa
(Thunb.) A. L. Schutte & B.-E. van Wyk, the inflo-
rescence-supporting leaves are smaller than the
other vegetative leaves and the inflorescence units
are congested. The flowers are totally concealed by
the leaves below the inflorescence units in X. guth-
riei and X. lanceolata. All the species have the
bracts fused with the pedicel at the base for a dis-
tance of 0.5 to 1.0 mm. Bracteoles are conspicuous
in X. canescens, X. elliptica, and X. phylicoides,
strongly reduced in X. cordifolia, X. reflexa, and X.
tecta (DC.) A. L. Schutte & B.-E. van Wyk, and
absent in the other species.
FLOWERS
Calyx. An attenuate calyx base is characteris-
tic of Xiphotheca. There is, however, one species
that has an intrusive calyx base (Schutte & Van
Wyk, 1993), but it is here regarded as a secondary
development. The upper two calyx lobes are in-
variably fused higher up than the lower three lobes.
Xiphotheca fruticosa, X. guthriei, X. lanceolata, and
X. reflexa are exceptional in having the carinal ca-
lyx lobe notably longer than the other lobes. The
shape of the calyx lobes is a significant taxonomic
character that varies from narrowly triangular and
acuminate to rounded and obtuse.
rolla. The corolla is yellow and PRENA
poe in Xiphotheca and tends turn
brown with age. Keel petals are obtuse, ыйды
pocketed, and auriculate. The pocket on the wing
petal is a thickened lobe, formed on the inside (ab-
92
Annals of the
Missouri Botanical Garden
axial side), but X. tecta is unusual in having a well-
developed pocket conspicuous also on the adaxial
side. Wing petal sculpturing is invariably present
in the upper basal area. The wing auricle is well
differentiated in X. canescens and X. elliptica, but
weakly so in the other species.
In Xiphotheca the stamens are dia-
delphous and the anthers almost uniform in shape
and size. The mode of attachment of the filaments
is alternately dorsifixed and subbasifixed.
Pistils. The number of ovules varies from two
to eight in the genus. These differences are useful
in distinguishing between some species.
FRUITS AND SEEDS
The significance of fruit characters as a taxonom-
ic character for Xiphotheca is obvious; the generic
name alludes to the shape of the pod. Pods are
sessile, laterally compressed, and constricted be-
tween the seeds in Xiphotheca. The seeds are ar-
illate and vary in color from uniformly green or
brown to green, mottled brown and brown, mottled
black.
CHROMOSOME NUMBER
Chromosome numbers of only three species have
so far been recorded: X. fruticosa, X. guthriei, and
Х. tecta. All have 2n = 18 (Dahlgren, 1967; Schut-
te, 1995). A basic chromosome number of x = 9
seems likely. This is in accordance with the pro-
posed base number for the tribe Liparieae (Gold-
blatt, 1981).
ALKALOIDS
Alkaloids have proved to be of great significance
at the generic level. In fact, it was one of the most
convincing characters used to motivate the rein-
statement of Xiphotheca as a genus (Schutte & Van
Wyk, 1993). The major alkaloids detected in Pries-
tleya DC. sect. Aneisothea DC. (now Xiphotheca)
were anabasine (a bipiperidyl alkaloid) and lupi-
nine (a bicyclic те alkaloid), while a
combination of lupanine- and sparteine-type alka-
loids (all tetracyclic кш гар alkaloids) and
minor quantities of ammodendrine е bipiperidyl
alkaloid) were located in Priestleya sensu stricto
(Van Wyk et al., 1991b). This offered additional
support to morphological evidence that Priestleya
is paraphyletic, which subsequently resulted in the
reinstatement of Xiphotheca (Schutte & Van Wyk,
1993). At the specific level, no clear pattern in
variation could be found, since the differences were
of a quantitative, rather than a qualitative, nature.
Table 1. Characters and character states used for the
cladistic analysis of the genus Xiphotheca. The fully re-
solved cladogram generated from this data set is shown in
Figure 1.
Taxa Character states
AMPHITHALEA 00000 00000 0
sce 013101. 110011
X. cordifolia 01101 1000? 1
Х. elliptica 01101 11001 1
fruticosa 10010 00100 0
X. guthriei 10020 00110 0
X. lanceolata 10020 00110 0
X. phylicoides 01001 10000 0
X. reflexa 10010 10110 1
X. tecta 01000 10000 1
chum
. Inflorescence supporting leaves: similar to others (0);
reduced in size (1).
2. In — па units: congested (0); not congested (1).
3. Inflorescences: borne on main stems (0); borne on lat-
eral twigs
4. Inflorescences: not concealed by leaves below inflo-
ence units (0); partly ыа br leaves below
1 nio rescence units ioi totally concealed by leaves be-
low inflorescence
. Peduncles: Ru y (0); recent (1).
Ü Bracteoles: absent hoe present (1).
7. Calyx lobes: acuminate (0); not acuminate (1).
8. taba lower lobe: as oe as the others (0); longer than
the other lobes (1
9. Ves ме роон (0); пої риЬезсеп! (1).
10. Wing petals: auricle weakly developed or absent (0);
ЖЕНЫ, auriculate (1
11. Ovule number: 2 (0); more than 2 (1).
INFRAGENERIC RELATIONSHIPS
Xiphotheca is defined by at least three unambig-
uous apomorphies and one subject to variation: (1)
the presence of bracteoles in most species; (2) the
fusion of the bracts with the base of the pedicel;
(3) the laterally oe and pods; and (4) the ac-
cumulation of an e as a major alkaloid
For the ا йа lin RE e was
chosen as outgroup, since it is the genus most
related to Xiphotheca (Van Wyk & Schutte,
- A data set was taedis. using 10 taxa an
11 queis (Table 1). Two fully resolved trees
resulted, both with a length of 15 and a consistency
index of 80. The one most favored is shown in Fig-
ure 1. In the other tree the positions of X. tecta and
X. phylicoides are switched. The cladogram indi-
cates two major clades: a Xiphotheca guthriei-group
(A), subtended by three synapomorphies, and a
Xiphotheca canescens-group (B) supported by one
apomorphy. These two infrageneric groups are giv-
en formal taxonomic rank below.
Volume 84, Number 1
1997
Schutte
Revision of Xiphotheca
Figure 1. Fully resolved sep of relationships in the genus Xiphotheca, based on the data set in Table 1 [dot,
ngle,
an apomorphy without homoplasy; rec an apomorphy
character;
TAXONOMIC TREATMENT
Xiphotheea Eckl. & Zeyh., Enum. Pl. Afric. Aus-
tral. 2: 166. 1836. TYPE: Xiphotheca rotun-
difolia Eckl. & Zeyh. NU PA designated b
Schutte & Van Wyk, 1993) [= Xiphotheca tecta
(Thunb.) A. L. Schutte & B.-E. van Wyk].
ie i m sect. =, DC., in Ann. Sci. Nat.
5, Prodr. 2: 121. 1825. TYPE: Priestleya i
iita. De. (lectotype, тения ated by Schutte & ыч
Wyk, 1993) [= де otheca elliptica (DC.) А.
Schutte & B.-E. у yk].
ооду shrubs or shrublets. Leaves alternate or
ы opposite or subopposite, simple, narrowly el-
liptic to almost circular, mostly flat, sometimes with
recurved margins, pinnately veined; petiole short,
= 1 mm long; stipules inconspicuous, less than 0.5
y with subsequent reversal or successive states of a multistate
=, a convergence; stars, Mm icti Ма (see text)].
mm long. /nflorescence axillary, 2-flowered, with the
two flowers opposite, aggregated into synflores-
cences of up to 20 flowers. Bracts linear to oblan-
ceolate, fused at the base with pedicel for 0.5-1.0
mm. Bracteoles minute, sometimes lacking. Corolla
yellow, longer than the calyx, glabrous. Calyx nar-
rowed to the base, rarely intrusive; upper two lobes
fused higher up than the lower three lobes; carinal
lobe sometimes longer than the upper four. Stan-
dard petal suborbicular to elliptic; apex emarginate.
Wing petals oblong, longer than the keel; the tips
imbricate; pocket developed as a thickened lobe
toward the inside. Keel petals widely obovate, with
weakly developed pockets, apex obtuse. Stamens
diadelphous, the vexillary filament free; anthers +
uniform in shape and size, alternately dorsifixed
and subbasifixed. Pistil sessile; style slender,
94
Annals of the
Missouri Botanical Garden
slightly upcurved, glabrous; ovary with 2 or more
ovules, densely sericeous to tomentose. Pods cori-
aceous, usually linear, sometimes obliquely oblong,
laterally compressed, constricted between the
seeds, 2 to many seeded, densely pubescent, to-
mentose, villous or glabrous. Seeds oblong-reniform;
hilum elliptic, surrounded by a fleshy collar-like
aril. Chromosome number 2n = 18. Nine species.
Xiphotheca is restricted to the fynbos region of
the Western and Northern Cape Provinces.
KEY TO THE SPECIES OF XIPHOTHECA
7. X. cordifolia
1. Leaves subopposite or alternate; calyx base not
Пие 2
. Inflorescences pedunculate Lnd
Inflorescences not pedunculate 5
. Ovary with 2 ovules
Fora a E SA yg
х es
4. Leaves narrowly elliptic, with flat margins; sin-
gle-stemmed tree-like shrubs up to 2.5 m tall;
nga. а X. canescens
5(2). Ovary with 5 or more ovules 6
5. wary with Zor 3 ovules с
6(5). Wing petal with pocket conspicuous on outer
surface; calyx shorter than keel, lobes triangu-
lar, tomentose; seeds green, not mottled |...
сае . X. tecta
6. Wing petal with pocket not conspicuous on out-
er surface; calyx + as long as the keel, lobes
oriens triangular, villous; seeds "тж ees
blac X. reflexa
е раан calyx lobe much longer than rdg tube
CUBE . lanceolata
7. Lower calyx lobe as long as or socio: than the
ube ee a ы
8(7). Leaves elliptic, silver, fene sericeous (long
silky appressed hairs) on both surfaces; Se
5-9 mm long; es file greenish pd mot-
tled dark brow X. fruticosa
8. Leaves пат old elliptic, green, = velu-
tinous (long soft upright hairs) on both surfaces,
densely so on плава bracts + 2.5 mm long;
seeds uniformly bro . X. guthriei
Section 1. Xiphotheca sect. Congestae A. L.
Schutte, sect. nov. TYPE: Xiphotheca reflexa
(Thunb.) A. L. Schutte & B.-E. van Wyk
Sectioni Xiphothecae similis, sed inflorescentiis parti-
alibus congestis et lobo calycis carinali longiori dif-
fert.
Section Congestae is similar to section Xiphoth-
eca but deviates in its congested inflorescence units
and in the carinal lobe of the calyx that is longer
than the upper four lobes. This section comprises
four species.
1. Xiphotheca fruticosa (L. А. L. Schutte &
B.-E. van Wyk, Taxon 42: 46. 1993. Lotus fru-
ticosus L., Syst. Nat. (ed. 10): 1179. 1759.
TYPE: South Africa. Western Cape, without lo-
cality, Anon. s.n. (lectotype, designated by
Schutte & Van Wyk (1993), S, Linnaeus Type
Herb. No. 5293.5).
Crotalaria lanata Thunb., Prodr. Pl. Cap.: 124. 1800.
TYPE: South Africa. Western Cape, *e Cap. b.
Spei," Thunberg s.n. (lectotype, designated by Schut-
te & Van Wyk (1993), UPS, Herb. Thunberg No.
Priestleya villosa , Prodr. 2: 122. 1825, nom. йер.
Xiphotheca Tm (DC.) Eckl. € Zeyh., Enum. Pl.
Afric. Austral. 2: 166. 1836. TYPE: South m
Western Cape, “Cap. de B. Esp.," Lambert s.n. (lec-
totype, designated by Schutte & Van Wyk (1993),
G-DC).
Single-stemmed, tree-like shrub up to 2 m tall,
not sprouting after fire, or sometimes, a many-
stemmed shrub up to 0.7 m tall, sprouting after fire;
woody rootstock sometimes present. Leaves alter-
nate, elliptic, flat, silvery, densely sericeous on both
surfaces, glabrescent. Inflorescences aggregated into
head-like synflorescences at tips of main branches,
somewhat concealed by supporting leaves; pedun-
cle absent. Bracts narrowly elliptic to linear, 5-9
mm long. Pedicel 2.5-3.0 mm long. Bracteoles ab-
sent. Calyx not intrusive at base; lobes acuminate,
shorter than the tube; carinal lobe slightly longer
than upper four; densely sericeous. Wing petals au-
pigens Le inconspicuous on outer surface.
2 to 3 ovules; densely tomentose. Pods
inflated, tomentose. Seeds pale greenish brown,
mottled dark brown. Figure 2.
Xiphotheca fruticosa is found on the Cape Pen-
insula-, Hottentotsholland-, Hex River-, and
Touwsberg Mountains, as well as on the mountains
near Montagu and hills near Bredasdorp (Fig. 3).
It grows in a sandy loamy soil at altitudes of 100
to 1200 m
This species resembles X. guthriei but differs in
longer bracts and long silky appressed, not veluti-
nous, hairs on the leaves.
Both sprouting and non-sprouting growth forms
occur in this species. Populations from the Bredas-
dorp-Elim area resprout after fire, while those from
the other localities do not. This difference in growt
form may be significant, but more fieldwork 18
needed before specific or infraspecific status can
be assigned with certitude.
Selected specimens examined. Non-sprouting form
SOUTH AFRICA. Cape Peninsula: Noord Hoek Moun-
tain, Barker 2080 (NBG). Cape Town: Devil's Peak, Bolus
3765 (SAM). Simons Town: Chapman's Peak, Pillans 5."
95
Volume 84, Number 1
1997
Schutte
Revision of Xiphotheca
‘uu ш LIN “EEE 19YIMOJ :А рог 129108 :S *$9Z9 sund пи MOET `чоио :0 1679 AMYN :q “ON FOZZ AMY W :6£09 107407, vy u's ffouomT Y ‘f 1 2692
snjog :H °9 :ggay 4940 73 *Э 998€ 244M9 :4 ‘q *g ‘ amyag Y 30]A сү “sunay ampu :д “(у “| имај олтешиц :4 ised Яшм :N YM “Я “(yey eui oi soqo] ладап) sao£qeo ij,
‘O ‘f “а ‘mara [eıa1e[ ur вләмор :S ‘N ‘J °9 ¿mala јетхеде *saago[ zy IN ‘H °9 ^g ју 'ртјогоиој у “AH "Psoonmaf "у O-N Px "ХЛ uym? оогоцту 4-ү ^6 amay
I
i
=
96 Annals of the
Missouri Botanical Garden
O Ae:
ex?
at : | (> > P
۹ 1
»
29" у du
4} 2
HEIGHT ABOVE SEA LEVEL
E Over 1500
30" ~ ET] 900 - 1500 m
5 (73 300 - 900 m
: = C Under 300 m
Vs E iu | 20 0 20 40 60 80 100km
эт : > к= ы |
ш ~ х o f
da > we’
ars = ~ T
LE ale А
FRL ~; a
3 LN = 3 à ғ
ET ~
vad 2 ae : Hm pde E ЫЕ: i
4 A el ea
ro ' = i. à Bit
2 коле BL Мга т ES
за" =
** r La e 20" 27 22'E 23° 24° 25° 26" 27
The known geographical distribution of Xiphotheca fruticosa (dots, non-sprouting form; squares, resprout-
Figu
ing form) Х. reflexa (triangles).
(BOL 51233). Montagu: Pypsteelfontein, Schutte 673-675
(JRAU). Ladismith: Touwsberg, Vlok & Schutte 155 (MO).
Sprouting form: SOUTH AFRICA. Elim: Koueberge,
kloof above Nuwepos, Oliver 5854 (NBG, PRE). Bredas-
о r slopes of mountain above Bredasdorp, Burgers
2708 (NBG, PRE), Vlok & Schutte 365 (MO).
2. Xiphotheca reflexa (Thunb.) A. L. Schutte &
B.-E. van Wyk, Taxon 42: 47. 1993. Crotalaria
тећеха Thunb., Prodr. Pl. Сар.: 125. 1800.
TYPE: South А са. Western Саре, “e Cap. b.
Esp.” Thunberg s.n. (lectotype, designated by
Schutte & Van Wyk (1993), UPS, Herb. Thun-
berg no. 16576).
Many-stemmed prostrate to straggling shrub up to
0.5 m tall, sprouting from a woody rootstock after
fire. Leaves alternate, ovate to lanceolate, flat, dense-
ly sericeo-tomentose on both surfaces, glabrescent.
Inflorescences scattered along main branches, partly
concealed by supporting leaves; peduncle absent.
Bracts linear, 5-8 mm long. Pedicel 2.5-3.0 mm
long. Bracteoles minute, caducous. Calyx not intru-
sive at base; lobes acuminate, much longer than the
tube; carinal lobe longer than the upper four, almost
as long as the keel petals; densely sericeous. Wing
petals auriculate; pocket inconspicuous on outer sur-
face. Ovary with 5-7 ovules; densely sericeous. Pods
laterally pis ae n — villous. Seeds brown,
mottled black. F
This species is also restricted to the southwest-
em Cape, from the Piquetberg in the north south-
eastward to the Caledon-Elim region in the south
(Fig. 3). Xiphotheca reflexa is usually found at low
altitudes (less than 300 m), growing in deep sandy
soils.
Characters distinguishing X. reflexa from X. lan-
ceolata are the sericeo-tomentose leaves and the
several-seeded (5—7) pods. Xiphotheca lanceolata
as a densely ае vestiture on ће
leaves, with rather stiff hairs on the margins and
midrib and 2- or 3-seeded im
Selected specimens examined. SOUTH AFRICA. Mal-
mesbury: Farm Bokbaai near Darling, Barker 10589
(NBG). Stellenbosch: Bottelary, прање: 12935 (NBG).
Malmesbury: near Hopefield, Compton 18928 (NBG).
Cape Town: lower slo e Mountain, Esterhuysen
15637 (BOL). Cape Peninsula: Оена Penfold 98
(NBG).
3. трн lanceolata (E. Mey.) Eckl. &
Zeyh., Enum. Pl. Afric. Austral. 2: 167. 1836.
S Lat lanceolata E. Mey., Linnaea 7: 150.
1832. TYPE: South Africa. Western Cape.
|
B
Volume 84, Number 1
1997
Schutte 97
Revision of Xiphotheca
HEIGHT ABOVE SEA LEVEL
20 0 20 40 60 BO 100km
prr rrr
Figure 4. The known geographical distribution of Xiphotheca lanceolata (triangles) and X. guthriei (dots).
“Kapsche Flüche bei Constantia,” Ecklon s.n.
(lectotype, designated by Schutte & Van Wyk
(1993), S).
Priestleya glauca T. M. Salter, J. S. Afr. Bot. 8: 256. 1942.
TYPE: South Africa. Western Cape, on lower slopes
of Hercules’ Pillar, Joostenberg, Pillans 6264 (lec-
totype, designated by Schutte & Van Wyk (1993),
BOL; isolectotypes, K, NBG).
Single-stemmed shrublet up to 0.6 m tall, not
sprouting after fire; woody rootstock absent. Leaves
alternate, narrowly elliptic, flat, densely appressed-
sericeous on both surfaces with long, rather stiff
hairs on margins and midrib, glabrescent. Inflores-
cences aggregated into head-like synflorescences at
tips of main branches, almost completely concealed
by supporting leaves; peduncle absent. Bracts lin-
ear, 5.5-6.0 mm long. Pedicel + 1.5 mm long.
Bracteoles absent. Calyx not intrusive at base; lobes
acuminate, much longer than the tube; carinal lobe
longer than the upper four; densely sericeous with
long, rather stiff hairs on lobes and main veins.
Wing petals auriculate; pocket inconspicuous on
outer surface. Ovary with 2 or 3 ovules; hirsute.
Pods laterally compressed; hirsute. Seeds раје
brown, mottled dark brown. Figure 2.
Xiphotheca lanceolata is a rare species that oc-
curs only on the granite hills in and around the
Cape flats (Fig. 4), at altitudes between 60 and 200
m. It is seriously threatened by urban development.
See discussion under X. reflexa.
Selected specimens examined. SOUTH AFRICA. Stel-
lenbosch: Faure, Barker 4121 (NBG). Somerset West: Ver-
gelegen, Compton 6410 (NBG). Paarl: top of granite hill
above Nooitgedacht, Dahlgren & Strid 4109 (LD); North-
ern slopes of Joostenberg, Pillans 6264 (BOL, NBG). Stel-
lenbosch: Sir Louwry’s Pass, Stokoe s.n. (SAM 64930).
4. Xiphotheca guthriei (L. Bolus) A. L. Schutte
& B.-E. van Wyk, Taxon 42: 46. 1993. Pries-
tleya guthriei L. Bolus, Ann. Bolus Herb. 4:
125. 1928. TYPE: South Africa. Western
Cape, hills near Elim, Guthrie 3866 (holotype,
BOL).
Single-stemmed shrub up to 0.3 m tall, not
sprouting after fire; woody rootstock absent. Leaves
alternate, elliptic to narrowly elliptic, flat, veluti-
nous on both surfaces, glabrescent. Inflorescences
aggregated into head-like synflorescences at tips of
main branches, almost completely concealed by
supporting leaves; peduncle absent. Bracts linear,
+ 2.5 mm long. Pedicel + 2 mm long. Bracteoles
absent. Calyx not intrusive at base; lobes triangular
acuminate, + as long as the tube; carinal lobe as
long as lateral lobes; velutinous. Wing petals not
98
Annals of the
Missouri Botanical Garden
auriculate; pocket inconspicuous on outer surface.
Ovary with 2 ovules; densely sericeo-tomentose.
Immature pods velutinous. Immature seeds uniform-
ly brown. Figure
Xiphotheca guthriei is limited to the surround-
ings of Bredasdorp, Elim, and Caledon (Fig. 4),
where it grows in loamy, clayey soil at altitudes
below 250 m. The survival of this species is threat-
ened by farming.
See discussion under Х. fruticosa.
Selected specimens examined. SOUTH AFRI-
CA. Bredasdorp: Kourivier, between Napier and
Stanford, Jordaan 976 (C); Farm Klein Uintjieskuil
just E of Viljoenshof, Oliver 4288 (PRE). Heidel-
aid hill N of Verkykerskop, Schutte 760 (JRAU).
Bredasdorp: ca. 3 km E of Viljoenshof, Vlok &
Schutte 4 (JRAU).
Section 2. Xiphotheca sect. Xiphotheca
This section differs from section Congestae in the
extended flowering units, in the inflorescences,
which are mostly pedunculate, and in the carinal
calyx lobe, which is as long as the upper four lobes.
It contains five species.
5. Xiphotheca phylicoides А. = Schutte &
B.-E. van Wyk, Taxon 42: 48. 1993. TYPE:
South Africa. Oudtshoorn Pen. lower north-
em slopes of Outeniqua Mountains on farm
Klein Moerasrivier, Vlok 2640 (holotype, PRE;
isotypes, B, BOL, JRAU, K, MO, NBG).
Many-stemmed shrub up to 1.2 m tall, sprouting
from a woody rootstock after fire. Leaves alternate,
elliptic to narrowly elliptic, with strongly revolute
margins, sparsely sericeous on adaxial surface,
soon becoming glabrous, densely sericeous on ab-
axial surface. Inflorescences borne along main
branches; peduncle 1.5-2.5 mm long. Bracts linear
to narrowly elliptic, 3-4 mm long. Pedicel 3.5—5.0
mm long. Bracteoles + 0.5 mm long, caducous. Ca-
lyx not intrusive at base; lobes acuminate, + as
long as the tube; carinal lobe as long as lateral
lobes; densely pubescent. Wing petals weakly au-
riculate; pocket inconspicuous on outer surface.
Ovary with 2 ovules; densely pubescent. Jmmature
pods laterally compressed; pubescent. Seeds un-
known. Figure 5
This species is known from only two localities on
the Outeniqua Mountains near Mossel Bay (Fig. 6).
Xiphotheca phylicoides is found in pebbly loamy
soil at altitudes between 530 and 800 m
The leaves of X. phylicoides are characteristic in
having strongly revolute margins.
Specimens examined. SOUTH AFRICA. Oudtshoorn:
= wer northern slopes of Outeniqua Mountains on farm
lein Moerasrivier, Schutte 801 (JRAU), Vlok 2437
ОБА, ), 2640 (В, BOL, JRAU, К, MO, NBG, PRE). Mos-
sel fee Attaquaskloof Nature Reserve, Vlok 2500
(JRAU).
6. Xiphotheca tecta (Thunb.) A. L. Schutte 4
B.-E. van Wyk, Taxon 42: 48. 1993. Liparia
tecta Thunb., Prodr. Pl. Cap.: 124. 1800
Priestleya tecta (Thunb.) DC., Prodr. 2: 122.
1825. TYPE: South Africa. Western Cape,
“Paardeberg, Picketberg, Hottentots Holland-
berg,” Thunberg s.n. (lectotype, designated by
Schutte € Van Wyk (1993), UPS, Herb. Thun-
berg Мо. 17009; isolectotype, 5).
didt Ages ee Eckl. & Zeyh., Enum. Pl. Afric
Austral. 2 836. Priestleya deren (Eckl.
& Zeyh.) don Linnaea 13: 469. 7
tecta var. rotundifolia . Cap.
2: 20. 1862. TYPE: South Africa. Western Cape, “In
lapidosis laterum monti
nie (Worcester),” Ecklon & Zeyher 1224 (lec-
totype, designated by Schutte & Van Wyk (1993), 5;
isolectotype, S).
ape ve polycarpa Eckl. & Zeyh., Enum. Pl. Afric
stral. 2 versa par э аў South Africa. Western
que “In locis lap s laterum montium prope
Klin па ај Ecklon & Zeyher 1225
(lectotype, designated by Schutte & Van Wyk (1993),
S; isolectotypes, S,
Priestleya stokoei L. Bolus, Ann. Bolus Herb. 4: 69. 1927.
TYPE: South Africa. Western Cape, Stellenbosch di-
vision, foothills of mountains near Lourensford, Som-
erset West, Stokoe 1375 (holotype, BOL).
Many-stemmed shrub up to 1 m tall, sprouting
from a woody rootstock after fire. Leaves alternate,
elliptic to almost circular, flat, often concave,
densely pubescent to tomentose on both surfaces,
glabrescent. Inflorescences borne along main
branches; peduncle absent. Bracts linear, 2.5-7.5
mm long. Pedicel 2—4 mm long. Bracteoles minute,
caducous. Calyx not intrusive at base; lobes acu-
minate to acute, slightly longer than the tube; ca-
rinal lobe as long as lateral lobes; densely pubes-
cent. Wing petals not auriculate; pocket
conspicuous on the outer surface. Ovary with
ovules; densely pubescent. Pods laterally com-
pressed; densely tomentose. Seeds green, not mot-
tled. Figure 5.
Xiphotheca tecta has a relatively wide distribu-
tion in the Western Cape, extending from Citrusdal
in the north to Somerset West in the south (Fig. 6).
It occurs on shale or granite soil at altitudes of 200
to 1350 m.
This species is unique in having a pocket on the
wing petals that is conspicuous on the outer sur-
ace.
3 шш ut OLIS 2162 тор `M 5069 102L A=], “9211 1001999049 :S ‘ZEOT I wouns Y ‘9p omuos X YOJA :0-М “292 mos : ‘26S emus 21-1 90025 SAMAST :Н-4 ‘108
omms т “0998 101A “IV "mag әлцеш Hy ‘smug mera :Д\ “И “Н “Я ‘speed Burm :A 0) *T ‘G "јој ayy о1 saqo] 1eddn) sao£qeo : YN *9 ‘D ХӘ ur apunped jo aouesaud oy 2100)
MOLA [FIT] ш SIOMOY i1, ‘O ‘f *g “mora jexeqe “вәлвә] :S “М TY “Y "ома "y “AS `зиәэзәиюә "y H-N DA "Y ‘W-1 "Prjofipaoo "y Нол "sepiooréud vooyioudiy "47v `S omit
5
2
e
X
5
с
0 б
52
50
Ax
є
Ф
а
=
=
= 2
+
со
Ф
Е ~
33
> =
ENT ENDS
Annals of the
Missouri Botanical Garden
ies c] Mn
е Sse
"oc aa
OR ON E
Ра 27
Figure 6. Тһе known geographical distribution of Xiphotheca tecta (dots) and X. phylicoides (triangles).
Selected specimens examined. SOUTH AFRICA
Paarl: Frenc
Schutte 714 (JRAU). Tulbagh: near Tul-
bagh Waterfall, Stokoe 1399 (BOL).
7. Xiphotheca cordifolia A. L. Schutte & B.-E.
van Wyk, Taxon 42: 48. 1993. TYPE: South
Africa. Western Cape, Worcester district, Hex
River Mountains, Milner Kloof, Esterhuysen
31640 (holotype, BOL; isotypes, K, 5
Single-stemmed, tree-like shrub up to 2.5 m tall,
not sprouting after fire; woody rootstock absent.
Leaves opposite, cordate, flat, pubescent only on ab-
axial surface, glabrescent. Inflorescences bome at tips
of lateral branches; peduncle 1.0-2.5 mm long.
Bracts not seen. Pedicel 2.0-2.5 mm long. Bracteoles
minute, caducous. Calyx intrusive at base; lobes
acuminate, slightly longer than the tube; carinal lobe
as long as lateral lobes; densely pubescent. Wing
petals not seen. Ovary with 4–6 ovules; densely pu-
bescent. Pods laterally compressed; densely pubes-
cent. Seeds uniformly brown. Figure 5.
The extremely limited geographical distribution
of X. cordifolia is illustrated in Figure 7. It has
been recorded only from the Hex River Mountains
north of Worcester in the Western Cape, where it
grows in rocky areas at streamsides at 1333 to 1666
m above sea level. The only known collections are
in the fruiting stage.
This species is characterized by its opposite, cor-
date leaves.
Specimens examined. SOUTH AFRICA. Worcester:
Hex River Mountains, Milner Kloof, Esterhuysen 32556
(BOL); Moraine Kloof, Esterhuysen 35642 (BOL).
8. Xiphotheca canescens (Thunb.) A. L. Schutte
& B.-E. van Wyk, Taxon 42: 46. 1993. Hypo-
calyptus canescens Thunb., Nov. Gen. Pl. 11:
153. 1800. TYPE: South Africa. Northern
Cape, “e Cap. b. Spei,” Thunberg s.n. (lecto-
type, designated by Schutte & Van Wyk
(1993), UPS, Herb. Thunberg No. 16339).
Priestleya schlechteri L. Bolus, Ann. Bolus Herb. 4: 125.
1928. TYPE: South Africa. Northern C
Schlechter 10943 (lectotype, designated by Schutte
& Van Wyk (1993), BOL; isolectotypes, BM, BOL,
GK LD, S, V, 2).
Single-stemmed, tree-like shrub up to 2.5 m tall,
not sprouting after fire; woody rootstock absent.
Volume 84, Number 1
1997
Schutte 101
Revision of Xiphotheca
x
С we = =
i
(Vi
s T Lu
=
8
HEIGHT ABOVE SEA LEVEL
1500 m
20 0 20 40 60 80 100km
tddi
The known geographical distribution of Xiphotheca canescens (triangles), X. elliptica (dots), and X. cor-
difolia (square).
Leaves alternate, narrowly elliptic, flat, densely pu-
bescent on both surfaces, glabrescent. Inflorescences
borne at tips of lateral branches; peduncle 1.0-2.5
mm long. Bracts linear, + 2 mm long. Pedicel 2-3
mm long. Bracteoles minute, caducous. Calyx not in-
trusive at base; lobes acute, shorter than the tube;
carinal lobe as long as lateral lobes; densely pubes-
cent. Wing petals distinctly auriculate; pocket incon-
spicuous on outer surface. Ovary with 5-8 ovules;
densely pubescent. Pods compressed; + glabrous.
Seeds uniformly brown. Figure 5.
Xiphotheca canescens is a rare and highly local-
ized species, known only from the area around
Nieuwoudtville in the Northern Cape (Fig. 7). It
occurs on shallow Table Mountain Sandstone in
rocky areas at altitudes of 660 to 780 m.
The species is closely related to X. elliptica, but
deviates in being a non-sprouter and having nar-
rowly elliptic leaves with flat margins. Xiphotheca
elliptica is a sprouter and has elliptic leaves, with
slightly recurved margins.
Selected specimens examined. SOUTH AFRICA.
Nieuwoudtville: Oorlogs Kloof, Compton 20892 (NBG); top
of Van ass, Goldblatt 2469 (NBG); 4 mi. W of
Nieuwoudtville, Lewis s.n. (SAM 64929); Farm Klein Ar-
endskraal, Van Wyk 1343 (JRAU), Farm Hotbergfontein,
Vlok & Schutte 46 (MO).
9. Xiphotheca elliptica (DC.) A. L. Schutte &
B.-E. van Wyk, Taxon 42: 46. 1993. Priestleya
elliptica DC., Prodr. 2: 122. 1825. TYPE:
South Africa. Western Cape, “Cap. de B.
Esp.,” Lambert s.n. (lectotype, designated by
Schutte & Van Wyk (1993), G-DC).
Ingenhoussia verticillata E. Mey., Comm. Pl. Afr. Austr. 1:
toitskloof, 3000-3500 ped.," Drége s.n. (lectotype,
designated by Schutte & Van Wyk (1993), P; isolec-
totypes, K, S).
Many-stemmed shrub up to 1 m tall, sprouting
from a woody rootstock after fire. Leaves suboppo-
site, elliptic, margins slightly recurved, densely pu-
bescent on both surfaces, glabrescent. Inflores-
cences borne at tips of lateral branches; peduncle
1.0–1.5 mm long. Bracts linear, 2.0-2.5 mm long.
Pedicel 3.54.0 mm long. Bracteoles minute, ca-
ducous. Calyx not intrusive at base; lobes acute,
shorter than the tube; carinal lobe as long as lateral
lobes; densely pubescent. Wing petals distinctly au-
riculate; pocket inconspicuous on outer surface.
Ovary with 5 or 6 ovules; densely pubescent. Pods
laterally compressed; densely pubescent. Seeds uni-
formly brown. Figure 5.
The distribution of X. elliptica is limited to the
Annals of the
Missouri Botanical Garden
mountains above Paarl, Stellenbosch, and Caledon
(Fig. 7). It grows on granite soil with an overlay of
Table Mountain Sandstone at altitudes ranging be-
tween 365 and 1400 m
See discussion under X. canescens.
Selected specimens ro SOUTH AFRICA. Cal-
and Rivier Sonder End,
: B
rg Mountain,
n. (SA ). Paarl: aimee. Mountains,
Tierkloof, a z^ (NBG).
Literature Cited
Bentham, C. 1843. Enumeration of — indig-
enous to southern Asia, and loeis and sou
rica. Hook.,
: G. Ben ‚р
Hooker (editors) урин тни) Wu 1 wx ai
Dahiya. R 1967. Chromosome numbers in some South
African genera of the tribe Genisteae s. lat. (Legumi-
nosae). Bot. Not. 120: 149-160.
De Candolle, A. P. 1825a. Notice sur quelques genres et
pum nouvelles de légumineuses. Ann. Sci. Nat. 4:
—, 18255. руде Systematis Naturalis Regni
sk o
— ——. 1826. lanin sur la famille des légumineuses.
Ecklon, С. F. & K. L. P. Zeyher. 1836. Enumeratio Plan-
(editors), Advances | in Legume бума кы 2. Kosal Bo-
tanic Gardens, Kew.
Harvey, W. H. 1862. Leguminosae. Pp. 1-285 in W. Н.
a & O. W. Sonder (editors), Flora Capensis, Vol.
"а gg Dublin
Holmgre n ER. W W. Keuken & E. K. RM 1981.
ap M ы: Regnum Veg.
Hutchinson, J. 1964. The Сеа of P ЗИМА. Plants,
Vol. 1. Oxford Univ. Press, Oxford.
Le Maitre, D. C. & J. J. Midgley. 1992. Plant reproduc-
tive ecology. Pp. 135-174 in R. M. Cowling (odia
The Ecology of Fynbos—Nutrients, Fire and Diversity.
Oxford Univ. Press, Cape Town.
Meyer, E. H. F. 1836. Commentariorum de Plantis Afri-
cae Australioris 1. Leopoldum Voss, Leipzig.
Polhill, R. M. 1976. Genisteae (Adans.) Benth. and re-
lated tribes (Leguminosae). Bot. Syst. 1: 143-368
. 198la. Tribe 27. Podalyrieae Benth. Pp. 3
397 in R. M. Polhill & P. H. Raven (editors), al
Ph.D. Thesis, Rand Afrikaans University, Johannes-
urg.
— 6; B.-E 1993. The reinstatement of
the genus Fita (Fabaceae). Taxon 42: 9.
Н;
and evolutionary importance in fynbos legumes. Pl.
Syst. Evol. 195: 243-259.
Van Wyk, B.-E. & A. L. Schutte. 1995. Phylogenetic re-
lationships in the tribes Podalyrieae, i and Cro-
talarieae. Pp. 283-308 in M. D. Crisp & J. J. Da
(editors), Advances in Legume пес 7. Royal Во-
tanic a Kew
. Н. Verdoorn & В. Greinwald. 1991а. Taxo-
mic г понео of alkaloids in the genus Liparia
(раска рапа) $. African J. Bot. 57: 344—347.
P. Bachmann. 1991b. Tax-
alkaloids in the genus
onomie signific ance of major
Priestleya. Biochem. Syst. € Ecol. 19: 595-598.
Walpers, С. С. 1839. je ва criticae in Leg-
uminosas Capenses. Linnaea 13: 449-543.
ње Cant SEE TOLL
ПРУ у
А REVIEW OF THE GENUS
ECCREMOCARPUS
(BIGNONIACEAE)'?
William G. D'Arcy?
GENTRY INVITATION SERIES
This is the first of a series of publications paying tribute to or using data left K the late Alwyn H. Gentry, who died
in an airplane crash in Ecuador on August
3. The Bignoniaceae had been his prim
nographic theme. From
a base at the Missouri Botanical Garden, Ein pine to many parts of the world дл зан field and herbarium
studies of this family. Some of his best known taxonomic work appeared in the Flora Neotropica series and in the floras
of Panama, Veracruz
Ecuador, and Venezuela. A summary of his
life and scientific contributions, with a complete list
of his publications, was provided by James S. Miller and collaborators in the Annals of the Missouri Botanical Garden
b
volume 83, number
Among materials left by Gentry is a computer database information from some 55,000 collections he had
examined from more than 122 herbaria around the world. He
a treatment of the Bignoniaceae of Colombia, which is to be published i in the Flora de
to explore and utilize the rich legacy left by Al Gentry. This Invitation Series is being
relying on material left by
e made to their work. Those interested should contact William G. D'Arcy at the Missouri
GD
rs are encourage
established for taxonomic publications by people who are г
to other contributions
Botanical Garden for details.—W
o left a series of unpublished manuscripts, including
olombia series.
Gentry, or who wish to pay antes
ABSTRACT
Eccremocarpus is a genus of three species that grow in = pan x: Anet South America. The plants are vines
with showy flowers. Morphology divides the genus into tw
Eccremocarpus and section Calampelis. This paper reviews "e work left lo the late Alwyn Н.
oups that are also geographically separated: section
Gentry and that of others
and reduces the number of species from former concepts to three. A key to the species and a map of their distribution
are provided.
The genus Eccremocarpus is distinct from other
Bignoniaceae in its dissected leaves, parietal pla-
centation, and aseptate, dehiscent capsules, and it
forms its own tribe, the Eccremocarpeae. It is also
distinct in its elevational range, which is well above
that of the rest of the family. Eccremocarpus in-
cludes three species forming two distinctive species
groups. One of these, section Eccremocarpus, oc-
curs in Colombia, Ecuador, and Peru. It includes
plants of variable appearance but only two clearly
distinguishable species, Eccremocarpus huainac-
capac and E. viridis. The other, section Calampelis,
with one species, E. scaber, occurs in Chile and
Argentina and its plants show less ied te
HISTORY
Eccremocarpus was first described in 1794 by
Ruiz and Pavón, who soon after (1798) described
two species, Eccremocarpus viridis from Peru, and
E. scaber from Chile. Eccremocarpus scaber was rec-
ognized in 1819 by D. Don as a separate genus,
Calampelis, which effectively lectotypified Eccre-
mocarpus with E. viridis. The first description and
figure of Eccremocarpus by Ruiz and Pavón is of E.
viridis and not of E. scaber. Other species described
by subsequent botanists, some in other genera, are
dealt wi А
Macbride (1961) treated the [iom in the Flora
of Peru, recognizing three speci
Eccremocarpus was revised b Sandwith (1965),
who hesitantly recognized six species and placed
them into two sections, sect. Eccremocarpus with
five species, and sect. Calampelis with only E. sca-
ber. Sandwith provided a description only for his
new E. vargasianus.
paper is number 1 of the GENTRY INVITATION SERIES, in acknowledgment of contributions to the study
! This
of the Bignoniaceae qu y Alwyn H
Financial support
m The
aid DEB-9509270) is e eee The Field Museum of Natural History, the
n D. and Сићепве T. MacArthur Foundation and the National Science Foundation
Royal Botanic Gardens, Kew, and
The New York Botanical Garden ger made their collections of Есстетосатриз available for this study.
3 Missouri Botanical Garden,
0. Box 299, St. Louis, Missouri 63166, U.S.A.
ANN. MISSOURI Bor. GARD. 84: 103-111. 1997.
Annals of the
Missouri Botanical Garden
Muñoz (1966) provided a description and illus-
tration of the Chilean Eccremocarpus scaber, reduc-
ing the variety sepium to synonymy.
The late Alwyn H. Gentry, whose aborted mono-
graphic studies of the Bignoniaceae this paper com-
memorates, followed the taxonomy of Sandwith. In
publication, Gentry treated one species, Eccremo-
carpus longiflorus, in his floras of Ecuador (1977)
and Colombia (in press), and he listed E. huain-
accapac, E. longiflorus, E. scaber, E. vargasianus,
and E. viridis for Peru (Brako & Zarucchi, 1993).
These species also appear as determinations in his
computerized database. However, Gentry made no
reference to one species recognized by Sandwith,
E. lobbianus Zahlbr. This element is known by
scant material in Europe, and it may not have been
studied by Gentry. Gentry provided a description
only for E. viridis (as E. longiflorus, see Gentry
1977).
Careful review of most of the material seen by
Sandwith and Gentry, and some additional collec-
tions not seen by them, has led me to the hesitant
conclusion that Eccremocarpus lobbianus, E. lon-
giflorus, Е. vargasianus, and E. viridis are variants
within a single species that should be known as E.
viridis. It cannot be known whether Gentry might
have continued to follow Sandwith’s concepts had
he lived to revise the genus as a whole, but in other
treatments, he did tend to have a narrower species
concept than the present author. Unfortunately,
none of the students of the genus as a whole, Sand-
with, Gentry, and the present writer, collected or is
known to have seen wild populations of Eccremo-
carpus, although Sandwith may well have seen cul-
tivated plants at Kew. This treatment departs from
the Sandwith/Gentry concept in recognizing only
three species, but in deference to the previous tra-
dition, indication is made of how taxa and collec-
tions were viewed by these earlier workers. In ci-
tations that follow, reviewers of specimens are
indicated by superscripts as: 5 N. У. Sandwith, © A.
H. Gentry, ' present author. No symbol means not
seen by any of these.
SYSTEMATICS
The tribe Eccremocarpeae was established by A
De Candolle (1845) to embrace the single genus
Eccremocarpus. This concept has been followed by
later workers (Sandwith, 1965; Gentry & Tomb,
1979 [1980].
Tribe Eccremocarpeae is characterized as having
a 1-locular ovary with two bifid, parietal placentas
(Gandhi & Thomas, 1983), a loculicidal capsule
that remains fused apically, and winged seeds. The
plants are vines with opposite, dissected leaves and
tendrils (Melchior, 1964: 456). Gentry and Tomb
(1979 [1980]) reported similarities in the pollen of
Eccremocarpus to that of Jacaranda of tribe Teco-
meae and to that of Tourettia of tribe Tourrettieae.
The fruit in tribe Tecomeae also is loculicidal.
Two groups are clearly identifiable in the genus.
Section Calampelis comprises the small-flowered,
relatively uniform Eccremocarpus scaber, which is
represented by abundant collections mainly from
Chile. Section Eccremocarpus comprises plants sim-
ilar to the type species, displaying larger flowers
and fruits than in section Calampelis. These plants
are found to the north of Chile in Peru, Ecuador,
and Colombia, and they are known by fewer, more
variable collections. Main differences between the
two groups are given in the key to species.
The reduction of section Eccremocarpus into two
species is done with some hesitation, especially as
it breaks with the Sandwith/Gentry tradition of rec-
ognizing four or five species. Judging from the few
color photographs seen, different morphs of E. vir-
idis look very different, an impression stemming
mainly from striking color differences in the caly-
ces and differences in flower dimensions. Corolla
length appears to differ considerably within the
same inflorescence. The varying flower colors re-
ported in cultivated plants of E. scaber invite sus-
picion that colors are variable in the northern spe-
cies, too. Separating characters noted in Sandwith’s
key, such as pubescence and flower length, display
continuous or overlapping patterns.
Eccremocarpus huainaccapac is closely related to
E. viridis and shares many characters with it, but
it is distinguished by its dense, glaucous-drying
overall pubescence and its slightly larger leaflets.
A much weaker case might also be made for rec-
ognizing Eccremocarpus vargasianus, but the evi-
dence seen does not commend this view.
GEOGRAPHY
Eccremocarpus is found in the Andes of South
America. Northern collections from Colombia, Ec-
uador, and Peru (E. viridis, E. huainaccapac) occur
mainly between elevations of 3200 and 3700 m,
and southern collections from Chile and southern
Argentina (E. scaber) occur mainly from 1000 to
1 m. This distribution is shown in the map of
Figure 1. To judge by collections seen, populations
are widely scattered and morphologically variable.
Similar patterns occur in many other wind-dis-
persed groups in Andean uplands, for example,
Mnioides (Asteraceae), Niphogeton (Apiaceae),
uya (Bromeliaceae), and Polylepis (Rosaceae).
س
Volume 84, Number 1
7
D’Arcy 105
Review of Есстетосагриз
SOUTH AMERICA
АЕ ЕЕЕ — и
Figure Map of western South кийа os dis-
tribution of Eccremocarpus species. Diam = Eccre-
d Bae cremocarpus нар
Circles = Ecer cremocarpus ier iiri. The collection of E. sca-
ber indicated in Peru is dubiously from a wild plant.
“We may generalize that wide-ranging species are
the rule rather than the exception for much of the
neotropical flora, especially in wind-dispersed
groups” (Gentry, 1979: 342)
TAXONOMIC TREATMENT
Eccremocarpus Ruiz € Pavón, Prodr. Fl. Peruv.
Chil. 90. 1794. TYPE: Eccremocarpus viridis
Ruiz & Pavón.
Koi D. Don, Edinb. Phil. ДЕ, 7: 89. 1829. ТҮРЕ:
ampelis scaber (Ruiz avón) Sweet = Eccre-
чар scaber Ruiz ns бп.
Vines, subfrutescent, stems without anomalous
vasculature, ridged and sulcate on drying, without
pseudostipules or interpetiolar glands. Leaves op-
posite, bipinnatisect or tripinnatisect, a much
branched tendril terminal on the leaf. Inflorescences
racemose, generally reduced to a few flowers. Flow-
ers with the calyx showy, often red or pink, cam-
slightly curved or ventricose, sometimes contracted
apically and almost urceolate, the lobes small, gla-
brous to pilose outside; anthers slightly exserted or
situated just below the mouth of the tube, the an-
ther medifixed and the thecae parallel or fixed near
the apex and divergent; ovary ovoid or conical, uni-
locular, the ovules multiseriate on two parietal pla-
centas; disc annular-pulvinate. Fruit an ovoid to
ovoid-ellipsoidal capsule, the calyx persistent,
unilocular, with two narrow valves that remain con-
nected; seeds plane, suborbicular, the narrow mem-
branous wings circling the body of the seed.
Etymology. Greek ekkremes = pendulous, and
karpos = fruit
KEY TO SPECIES
1. Flowers less than 3 ст long; calyx less than "i
near the base; plants from Chile, rare or not in
3
Peru. Section — лон . E. scaber
„ ly
rl
2
38
JH
<
3
3
=
5
3c
i йай: fruit base pod in the ca-
eafy ан of stems pilose; pubescence most-
i rabia hairs to 4 mm long; leaves op-
posite; leaflets mostly sessile or held close to
the rachis, entire or gros ма lobed: with one prin-
cipal nerve from ih bas ase
nerves; plants from Peru, Ecuador, or Colombia.
Section Eccremocarpus.
106 Annals of the
Missouri Botanical Garden
2. Plants tomentose; stamens five -..-------------------- al Servicio de la Botánica en la Universidad del
2. Plants glabrate; ra
2. E iris Guiada E. lobbianus
E. longiflorus, E. vargasi iii)
Eccremocarpus sect. ERN MU
1. Eccremocarpus huainaccapac Vargas, Bol.
Soc. Peru. Bot. 1: 15. 1948. TYPE: Peru. Cuz-
co: Quesser-huailla, 3 m, largas
3034 (holotype, CUZ; isotypes, KS, 05“).
Vine, the stem with alternating ridges and canals,
pilose with copious erect weak, mostly eglandular
hairs of varying length to 2.5 mm. Leaves opposite,
generally 3—6 cm long, bipinnate with opposite pin-
nae, each with a terminal, ternately branched ten-
dril and usually three primary alternate pinnae,
leaflets subentire or lobed to pinnatisect and ap-
pearing 3-foliolate, mostly oblique and appearing
rhombic, the terminal leaflet largest, 8-12 X 3-5
mm wide, proximal leaflets often ovate or elliptical,
pilose with weak simple, light colored multicellular
hairs to 1 mm long, more so beneath, discolorous,
dark above; petiolules 1-2 mm long, pilose; petiole
2—6 cm long, glabrate or granular-pubescent. Inflo-
rescences leaf-opposed, racemose, several to many
flowered, to 14 cm long, pubescent with simple
gland-tipped hairs; bracteoles ca. 10 mm long,
ovoid, apically obtuse; pedicels 5-15 mm long, pi-
lose. Flower buds narrowly ovoid; calyx campanu-
late, 8-10 X 13-15 mm, 5-dentate, split 4-4 way
down, pilose with gland-tipped hairs; corolla ba-
sally campanulate, 40—45 mm long and 2-3 mm
wide within the calyx, above cylindrical, pilose
overall outside, minutely pilose basally within, 7—
mm wide at the mouth, the lobes green, obtuse,
ca. 4 mm long, glabrate; stamens 5, unequal, the
filaments inserted ca. 7 mm from the base, free,
sparingly pubescent at point of insertion, the an-
thers unequal, 5-10 mm long (in the same flower),
dorsifixed near or above the middle, thecae sepa-
rate but parallel below the insertion point; style ex-
serted ca. 4 mm, the lower stigma lobe reflexed,
ovary sessile, ovoid, with two pronounced longitu-
dinal ridges, 4 mm long, the surface granular; disc
annular-pulvinate with a'few minute distal tri-
chomes, ca 1.5 mm tall, ca. 6 mm wide. Fruit not
known. Figure: Vargas (1948: 15, 16).
In the one flower examined (Nufiez & Galiano
13414), the shorter stamens may lack pollen. The
stamens were unequal in size, and the corolla was
exserted from the calyx by only 1.5 ст, suggesting
that the flower was not yet fully expanded into an-
thesis.
Vargas (1946: 47) listed the species in Diez Afios
Cuzco, but did not provide a description, and the
name was not validly published until 1948. In the
1946 publication, Vargas gave the type locality as
Qquesser-huailla, Ckoricocha, but the specimen at
US reads Igneser-Huailla.
Eccremocarpus huainaccapac is similar in form
to E. viridis, but its overall pubescence is strikingly
distinct. The name of this species honors the last
Inca king, Huainacapacc (sic). Vargas (1948) sup-
plied the common name “Chucchucha.”
Distribution. This species is known from Peru,
in the vicinity of Cuzco and Apurimac between
3100 and 4100 m elevation. It has been collected
in flower in December and August, and the fruit is
unknown.
All known specimens (E. ر U. Cuz-
co uzco, 24 Dec. 1942
(A), Vargas 3034 (K*, U
cha, Yanacocha, 13%15'S, 72716' Y 4150 m, (fl), Nuñez &
Galiano 13414 (МОС). Apurimac: Grau Province, Inter-
andean valley of Coyllurqui, 13°50'S, 72°25'W, 3165 m,
26 Aug. 1991 (sterile), Nuñez et al. 14135 (MO); Ay-
maraes, 160 km from Challhuanca towards Puquio, dist.
Cotarosi, 4050 m, 7 Jan. 1962, Saunders 776 (К°).
2. Eccremocarpus viridis Ruiz & Pavón, Syst.
Veg. Fl. Peruv. Chil. 157. 1798. TYPE: Peru.
Near Muña, Ruiz s.n. (holotype, MA-2* = pho-
to, F-29234, F-fragment).
аа ле longiflorus Humboldt & Bonpland, Pl. Ae-
n. 1: 229. 1808. TYPE: Ecuador. Loja: Humboldt
& B onpland s.n. (holotype, Р = photo, F-39401).
iie pr obbianus Zahlbr, Ann. Nat. Hofmu
a, Lo a 1897. TYPE: Peru Boned. Lobb
sc tle WS fide Sandwith, 1965: bb
E Шер КУ). PARATYPE: Maclean s.n. A pe,
Cuatrecasas, Trab. Mus. Nac.
type, F', isotypes, E.
Eccremocarpus vargasianus Sandwith, Kew Bull. 19: 406.
ased on E. viridis sensu Vargas, Bol. Soc.
Peru. 1: 15. 1948, non Ruiz & Pavón.
Cosco: Vargas 5956 (CUZ?, K, isotypes MO-2*).
Vine climbing and twining, 2-4 m long, semi-
woody, the stem with alternating ridges and canals,
glabrate with sparse inconspicuous ascending
eglandular hairs, erect gland-tipped hairs and re-
duced hairs present on emerging growth, sometimes
persistent at the nodes. Leaves opposite (sometimes
one reduced or suppressed), generally 3—6 cm long,
bipinnate with opposite pinnae, each with a ter-
minal, mostly 3 times branched, wiry tendril, usu-
y with four primary alternate pinnae, leaflets |
ovate or elliptical to cordate or rhomboid, 3-7(-9)
mm long, 1.5-25 mm wide, glabrate to short- glan-
Volume 84, Number 1
1997
D’Arcy
Review of Eccremocarpus
dular-pilose; petiolules 2-5 mm long, often pubes-
cent; petiole 2-6 cm long, or glabrate or granular-
pubescent. Inflorescences opposite the leaves,
racemose, several flowered, to 16 cm long, pubes-
cent with short, simple, erect, gland-tipped hairs;
bracteoles 4–10 mm long, deltoid to linear, apically
acute; pedicels 5-15 mm long, the flowers twisting
to one side. Flowers with the calyx red or pink,
campanulate, 8-10 х 13-30 mm, 5-lobed, split %—
way down, glabrous to pilose with simple or
gland-tipped hairs, sometimes nervate, the lobes
lanceolate and short-acuminate to ovate and round-
ed; corolla mostly yellow, green at the tip, the tube
cylindrical, broader at the base, slightly curved, 3–
8(-12) X 1-3 cm, slightly contracted at the mouth,
ca. 4 mm wide at the mouth, glabrous or pubescent
with short, erect, sometimes glandular hairs, the
lobes rounded, porrect or recurved, glabrate, mostly
with dense minute peltate trichomes, appearing
whitish in bud; stamens four, staminode wanting,
equal, inserted about 20 mm from the base of the
tube, the filaments 50-55 mm long, anthers dorsi-
fixed, inserted on the filament about halfway up,
the thecae parallel, 7-10 mm long, slightly glan-
dular pubescent; style ca. 8 cm long, the stigma
situated at the corolla mouth, surrounded by the
united anthers, ovary narrowly ovoid, sessile, mi-
nutely glandular papillose, 8-10 mm long, the disc
annular-pulvinate, glabrous. Fruit ovoid or ellip-
soid, 20-45 X mm, with 4 dark grooves,
dehiscing into two papery valves; the base enclosed
in the calyx; fruiting calyx persistent, slightly
spreading; seeds suborbicular, the body narrow,
surrounded by a wing. Figures: Figure 2; Humboldt
& Bonpland (1805-1817, vol. 1, fig. 65); Gentry
(1977: 71 (as E. longiflorus)).
The characters used by Macbride (1961) and
Sandwith (1965) to separate elements of this spe-
cies into distinct taxa, corolla length and pubes-
cence, are not diagnostic. There is great variation
in dimensions and general appearance of flowers of
this species, but the variation in most characters
appears to be continuous, leading to the conclusion
that a series of widely separated populations rep-
resent conspicuous variants of a single polymorphic
species. Perhaps most conspicuous is corolla size
and width in relation to calyx length. In some spec-
imens (Prieto P-151, Davies 220), the corolla (10
cm long) greatly exceeds the calyx and 18 narrowly
tubular throughout. Such a flower was illustrated as
the type of Eccremocarpus longiflorus. In other
cases, such as the flower illustrated as the type of
E. viridis, the corolla is relatively broader, hardly
or not exserted from the calyx. On one collection
(Davis 220), two flowers have long (9-11 cm) co-
rollas and two have short (6 cm), barely exserted
ones. Exsertion of the stamens is also variable;
some collections (McPherson 13159) have the sta-
mens included in the corolla mouth and in others
(Vargas 5956) they are clearly exserted. Stamens
are subequal in length. In Vargas 5956, the type of
E. vargasianus, the longer pair of stamens is only
about 1 mm higher than the shorter ones. In this
specimen, the corolla is narrowest below the mid-
dle. In other collections, the corolla is narrowest
above, near, or below the middle, and in some col-
lections, it is evenly cylindrical, either straight or
curved, for its entire length.
A difficulty in interpreting the significance of
flower features in herbarium collections is the like-
lihood that collections were made at different stages
of flower opening, as noted under E. huainaccapac
above and by Sandwith (1965: 147). The material
underlying Е. lobbianus and the original illustration
of E. viridis may represent young flowers and not
the dimensions and shapes of flowers when they are
fully unfolded. The type material of Eccremocarpus
viridis at Madrid and Chicago lacks flowers (Sand-
with, 1965) and an indication as to where it was
collected. The species concept of Sandwith was
therefore shaped largely by the two other collec-
tions he cited, which are thought to come from
Мића in the Huanaco Department of Peru. Thus, if
Eccremocarpus viridis were to be divided into two
or more species, it is not clear what the separated
elements should be called.
Distribution. This species is found in Colom-
bia, Ecuador, and Peru mainly between 2700 and
600 m elevation. Most Colombian collections are
from the Cordillera Central. Collections from the
northern part of the range, Colombia and Ecuador
and northern Peru, are much more uniform in char-
acter than those from parts of Peru from where most
synonyms were described.
This species appears to flower and fruit through-
out the year.
Representative specimens (Eccremocarpus viridis). CO-
LOMBIA. Caldas: Nevado Del Ruiz, Carr. Term
09); Tolima Las Mesetas
13 May 1932, Cuatrecasas 2743 (COLS, eu DOR.
Napo: M 1977, de Vries s.n. (AAU*). PERU. Cuz-
Urubamba, Penas a Kosniriti, 3600 m, 23 Mar. 1946.
Vargas ET (K*e, MO“).
Eccremocarpus sect. Calampelis (D. Don) A.
DC., Prodr. 9: 238. 1845. Calampelis D. Don,
Edinb. Phil. Jour. 7: 89. 1829. TYPE: Cal-
ampelis scaber (Ruiz & Pavón) Sweet = Ессте-
mocarpus scaber Ruiz & Pavón.
Annals of the
Missouri Botanical Garden
108
e 2. Var HR e ag viridis Ruiz & Pavón. Inflorescence and leaves. After Vargas 5956 (type collection of E.
Figure
vargasianus Sandw.
3. Eccremocarpus scaber Ruiz & Pavón, Syst.
Veg. 157. 1798. Calampelis scaber es T Pa-
vón) Sweet, Brit. Fl. Gard. ser. 130;
1831. [As “scabra.”] TYPE: Chile. M Col-
chagua, Rancagua, and San Jacob, Ruiz & Pa-
vón 1798, (holotype, MA, photo, 029233, F).
Eccremocarpus sepium Bert. : *Merc. Chil. 1829" cf.
Bull. Ferussac, 20. 111. . Ec cabanas se scaber
var. [b] saepium (Bert.) se DC., Prod., 9. 1845.
ТҮРЕ: oe Sepibus secus, vias р аз унаа
5. Yag uillota, Bertero 965 (BM*, Е, ЕР,
MICH*, МО’, Ре). [Accepted as a synonym by Muñoz
966).
Eccremocarpus scaber var. aurea [sic] Benary, Gartenflora
РВК?
22: 608. 1903. TYPE: cult. Нон. Schónbrunn (?W,
not seen).
Eccremocarpus scaber var. carmineus [sic] Spigolatore,
ull. R. Soc. Toscana Orticultura 29(2): — ч 22.
1904. TYPE: cultivated in ?France, not loc
Eccremocarpus scaber carmineus Pynaert, Rev. Hort. Belge
31: 55. 1905. TYPE: cult. Belgium, not locat
ccremocarpus scaber var. roseus Huxley et al., New Roy
Hort. Soc. Dict. Gard. 2: 122. 1992. TYPE: Not lo-
cited:
Vine, climbing and twining, 2-6 m long, basally
Fide the stem drying with alternating ridges and
s, puberulent with short, erect, often gland-
сто hairs and occasional weak hairs to 2 mm
Volume 84, Number 1
1997
D’Arcy 109
Review of Eccremocarpus
long. Leaves opposite (sometimes one reduced or
suppressed), generally 2-6 cm long, bipinnate with
opposite pinnae, each with a terminal, mostly ter-
nately-branched wiry tendril, usually with four pri-
mary mostly opposite pinnae, these with 3-5 leaf-
lets, leaflets ovate to rhomboid, often oblique, 8—
30 X 7-15 mm, basally cordate or truncate, api-
cally obtuse, margins entire or dentate, shiny-gray-
ish to dark green above, glabrate to pilose with
short glandular hairs, mostly appearing as palmate-
ly 2-5-nervate; petiolules 2-5 mm long, often pu-
bescent; petiole 2-6 cm long. Inflorescences oppo-
site the leaves, sometimes appearing terminal,
racemose, several to many flowered, to 25 cm long,
pubescent with simple, mostly gland-tipped hairs;
bracteoles 4—10 mm long, deltoid to linear and api-
cally attenuate; pedicels 5-30 mm long. Flowers
with the calyx red or orange (green), campanulate,
ca. 8 X 4-8 mm, 5-dentate, unequally split 15—
way down, puberulent with gland-tipped hairs;
sometimes conspicuously nerved, corolla red or or-
ange, sometimes yellowish near the tip, the tube
subcylindrical, 20-25 X 2-3 mm wide at the base,
expanded about % way up on one side to 5 mm
across, contracted at the mouth, the lobes rounded,
mostly recurved, glabrate, ca. 4 mm wide at the
mouth; stamens four, unequal or (Mufioz) equal, 8
mm long, the filaments inserted about halfway up
the corolla, then free or (Mufioz) united to about
the middle of the thecae; anthers 2.4 mm long, the
anther thecae basally divaricate or (Mufioz) parallel
and united, the lobes of the stigma slightly expand-
ed when closed (dried), slightly shorter than and
surrounded by the anthers; ovary sessile, ovoid to
conical, smooth except for two longitudinal ridges;
disc annular-pulvinate, ca. 1 X 2.5 mm across, red
(Muñoz). Fruit short-stipitate, ellipsoidal, 30-40 X
15-20 mm, glandular, with 4 dark grooves, dehisc-
ing into two papery valves; fruiting calyx persistent
and slightly curved away from the fruit base; fruit-
ing peduncles 20-30 mm long; seeds 3-3.5 mm
across, suborbicular, the body dark brown, ovoid,
surrounded by a shiny, hyaline wing. Figures: Ed-
ward's Bot. Reg. (1825: t. 939*); Sweet, Brit. Fl.
Gard. (1831: t. 30); Maund (1831*: 289*); Louden
(1844: 1263*); Schumann (1894: fig. 93a, b); Ben-
ary (1903: 609); Spigolatore (1904: 340); Pynaert
& Pynaert (1905: 55); Muñoz (1966*); Нау &
Synge (1969: 247*); Hoffman J. (1978: 132*); Graf
(1986: 184*); Tsukamoto (1988 vol. 3: 339%); Graf
(1992: 270*); Huxley (1992: 1: 352); Belmonte et
al. (1994). (* colored illustration.)
Two other names, Tourretia scabra Dombey and
Dombeya nodiflora L'Her., were noted in the syn-
onymy of this species by De Candolle (1845), but
they were not validly published.
Distribution. Low cordillera in Chile from
Aconcagua to Valdivia, mainly between 1000 and
1800 m, but ranging from 300 to 3000 m elevation.
The species is also found in Chubut and Rio Negro,
Argentina. Gentry's database recorded two collec-
tions, Chavez 3467 and Shepard 9, from Cuzco and
Puno Departments in Peru. The Chavez collection
was labeled as a cultivated plant, and because
Puno is so distant from the range of the species in
central and southern Chile, the Shepard collection
is also assumed to have been cultivated. Eccremo-
carpus scaber is occasionally cultivated outdoors in
Europe as a curious ornamental. The species has
been collected in flower in South America mainly
from October to March, but plants in southern Eu-
ropean gardens are said to bloom continuously (Spi-
golatore, 1904).
Plants of this species in European cultivation
have displayed considerable variation of flower col-
or, e.g., calyx green, corolla rose, dark red, scarlet
to deep orange-red, orange, or golden (Loudon,
1844; Benary, 1903; Pynaert € Pynaert, 1905), al-
though this variation has not been noted by Chilean
writers (Muñoz, Hoffman, Navas). In addition to the
names noted in synonymy under this species, Hux-
ley (1992: 122) noted Eccremocarpus Anglia Hy-
brids, with flowers yellow, orange, pink, scarlet, and
crimson, which are here considered to be forms of
E. scaber.
Pollination of Eccremocarpus scaber is by hum-
mingbirds as evidenced by notes on collections (El-
liott 247) and the common name Chupa-chupa
(Behn s.n., Elliott 247, Kausel 1673), a Spanish
word for suck, as hummingbirds seem to do when
their bills enter the flowers. A study of nectar and
nectaries in Eccremocarpus scaber by Belmonte et
al. (1994) noted pollination by Giant Humming-
birds (Patagonia gigas gigas
The climbing ability of this pales by means of
tendrils with tactile responses has long been sub-
ject of study (Darwin, 1891; Junker € Reinhold,
1975; Tronchet, 1977; Junker, 1977).
Representative specimens (Eccremocarpus sca-
ber). ARGENTINA. Chubut: entre El Bolsón y Lago
Puelo, 42°05’S, 71?38'W?, 13 Nov., "m Peg (КУ).
CHILE. Colchagua: Cuming 21 (ВМ). Curico: Curico,
Los won (Andes de Curico), 35°01’S, 70°48’ W, 1000
m, 20 Jan. 1942, Aravena 33365 (МО“). Santiago: ТИП,
33°55 s. 2057 W, 840 m, 9 bus Pod Behn 21492 (KS).
ivia: Buchtien s.n. (LS aiso: Quebrada nn
33S, 71°W, 1 Mar. 1952, Mira 6474. (F, MO“, MO“).
PERU. Puno: Lake Titicaca, 15*48'S, 69°24’ W, 3125 m,
26 Nov. 1919, Shepard 9 (NY“).
110
Annals of the
Missouri Botanical Garden
REJECTED NAMES
Eccremocarpus ruber Regel, Cat. Pl. Hort. Aksakov.
1860. Nomen nudum.
Literature Cited
Belmonte, E., L. Cardemil & M. T. K. Arroyo. 1994. Flo-
ral nectary structure and nectar composition in Eccre-
mocarpus scaber (Bignoniaceae), a hummingbird-polli-
nated plant of central Chile. Amer. J. Bot. 81(Apr.
Benary, B. 1903. Neuheiten neuer Samen und Pflanzen
fiir 1903/04. со 22:
Вгако, L. & J. L. Zarucchi. 1993. Сиви of the Flow-
ering Plants and Gymnosperms of Peru/Catálogo de las
el Perú. Monogr. Syst.
Darwin, C. 1891. The Movements and Habits of Climbing
Plants. John Murray, London.
De Candolle, A. 1845. Bignoniaceae. /n: Prodromus Sys-
tematis Naturalis 9: 142-248 [Eccremocarpus pp. 238-
239].
Edwards, S. T. 1825. Bot. Reg. 11: t. 939. [Eccremocarpus
scaber.
Gandhi, K. N. & R. D. Thomas. 1983. Placentation in
йа асем а аз Жы акы р Catalpa, Millingtonia,
remocarpus plant structure. Phytologia
Gentry, А. Н. 1977. Fam. 178. Bignoniaceae. In: G. H
ling & B. Sparre, Flora of Ecuador. Opera Bot. 7 [Ec-
cremocarpus pp. 70—72].
1979. Distribution patterns of weg Big-
m-Nielsen, "Tropical
ra de Co lombia.
cations of вом palynology. Ann. Missouri Bot
d. 66:
Graf, A. B. 1986. Тра. Roehrs, East Rutherford, New
Jersey.
———. 1992. Hortica. Roehrs, East Rutherford, New
Jersey.
Hay, R. & P. M. Synge. 1969. The Dictionary of Garden
Plants in Colour. Ebury Press, London.
Hoffman J., A. 1978. Flora Silvestre de Chile: Zona Cen-
ud. "
npland. 1805-1817. Plantae Ae-
; 1 in a Aide e aux régions équinoctiales
du Nouveau dian fait en 1977-1804, Partie 6,
Botanique.
TM A. 1992. Bignoniaceae 1: 350—352; Eccremocar-
: 122. In: The New Royal Horticultural Society
Dicti tionary of Gardening. Macmillan, London
Junker, S. 1977. Ultrastructure of tactile papillae on E.
rils of иы scaber R. et Р. New Phytol. 7
610
€ 1. Reinhold. 1975. A scanning electron mi-
croscopic no of the surface of нн) tendrils [Ec-
cremocarpus scaber, Bryonia dioica}. J. Microscopie
Biol. Cell. 23: MTS 180.
Loudon, J. C. 1844. Of the half- hardy ligneous plants of
the order Bignoniscsále P. 1263 in Arboretum et Fru-
ticetum Britannicum. Longman et al., London.
Macbride, J. F. 1961. Bignoniaceae. Field Mus. Bot.
13(5c-1): 3-105.
Maund, B. 1831. The Botanic Garden 4 [VII] no. 289.
Melchior, H. 1964. A. Engler's Syllabus der Pflanzenfam-
Шеп. Gebriider Borntraeger, Berlin
uñoz, C. D. 1966. Flores Silvestres de Chile. Ed. Univ.
Chile Стара PP: 135-136].
Pynaert, C. & L. Pyna 1905. Les Eccremocarpus sca-
r. Rev. eden bes aaa 31:
Ruiz Н. & J. Раубп. 1794. Eccremocarpus. Prod. Fl. Pe-
ruv. Chil. 90.
. 1798. Syst. Veg. Fl. Peruv. Chil.
Sandwith, N. Y. 1965. Contributions to the flora of trop-
ical America: LXXI: Notes on Bignoniaceae: xxvii: А
synopsis of Eccremocarpus. Kew Bull. 19: 144-151.
Ter K. Bignoniaceae. /n: А. Engler & K.
antl. Die natürlichen eye 4(3b): 189-
e [Eccremocarpus P 244.
Spigolatore, L. 1904. I C erid 210, Soc. Tose. Отис.
Oct 1904: 338-341.
Sweet, R. 1831. Brit. Fl. Gard. ser. 2, 1: t. 30.
ae A. 1977. La Sensibilité des Paks Masson,
Pari
Die рана Y. 1988. Grand Dictionary of Horticulture,
vol. 1: 339. Ha ll Tokyo.
Vargas, C. y Bignoniaceae. P. 47 in Diez
Años al Servicio de н {инен en la Universidad del
Cuzco. Univ. Cuzco, Peru.
1948. El Género Eccremocarpus. Bol. Soc. Peru
Bot. 1: 14–16.
LIST OF COLLECTIONS
Specimens are listed alphabetically by principal соПес-
tor and number, det b denti-
fications are indicated by numbers: 1, Eccremocarpus
uainaccapac; $ E. viridis; 3, E. scaber. Differing
identifications made by previous н follow in ys
theses: the species epithet and the workers’ initials: G: A
H. Gentry, S: N. 5. Sandwith.
Without collector, number
number (MO')
Y) 3
F) 3. Without collector,
3 EUROPE. Without collector, numbe
Anderson s.n. (MO') 3 EUROPE. England. Aravena
33316 (мог) 3 CHILE. Curico. Aravena 33365 (МО“)
3 CHILE. Curico. Ball s.n. (МҮ) 3 CHILE. Santiago.
Behn s.n. (F) 3
Рс) 3 CHILE. Buchtien s.n. (15) З CHILE
Calvert s.n. (BM?) 3 CHILE. Valparaiso. Canby s.n.
(NY) З CHILE. Chavez 3467 (MO*) З PERU. Cuzco.
Cleef & Hart 2500 (МОГ, U^) 2 (longiflorus-G) COLOM-
BIA. Caldas. Comber 1011 (K') 3 CHILE. Cook & Gilbert
1237 (US?) 2 (longiflorus- perm PERU. Cuzco. Con-
treras? & Veitch 218 (KS) 2 PERU. Crovetto 3276 (Е) З
ARGENTINA. Ch 01е, K**) 2
ee aa S) COLOMBIA. Tolima.
(COLS, F-3') 2 (longiflorus-G) COLOMBIA. Caldas. Cua-
irécdsaó 9351B (COL?)
Caldas. Cuatrecasas 20400 (F-
Cum. ез 21 (ВМ) 3 CHILE.
(К) 3 CHILE. Valparaiso.
Davis (Davies) s.n. (MO“,BMS*) 2 PERU. Huánuco.
Davis comm. Veitch 220 (КУ) 2 PERU. Huánuco. Dawe
860 (KS, NY“) 2 (longiflorus-G,S) COLOMBIA. De Barba
487 (F) 3 ARGENTINA. Chubut. Devia & Prado 1880
3) 2 COLOMBIA. Valle.
Colchagua. Cuming 615
ЭЕ ааа E Wai
Volume 84, Number 1
1997
D’Arcy 111
Review of Eccremocarpus
(MOS ex Tul 2 (longiflorus-G) COLOMBIA. Valle. de
U*) 2 (longiflorus-G) ECUADOR. Napo.
. Santi
Ellenberg 4867 (МОС) 2 (v Ят АЙЕЛ" ') PERU. Elliott
419 "ge p CHILE. Elliott 247 (K') 3 CHILE. Elwes s.n.
(K) 3
Fournier s.n. 1. (Рх) 2 (longi G,S) ECUADOR. Pi-
chincha. Frbr. Bert. Jime. ? s.n. (МОС) З CHILE. Freire
et al. 656 (ОСА) 2 (longiflorus-G) ECUADOR. Chim-
razo. Frödin s.n. (ЖҮ HILE. Santiago. Fródin
635 (BM*) 3 CHILE. Aconcagua.
Gay s.n. (KS) 2 (longiflorus-S) ECUADOR. Gay s.n. (Ре)
3 CHILE. Santiago. Gay s.n. (Р) З CHILE. Valdivia.
Gay 20 (P*) З CHILE. Santiago. Goodspeed 16853 (МО!)
3 CHILE. Aconcagua. Goodspeed 23344 (K') 3 CHILE.
Aconcagua.
Halpin s.n. (CLEMS®, КУ) 3 CHILE. Hartweg 148 (К)
2 (longiflorus-G,S) ECUADOR. Loja. Harvey s.n. (KS) 3
HILE. Hastings 171 d 3 CHILE. Hirsch P1022 (К)
2 (vargasianus-S) PERU. Urubamba. Hort. Vilmourin s.n.
me 1857 (МО) 3 EUROPE. = Humboldt s.n. (KS,
Рао) 2 (longiflorus-G,S) PER oja.
Jamaa sn (K 2 артат ECUADOR. ^
chincha. Jameson 56 (NY*) 2 (longiflorus-G) ECU
DOR. Pichincha. Jameson 186 (K**) 2 longiora)
ECUADOR. Pichincha. Jameson 286 (BMS ur "Б, NYS,
Ре) 2 (longiflorus-G,S) ECUADOR. Pichine
Karsten s.n. (WS) 2 (longiflorus-S) COLOMBIA. Pichin-
cha. Karsten s.n. (WS) 2 tora ECUADOR. Cun-
dinamarca. Kausel 1673 (F') 3 CHILE. Santiago. King
589 (BM*) 3 с King 712 (BM*) 3 CHILE. Kuntze
s.n. ا 3€
5 eer S.A. California. Lehmann 3149
2 (longiflras-G. S) COLOMBIA. Caldas. Lobb
Fen 2 ps Porco RU. panes et al. 48
ex-MEDEL) 2
2 (lobbianus-S) PERU. Macrare? s.n. (K') 3 тј
thews 3176 (BM*, К!) 2 (longiflorus-G,S) Р
nm :
Amazo
Pherson 131 59 (MO) 2 (вовсе С) COLOM ВІА. Ап-
i CHILE. Mexia 7887 (ВМ“,
. Curico. Meyen s.n. (Р) З
CHILE. Meyer 9422 (KS) 3 ARGENTINA. Chubut. Mid-
dleton s.n. (BM*) З CHILE. Molau & Ohman 1635 (GB*
= photocopy МО) 2 (vargasianus-G) PERU. Cuzco.
Montero 67a (MO*) З CHILE. Colchagua. Montero 260
(К) 3 CHILE. Santiago. Montero 507 (МО“) З CHILE.
Santiago. Morrison 16853 (МО) З CHILE. Aconcagua.
Nuñez & Galiano 13414 (MO) 1 PERU. Cuzco. Nuñez
& Luna 8841 (F', MO“) 2 (vargasianus-G!) PERU. Cuz-
co. Nuñez et al. 14135 (MO*) 1 PERU. Apurimac.
Ollgaard et al. 38199 (ААО) 2 (longiflorus-G) EC-
UADOR. Chimborazo. Úllgaard et al. 98155 (ААО?) 2
(longiflorus-G) ECUADOR. Carchi. Ortiz s.n. (AAU*,
ОСА) 2 (longiflorus-G) ECUADOR. Imbabura. Ortiz 30
(AAUS, NY!) 2 (longiflorus-G) ECUADOR.
Pearce 533 (K*) 2 PERU. Huánuco. Pearce 823 (КУ)
2 (vargasianus-S) PERU. Penland & Summers 1080 (F')
2 ECUADOR. Azuay. Pennell 12262 (Е, GH, NY“) З
ug O’ Higgins. Philippi s.n. (HB*) 3 CHILE. San-
ago. Poeppig 2 (BM*) З CHILE. Prance 26595 (050) 2
nme ibis С) ECUADOR. Napo. Prieto P-151 (NY“=
MO, photocopy) 2 (longiflorus-G,Wurdack) ECUADOR.
Canar. Purdie s.n Wing K-25) 2 (longiflorus-G,S) CO-
LOMBIA. Risa
ied & Hirsch Р. 1022 =? 2 о PERU.
a. Raddin s.n. (F') 3 ago. Reed
s.n. K): 3 CHILE. Maule. ed eL (MO) 3 NEW
ZEALAND. Ruiz & Pavón 5/14 (MA-3°, BM) 2 PERU.
са & Pavón s.n. (BMS, Е ex MA, ЕР) 3 CHILE. San-
a.
andere 288 (К) 3 CHILE. Saunders 776 (КУ) 1
PERU. Apurimac. Schlatzer s.n. (ААО) З CHILE. San-
tiago. Schmidt s.n. (HB*) 3 dp Shepard 9 (NY) З
PERU. Puno. Simpson s.n. (P*) З CHILE. Sodiro s.n. (Р5©)
2 (longiBorus-C; 5) COLOMBIA. на іпатагса.
e COLS) 2 (longiflorus- G,S) COLOMBIA.
LOM
ge 3034 (KS, USS) 1 PERU. Cuzco. п 5956
(К°, МОС) 2 (vargasianus-G,S) PERU. Cuz peas
a
У у
644 (МОГ) 2 p us-G) PERU. Gunde: Vieillard
F (РОУ 3 CHIL aiso.
Weberbauer (F?) : longiore. Macbride, S) PERU. Ca-
jamarca. Weberbauer 4938 (?) 2 yn pitt Macbride,S)
ECUADOR. Werdermann 482 (BM* MO“, US) 3
CHILE. Santiago. Wildenow s.n. on 3 CHILE.
Zóllner 6486 (L°) 3 CHILE. Aconcagua. Zóliner 9350
(МО) 3 CHILE. Valparaiso. Zöllner 11052 (МО) З
CHILE. Santiago.
INDEX
Calampelis 105, 107, 108
caber 105, 107
scabra 108
Dombeya
nodiflora 109
Eccremocarpus 105, 106
uainaccapac 106
lobbianus 106
mutisiana 106
scaber 105, 107, 108
scaber carmineus 108
scaber var. aurea 108
scaber var. слано 108
scaber var. 108
scaber var. [b]. f TE 108
sepium 108
vargasianus 106
viridis 105, 106
Tourretia
scab 109
HOT-SPOTS ANALYSIS FOR
CONSERVATION OF PLANT
BIODIVERSITY IN THE
MEDITERRANEAN BASIN!
Frédéric Médail? and Pierre Quézel?
ABSTRACT
e to the increase of human impact on the world scale, there is an urgent need to identify the sectors of the pne
biodiversity that are also the most endangered. Examination of the plant biodiversity of the five regions wit i-
terranean climate (SW Australia, the Cape region of South Africa, California,
in) cle arly demonstrates their key role in the world context. The delimitation and definition of 10 red alert
the Mediterranean basin and in Macaronesia are explained in detail. The 10 sectors
or “hot-spots” situated in
mediterranean Chile, and the Mediter-
identified are: the Canary Islands and Madeira, the High and Middle Atlas mountains, the Baetic-Rifan complex, the
aritime and Ligurian Alps, the Tyrrhenian Islands,
editerranean
berian Peninsula
Southern and Central Greece, Crete, mene? and Cyprus, the
Syria-Lebanon- Israel area and, lastly, the Cyrenaic Mediterranean. There are two main cen
asin in the west that pisc the I and Moroc
includes "Turkey and Gases. This analysis demonstrates the uniqueness and fragility of do island. чена
ev AME in the
in the East that
RÉsUMÉ
L'augmentation de l'impact anthropique à l'échelle du Globe nécessite d'identifier de facon urgente les secteurs de
plus haute biodiversité, qui présentent également les plus grandes menaces
. L'examen de la biodiversité végétale des
cinq régions soumises au bioclimat méditerranéen (S.W. de l'Australie, région du Cap en Afrique du sud, Californie,
Chili méditerranéen et bassin méditerranéen) souligne nettement leur róle
clé, par rapport au contexte mondial.
es Alpes maritimes et ligures, les iles tyrrhéniennes, le sud et le centre de la Grèce, la Crete, l'Anatolie et Chypre,
l'ensemble Syrie-Liban-Israel, et, enfin la Cyrénaique méditerranéenne. Deux
sur le pourtour méditerranéen: un
tal qui comprend la péninsule ibérique et le Maroc,
póles principaux de biodiversité existent
et un oriental englobant
ciden
la Turquie et la Grèce. Voriginalité des ensembles insulaires et sms fragilité se arate de cette analyse.
A few years ago, addressing increasing human
impact on a world scale, the scientific community
sounded an alarm, pointing out the harmful effects
that would result from a decrease in biodiversity
(e.g., Wilson, 1988; Lubchenco et al., ; Sol-
brig, 1992; Chauvet & Olivier, 1993; Levéque,
1994). This issue reached its pinnacle with the
Convention on Biological Diversity, signed during
the Rio Summit in June 1992, with the adoption of
Agenda 21.
The accelerated decrease in the surface area of
tropical forests (e.g., Myers, 1986; Forget, 1994)
monopolized attention because this biome is seen
as the world’s principal pole of biodiversity (Gentry,
1982, 1988). Alpha diversity thus reaches record
levels in Amazonia, where, in Ecuador—in a 1-ha?
plot—Valencia et al. (1994) listed no less than 473
different species of phanerophytes with stems at
least 5 cm in diameter, while Duivenvoorden (1994)
observed 310 higher plants in a plot of 0.1 ha in
olombia.
In this context, identification of areas of major
biodiversity is very desirable, considering that rates
of extinction are unprecedented (Stanley, 1987).
e current rate of disappearance of species is
1,000 to 10,000 times greater than that of major
geological periods of mass extinction (Wilson,
1988). Myers (1988, 1990) developed an analytical
methodology that enabled him to define red alert
areas of biodiversity (known as “hot-spot areas”).
A “hot-spot” is a sector with an exceptional con-
centration of species and a high rate of endemism
! We are grateful for helpful comments by D. ee P. Ponel, A. Strid, T. Tatoni, and R. жор and for
go Adan from S. Pignatti concerning biodiversity areas i
n Italy. We also thank M. Field for linguistic advic
iversité d'Aix-Marseille III, Faculté des Sciences de Marseille-St Jéróme, Institut Méditerranéen d'Ecologie et
de Paléoécologie, C.N.R.S, URA 1152, Laboratoire de botanique et d'écologie méditerranéenne, Case 461, F-13397
Marseille cedex 20, France
ANN. MISSOURI Bor. GARD. 84: 112-127. 1997.
AAA AO PUR em rig dapes c LEA Vg USING a EGER
Volume 84, Number 1
1997
Медан! 4 Quézel 113
Mediterranean Plant Biodiversity
that is in great danger of destruction. Species rich-
ness and endemism are two attributes of biodiver-
sity commonly used in Conservation Biology be-
cause they reflect the complexity and uniqueness
of ecosystems (Caldecott et al., 1996), and also be-
cause data for them is relatively easy to obtain on
a global level. By limiting himself to the plant com-
position, Myers (1988) first defined 10 hot-spots in
tropical forests, then four others in the same biome
and four in the mediterranean bioclimate (SW Aus-
tralia, the Cape region of South Africa, California,
and part of Chile: Myers, 1990). This author had
already emphasized the important role played by
the circum-Mediterranean area as a reservoir of
plant biodiversity. However, he cautiously refrained
from regarding the entire Mediterranean basin as a
hot-spot, arguing that it covers too large an area
and that there is insufficient knowledge concerning
certain parts of this area. Davis et al. (1994) re-
cently drew up a list of 250 centers of plant diver-
sity for the entire world, but this remarkable syn-
thesis does not give details of all the Mediterranean
hot-spots.
Therefore, more detailed analyses on the scale
of the Mediterranean basin are necessary in order
to try to establish a comparative study of the flo-
ristic richness and endemism of the various coun-
tries around the edge of the Mediterranean. This is
possible for the total number of taxa that are pres-
ent, i.e., 25,000 species (Quézel, 1985) or 30,000
species and subspecies (Greuter, 1991), but also for
the endemic species (around 12,500). Even if these
estimates are often imprecise, they can neverthe-
less provide precious information on both the lo-
cation of the main centers of endemism and the
situation of the areas of high biodiversity. They can
also give an idea of the level of threat posed to any
part of the basin, particularly after the work carried
out in this direction by the International Union for
Conservation of Nature (I.U.C.N., 1980, 1983).
SITES AND METHODS
LIMITS OF THE MEDITERRANEAN REGION
The question of the limits of the Mediterranean
region has been approached in various ways, in-
cluding from the point of view of floristic methods,
the analysis of vegetation structures, climatic inter-
pretations, and, lastly, bioclimatic criteria. Al-
though the validity of the Mediterranean isoclimatic
area (Daget, 1977) is often accepted, it seems more
realistic to use more traditional boundaries. There-
fore we adopted (Figs. 1, 2) the limits drawn in the
Natural Vegetation Map of the Countries of the
Council of Europe (Conseil de l'Europe, 1987) and
in the map of the vegetation of the eastern Medi-
terranean (Quézel & Barbero, 1985) for the coun-
tries of the northern Mediterranean, and the
100-mm isohyet, which remains a standard for de-
fining the southern boundary between the mediter-
ranean bioclimate and the Sahara, even if the
150-mm value would be locally more exact nowa-
days (Quézel & Barbero, 1993). The defined Med-
iterranean region thus covers an area of around
2,300,000 km? (Quézel, 1985). The archipelagos of
the Canaries and Madeira have also been taken into
account because, from a biogeographical viewpoint,
these islands are now considered as a superprov-
ince of the Canaries subregion, which is itself in-
corporated in the Mediterranean region (Rivas-Mar-
tinez et al., 3
ESTIMATION OF THE FLORISTIC RICHNESS AND
ENDEMISM OF THE MEDITERRANEAN REGION
One of the main difficulties in estimating Medi-
terranean biodiversity is that there is rarely any
direct correspondence between the biogeographical
boundaries and the political boundaries of a state,
while the floristic assessments are nearly always
prepared according to the latter criterion. However,
an attempt has been made by one of us (Quézel,
1985) and these results were later accepted by sev-
eral authors, including Greuter (1995) and Hey-
wood (1995). Precise assessments for the Turkish
Mediterranean region still have to be established
because its separation from the Irano-Touranian
area is still a matter of debate; these communities
share unquestionable biogeographical and biocli-
matic affinities.
The inconsistent floristic knowledge of the dif-
ferent countries of the Mediterranean perimeter is
also an obstacle. Some sectors—Italy, Turkey, An-
dalusia, Crete, and Corsica—are classified accord-
ing to modern floras, while others still do not have
complete floras (Morocco, Spain, the Balkans, Al-
bania). The taxonomic levels also vary greatly, ac-
cording to whether taxonomists adopt a restrictive
approach or take the taxonomic breakdown further
than is normally accepted. In addition to the rec-
ognition and fluctuating taxonomic status of some
taxa, there is the problem of whether or not to take
into account highly polymorphic groups, which are
often apomictic, such as Hieracium, Taraxacum,
Achillea, Rosa, Rubus, and Limonium. Thus, in Sar-
dinia, no less than 23 endemic species of Limonium
were recently described (Arrigoni, 1976-1991),
i.e., 22% of the total local endemic flora. Further-
more, some assessments made take into account
only the species, while others consider the species
Annals of the
114
Missouri Botanical Garden
вале ивопемопројј OY} JO sirurT
"urseq ивәиешәпрәу ur шеншориз juejd jo ваопортош y3ry щим 8101298 peoryde1309301g
`1 ona
%07 > 9161 usruapua > %01 SS 0T < ILI шѕтшәриә ae
115
Медан 4 Quézel
Volume 84, Number 1
1997
Mediterranean Plant Biodiversity
"воле ивопемонпрој 241
JO siu оу) SIUYIP ош хоп au 'отепола о) ивопвмопројј :01 19918] -попвао | ешАС :6 *snidé?) pue er[ojeuy :g 791917) :/ ‘222215 [&ju27) pue шәщцпос :9 "SPUB|SI пето мај, °C
"за ју ueunzr] pue әшциеу + 'xo[duioo uejry-oneseg :g ^surejunojy sepy 2[PPIN pue Чан :2 'вов ето пцоле ивлорвуиј pue ѕәшеив”) :ү “sjods-joy uiseq ивопемопроџ ‘g әл
ele
F d ^,
116
Annals of the
Missouri Botanical Garden
and subspecies, or even the varieties and doubtful
forms, without necessarily specifying this clearly.
Since it is impossible to ensure uniformity of the
results, we have indicated the sources used in the
published tables, on the understanding that, the es-
timation of the number of endemics and subendem-
ics (endemics common to several countries, sensu
Pignatti, 1982), was problematic or even impossible
in some cases.
The definition and delimitation of endemic taxa
are often subjective and quite difficult because au-
thors usually characterize endemic as a species re-
stricted to a politically defined territory without bio-
geographical consideration. According to Strid (pers.
comm.), biogeographers ought to develop a classifi-
cation of endemics reflecting real distribution area
and degree of ecological specialization. Mountain
endemism includes the endemic species and sub-
species regularly found above an altitude of ca. 1600
m, i.e., in the Mountain-Mediterranean, Oro-Medi-
terranean, and Alti-Mediterranean zones.
However, the data for the Mediterranean and the
Macaronesian islands are much more precise, fol-
lowing recent well-documented works (Shmida &
Werger, 1992; Gamisans & Jeanmonod, 1993; Tur-
land et al., 1993; Press & Short, 1994) and the
results of a recent conference (held in 1993) de-
voted to the flora of the islands of the Mediterra-
nean.
RESULTS
BIODIVERSITY OF THE MEDITERRANEAN
BIOME IN THE WORLD CONTEXT
According to the recent world overview study by
Davis et al. (1986) for the I.U.C.N., the five regions
with a mediterranean climate (the Mediterranean
basin, California, the South African Cape, SW Aus-
tralia, and mediterranean Chile) have remarkable
biodiversity. Table 1 shows the results reworked by
Myers (1990), particularly after Goldblatt’s (1978)
estimates for the Cape area or those of Raven and
Axelrod (1978) for mediterranean California. Flo-
ristic patterns in mediterranean-type ecosystems of
southwestern Australia are still unclear: Hobbs et
al. (1995) made an estimation of 8000 species in
the Southwest Botanical Province, with about 75%
endemism; we keep this number but we stress that
it is double Myers’s estimate.
Thus, the mediterranean biome, only 2% of the
world’s surface area, contains 20% of the total
world floristic range. The estimated total number of
mediterranean plants varies between 45,000 and
‚000 species (Heywood, 1995). This difference is
mainly due to the inclusion of either the South Af-
rican Cape (90,000 km?) or the whole of South Af-
rica (2,573,000 km?). From the biogeographical
viewpoint, it would be preferable to take the value
of 45,000 species. Thus, South Africa, SW Austra-
lia, California, and mediterranean Chile contain
8% of the plant biodiversity in 0.3% of the earth’s
surface area (Myers, 1990).
Despite its considerable area, the circum-Medi-
terranean world is a major contributor to the bio-
diversity of the mediterranean biome, since 10% of
the higher plants are found there in 1.6% of the
earth’s surface. It is interesting to compare this cir-
cum-Mediterranean biodiversity with that of other
regions, both temperate and tropical. Thus, al-
though it covers one-tenth of the area of the former
U.S.S.R., the Mediterranean basin contains 4,000
more species. Similar comparisons with the United
States, Europe, and China are clearly in favor of
the circum-Mediterranean region. Even certain
tropical or subtropical countries such as India or
Zaire have less biodiversity, while Brazil has twice
as many species but covers an area four times
greater. Note again that the whole of tropical Africa
contains only around 30,000 plant species (Good,
1974) in an area four times greater than the circum-
Mediterranean region.
Calculation of the number of species per 1000
km? provides an evaluation that is independent of
surface area. For the mediterranean biome, we ob-
tain values of between 95.5 species for 1000 km?
in the Cape region to 10.8 for the Mediterranean
basin (Table 1). This can be compared with: 1 spe-
cies/1000 km? in Europe, 3.1 in China, 4.7 in Zaire
and in India, 6.5 in Brazil, 40 in Colombia, and 90
in Panama.
The number of endemic species in the Mediter-
ranean region also reaches very high values, usu-
ally at least equal to those found in most tropical
areas of the world. Only islands such as Borneo,
Cuba, New Caledonia, Hispaniola, New Zealand
(Gentry, 1986), or high-altitude habitats are richer
in endemic species, but in relatively limited areas.
BIODIVERSITY OF THE WORLD’S MEDITERRANEAN
CLIMATE REGIONS
In relation to the other mediterranean climate
regions, the Mediterranean basin has the greatest
variety of species, both general and endemic, but
in a much greater surface area (84% of the total).
Despite the considerable differences in surface
area, the two mediterranean climate regions of the
northern hemisphere contain a virtually equal per-
centage of endemic species (Table 1). Furthermore,
mediterranean California has more or less the same
MEER t LL
сиса о оваа ова
Volume 84, Number 1
1997
Médail & Quézel 117
Mediterranean Plant Biodiversity
Table 1.
Hobbs et al., 1995; 4: Bond & Goldblatt, 1984).
Plant biodiversity of the world’s five mediterranean climate regions (1: Quézel, 1985; 2: Myers, 1990; 3:
Approximate Number of Endemic species
b
number species per Approximate
Areas (km?) of species 1000 km? number %
North hemisphere
Mediterranean basin (1) 2,300,000 25,000 10.8 12,500 50
California Floristic Province (2) 324,000 4,450 13.7 2,140 48
Austral hemisphere
Mediterranean Chile (2) 140,000 2,900 20.7 1,450 50
S.W. Australia (3) 112,260 8,000 71 6,000 75
Cape Floristic Region (South Africa) (4) 90,000 8,600 95.5 5,860 68
Total 2,966,260 48,950 16.5 27,950 57
(%/World) (2%) (20%)
surface area and general biodiversity as Morocco,
but California is four times richer in strictly local
endemics. However, the situation is reversed if, in
Morocco, we consider all circum-Mediterranean en-
demics in the broad sense. The case of mediterra-
nean regions in the southern hemisphere is even
more remarkable, since, due to the long geograph-
ical isolation of these areas, the endemic richness
reaches up to 70% in South Africa (the Cape re-
gion) and in SW Australia. The mediterranean re-
gion of the Cape, particularly the Cape Peninsula
е et al., wee d an extraordinarily rich
flora: around 8600 species (Goldblatt, 1978; Bond
& Cuba 1984), ме ing 5860 endemics, are
ound there, in a surface area equal to that of
Greece, which makes it one of the most remarkable
hot-spots in the world.
BIODIVERSITY OF THE MEDITERRANEAN BASIN AND
MACARONESIA
Despite the above-mentioned various obstacles
to the establishment of an accurate assessment of
circum-Mediterranean biodiversity, and with cer-
tain reservations regarding the accuracy of the val-
ues quoted, certain points can be stressed (Tables
2 and 3):
—The Mediterranean parts of Turkey and Spain are
found to have the richest variety (around 5000
species), followed by Greece, Italy, France, and
Morocco (with more than 3000 species).
—The percentage of endemics is highest in Turkey
(31%), then Morocco (21%), Spain (19%),
reece, Syria- on, and Italy. Greece is an
interesting case: endemism is indeed high, par-
ticularly in the mountains, with 27% (Strid,
993), and it would be even higher if all the
Aegean perimeter were taken as a single entity.
Similarly, evaluations that take into account the
Ibero-Moroccan complex would at least double
the number of endemics in these sectors. Fur-
thermore, assessment of endemics by country or
by biogeographical zone never or rarely includes
the endemics of the entire Mediterranean basin,
which clearly underestimates the indicated rate
of endemism.
—The specific nature of the islands included with-
in the Macaronesian complex, where, for histor-
ical reasons (Suning, 1979; Bramwell,
Quézel, 1995) and due to the limited amount of
competition, the percentage of endemism reach-
es very high levels in Madeira (26%) and in the
Canaries (38%), while there is a rather limited
specific richness. The same situation is found,
but to a lesser degree, in the islands of the Med-
iterranean Sea, where endemism is generally
around 10% but the overall range of flora is
greater, with between 1500 and 2500 taxa
o
eS
A
Therefore, in simplified terms, there are two ma-
jor centers of biodiversity in the Mediterranean ba-
sin: a western center, including the Iberian penin-
sula and Morocco, and an eastern pole that
encompasses Turkey and Greece. The islands of the
Mediterranean Sea and, to a greater degree, the
Macaronesian islands, have a very high rate of en-
demism
MAXIMAL BIODIVERSITY AREAS OF THE MEDITERRANEAN
BASIN AND MACARONESIA
The circum-Mediterranean region is—according
to the geographical zones of which it is composed—
considerably varied in both its number of species
and its number of endemics. Thus we should use
biogeographical concepts in order to define circum-
Mediterranean hot-spots, and political boundaries
should be ignored.
Initially, we attempted to determine the biogeo-
Annals
118 of the
чиен Botanical Garden
Table 2. Plant biodiversity of the countries from the Mediterranean basin (islands excluded). The figures indicate
the number of nonet except for the values marked with an asterisk, which also include subspecies. Unless otherwise
indicated, sources are: (a): Davis et al. (1986) and (b): ues i i Other sources: (c): original observations, (d):
sea: (1991), (e): кезеке & Ibn Tattou (1995), (0): Enriquez-Barroso & Gomez-Campo (1991), (g): Bartolo et al.
(1977), (h): Boulos (1995), (i): Davis et al. (1994), (j): Tam (1988), do: LU.C.N. (1980), (1): Olivier et al. (1995), (m):
Gomez-Campo et al. (1984).
Approximate number
ко т of species
a Number and approximate
Total areas ranean In Mediter- % ог ДА alani
(а) bioclimate ranean region a P
Countries (km?) (b) (km?) Total (a) (b) Number %
Могоссо 659,900 300,000 4,200 *3,800 *900 (e) 21.4
Algeria 2,381,000 300,000 3,150 2,700 *320 (d) 10.1
Tunisia 164, 100,000 1,800 1,600 #39 (f) 2.1
Libya 1,759,000 100,000 1,600 1,400 *140 (g) 8.7
Egypt 1,000, 15,000 2,060 1,100 61 (h) 3
Israel 20,700 10,000 2,200 2,000 *165 (i) T
Jordan 97,600 10,000 2,200 1,800 (c) 145 (i) 6.6
yria 185,000 50,000 3,100 2,600 (c) *395 (1) 12.7
Lebanon 10,400 10,000 2,600 ,600 311 (i) 12
Turkey 779,000 480,000 8,600 5,000 2,651 (j) 30.8
Continental Greece 107,000 90,000 5,700 *4.000 *742 (i) 13
Albania 28,700 20,000 3,000 2,200 46 (k 1.5
Former Yugoslavia 255,000 40,000 5,000 2,500 *320 (i) 6.4
Continental Italy 251,400 200,000 4,870 (c) 3,850 (c) 570 (c) 11.7
Continental France 549,600 50,000 ,800 3,200 180 (1) 3.7
Continental Spain 504,000 400,000 6,720 (d) 5,000 1,286 (d) 19.1
Portugal 91,000 70,000 2,600 (d) 2,500 114 (m) 4.4
Table 3. Plant biodiversity of the Mediterranean and Macaronesian islands. The figures indicate the numbers of
species and subspecies, except = the values marked with an asterisk, which also include varieties. Rare and threatened
a correspond to Ex, E, V, R, 1, and K categories of LU.C.N. Sources: (a): Davis et al. (1986), (b): Gamisans $
Jeanmonod (1993), (c): Bočchieri (1995), (d): Raimondo et al. (1994), (e): Lanfranco (1995), (f): Turland et al. (1993),
(g): Alziar (1995), (h): Schmida & Werger (1992), (i): Dalgaard (1994), (5): LU.C.N. (1980), (k): Original chee
(1): Médail & Verlaque (1997), (m): Polunin (1987), (n): Mus (1995, modified), (о): Verlaque pers. comm., (p): Leon
al. (1985), (а): LU.C.N. (1983).
threatened
rof Endemism ad
indigenous in total Endemism 7 POTERNE е
Атеа species and flora in total species Tare
(km?) subspecies s. str. flora s.l. s. str. s.l. dudit
Balearic Islands 5,014 1,450 (a) 94 (j) 180 (o) 6.5 12.4 59 (n)
Corsica 8,748 2,354 (b) *130(b 270(Б) 5.5 11.5 90 (0)
Sardinia 24,090 2,054 (c) 106 (k) 200 (k) 5:2 97 39 (р)
Sicily 25,708 2,700 (d) 260 (о) 310 (о) 9.6 11.5 654 (d)
Malta 316 700 (e) 16 (e) 32 (k) 2.3 4.6 12 (p)
Crete (incl. Karpathos) 8,700 1,706 (f) 171(f 200(m) 10 11.7 119 (q)
Cyprus 9,250 1,620 (g) 130(g 170 (2) 8 10.5 69 (р)
Canaries 7,273 1,582 (h (h) (k) 31.8 37.9 432 (k)
Madeira Archipelago 796 670 (i) 113 (i) 175 (i) 16.9 26.1 137 (q)
Volume 84, Number 1
1997
Médail & Quézel 119
Mediterranean Plant Biodiversity
graphical sectors where the rate of endemism is
more than 20%; this figure is generally considered
by biogeographers as delimiting a high level of en-
demism (Greuter, 1991; Quézel, 1995). Such рег-
centages are not exceptional, particularly in the is-
land and mountain areas, as demonstrated by
Gomez-Campo et al. (1984) for the Iberian penin-
sula, where values of more than 50% are reached
in the Baetican cordillera. All these zones were
mentioned in Figure 1, where other areas can be
distinguished, such as the Canary Islands and Ma-
deira, the Middle and High Atlas ranges and the
Rif in Morocco, the Sierra de Estrela in Portugal,
as well as the upper mountain areas of Corsica,
Sicily (Madonia and Etna), and probably Sardinia
(Gennargentu mountain) in the central Mediterra-
nean. Further to the east, endemism is very high in
the upper mountain areas of southern Greece and
of Crete, the Taurus ranges of Anatolia, the Troodos
mountain of Cyprus, and the summits of the Leb-
anon and the Anti-Lebanon.
Other areas with a rate of endemism of around
10% deserve mention (Fig. 1): Sierra de Monchique
in southern Portugal, Sierra de Gredos in the cen-
ter-west of Spain, the hills of Teruel and Catalonia
in the northwest, the Balearic Islands, the eastern
Pyrenees, the Maritime and Ligurian Alps, the
whole of Corsica and Sicily, some parts of the Ital-
ian peninsula (Pignatti, in litt. 1994), the coasts of
Montenegro and Albania, the Pindus range and
mount Athos peninsula in Greece, Cyprus an
Crete considered together, the Mediterranean Cyr-
enaic, and the coastal Tell Atlas of Orania and Ka-
bylia in Algeria.
Another way of estimating biodiversity is to iden-
tify zones of great floristic species richness, which,
together with a study of endemism, would help to
define hot-spots. In the present case, we selected
geographical zones in their own right that cover a
small or medium surface area, where the floristic
variety is greater than 2000 species per 15,000
km?. In the circum-Mediterranean region, the sec-
tors that satisfy these criteria are certainly numer-
ous, although they are sometimes difficult to define
because of insufficient floristic knowledge for pre-
cise zones. However, the following can be included
in this group: the Baetic-Rifan complex, the eastern
Pyrenees, the Maritime and Ligurian Alps, Corsica
and Sicily, the central Appennines and Calabria,
the Dalmatian coast, Albania, Greece, Crete, the
Amanus-Taurus region, the Syrian-Lebanese coast,
Israel, and the coastal areas of Orania and Kabylia.
THREATS TO MEDITERRANEAN PLANT DIVERSITY
There are two very different situations in the cir-
cum-Mediterranean region relative to human im-
pact (Barbero et al., 1990). The non-coastal zonal
ecosystems of the northern part of the basin
(France, Italy, and Greece) are markedly free of
disturbance, and this leads to the extension of pre-
forest and forest areas dominated by expansionist
species. These include the Pinus species, which
have remarkable ecological and biological plastic-
ity and very high dissemination capacities (Barbero
& Quézel, 1989). This extension of phanerophytes
and also of chamaephytes tends to cause regression
of formations of reduced vegetation cover, particu-
larly the pastures, which are among the ecosystems
that are the richest in Mediterranean taxa and in
endemics. Therefore the collapse of the agro-sylvo-
pastoral system of previous centuries causes major
modifications to the structure and architecture of
phytocenoses, with standardization of the flora and
fauna. The Mediterranean elements tend to be re-
placed by medio-European species that are more
ubiquitous. On the other hand, there is strong hu-
man pressure on a large part of the coastal and
juxta-coastal areas, which poses a serious threat to
halo-psammophile and palustral species. Island
ecosystems are also very affected by the develop-
ment of mass tourism.
The areas in the southern part of the Mediter-
ranean basin (in particular North Africa) are sub-
jected to the major impacts of the constant increas-
es in population and in livestock, which totally
destructure the ecosystems and the soils, causing
strong erosive phenomena and very poor regener-
ation. This population explosion of the southern
countries remains the major problem: their total
population was 40 million in 1900, is now 290 mil-
lion, and will probably reach 370 million by the
year 2000 (Le Houérou, 1991).
Various recent works have tried to assess the
risks incurred by the Mediterranean flora (Gomez-
Campo, 1985; Quézel & Barbero, 1990; Ramade,
1990). The I.U.C.N. organization was a pioneer, and
Leon et al. (1985) published a general assessment
for all circum-Mediterranean flora and endemics
(Table 4). Naturally, these results are open to ques-
tion because they were established by various re-
searchers who did not all have the same conception
of the threats and the various categories proposed
by LU.C.N. to assess the risks. In addition, the ex-
act situation of several species in several countries
is unknown. However, according to this overview,
53% of the endemic species, i.e., 1529 taxa, are
endangered (excluding Syria, Lebanon, and Tur-
key). One should be cautious before proposing val-
ues for the entire flora, but an analysis of the Al-
gerian flora (Mathez et al., 1985) showed that
almost 50% of the species had not been observed
120
Annals of the
Missouri Botanical Garden
Table 4. Threats affecting Mediterranean endemic
plants, in countries where data are available (Leon et al.,
1985, modified). Other sources: (a): Mus (1995), (b):
Bramwell (1990), (c): Montmollin & latrou (1995), (d):
Olivier et al. (1995), (e): Conti et al. (1992), (f): Raimondo
Country Endangered
or and are
island Extinct vulnerable unknown
Albania 0 3 19
geria 1 эз 80
Balearic Islands (a) 1 18 29
Canaries (b) 1 243 162
orsica 1 32 3
Crete (c) 0 18 67
Cyprus 0 19 50
Egypt 2 18 48
France (d) 0 55 21
reece 5 61 435
Israel 0 3 10
Italy (e) 0 98 108
Libya 0 20 42
Madeira 0 47 61
Morocco 0 4 238
Portugal 2 22 34
Sardinia 0 8 10
Sicily (f) 4 107 161
in 1 32 202
Tunisia 0 1 1
Former Yugoslavia 1 Ў; 119
again for 20 years. The situation is also very wor-
rying in Macaronesia, where more than 20% of the
flora is endangered (LU.C.N., 1983). Leon et al.
(1985) listed only 16 species that have become ex-
tinct in the Mediterranean region since the begin-
ning of the century. Recently, Greuter (1994) in-
dicated that 37 species and subspecies of vascular
plants of the Mediterranean area are presumed to
be extinct. Nevertheless, these results are infinitely
too optimistic to have any real significance, partic-
ularly for the countries of the southern Mediterra-
nean. Furthermore, if we could really take into ac-
count the erosion of the different habitat types and
of the number of populations of rare species, the
situation would be undoubtedly worse that it seems.
DISCUSSION
The results and analyses mentioned above can
be used to identify sectors of the greatest general
biodiversity with the greatest number of endemics
that appear to be the most endangered, i.e., the
“hot-spots” (Fig. 2) as defined by Myers (1988,
). We distinguished 10 sectors, which clearly
fit Myers's definition, in the circum-Mediterranean
egion and we summarize the main threats to these
hot-spots (Table 5). Some other zones (the Dalma-
tian coast, the eastern Pyrenees, and some parts of
the Italian peninsula, or even Kabylia) could have
been defined as hot-spots, but the data concerning
them is too incomplete.
ARCHIPELAGOS OF THE CANARIES AND MADEIRA
These islands of quite recent volcanic origin (5
to 7 million years: Carracedo, 1980) with a surface
area of 8100 km?, which are not connected to the
continent, were apparently colonized by the long-
distance transport of diaspora (Shmida & Werger,
1992). The initial absence of competition allowed
strong adaptive radiation (Lems, 1960; Bramwell,
1975) and the establishment of a unique post-Mio-
cene flora with a rich variety of endemics (Hum-
phries, 1979; Bramwell, 1976, 1985; Quézel,
1995). Thus, of the total number of species 38%
are endemic in the Canaries (Schmida & Werger,
1992) and 26% in Madeira (Dalgaard, 1994). But
this unique flora is endangered (27% of the endem-
ic flora in Madeira and 41% in the Canaries, ac-
cording to the LU.C.N., 1983). The Canaries ar-
chipelago is subjected to drastic human impact in
its low-altitude zones by a considerable tourism in-
frastructure and the development of banana plan-
tations. Infracanarian formations of cactoid Eu-
phorbia and (to a greater extent) laurel woods have
clearly regressed (Santos, 1990). Thus, the Gran
Canaria laurisilva now covers less than 1% of its
original area, while 90% of Tenerife’s laurel woods
s disappeared (Bramwell, 1990). Trees such as
oenix canariensis . and Dracaena drago (L.)
= are only found together in a residual position, от
where they have been planted. Competition from
allogenic plants (Agave, Opuntia, and Acacia) is
also a serious problem. In the main island of the
Madeira archipelago, the laurisilva covered some
60% of the total area but is now reduced to about
16% (10,000 ha) according to Press and Short
1994).
HIGH AND MIDDLE ATLAS MOUNTAINS
Covering an area of around 50,000 km?, the High
and Middle Atlas ranges are populated by numer-
ous endemics—306 and 237, respectively—ac-
cording to Enriquez-Barroso and Gomez-Campo
(1991). The high rate of endemism is explained by
the long isolation of these massifs and their high
altitude. But, in addition to this mainly residual
flora (Quézel, 1957), there is quite a large number
of schizoendemics that demonstrate the role of the
Atlas mountains in the neo-speciation process (Gal-
Volume 84, Number 1 Медан 4 Quézel 121
1997 Mediterranean Plant Biodiversity
= = 5 å land, 1988). This general region is greatly endan-
Ec 3 5 + + + gered by anthropization, particularly by overgraz-
5 3 7 ing, land clearance, and anarchical deforestation
ж that lead to desertification, even in the pastures of
d spiny xerophytes in clumps at high altitude. The
- 3 a Y ai balance was disturbed in the 1950s and, for ex-
ms t ample, land clearance of the Azilal Province alone
now reaches 3000 to 5000 hectares per year, with
пи annual erosion of between 5 and 10 т? per hectare
See) +++ +++ (Estrade, 1988).
54 E +++ +++
' THE BAETIC-RIFAN COMPLEX
EI
a b- E ЕЁ 5 + Ea Andalusia and the Rif, linked together until the
5 [335883 : d of the Tertiary era, are grouped together in the
ЖТ Е end of the ry era, are grouped tog
E same hot-spot because they have great floristic,
5 3 2 Б ecological, and bioclimatic affinities. Around 75%
$ 3 E E | T + 1 zw of their total of 3500 species are common to the
+ 5 E E + two regions (Valdés, 1991).
њи ~ i Rif and Tell Coastal Ranges of western Alge-
El 3 x ria. These two sectors of North Africa have a rich
E т + de t A i H t variety of endemics. In the Moroccan Rif range,
E E T there are at least 190 endemics, including 50 that
8 5 are strictly limited to this zone (Enriquez-Barroso
= "m i & Gomez-Campo, 1991). Unfortunately, this region
t 3 E yd pos t is subjected to the relentless loss of natural vege-
ү 55 4 T tation, particularly due to the illegal growing of
3 < = hashish. Between 1966 and 1986, the wooded areas
ЈЕ ЗА and scrublands of the central-western Rif respec-
5 E " T: tively decreased by an average of 4296 (989 ha/
K Bitt is + T H 1 + + year) and 38% (2852 ha/year), while cultivated
ps 8 ы т land increased Ьу 93% ог 3847 ha/year (Boukil et
E al., 1987).
3 In Algeria, the Oranian and coastal sections of
B E : Т t Н b н ^ the Tell Atlas have, respectively, 91 and 89 endem-
E 5 EU ДИНАН ics (Enriquez-Baroso & Gomez-Campo, 1991). The
| 5 high rate of endemism reflects the Iberian and Mo-
| = roccan influences (Quézel, 1964a). There is also
| = E چ و مو E strong human pressure, and habitats disappear at a
| s = + е ++ rate that is difficult to estimate (Mathez et al.,
| >
| = © Andalusia. This region of southeast Spain,
| = F 87,267 km? in area, is unquestionably one of the
| pe Е А most important hot-spots of the Mediterranean ba-
| = E 2. $ sin. The complexity of the geologic and climatic
| 5 its ^h 2 history, but also the great diversity of habitats and
| E B = le 3 = а 8 E substrata (including serpentines, dolomites, and
| po To E & E Б= = gypsum), explain this high biodiversity (Valdés,
| 5 > = $33 э ы Е g 1993). Hernández-Bermejo et al. (1993) listed 484
| É 5 = E 5 са 5245 strictly endemic taxa and 465 subendemics in An-
| à $8 T t EE 3 - E dalusia, and Gomez-Campo et al. (1984) obtained
2 3 ZEEE = $ aE percentage of endemism greater than 50% in the
8 SK SEES EEE Serrania de Ronda and in Sierra Nevada. The latter
massif contains 177 endemics, including 66 that
122
Annals of the
Missouri Botanical Garden
are exclusive (Gomez-Campo et al., 1984). Due to
this exceptional biodiversity and the serious threats
(600 rare taxa, including 68 in danger of extinction,
according to Hernández-Bermejo et al., 1993), the
local authorities have just recently put in place an
integrated conservation plan (Hernández-Bermejo
& Clemente-Muñoz, 1993) that aims to limit human
activities in the mountain and coastal areas.
MARITIME AND LIGURIAN ALPS
The southwestern end of the alpine range is a
noteworthy pole of biodiversity, since around 31
species and subspecies, including almost 140 en-
demics, are found in this 9500 km? area (Médail &
Verlaque, 1997). This high degree of richness is
due to (a) the survival of ancient taxa (much of the
region was not affected by the Wiirm periods of
glaciation) facilitated by the diversity of habitats
and ecological niches (with a strong altitudinal gra-
dient in a small distance) and (b) the establishment
of more recent taxa facilitated by the continuity of
the alpine range. With the increase in population
and the explosion of tourism along the entire coast-
line, the most serious dangers again threaten the
species of the coast and low altitudes; one example
is the disappearance of Silene sericea All., an en-
demic Tyrrhenian psammophilous species.
TYRRHENIAN ISLANDS
The Balearic Islands (except for the Pithyuses,
which are biogeographically linked to the Iberian
peninsula), Corsica, Sardinia, and Sicily are the
remnants of areas that once belonged to the Pro-
toligurian massif (Alvarez, 1976), a Hercynian for-
mation that was fragmented in the Oligo-Miocene,
causing migration of the Corso-Sardinian micro-
plate. Only a part of Sicily (Peloritan massif) ad-
joined the Protoligurian massif in the south, but
this island was included in this hot-spot for prac-
tical reasons. As a result, these islands have several
floristic affinities, even if this Tertiary isolation con-
tributed to the differentiation of neo-endemics that
are specific to each area.
Balearic Islands. This archipelago has a rela-
tively limited specific richness, with 180 endemic
taxa of which 48 are endangered (Mus, 1995), but
only 1 is known to have become extinct: Lysimachia
minoricensis Rodr., which has, however, been con-
served ex situ. Tourism and the changes of land use
have seriously altered and modified the habitats,
particularly coastal communities, low-altitude
scrublands, and pine forests. According to a recent
study (Mus & Mayol, 1993), 5% of the overall flora
of the Balearics is seriously endangered.
Corsica. The flora and vegetation of this island
are now well known, particularly after the works of
Gamisans (1976-1978, 1991), Gamisans and Jean-
monod (1993), and Contandriopoulos (1962, 1990).
In an area of 8748 km?, 2354 indigenous taxa аге
found, with 270 endemics and subendemics, i.e.,
an 11.596 rate of endemism. The mountain flora
includes 39% endemics. Although Corsica's flora
seems to be relatively little endangered in the me-
dium and upper altitudes, the situation is clearly
more worrying in the coastal fringe, where three
quarters of the endemics are endangered. It should
be pointed out that 6 of them will probably become
extinct, including 2 endemics specific to Corsica,
as well as the extremely rare Naufraga balearica
Constance & Cannon.
Sardinia. Although it is floristically rather poor
(ca. 2050 taxa for 24,100 km?), Sardinia deserves
to be included in this study because its well-doc-
umented endemism (Arrigoni, 1976-1991) is sig-
nificant (106 endemics s. str.) and there are con-
siderable threats. The low-altitude areas are again
the most endangered, but conservation measures
are still quite parsimonious, and there are only few
reserves (Bocchieri, 1995).
Sicily. Due to its geographical position and
contrasting paleogeographical origin, Sicily is an
area of major botanical interest, with around 2700
taxa and 310 endemics and subendemics (Ver-
laque, pers. comm., according to Raimondo et al.,
1994, modified). The flora of the hills of Madonia
alone includes 50% of Sicily's species and 40 en-
demics (Brullo et al., 1995) in less than 246 of the
island's area. The Etna district contains 21 endem-
ics (Brullo et al., 1995). There are also endemic
trees, which is uncommon in the Mediterranean ba-
sin: Abies nebrodensis (Lojac.) Mattei, which is se-
riously endangered, Betula aetnensis Raf., Celtis
aetnensis Torn., Pinus nigra Arnold subsp. laricio
Maire, and the recently discovered Zelkova sicula
Di Pasquale, Garfi & Quézel. The human impact is
mainly responsible for the probable extinction of
29 taxa, including 4 Sicilian endemics: Allium per-
mixtum Guss., Anthemis abrotanifolia (Willd.)
Guss., Carduus rugulosus Guss., and Limonium ca-
tanense (Tinbeo) Brullo (Brullo et al., 1995).
SOUTHERN AND CENTRAL GREECE
The Peloponnese, with an area of ca. 21,000
km?, para a total of 2400 species, including 300
Greek endemics, i.e., 12.5% endemism (latrou,
1986). Strid (1993) њивама 26.5% orophile еп-
demism in the Peloponnese, i.e., 143 endemics;
о ас ва SEO NE о о са ~ овај а а оса
Volume 84, Number 1
1997
Médail & Quézel 123
Mediterranean Plant Biodiversity
this reaches 38% if we include the Balkan endem-
ics. The hills that contain the greatest biodiversity
are the Taygetos, Chelmos, Parnon, and Killini
mountains. The flora of the Peloponnese was af-
fected mainly by north-south migrations and has
strong links (130 endemics in common) with the
central Sterea Ellas region, which were joined to-
gether only 900,000 years ago. In this hot-spot we
have included the southern part of this region, from
the Karpenision belt, where, in an area virtually the
same as that of the Peloponnese, the rate of endem-
ism and the floristic richness are comparable, par-
ticularly in the mountain massifs: Parnassos, Giona,
Vardoussia, Panaetolicos, Dirphis; rupicolous en-
demism is more than 10% in almost all of this area,
and reaches 20% locally (Quézel, 1964b).
When Greece joined the European Union, its
economy swung from an agro-pastoral system to a
productivist market economy, which will cause pro-
found changes in the structure and architecture of
the ecosystems, and therefore in biodiversity (Mar-
garis, 1992).
CRETE
Crete is well documented from floristic (Turland
et al., 1993) and phytoecological viewpoints (Bar-
bero & Quézel, 1980; Zaffran, 1990). Spared by
Quaternary glaciations and isolated for around 5
million years, this island has a unique flora, with
171 endemics out of a total of 1706 species and
subspecies (Turland et al., 1993) in an area of 8700
km?. The orophile flora of the three main massifs
(Levka Ori, Psiloritis, and Dhikti) is very poor, but
of the 217 taxa listed, 44% are endemic (Strid,
1993). In his analysis of the vegetation of Crete’s
mountains, Zaffran (1982) obtained comparable re-
sults: 25 to 40% of endemics for the rupicolous
vegetation and 20 to 30% for stripped pastures.
The early colonization by man, about 8000 years
ago, profoundly altered the natural balance, with
exportation of timber and olives to Egypt between
2500 and 1500 B.C. (McNeely, 1994). Crete still
bears the marks of thousands of years of intensive
exploitation; forest patches are rare, while the
scrublands have a lesser richness and floristic di-
versity.
ANATOLIA AND CYPRUS
Cyprus. The island of Cyprus, with an area of
9250 | km?, includes 1620 taxa with 170 endemics
and subendemics (Alziar, 1995). The most inter-
esting sector is the Troodos massif, where serpen-
tinicole endemism is highly developed, with unique
formations of Cedrus brevifolia (Hooker fil.) Dode
and Quercus alnifolia Poech (Barbero & Quézel,
1979) and at least 45 local endemics. The entire
island is subjected to major erosion (Tsiourtis,
1993), and mass tourism is developing rapidly on
the south coast.
Anatolia. Southern Anatolia, especially the
Taurus and Amanus areas, is one of the major cen-
ters of biodiversity and endemism in the circum-
Mediterranean area. Biogeographical spectra in
studies of the vegetation (Quézel, 1973) show that
orophile endemism in the Taurus area reaches 20—
40% in rupicolous associations, 10-20% in
stripped pastures, and 35-70% in culminating
screes, with a total of 250 endemics among the 650
species presented in the surveys. Davis et al.
(1994) mentioned 2500 species including 250
Turkish endemics in the eastern part of the Taurus
area and 3365 species in southwest Anatolia, in-
cluding 675 Turkish endemics.
Even if the situation of the natural environments
is better than in some other areas of the Mediter-
ranean, the human impact is nevertheless percep-
tible in the overall Amanus/Taurus area. The nu-
merous steppes are the result of continuous human
activity, access routes, crop fires, and pasture fires.
Land clearance over several centuries has mainly
affected the forests of Pinus nigra Arnold subsp.
pallasiana (Lamb.) Holmboe, Cedrus Шат A.
Richard, and Quercus cerris L. (Akman et al.,
1991), but also the scrublands of endemic Astrag-
alus (Demiriz & Baytop, 1985). However, one of the
major problems is the intensive harvesting of main-
ly endemic bulb plants (Oldfield, 1990) for deco-
rative and culinary purposes. Thus, according to
Sezik (1989), 57 million tubers belonging to 38
species of orchids are picked annually and are used
to prepare “salep,” a milk drink.
SYRIA-LEBANON-ISRAEL
The summits of the Lebanon and the Anti-Leb-
anon ranges (Slenfe-Qammoua, Quriet es Sauda,
Ehden Sannin, and Mount Hermon) and their pe-
ripheries are areas rich in endemics. One hundred
species specific to mount Hermon and the Anti-
Lebanon range have been counted (Zohary, 1973),
while Auerbach and Shmida (1985) mentioned 47
endemic taxa on the coast of Israel, i.e., a rate of
27%. Nevertheless, the conflict in this region has
seriously degraded the phytocenoses of the region,
e.g., the formations of Cedrus libani. However, pre-
cise and recent data on the evolution of the natural
environments are not available.
124
Annals of the
Missouri Botanical Garden
MEDITERRANEAN CYRENAIC
Even if the northern part of the Cyrenaic region
does not have a rich flora (1300 species), it still
contains no less than 81% of Libya’s flora, in only
1.3% of the country’s area. The rate of endemism
is around 10%, with 137 endemic taxa including
84% of essentially Mediterranean origin (Bartolo et
al., 1977, modified). The Djebel Akdhar plays a
key role in the area’s biodiversity, but the surface
area of the natural environments of the “Green
Mountain” has been reduced by half (or about
250,000 hectares) in half a century (EP Hamrouni,
1990). Tree-cutting, repeated fires, overgrazing, and
mechanical land clearances in low altitudes to ex-
tend the orchards and vineyards have strongly al-
tered this Mediterranean enclave.
CONCLUSION
Like the four other mediterranean areas of the
world (Cody, 1986), the circum-Mediterranean re-
gion is one of the world's major centers for plant
differentiation. The assessments mentioned above
show that, particularly for the Old World, this re-
gion ranks among the most important areas of plant
biodiversity. The Dd region
alone contains as many species as Europe and
Northern Africa IE Similarly, its high level
of endemism is remarkable, and is often equal to
that of the world’s tropical regions.
In the Mediterranean basin, this richness is not
uniformly distributed and the areas of maximum
biodiversity, particularly the 10 hot-spots defined
here, mainly trace their origins to historical and
paleogeographical factors (Pons & Quézel, 1985).
The Mediterranean region is an area of refuge but
also floral exchange and active speciation (Quézel,
1978, 1985, 1995). In the occidental part of the
Mediterranean basin, the maximal biodiversity ar-
eas coincide notably (Verlaque et al., in press) with
the “Protoligurian massif” and the “Orogenic Belt,”
which stretched from the Alps to the Baetic moun-
tains during the Oligocene (Alvarez, 1976). The
phenomena of stress (Stebbins, 1952), competition,
and also the diversity of habitats, with a varied ped-
ological component, explain the region’s great rich-
ness and its endemism, which is particularly im-
portant in the high mountains and in the islands.
However, this biodiversity is now seriously en-
dangered by radically different phenomena that af-
fect the north and south of the basin. The southern
countries are subjected to reckless human activity,
with overexploitation by man and livestock in all
available areas, following an exponential growth in
population. On the northern perimeter, however, the
virtual abandonment of the agro-sylvo-pastoral sys-
tem is causing uniformization of the ecosystems and
flora, linked to a general biological upsurge, while
the coastal areas are ravaged by the spread of mass
tourism. In order to counteract this erosion of the
phanerogamic biodiversity, it would be necessary to
refine the assessments presented in this work and
to develop integrated conservation strategies for the
identified hot-spots (Falk, 1990), both on the re-
gional level (as has been started in Andalusia, the
Balearics, and Corsica) and on national and inter-
national levels. We must also emphasize that threat-
ened habitats in need of conservation (e.g., wet-
lands and coastal habitats) are not necessarily
hot-spots of biodiversity. Thus, it is useful to make
the distinction between a general need for conser-
vation of endangered habitats, and a more specific
need for conserving centers of plant richness and
endemism through hot-spots.
In addition, it would be instructive to survey the
hot-spots of other taxonomic groups besides plants
in order to determine whether the various defined
zones coincide. In Great Britain, Prendergast et al.
(1993) did not find very close relationships between
the hot-spots of the different groups, and this
should be tested in the Mediterranean area.
Literature Cited
Akman, Y., P. Quézel, M. Barbero, O. Ketenoglu & M.
191-425
Alvarez, W. 1976. А nne continuation of the Alps.
Geol. Soc. Amer. Bull. 8
Alziar, G. 1995. La flore А nc Ecol. Medit. 21: 47-
n».
Arrigoni, P. V. 1976-1991. Le piante endemiche della
Sardegna. Boll. Soc. Sarda Sci. Nat. 16-27.
Auerbach, M. & A. Shmida. 1985. Harmony among en-
demic littoral plants and adjacent floras in Israél. J.
Biogeogr. 12: 175-187.
Barbero, M. & P. Quézel. 1979. Vulp Up: à T
des groupements forestiers de Chypre. Doc. Phytos
nouv. ser. 4: 9-34 + 1 tab. h.-
& ———. 1980. La végétation forestière de Сте-
te. Ecol. Medit. 5: 175-210.
& 98
9. Structures, architectures fores-
tiéres à sclérophylles et о des incendies. Bull.
col. 20: 7-14.
. Bonin, R. Loisel & P. Quézel. 1990. Changes
and eR RRO of forest ecosystems caused by human
activities in the western part of the mediterranean bas-
sin. Vegetatio 87: 151-173.
Bartolo, G., rullo, A. Guglielmo & C. Scalia. 1977.
Салсын fitogeografiche sugli endemismi della
Cirenaica settentrionale. Archiv. Bot. Biogeogr. Ital. 53:
–154.
Bocchieri, E. 1995. La connaissance et l'état de conser-
а а ава ако а ТҮ" E пас оваа
Моште 84, Митбег 1
1997
Медан 4 Quézel 125
Mediterranean Plant Biodiversity
vation de la flore de la Sardaigne. Ecol. Medit. 21: 71–
81
Bond, P. & P. Goldblatt. 1984. Plants of the Cape Flora.
. S. African Bot., Suppl. 13: 1-455.
Воші, A Е, A. Blali & M. и Каш. т. nce ние
] dégradati
ers tun de l'Agriculture et de е; емнен Ааай
re, Centre régional де l’inventaire et des aménagements,
Tetouan.
Boulos, L. 1995. Flora of Egypt. Checklist. Al Hadara
ubl., Cairo.
Bramwell, D. 1975. Some morphological aspects of the
adaptative radiation of Canary Islands Echium species.
Anal. Inst. Bot. Cavanilles 32: 241-254
1976. The endemic flora of the Canary Islands;
Distribution, Relationships and Phytogeography. Pp.
207-240 in G. Kunkel (editor), Biogeography and Ecol-
ogy in the Canary Islands. W. Junk, The Hague.
1985. Contribución a la ie PRI de las Islas
Canarias. Bot. o IV Ci.
nserving ейнае іп уче eres Is-
Ф
. Spampina 995. Con
colon! sc a sulla flora della Sila Ecol.
Medit.
Glo di M. D. imis, T. H. Johnson & B.
G 1996. rities for conserving global
species s and nic as Biodiv. Conserv.
Carracedo, 1. 0 1980. Atlas básico de Canarias. Inter-
insular Can
Chauvet, M. & L ; Oliv ier. 1993. La biodiversité, enjeu
planéteits—Présbrpet notre patrimoine génétique. Sang
de la yu Paris.
Cody, M. L. 1986. Diversity, rarity, and conservation in
mediterranean-clima
Conseil de l'Euro
relle des états us des Communautés européennes
et du Conseil de l'Europe. Publ. no. Eur. 10970, Com-
mission des Communautés Européennes, Luxembourg.
Contandriopoulos, J. 1962. Recherches sur la flore en-
démique de la Corse et sur ses origines. Ann. Fac. Sci.
T 32: 1-354.
990. Spécificité de l'endémisme corse. In: Bio-
paid aspects of insularity. Rome, 18-22 May
1987. Atti Accad. Naz. Lincei 85: 393-416.
Conti, F., A. Manzi & F. Pedrotti. 1992. Libro rosso delle
piante d'Italia. WWF Italia, Roma.
Cowling, R. M., I. A. W. MacDonald & M. T. Simmons.
1996. The Cape Peninsula, South Africa: Physiograph-
ical, biological and historical background to an extraor-
dinary hot-spot of biodiversity. Biodiv. Conserv. 5: 527—
Daget, P. 1977. Le bioclimat méditerranéen, caracteres
généraux, — de caractérisation. Vegetatio 34: 1-20.
Dalgaard, V. Checklist of chromosome —
counted in Makita flowering plants, with notes on
polyploidy, life form, endemism and و فان
J. Bot. 14: 241-255.
Davis, P. 1 ora of Turkey and the east Aegean 15-
lands supplement. Vol. 10. Edinburgh Univ. Press, Ed-
inbu
Davis, S. D. V. H. Heywood & A. C. Hamilton. 1994.
Cen ant Diversity. A Guide and Strategy for
their lel rawi e Volume 1. Europe, Africa, South
West Asia and the Middle East. WWF & I.U.C.N.
LU.C.N. reer Unit, Cambridge
Gregerson, L. Henson, C. J.
Leon, pi чије H. Synge & J. Zantovska.
1986. ede in ree What do we know? 1.U.C.N.,
Delis H. & T. Baytop. 1985. The Anatolian assum
1 ant Con-
servation in the Mediterranean area. pidum 7, W.
Junk, Dordrecht, Boston, Lancas
Duivenvoorden, J. F. 1 Gaohi ~ species counts
in the rain forests of the middle Caqueta area, Colom-
bian Amazonia. Biodiv. Conserv. 3: 685-715.
El'Hamrouni, A. 1990. Approche bioclimatique et bio-
géographique du Djebel Akhdar Cyrénaique, Libye.
Ecol. Medit. 16: 163-167.
Enriquez-Barroso, A. & C. Gomez-Campo. 1991. Les
plantes endémiques de l'Afrique du Nord-Ouest: Al-
gérie, Maroc et Tunisie. Bot. Chron. 10: 517—520.
Estrade, R. 1988. Les causes économiques de la dégra-
dation du milieu naturel, le cas de la province d'Azilal.
Séminaire natl. sur l'aménagement des bassins versants.
gres de l'Agriculture et de la Réforme A
Falk, 1990. Integrated strategies for гае й
а са diversity. Ann. Missouri Bot. Gard. 77:
384
Fennane, M. & M. Ibn Tattou. 1995. Га Ноге rare des
hautes montagnes maroc € in Actes
.R.P.E., Provence-Alpes-
Cóte dena Colloque scientifique Emr Bio-
Mes
Forget, P M. 1994. Les foréts tropicales en sursis. La
Recherche 270: 1154—1162.
Galland, N. 1988. Recherche sur l'origine de la flore or-
ophile du Maroc. Etude caryologique et cytogéogra-
phique. These Fac. Sci. Univ. Neuchátel. Trav. Inst. Sci.
Univ. Mohammed V, Sér. Bot. 35: 1-168.
Gamisans, J. 1976-1978. La végétation des montagnes
corses. Phytocoenologia 34: 425—498; 41: 35-131; 42:
133-179; 43: 317-376; 44: 371-432.
991. La végétation de la Corse. /n: D. Jean-
monod & H. M. Burdet (editors), Compléments au Pro-
drome de la Flore Corse. Conservatoire & Jardin bota-
nique de Geneve, Geneve.
——— & D. Jeanmonod. 1993. Catalogue ay SET
vasculaires de la Corse seconde édition. / Jea
ve.
ге of neotropical plants species
— Evol. Biol. 1 1-84.
in tropical versus temperate ا
plant eR vat Pp. 153-181 in M. E. Soulé (editor),
Conservation Biology. The Science of Scarcity and Di-
Sinaveras Publ., Sunderland. —
. Tree species richness of upper Amazonian
ore is: Pro. Natl. Acad. Sci. U.S.A. 85: 156–159.
Colla P. 1978. An analysis of the flora of Southern
Africa: Its ero: relegando and origins.
Ann de
Gemei-Campo, C. (Editor). 1985. Plant Соини i in
the Mediterranean area. Geobotany 7, W. Junk, Dor-
drecht, Boston, Lancaster.
6 1: Bernnidex-De-Castro, M. J. Cagiga & M. D.
Sashes: Ydlénio. 1984. Endemism in ref Iberian Pen-
insula and Balearic Islands. Webbia 38: 709-714
126
Annals of the
Missouri Botanical Garden
Good, R., D'O. 1974. The tees of Flowering Plants.
4th ed. Longman, Lon
Greuter, W. 1991. Botanie diversity, endemism, rarity,
and extinction in the Mediterranean area: An analysis
based on the Ded volumes of Med-Checklist. Bot.
Chron. 1
994. ак in Mediterranean areas. Philos.
d B 344: 41-46.
995. Origin and pecularities of Mediterranean
island nais Ecol. а 21: 1-10.
Hernández-Bermejo, . & M. Clemente-Muñoz. (Edi-
tors). 1993. тсң де la Fora en Andalucía. Junta
de Andaluc
aa AEE & M. Clemente-Muñoz. 1993.
Catálogo general de las especies de recomendada pro-
tección en Andalucía — raras y amenazadas
de extinción. Pp. 43-62 in J. E indes -Bermejo &
M. Clemente- Muñoz (editors), Fracción de la Flora en
Anil Junta de Andaluc
Heywood, V. H. The n flora in the
context of world biodiversity. Ecol. Medit. 21: 11-18.
Hobbs R. J., R. H. Groves, S. D. Hopper, R. J. Lambeck,
B. B. Lamont, S. Lavorel, A. R. Main, J. D. Majer &
D. A. Saunders. 1995. Function of biodiversity in the
arp y ecosystems d southwestern Austra-
lia. Pp. 233-284 in G. W. Davis & D. M. Richardson
(editors), Mediterranean-Type Кы ны The Func-
Sian
tion of Biodiversity. Ecological Studies, Vol. 109.
Springer-Verlag, Berlin, Heidelberg
реак, C. J. 1979. Endemism ы evolution in М
aronesia. Pp. 171-199 in D. Ремо) (editor), iat
id Islands. Academic Press, Londo
Iatrou, G imvoli sti meleti Ке endemismou tis
chloridas tis Peloponnissou. Thesis Inst. Bot. Univ. Pa-
tras.
LU.C.N. 1980. First preliminary draft of the list of rare,
threatened and endangered plants for the countries of
North Africa and the Middle East. Mimeo, I.U.C.N.,
Kew.
servation Monitoring Centre. 1983. List of
rare, option and endemic plants in Europe. 2nd ed.
Nature and Environmental Series, 27. Council of Eu-
rope, Strasbourg.
Lanfranco, E. 1995. The Maltese flora and conservation.
Ecol. Medit. 21: pe
Le Houérou, H. N. ids Méditerranée en l'an 2050:
Impacts E E éventuelle évolution clima-
tique et de la démographie sur la végétation, les éco-
systémes et l'utilisation des terres. La Météorologie: 4—
37.
Lems, K. 1960. Botanical notes on the Canary Islands.
II. The evolution of plant forms in the islands: Aenium.
Ecology 41: 1-17.
Leon, C., G. Lucas & H. Synge. 1985. The value of in-
formation а saving threatened mediterranean plan
Pp. 177-196 in C. Gomez-Campo (editor), Plant pa
servation in the Mediterranean area. Geobotany 7, W.
Junk, penis Boston, Lancaster.
Leveque, С. . Le concept de biodiversité: de nou-
veaux dee sur la nature. Nature, Sciences, Sociétés
3-254.
Leba, L., A. M. Olson, L. R, Brubaker, S. R. Car-
лаик! M. M. Holland, 5. P. Hubbell, 5. A. Levin, J.
A. MacMahon, P. A. Matson, J. M. Melillo, H. A. Moo-
ney, C. H. Peterson, H. R. Pulliam, L. A. Real, P. J.
Regal & P. G. Risser. 1991. The Sustainable Biosphere
Initiative: An ecological research agenda. Ecology 72:
371-41
992. Régions de montagne en Gréce.
ture et transformation des terres dans le bassin médi-
terranéen. Icalpe, Le Bourget-du-
Mathez, J., P. Quézel & С. Raynaud. 1985. Тће Марћгеђ
countries. Pp. 141-157 in С. Gomez-Campo cm
Plant Conceal in the Mediterranean area.
any 7, W. Junk, Dordrecht, Boston, Lanca
i Lessons from the fepe rais and
biodiversity. Biodiv. Conserv. 3: 3-20.
Médail, F. & R. Verlaque. 1997. Ecological characteris-
tics and rarity of endemic plants from S.E. France and
veri xa Implications for biodiversity conservation. Biol.
erv., in press.
нара B. & G. A. latrou. 1995. Connaissance et
WD. de la flore de l'ile de Crete. Ecol. Medit.
21:
Mus, и oa: Conservation of flora in the Balearic Is-
lands. Ecol. Medit. 21: 185-194.
& J. Mayol. 1993. Plans de conservacio dels ve-
eral D'Estructures Agraries i Medi Natural, Servei de
Conservacio de la Naturalesa, Palma de d
Myers, N. aa Ae deforestation and a mega-ex-
tinction. Pp. M Soulé (editor), Ca
vation р The Science of Scarcity and Diversity.
Sinaveras Publ., Sunderland.
988. nad biotas: “Hot-Spots” in tropical
forests. The Environmentalist 8: 187—
——. 1990. The biodiversity с allenge: Expanded
Hot-Spots analysis. The Environmentalist 10: 243-256.
Oldfield, The international trade in bulbs.
T.R.A. FELC. (Int.) Bull. 11: 34-46.
Olivier, L., J. P. Galland, H. Maurin & J. P. Roux (Editors).
995. Livre rouge de flore menacée de France. Tome
Espèces prioritaires. Museum National d'Histoire
Naturelle & Conservatoire Botanique National de Por-
querolles, Pari
Pignatti, S. 1982. Flora d'Italia. Edagricole, Bologna, 3
vol.
e
Polunin, O. 1987. Flowers of Greece and the Balkans, à
Field Guide. Oxford Univ. Press, Oxford & New York.
Pons, A. & P. Quézel. 1985. The history of the flora and
Pp.
po (editor), Plant Conservation in the
rea. Geobotany 7, W. Junk, Dordrecht, Boston, Lan-
aster.
Prendergast, J. R., R. M. Quinn, J. H. Lawton, B. C. Ev-
rsham & D. W. Gibbons. 1993. Rare species, the co-
r of diversity hot-spots and conservation strat- —
egies. Nature = 335-337. a
Press, J. R. & M. J. Short. 1994. F eet E Madeira. The
Natural History Museum, HMSO, London.
Quézel, P. 1957. Peuplement végétal des кибе montag-
nes d'Afrique du Nord. Encycl. Biogéogr. Ecol., 10.
Lechevalier, Paris.
а. Lendémisme dans la flore de l'Algérie.
C. R. Soc. Biogéogr. 361: 137-149. :
. Végétation des hautes montagnes де —
Grice méridionale. Vegetatio 12: 279—
Volume 84, Number 1
1997
Médail & Quézel
Mediterranean Plant Biodiversity
127
1973. Contribution à l'étude ode
du Tourue. Phytocoenologia 1: 131-222
978. Analysis of the flora of the mediterranean
ES ud Africa. Ann. Missouri Bot. Gard. 65: 479—
1985. Definition of the Mediterranean region
and the origin of its flora. Pp. 9-24 in C. Gomez-Campo
(editor), Plant Conservation in the M вијка Area.
Стори 7, W. Junk, Dordrecht, Boston, Lancaster.
5. La flore du bassin méditerranéen: Origine,
mise en аи endémisme. Ecol. Меди. 21: 19–39.
& M. Barbero. 1985. Carte de la végétation po-
tentielle de la région пи гај Feuille по. 1
Méditerranée Orientale. C.N.R.S.,
& Les foréts pê А
Problèmes posés par leur signification historique, écol-
ogique et leur conservation. Acta Bot. Malac. 15: 145-
178.
—— — € ——. 1993. Variations iouis au Sa-
hara et en Afrique sèche depuis le Pliocéne: Enseig-
nements de la flore et de la végétation lía. Bull.
Ecol. 24: 191-202.
Raimondo, F., L. Gianguzzi & V. Ilardi. 1994. Inventaria
delle specie “а rischio" nella flora
la Sicilia. Quad. Bot. Ambientale Appl. 3: Paride
990. Conservation des écosystèmes médi-
Plan Bleu, 3. P.N.U.E.-P.A.
‚ €: D. L Axelrod. 1978. Origin and n.
ships el the California flora. Univ. California Publ. Bot.
Riva aeai S., W. Wildpret, T. E. Diaz, P. L. Perez
De Paz, M. Del Arco & O. Rodriguez. 1993. Excursion
guide. Outline vegetation of Tenerife Island Canary Is-
lands. Itinera Geobotanica 7: 5-167.
. Bosques de laurisilva en la region ma-
. Nature and Environmental Series, 49.
Council of Europe, Strasbourg.
Sezik, E. 1989. Turkish orchids and salep. Pp. 181-189
in Orchidées botaniques du monde entier. Soc. Fr. Or-
chidophilie, Paris.
Shmida, A. & M. J. A. Werger. 1992. Growth form di-
versity on the Canary Islands. Vegetatio 102: 183-199
Solbrig, O. T. 1 1992. Тће IUBS- SCOPE-UNESCO program
S ове aspects of the Greek
mountain ee Fragm. Florist. Geobot. Suppl. 2: 411–
433.
Suning, P. 1979. Origins of the Macaronesian flora. Pp.
3—40 in D. Bramwell (editor), Plants and Islands. Ac-
ademic Press, London.
Tsiourtis, N. 1993, Retenues et contróle de la sédimen-
25 in Actes des 5émes Ren-
the Cretan Área. Annotated Checklist and Atlas. The
Natural History Museum, HMSO, London.
Valdes, B. 1991. Andalucía and the Rif. Floristic links
and a common flora. Bot. Chron, 10: 117-124,
i Origen y génesis de la flora Andaluza.
Pp. 23-28 in J. E. Hernández-Bermejo & M. Clemente-
Mufioz (editors), Va a de la Flora en Andalucía.
Junta de Andaluc
Valencia, R., H. nem & G. Paz y Mino. 1994. High
tree alpha-diversity in Amazonian Ecuador. Biodiv.
Conserv. 3: 21-28.
Verlaque, R., F. Médail, P. Quézel & J. F. Babinot. En-
démisme végétal et paléogéographie dans le bassin
méditerranéen. Geobios, in press.
Wilson, E. O. (Editor). 1988. Biodiversity. National Ac-
ademic Press, Washington
Zaffran, J. 1982. Contribution à la flore et à la végétation
de la Crète. 2 vol. Publications Univ. Provence, Mar-
seille.
. Contributions à la flore et à la не are
on la ска Publications Univ. Provence, Аіх-еп-
Coban M. 1973. Geobotanical meres of the Middle
East. Gustav Fisher Verlag, Stuttg,
SURVEY OF CHROMOSOME
NUMBERS IN RUBUS
(ROSACEAE: ROSOIDEAE)!
Maxine M. Thompson?
ABSTRACT
A literature survey was made of chromosome number counts for Rubus species. Numbers are presented for 387
species, representing about 40 percent of the total number, and including 11 of the 12 subgenera. The basic number
is universally 7 and ploidy levels include 2x, 3x, 4x, 5x, 6x, 8x
Әх, 10x, 11х, 12x, 14x, and, questionably, 13x and
8x. In a few species, more than one chromosome number has been reported. Attempts were made to provide the
currently accepted taxonomic designation for each species as well as the outdated synonyms used in original publi-
cations. The objective was to provide a summary of available information on chromosome numbers of Rubus species
and to point out the gaps that need to be addressed.
Rubus is a large and important genus that in-
cludes an estimated 900 to 1000 species widely
distributed throughout the world. Representatives
are found on all arable continents as well as on
oceanic islands, and many species provide an im-
portant food resource for both humans and animals.
Early cytological studies demonstrated that poly-
ploidy has played a significant role in the evolution
of this genus (Gustafsson, 1943). The basic number
is universally 7 and, currently, ploidy levels are
known to range from 2x to 14x, and possibly 18x.
The most recent survey of Rubus chromosome num-
bers was included in Chromosome Numbers of
Flowering Plants (Fedorov, 1969). Since that time,
counts for many more Rubus species have been
published in widely scattered reports. Knowledge
of the chromosome numbers of species is important
to botanists studying cytotaxonomy, evolution, and
phylogenetic relationships and to plant breeders
utilizing interspecific hybridization as a breeding
procedure.
The exact number of species is unknown because
the only comprehensive world taxonomic treatment
was published more than 80 years ago (Focke,
1910-1914). During the past eight decades, many
new species have been described and, as new ev-
idence has accumulated, many nomenclatural
changes made. Recent regional taxonomic studies
such as those by Davis (1990), Davis et al. (1967—
1970), Edees and Newton (1988), Hogdon and
Steele (1968), Kalkman (1984, 1987), Weber
(1972, 1981), Weber and Maurer (1991), Yü
(1985), and Zandee and Kalkman (1981) have
helped clarify the identity of many species in the
authors’ respective regions. Excluding the subge-
nus Rubus, there are approximately 335 species.
For subgenus Rubus in Europe, Edees and Newton
(1988) gave 300 species in Britain alone (not in-
cluding the numerous continental species), whereas
for this subgenus in eastern North America, Davis
(1990) claimed 198 species. Thus, current esti-
mates of the total number of species in the genus
Rubus may reach 900 to 1000. A comprehensive
taxonomic study of this genus is needed to deter-
mine more accurately the number of species.
Chromosome number counts for 387 species,
representing about 40 percent of the species in the
genus, are presented in Table 1, at the end of this
article. An additional 75 taxa in subgenus Rubus
whose names are invalid or whose identities are
uncertain are also given because, although not ac-
curately named, they are representatives of the Eu-
ropean blackberry flora. Although voucher speci-
mens are reportedly available for some of the
species counted, 1 have made no effort to verify
their identity, except for those counted by myself.
However, by consulting recent literature and
through personal communication with taxonomists
specializing in Rubus, I have attempted to provide
currently accepted names for the species reported.
The originally published name is also included so
that, in the event that my re-naming is incorrect, à
Rubus taxonomist could readily identify the spe-
cies. The number of counts for a species varies from
o e
1 The author gratefully acknowledges the assistance of the following Rubus specialists who kindly reviewed specific
subgenera and made valuable nomenclatural
.
suggestions in the early stages of preparation of this manuscript: Heinrich
E. Weber for the European blackberries; Mark Widrlechner for North American blackberries; and Naohiro Naruhashi
for the subgenus Ideaobatus
22715 NW Frazier Crick Rd., Corvallis, Oregon 97330, U.S.A.
ANN. Missouri Bor. Garp. 84: 128-164. 1997.
Volume 84, Number 1
1997
Thompson 129
Chromosome Numbers in Rubus
one to several. In cases where different chromo-
some numbers are reported for a species, these may
be explained by one of the following: one of the
counts may represent a variant individual resulting
from spontaneous doubling of chromosomes at some
early stage of development or from the union of an
unreduced gamete with a reduced gamete, a com-
mon phenomenon in this genus; they may actually
represent populations within a species that have
different ploidy levels; or they may be the result of
mistaken identification by an author.
In the discussion and in Table 1, the species,
chromosome numbers, and reference citations are
arranged according to the subgenera established by
Focke (1910-1914), except for Micranthobatus,
whose species have been separated out of Lampo-
batus by Kalkman (1987), and Dalibarda. Since
most of the species placed in subgenus Dalibarda
by Focke (1910-1914) have been transferred to
subgenus Cylactis and R. dalibarda, itself, has been
moved to a different genus, as Dalibarda repens L.
(Bailey, 1941), the continued acceptance of Dali-
barda as a subgenus is questionable. In Table 1, the
currently accepted name is given first, synonyms
are indented under the valid name, and names in-
dented, in parentheses, and preceded by “as” in-
dicate that in the original publication, identification
was incorrect but, subsequently, the plant was re-
identified. Eleven of the 12 subgenera are repre-
sented: only Lampobatus sensu stricto remains to
be studied.
SUBGENUS CHAMAEMORUS
The chromosome number of R. chamaemorus, the
only species in this subgenus, is well established
with 18 hexaploid counts.
SUBGENUS CHAMAEBATUS
Of about six species in this subgenus, three have
been counted: R. calycinus and R. pectinellus are
hexaploid, whereas R. nivalis is diploid.
SUBGENUS COMAROPSIS
Only one of the two species in this subgenus has
been counted: R. geoides is tetraploid.
SUBGENUS CYLACTIS
There are chromosome number counts for 10 of
the appproximately 16 species now included in this
subgenus. Six of these, R. lasiococcus, R. pedatus,
R. pubescens, R. pseudojaponicus, R. stellatus, and
R. subarcticus are diploid. Rubus arcticus which,
with several counts, is no doubt predominately dip-
loid, was found to have a distinct triploid popula-
tion (Johnson & Packer, 1968). Zhukova (1980)
also reported triploidy for this species. It is not un-
common that diploid species give rise to an occa-
sional triploid or tetraploid individual which may,
through vegetative reproduction or apomictic seeds,
spread to form colonies or populations of consid-
erable extent. In the case of R. humulifolius, more
information is necessary to determine whether this
species is mainly diploid (two counts) or tetraploid
(one count). However, for R. nepalensis, the three
tetraploid counts, as compared to only one diploid,
suggest that it may be principally a tetraploid spe-
cies. With 14 counts, R. saxatilis is well established
as a tetraploid species.
SUBGENUS OROBATUS
Of the approximately 18 species in this exclu-
sively South American subgenus, 6 have been
counted and all are hexaploid. These include the
relatively weak-growing plants of R. acanthophyl-
lus, R. coriaceus, and R. glabratus, as well as the
very vigorous species R. macrocarpus, R. nubige-
nus, and R. roseus, which is also vigorous and has
robust canes to 6 m or longer. This high chromo-
some number contrasts with the consistent diploid
number in the blackberries (subg. Rubus) that oc-
cur in the same Andean region as the Orobatus
species.
SUBGENUS DALIBARDASTRUM
Three of the 10 to 12 species in this subgenus
have been counted and all are polyploid; R. tricolor
and R. tsangorum are tetraploid, whereas R. am-
phidasys is hexaploid.
SUBGENUS MALACHOBATUS
When Fedorov (1969) published his Rubus chro-
mosome number survey, only seven species in the
Malachobatus had been counted. Currently, 40
(31%) of the estimated 127 species have been
counted. Thus far, all species in this subgenus are
polyploid: there are twenty-seven 4x, five 6x, six
8x, and two 14x species. Although there are seven
diploid counts, this ploidy level is not consistent
with other information. Four of the species report-
edly diploid were found to be high polyploids by
other investigators: R. fairholmianus was octoploid
(Nybom, 1986); R. gardnerianus was octoploid (Gill
et al., 1984; Singhal et al., 1990); R. rugosus was
both octoploid (Iwatsubo & Naruhashi, 1992) and
14х (Nybom, 1980, 1986; Thompson, 1995a); and
R. hayata-koidzumi was tetraploid (Iwatsubo & Na-
130
Annals of the
Missouri Botanical Garden
ruhashi, 1993; Thompson & Zhao, 1993). Of the
other three doubtful diploid species, two have sole
counts, namely, R. fulvus and R. micropetalus by
Subramanian (1987), and the third, R. paniculatus,
has two counts (Malik, 1965; Mehra & Dhawan,
1966). The identity of all species with diploid
counts needs to be verified and their counts recon-
firmed before it can be concluded that diploid spe-
cies occur in this predominately polyploid subge-
nus.
Omitting the discrepant diploid counts, there ap-
pear to be three species in which two different
numbers actually do occur. Naruhashi and Iwatsu-
bo (1993) reported that two forms, both hexaploid
and octoploid, are commonly found in both R. hak-
onensis and R. buergeri. For R. rugosus, there is one
octoploid and three 14x counts. These two different
counts for R. rugosus, along with the ploidy levels
reported for its close relative, R. moluccanus (4x),
and its allies R. multibracteatus (4x), R. hillii (6x),
R. fairholmianus (8x), possibly R. indicus (8x), and
R. glomeratus (14x), suggest that this group of spe-
cies represents a polyploid complex. Further cyto-
taxonomic studies are necessary to clarify the re-
lationships among these species.
Thus far, there is no evidence for apomixis
among the Malachobatus polyploid species. Ny-
bom’s (1986) demonstration of sexual reproduction
in two species, R. fairholmianus and R. rugosus,
needs to be expanded to other species in order to
determine if this is the predominate mode of repro-
duction in this subgenus. The existence of relative-
ly clear-cut species boundaries supports the con-
cept that hybridization and apomixis have not
played an important evolutionary role, if any, as
they have in Rubus, the other major polyploid sub-
genus.
SUBGENUS ANAPLOBATUS
In this small subgenus of possibly five to six spe-
cies, the four species counted are all diploid. An-
oplobatus is closely related to the predominately
diploid /daeobatus. The sole triploid report for R.
deliciosus by Longley (1924) was no doubt an ab-
errant individual or possibly an interspecific hy-
brid.
SUBGENUS IDAEOBATUS
Counts have been reported for 70 (52%) of the
estimated 135 species in this subgenus. Fifty-five
of these are reported only as diploids; nine of them
are reported as both diploid and other ploidy levels;
four are reported as tetraploid only; one as ca. 13x;
and one as 18x. The nine species that are reported
as diploid plus other counts are probably basically
diploid, with the other counts, triploid or tetraploid,
merely cytological aberrant individuals or colonies.
The single triploid count for R. strigosus, which is
a well-established diploid species, as well as the
sole triploid counts for the diploid species R. par-
vifolius and R. yoshinoi, represent aberrant individ-
uals which arose from union of a reduced and an
unreduced gamete (Naruhashi & Iwatsubo, 1993).
Spontaneous chromosome doubling in zygotes or in
somatic tissues to form tetraploid individuals is also
not uncommon. The tetraploid counts for the basi-
cally diploid species R. foliolosus, R. hypargyrus,
and R. niveus are most likely aberrant individuals
of this type. The chromosome numbers above dip-
loid reported for the South African species R. ape-
talus (2x and 4x), R. longipedicellatus (2x, 4x, and
5x), and R. pinnatus (2x and 4x) have been ex-
plained by Spies and DuPlessis (1985) as having
arisen through interspecific hybridization and intro-
gression, accompanied by doubling of chromo-
somes. The primary basic number for these species
is considered to be diploid. Altogether, it appears
that 63 (90%) of the 70 Idaeobatus species counted
are basic diploid species.
The four species for which only tetraploidy has
been reported include R. leucocarpus, R. nishimu-
ranus, R. probus, and R. sachalinensis. These spe-
cies arose recently enough that their progenitors
have, with a certain degree of certainty, been as-
certained or can be postulated. With no knowledge
of the chromosome number, Focke (1910-1914)
considered R. leucocarpus to be a subspecies of R.
niveus Thunb. and mentioned that it differs from
the typical species in its larger flowers and fruits
and its more robust plant, traits that are character-
istic of autotetraploids. Thus, it is probable that this
species arose directly from R. niveus. The suspect-
ed hybrid origin of the tetraploid R. nishimuranus
has been confirmed by Naruhashi (1976) and Na-
ruhashi and Iwatsubo (1993), who concluded that
this species is an allopolyploid derived from a cross
of R. trifidus X R. hirsutus. The third tetraploid
Idaeobatus species, R. probus, was recently report-
ed (as R. muelleri) by Thompson (1995a), who made
counts on three plants from Australia; two plants
from one source and one plant from another. Be-
cause this species is so similar to R. fraxinifolius,
Australian botanists have disagreed as to whether
it should be included in that species or considered
a separate species. The tetraploid count of R. pro-
bus provides another trait, in addition to minor тог
phological traits, that may help to distinguish these
two species; that is, if R. fraxinifolius is diploid, as
suspected. It is not possible to claim, although in-
—— ——p——mr
Volume 84, Number 1
1997
Thompson 131
Chromosome Numbers in Rubus
teresting to speculate, that R. probus may be an
autopolyploid derivative of R. fraxinifolius. The
fourth tetraploid /daeobatus species, R. sachalinen-
sis, which is widely distributed in northeast Asia,
was demonstrated by Rozanova (1939) to have aris-
en through autopolyploidy from R. idaeus. In north-
east Asia, К. idaeus varieties melanolasius and
sachalinensis (both now included in R. sachalinen-
sis) are tetraploid. However, the forms to which
these varietal names have been applied in north-
western North America (included here as synonyms
of R. strigosus) are diploid. Although not given in
Table 1, several polyploid raspberry cultivars (R.
idaeus) have arisen spontaneously, e.g., the triploid
‘November Abundance’, ‘Belle de Fontenay’, and
‘All Summer’ (Darrow, 1937) and the tetraploid *Co-
lossus’, ‘Hailshamberry’, and ‘LaFrance’ (Jennings,
1988)
e most puzzling reports for this almost uni-
versally diploid subgenus are the very high ploidy
counts reported for two Rubus sp. in the mountains
of New Guinea by Borgmann (1964). Fortunately,
Borgmann (1964) cited voucher specimens for
these unknown Rubus species and C. Kalkman, of
the Rijksherbarium in Leiden, the Netherlands,
very kindly provided me with the identification of
these as well as Borgmann’s (1964) other unknown
species with chromosome counts (C. Kalkman,
pers. comm.). The /daeobatus species are R. archi-
boldianus (2n = ca. 91) and R. lorentzianus (2n =
126). Both of these counts are highly suspect. Zan-
dee and Kalkman (1981) mentioned only nine
Idaeobatus species in New Guinea. Based on their
relationships, six of these would be expected to be
diploid; R. fraxinifolius and R. chrysogaeus are re-
lated to R. rosifolius (2x); and R. ferdinandi-muel-
leri, R. papuanus, and R. montis-wilhelmi (2x) are
closely related enough to be considered a species
complex. The chromosome number for the ninth
species, R. macgregorii, has not been reported.
These extraordinarily high chromosome numbers
definitely need confirmation, as well as counts
made for the other species in New Guinea, before
an attempt can be made to hypothesize how two
species with such high ploidy levels of 13x and 18x
could have evolved from the surrounding species
that are most likely diploids. It is possible that cy-
tological preparations of these Rubus sp. may have
been mislabeled. If the count really was of a Rubus
species, this high number may belong to an un-
known Malachobatus species because polyploidy is
common in this subgenus, with levels reaching 14x
98
(2n =
SUBGENUS MICRANTHOBATUS
Of about 12 species in this subgenus, chromo-
some numbers have been counted for 5: R. cissoi-
des, R. parvus, R. schmidelioides, and R. squarrosus
from New Zealand and R. royenii var. hispidus from
New Guinea are all tetraploid.
SUBGENUS RuBUS (EUROPEAN BLACKBERRIES)
The major challenge in this subgenus is not cy-
tological but rather taxonomical. For over 200
years, and continuing today, numerous taxonomists
have described and named a few thousand species
in this very complex group. The massive array of
variation that exists results from the reproductive
processes characteristic of this group. Except for
the few sexual diploid species, most species are
facultative apomicts that can freely hybridize, even
with rather distantly related species. In addition to
sexual reproduction, segregating offspring may
arise through a subsexual process involving normal
meiosis in the embryo sac, followed by fusion of
two of the resultant haploid cells. However, the pre-
dominate mode of reproduction is pseudogamy. Al-
though pollination is necessary to initiate devel-
opment, the embryo is strictly maternal, which
provides a mechanism for wide distribution of a
uniform genotype. In this subgenus, there is an
equilibrium system in succeeding generations with
alternating apomictic and sexual reproduction,
which provides for both segregation and rapid, ex-
tensive dispersal of a single genotype. Depending
upon its adaptability, this “clone” may occur as a
local variant in a limited area only or it may be-
come very widespread, thus simulating a true spe-
cies. Dispersal is accelerated further by the uni-
versal propensity for vegetative propagation. These
processes have combined to cause a blurring of
conventional species boundaries, which accounts
or the numerous nomenclatural discrepancies
among taxonomists and the thousands of names that
have been proposed and subsequently discarded.
Although there are some species that are widely
distributed and morphologically distinct enough to
be recognized by all botanists, there are many more
forms where boundaries are not so clear-cut. It is
unlikely that an exact number of species will ever
be universally accepted.
Plants with aberrant chromosome numbers may
arise through parthenogenesis to give haploid
lants, by meiotic disturbances that result in aneu-
ploids, from unreduced gametes that give rise to
offspring with increased chromosome numbers, or
by spontaneous doubling of chromosomes in so-
matic cells.
132
Annals of the
Missouri Botanical Garden
Many of the chromosome counts presented in Ta-
ble | were made many years ago when a different
classification and nomenclatural system was in
vogue. Heslop-Harrison’s (1953) major cytological
study of this group relied upon the species desig-
nations of W. C. R. Watson, the authority for British
Rubus taxonomy at that time (Watson, 1958). Sub-
sequently, Edees and Newton (1988) reassessed
this genus in Britain and have discarded, for vari-
ous reasons, many species names applied by Wat-
son. Therefore, many of Heslop-Harrison’s chro-
mosome counts were made on taxa whose identity
is now questionable. Gustafsson (1943) summarized
his and others’ counts for European blackberries
and, although some of his Scandinavian species
have been revised, in several cases the current syn-
onymy is known so the counts can be presented for
validly named species. In other cases, names were
applied to local forms without proper documenta-
tion; these have been relegated to uncertain identity
or invalid species names. Principal authorities used
to establish current nomenclature in the European
blackberries were Weber (1972), Weber (1981), H.
E. Weber (pers. comm.), and Edees and Newton
(1988), although I take full responsibility for all
errors in interpretation.
Chromosome numbers are reported here for 194
species of European blackberries with my concept
of currently accepted nomenclature (while recog-
nizing that there may not be general agreement
among taxonomists about these names). Since the
cytological situation differs somewhat in the two
major sections of this subgenus in Europe, they will
be treated separately.
In section Rubus, chromosome counts of 154
species are summarized. Only three (2%) basic dip-
loid species have been documented; R. canescens,
R. ulmifolius, and its close relative, R. sanctus.
Six (4%) species are reported to be triploid; R.
divaricatus (as R. nitidus complex), R. grabowskii
(as R. thrysoideus complex) and its close relatives,
R. elatior and R. montanus, are all well established
as triploid species (Gustafsson, 1943). The sole
count of tetraploid for R. divaricatus is either an
error or an aberrant individual. Rubus brevistami-
nosus and R. hylophilus, with only one triploid
count each, need confirmation.
With 125 species counted only as tetraploid, this
is clearly the predominate ploidy level in this sec-
tion. Intraspecific chromosome number variation is
reported for 14 species, all of which have at least
one tetraploid count. The one diploid count in R.
infestisepalus was interpreted by Heslop-Harrison
(1953) to be a haploid variant in this tetraploid
species. The triploid count for R. fissus, reported by
Beijerinck (1956), was made on a single plant and
contrasts with the other tetraploid counts for this
species. Rubus leucostachys, with one triploid count
and one tetraploid, remains to be clarified. Six spe-
cies are given as both tetraploid and pentaploid; R.
drejeri, R. formidabilis, R. hartmanii, R. lentigino-
sus, R. milesii, and R. pedemontanus. Except for R.
pedemontanus, the chromosome numbers of these
species need to be reappraised. Although Gustafs-
son (1943) considered R. pedemontantus (as R. bel-
lardii) to be a uniform, widespread, and well-estab-
lished pentaploid species and dismissed the
tetraploid count in Maude (1939) as an error, two
additional tetraploid counts have been subsequent-
ly reported. Czapik (1987) specifically mentions
two chromosome races for R. bellardii in Europe.
Also, it may be that the two different counts given
for R. pedemontanus can be accounted for by the
fact that R. bellardii is synonymous only in part
with R. pedemontanus and that 1 have erroneously
synonymized all R. bellardii reports. Four species
are reported as both tetraploid and hexaploid: R.
infestus, R. newbridgensis, R. nitidiformis, and R.
pyramidalis var. parvifolius. Heslop-Harrison
(1953) explained that the hexaploid count for K.
pyramidalis var. parvifolius was clearly an aberrant
seedling, probably resulting from an unreduced ga-
mete, because determinations from the adult plant
were all tetraploid. It is possible that a similar ex-
planation may account for the single hexaploid
counts for R. infestus, R. newbridgensis, and R. ni-
tidiformis. In the absence of any well-established
hexaploid species in this section, it is most likely
that the primary number for all species where these
two different counts are given is basically tetraploid
and that other counts are due to occasional aberrant
plants or to errors in identification. Additional
counts are necessary to verify this supposition. Al-
together, if species with both tetraploid and other
counts (except for the triploid R. divaricatus) are
added to those with only tetraploid counts, 139 of
the 154 (90%) of the species in this section are
tetraploid.
Five (3%) of the species are reported to be pen-
taploid: R. anglocandicans, R. marshallii, R. kol-
lundicola, R. pedemontanus, and R. vestervicensts.
There are only single counts for the first two species
and, in fact, Heslop-Harrison (1953) suspected that
the single plant of R. marshallii counted was ac-
tually a large form, not typical of the species. Thus,
these two species need confirmation. The other
three have been well documented by Gustafsson
(1943) as pentaploid. Of these pentaploid species:
only R. pedemontanus is widely distributed in Eu-
E
Е.
Volume 84, Number 1
Thompson 133
Chromosome Numbers in Rubus
rope; the others are local endemic forms in Britain
and in Sweden.
The only hexaploid count for a species in the
section Rubus is a single count for R. bloxamianus
in Maude (1939) which, without confirmation, may
be considered an aberrant individual or an error.
Both Gustafsson (1943) and Heslop-Harrison
(1953) concluded that, except for a possible aber-
rant individual, hexaploid species are not found in
this section. This conclusion has been substanti-
ated in subsequent studies.
In section Corylifolii, as in section Rubus, tetra-
ploidy is the predominate chromosome number.
However, this section differs cytologically in sev-
eral respects: there are no diploid species, there
are relatively more pentaploid species, there are a
few hexaploid species, a few species have two cy-
totypes, and there are a few aneuploids.
The lack of diploids is to be expected since Lid-
forss (1914) postulated that species in this section
were primary and secondary hybrids and segrega-
tion products of crosses between the tetraploid R.
caesius, a species currently placed in its own,
monotypic section, Caesii, and species in section
Rubus. The Corylifolii species are not as numerous
nor as widely distributed as species in section Ru-
bus and species boundaries are even less distinct,
as evidenced by the relatively large number of va-
rieties described for some species (Gustafsson,
1939).
Of the 38 species counted, 17 (45%) are given
as tetraploid only and 8 (21%) as pentaploid only.
Four (10%) are well documented as hexaploid only.
The fact that the hexaploid species are few in num-
ber and have rather limited distribution suggests
that this ploidy level is not well developed. Gus-
tafsson (1939) presented evidence that in nine
(24%) of the Corylifolii species, populations do oc-
cur with different chromosome numbers. His results
are creditable not only because of his knowledge
of the species identity, but also because he made
several counts on samples from different popula-
tions. Rubus camptostachys was 4x, 6x and aneu-
ploid (+ 30); R. dissimulans varieties were 4x, 6x
and aneuploid (+ 42); R. eluxatus was 4x and
aneuploid (+ 28 and 6x + 3); R. fasiculatus was
Ах, 5x, бх, and aneuploid (4х + 1); R. gothicus was
Ах, 5х, and + 28; R. lidforssii was 4x and 5x; К.
lindbloomii was 4x and 6x; R. nemorosus was 4x
and 5x; and R. norvegicus was 4x, 5x, and 6x. The
counts of both tetraploid and hexaploid reported for
R. conjungens by Heslop-Harrison (1953) were not
explained as representing different populations, so
it is not clear if the hexaploid count reflects a dif-
ferent cytotype or merely an aberrant individual.
Counts for 75 "species" in both sections Rubus
and Corylifolii classified as *doubtful determination
and/or taxonomic status" are presented here, but
separately because, regardless of their true identity,
these data add to the cytological picture of the
group as a whole in Europe. These include many
counts for British plants to which valid continental
species' names had been erroneously applied, local
forms that had originally been given species rank
but are no longer accepted as such, aggregate spe-
cies such as R. hirtus, as well as forms to which
names were applied but never documented.
SuBGENUS RUBUS (NORTH AMERICAN
BLACKBERRIES)
Chromosome numbers in the North American
blackberries were studied most intensively in the
East by Longley (1924) and Einset (1947), and in
the West by Fischer et al. (1941), Darrow and Lon-
gley (1933), Brown (1943), and Zielinski and Galey
(1951). In comparing the blackberries of eastern
North America with those of Europe, some similar-
ities and differences may be mentioned. One sim-
ilarity is the important evolutionary role of inter-
specific hybridization and facultative apomixis so
that species boundaries are difficult, if not impos-
sible, to define accurately. Another commonality is
the significance of polyploidy in speciation, al-
though the eastern North American group repre-
sents an even wider range of ploidy levels, includ-
ing 2x, 3x, 4x, 5x, 6x, 7x, 8x, 9x and even one
aneuploid (5x + 1) species. Also similar is the
common association of facultative apomixis (pseu-
dogamy) with polyploidy (Einset, 1951). One sig-
nificant difference is the existence of several, ap-
parently sexual, diploid species compared to only
three diploids among the several hundred species
of blackberries in Europe. Also, whereas tetraploid
is the dominant ploidy level in the European black-
berries, this number appears to be relatively less
common in the North American species. From the
limited counts available for eastern North American
species, it appears that, except for the section Fla-
gellares where ploidy levels higher than tetraploid
are common, the most frequent numbers reported
are diploid and triploid.
The most recent attempts to clarify the taxonomy
of the eastern North American blackberries were
those of Davis et al. (1967-1970), updated by Da-
vis (1990), and Hogdon and Steele (1968). Species
listed in Table 1 follow the Davis et al. (1967—
1970) treatment (with modifications suggested by
Mark Widrlechner, pers. comm.) because it in-
cludes all of the eastern North American species,
Annals of the
Missouri Botanical Garden
whereas Hogdon and Steele (1968) dealt only with
those of the New England states. It is suspected
that the widely varying chromosome numbers re-
ported for some species can be partly attributed to
misidentification of the plants. However, that vary-
ing chromosome numbers may exist in wild popu-
lations is demonstrated by Einset's (1951) compre-
hensive study of chromosome numbers of offspring
following self-pollination, controlled cross-pollina-
tions, and open pollinations of plants with estab-
lished numbers. An individual polyploid plant (3x,
4x, 5x, 5x + 1, 7x, or 9x) produced not only apo-
mictic offspring resembling the maternal plant in
morphology and chromosome number but also par-
thenogenetic haploids and true hybrids with vary-
ing chromosome numbers as a result of different
combinations of reduced and unreduced gametes.
The extent to which these deviant chromosome
number forms survive and spread in nature de-
pends upon their level of apomixis and/or success
of vegetative propagation, their adaptability, and
their ability to compete as compared to the species
norm from which they were derived.
Chromosome counts have been reported for 45
(22%) of the approximately 200 species of eastern
orth American blackberries. However, it is clear
from Table 1 that the information is scanty, with
only a single count for most species, and that there
is a very inconsistent pattern in the numbers re-
ported for some species. More cytological infor-
mation is necessary to determine how much valid
intraspecific variation in chromosome number ac-
tually does exist. Unfortunately, there have been no
significant cytological studies to expand and cor-
roborate the early work of Longley (1924), many of
whose plants have suspect identity. The following
is an attempt to summarize the limited cytological
information for each section.
In section Alleghenienses, chromosome counts
are 2x, 3x, and 4x. Rubus allegheniensis appears to
be a well-defined diploid, sexual species. Aalders
and Hall’s (1966) diploid count for R. allegheniensis
represented 22 different clones in southwest Nova
cotia. These authors counted chromosomes and
studied morphological traits of 470 blackberry
plants collected throughout Nova Scotia. They con-
cluded that there were two distinct diploid species,
R. allegheniensis and R. hispidus, and that 355 of
the plants studied were triploid hybrids and the
remaining 52 were 4x, 5x, or 6x, probably derived
from further hybridizations and unreduced gametes.
The polyploids all appeared to be facultatively apo-
mictic, which would account for the perpetuation
of so many triploid clones. Based on morphological
criteria, these triploids were interpreted as having
been derived from hybridization of the two basic
species, with one of them contributing an unre-
duced gamete. This study demonstrates the cyto-
taxonomical complexity facing a botanist attempt-
ing to classify the blackberries into discrete
species. The two triploid counts for R. alleghenien-
sis by Longley (1924) and Einset (1947) most prob-
ably were from aberrant plants arising from an un-
reduced gamete. It is interesting to note that,
although R. allegheniensis is considered to be a
good diploid species, several allegheniensis-type
tetraploid cultivars (e.g., ‘Lawton’, ‘Ancient Briton’,
‘Snyder’, ‘Taylor’, and ‘Eldorado’) have been se-
lected directly from wild populations, although
some have been speculated to be interspecific hy-
brids. In fact, the lineage of the modern eastern
upright blackberry cultivars, all tetraploids, traces
back to these early wild selections. Obviously, tet-
raploid forms do appear in nature and, at least
some of them, are superior enough in horticultural
traits to have been selected for human usage. Be-
cause they do not appear to have become a signif-
icant part of the population, the question arises
about their relative adaptablility in nature as com-
pared to the diploids.
In section Arguti, 2x, 3x, 4x, 5x, 5x + 1, and 6x
counts have been reported for various species. I
will make no attempt to summarize these because
I suspect that the erratic cytological picture reflects
misidentified plants in too many cases. It is prob-
able that R. argutus is a diploid species, with an
occasional triploid aberrant.
The section Canadenses is represented here by
R. canadensis. Craig (1960) reported triploidy in 13
individual clones collected in seven regions of New
England, New Brunswick, and Nova Scotia. With
three other authors’ triploid counts and Einset's
(1951) demonstration of apomixis in triploid R.
canadensis, it appears that this cytotype prevails in
a significant part of the range of this species. The
three reports of diploid forms indicate that there
are also diploid populations.
In section Cuneifolii, there are counts of 2x, 3%
and 4x. Rubus cuneifolius is a diploid species.
Based on meiotic pairing relationships, Spies and
DuPlessis (1985) interpreted their individual trip-
loid and tetraploid plants as autopolyploids of the
diploid species. These authors made the same con-
clusion about the triploid and tetraploid R. pascuus
plants although, in this case, no diploid species was
observed.
Section Flagellares is the only group in eastern
North America in which most species have been
consistently reported to be polyploid. I suspect that
Longley's (1924) diploid counts for R. recurvicaulis
Volume 84, Number 1
1997
Thompson 135
Chromosome Numbers in Rubus
and triploid for R. biformispinus and R. multiformis
were based on misidentified plants. Because many
of Longley’s (1924) chromosome counts are incon-
sistent with those of others, the identifications of
his plants are highly suspect. The remainder of the
counts for this section are 4x, 5x, 6x, 7x, 8x, and
9x, which, if these plants were correctly identified,
is strongly suggestive of a polyploid complex sim-
ilar to that of R. ursinus in western North America.
In the section Hispidi, 2x, 3x, 4x, 5x, and 8x
counts are reported. Aalders and Hall’s (1966) dip-
loid counts for 41 clones of R. hispidus collected
widely in southwestern Nova Scotia, along with
Thompson’s (1995a) diploid count from a North
Carolina plant, suggest that this is a widespread
basic diploid species. It is most likely that Lon-
gley’s (1924) pentaploid and octoploid counts for R.
hispidus were mistakenly identified plants, possibly
belonging to section Flagellares, which has a trail-
ing plant growth habit like the Hispidi but, unlike
Hispidi, is characterized by higher ploidy levels.
There is one count each for R. huttonii (4x), R. plus
(3x), R. signatus (3x), and R. tardatus (3x).
In section Setosi, chromosome numbers reported
are 2x, 3x, 4х, and 5x. Rubus setosus has both dip-
loid and triploid counts, but with such a limited
sampling it is not possible to determine the relative
abundance of each of these ploidy levels. There is
one count each for R. dissensus (3x), R. glandicaulis
(3x), R. clandestinus (4x), R. hanesii (4x), R. miscix
(3x), R. notatus (2x), and R. wisconsinensis (5x).
For section Triviales, the only species counted,
R. trivialis, has been consistently given as diploid
by six investigators. There seems to be no question
about this being a widespread, basic diploid spe-
cies.
Although limited cytological information is avail-
able for the eastern North American blackberries,
a few tentative conclusions are warranted. It ap-
pears that, in each section, there is at least one
easily recognized species, i.e., the one after which
the section is named. Seven of these, R. alleghe-
niensis, R. argutus, R. canadensis, R. cuneifolius, R.
hispidus, R. setosus, and R. trivialis, are basically
diploid but may have triploid or tetraploid individ-
uals or populations as well. By contrast, the eighth
one, R. flagellaris, is polyploid, with the predomi-
nate chromosome number unknown due to discrep-
ant counts and insufficient sampling. Because of
the discrepancies in chromosome numbers, even
within species, counts need to be made on popu-
lation samples rather than individual plants. Such
counts made in conjunction with a taxonomic re-
appraisal would greatly contribute to the clarifica-
tion of the blackberry species of eastern North
America.
The section Ursini, represented only by R. ursi-
nus, is geographically isolated from all other North
American blackberries. It occurs only in the west-
ern parts of the Pacific States from southern Cali-
fornia to southern British Columbia and in western
Idaho. Brown (1943) made an intensive cytotaxo-
nomic study of this species in the southern half of
its range and concluded that there were two main
chromosome numbers: octoploidy was dominant in
most of California, whereas 12x was dominant from
northern California to southern Oregon, and pre-
sumably northward. The limited numbers of plants
found with 9x, 10x, or 11x, were assumed to have
been derived from the two main chromosome types
through hybridization. The relatively few odd-
ploids found suggested that apomixis is not an im-
portant mode of reproduction in this species. The
lack of consistent morphological traits associated
with chromsome number led Brown (1943) to con-
clude that this western blackberry population con-
sists of only one species with six different ploidy
levels.
SUBGENUS RUBUS (SOUTH AMERICAN
BLACKBERRIES)
Of Focke's (1910-1914) four sections of Rubus
in South and Central America, 1.е., Dissitiflori, Xer-
ocarpi, Duri, and Floribundi, chromosome counts
have been reported for species only in the Flori-
bundi. Thus far, the information is very limited,
with only one diploid count each for R. adenotri-
chos, R. robustus, and R. urticifolius, and two for R.
bogotensis. Should further studies confirm the lack
of polyploidy in these blackberries, this cytological
situation is a striking contrast to that of this sub-
genus in North America and Europe.
PROPOSED NATURAL INTER-SUBGENERIC HYBRID
Tetraploid R. glaucus, both cultivated and in the
wild, is widespread in the northern Andean coun-
tries and in Central America. Darrow (1952) first
suggested that this species was an allopolyploid
originating from hybridization between a black
raspberry and a South American blackberry. Jen-
nings's (1978) studies on anthocyanin pigments in
R. glaucus and its putative parents provided sup-
port for this concept.
In conclusion, this compilation provides scien-
tists working with this genus convenient access to
the chromosome numbers of species published to
date. The current state of knowledge is presented,
136
Annals of the
Missouri Botanical Garden
and the gaps that need further investigation are
made evident. Although counts have been made on
an estimated 40 percent of Rubus species, many of
these are single counts that need confirmation, and
the chromosome numbers of the remaining 60 per-
cent of the species need to be determined. It is
hoped that this paper will stimulate an interest in
additional cytological and taxonomical studies that
will contribute to the elucidation of the nature of
the species in this large and complex genus.
Literature Cited
Aalders, L. E. & I. V. Hall. 1966. A cytotaxonomic survey
of the native blackberries of Nova Scotia. Canad. J.
enet. Cytol. 8: 528-532.
Allander, H. 1941. Om den Svenska S.K. Rubus nemo-
rosus. Svensk Bot. Tidskr. 35: 287-295.
Bailey, L. H. 1941. Species batorum. Dg or Rubus
in North America. I. Gentes Herb. 5:
Bammi, R. K. 1965a. ‘Complement AN ina
natural hybrid between Rubus procerus Muell. and R.
laciniatus Willd. Nature 208: 608.
об. Cytogenetics of Rubus. ТУ. 1 achytene
morphology of Rubus parvifolius L. chromosome com
plement. Canad. a hee Cytol. 7: 254—258.
& H. ‚шад Bà seis of Rubus. V.
pn Muell.
Natural жм inet betw
Cha mosome
some flowering plants of West Кана. II.
Cod: ег 22: 41-51.
eek, А. van де. 1981. Batologische notities 2. Nieuwe
gegevens over e us. Gorteria 10: 147—150.
Beijerinck, 56. Rubi Neerlandici. Verh. Kon. Ned.
kad. татаас Afd. Natuurk., Tweede Reeks 51: 1–
Belyaeva, V. A. & V. N. Siplivinskiy. 1975. Khromosom-
ак chisla i taksonomiya nekotorykh vidov Baykalskoy
ry. Bot. Zurn. SSSR 60: 864-872.
. 1981. Chromosome number reports
LXXIII. Taxon 30: 859
Berger, X. 1953. Untersuchungen über die Embryologie
partiell Weser бара Rubusbastarde. Ber. Schweiz.
Bot. Ges. 63
Beuzenberg, E
& J. B. Hair. 1983. Contributions to a
1994. omosome numbers in
и ambles (Rubus L., Rosaceae). II. Arbor. Kórnickie 39:
87.6):
———. 1995. Chromosome numbers of Polish brambles
(Rubus, Rosaceae). Willdenowia 25: 267-271.
Borgmann, E. 1964. Anteil der Polyploiden in der Flora
des Bismarckgebirges von Ostneuguinea. Z. Bot. 52:
8-172.
Britton, D. M. & J. W. Hull. 1957. Ch
of Rubus. dcr ^ Hort. Digest 11: 58—60.
Brown, S. W. . The origin and nature of variability
in the рее Can blackberries (Rubus ursinus Cham.
& Schlecht. and R. lemurum sp. nov.). Amer. J. Bot. 30:
h
686—697.
Сһотіѕигу, М. 1927. Beitrag zur Keimfahigkeit und Zyt-
ologie des Pollens einiger Prunus—und Rubussorten.
hia Bot. 9: 6 6.
Christen, H. R. 1950. Untersuchungen über die Embryol-
ogie pseudogamer und sexueller Rubusarten. Ber.
Schweiz. Bot. Ges. 60: 153-198.
Craig, D. L. 1960. Studies on the UN and the pais
ing virt of Rubus canadensis L. Canad. J. Gene
Cytol. 2
Crane, M. » 1935. Blackberries, hybrid berries, and au-
tumn-fruiting uvis p. 121-128 in Cherries and
Soft Fruits Conf.,
C
Czapik, R. 1987. foede in vat bellardi Weihe. Acta
Biol. Cracov., Ser. Bot.
Dale, A. & R. Ingram. 1981. RN numbers of
some Tub eei blackberries. Hort. Res. (Edin-
burgh) 21:
Darrow, С. ^s a Blackberry EM improve-
ment. Pp. apes in Yearb. Agri
t Rubus glaucus. The fei blackberry
“A Central em northern South America. Ceiba 3: 97-
A. E. Longley. 1933. Cytology and breeding
of Rubus macropetalus, the Logan, and related black-
315-330.
me chromosome nu rs in British
species of Rubus. Mem. & Proc. Manchester Lit. Soc.
Datis, Н; ru 1990. Studies in Rubus. Castanea 55: 22-
30
Fuller & T. Davis. 1967-1970. Contri-
butions toward the revision of the Eubati of eastern
North America. Castanea 32: 20-37; 33: 50-76, 206-
240; 34: 157-179, У 35: 176-194.
Dmitrieva, $. А. & У. I. Parfenov. 1985. Karyyalagich-
naya kharaktarystyka пева vidov karysnykh ras-
lin flory Belarusi. liv, а Nauk Belorussk. SSR. Ser.
Biol. Nauk (Baku) 6:
Едеез, Е. 5. & A. ат 1988. Brambles of the British
Isles. The d Society, London
Einset, J. 1947. Chromosome stüdies in Rubus. Gentes
Herb. 7: v 192.
1951. Apomixis in American polyploid black-
bemes Amer. J. Bot. 38: 768-772.
—— & C. Pratt. 1954. Hybrids between blackberries
and ла raspberries. Proc. Amer. Soc. Hort. Sci. 63:
Engels, T. 1979. Chromosome numbers of vascular
n Norway, including Svalbard. Opera Bot. 52:
Faasen, P. van & P. Nadeau. 1976. IOPB chromosome
mber reports LI. Taxon 25: 157.
Floris A. (Editor). 1969. Chromosome Numbers of
lowering Plants. Komarov Bot. Inst, Acad. Sci.
U.S.S.R., St. Peters
Fischer, H. E., G. M. Darrow & G. F. Waldo. 1941.
ther chromosome studies of some varieties of bladi bet
ies. Proc. Amer. Soc. Hort. Sci.
Focke, W. O. 1910-1914. Species Ruborum. Mono.
phiae generis Rubi Prodromus. Biblioth. Bot. 17 (72
part I): 1-120 (1910); 17 (72, Part II): 121-223 (1911);
19 (83): 224-488 (1914). Stuttgart.
Gill, B. S., 5. 5. Ви, M. 5. Sidhu E V. K. Singhal. 1984.
IOPB chromosome number reports LXXXIV. Taxon 33:
Gilli, A. 1969. Die Zytologie von Rubus gemi Focke
und Rubus candicans Wh. Oesterr. Bot. Z.
432.
Volume 84, Number 1
1997
Thompson 137
Chromosome Numbers in Rubus
cow À. 1933. Chromosomenzahlen in der Gattung
us. Hereditas 18: 77—80.
1939. Differential "oid pain within the black-
héros. gee 25:3
The genesis ems -— ii n blackberry
flora. aa Тек Lund. 39: 1
Haskell, G. 1960. Genetics. Pp. a in Annual Repo
Scott. Hort. Res. Inst. for 1959-1960, мека А,
Scotland.
. B. Paterson. 1966. Chromosome number of
a sibs Antarctic и Машге 211: 759.
N. 961. Developmental sequence of
chromosome Pob | in a cytologically unstable Rubus
hybrid. Genet. Res. 2 24.
Heslop-Harrison, Y. 1953. Cytological gerent in the ge-
nus Rubus L. 1. Chromosome numbers in the British
ele. ubus subgenus Eu-
batus in New England. A conspectus. Rhodora 68: 474—
13.
Hsu, C. C. 1968. Prelimin on the
vascular plants of Taiwan (II). Taiwania 14: mes
— Mn — e, Y. Araki & N. Naru-
hashi. hromosome иий е of Rubus Xpseudo-
sieboldi Rosaceae) = Jap. Bot. 67: 10-14.
ashi. 1991. Karyomorphological and
—1 studies of Rubus parvifolius, R. coreanus,
R. Xhiras — А Педро Cytologia 56: 151-156.
. Cytotaxonomical studies of Ru-
bus (Rosaceae) L y sacar numbers of 20 species
and 2 natural еме J. Јар. Bot. 67: 270-275.
& ytotaxonomical studies of Ru-
bus (Rosaceae). T. ¡cri numbers of 21 species
and 6 natural — J. Jap. Bot. 68: 159-165.
Weber. 1995. Chromosome
mbers of Putas vibus (Rubus subg. Rubus,
подом Pl. Syst. Evol. 198: 143-149.
Jennings, D. L. 1978. The blackberries of South Ameri-
— Ап unexplored reservoir of germplasm. Fruit Var.
J. 32: 61-63
1988. Raspberries and Blackberries. Their
Breeding, Diseases and Growth. Academic Press, Lon-
don.
Jinno, = 1951a. Chromosome numbers in Rubus. I. Кго-
mosomo 9-10: 360-361. [In Japanese, with English
su nies
. 1951b. The chromosomes in Rubus. ЇЇ. Jap. J.
Genet. 26: 133-135. [In Japanese, with English sum-
. 1958. Cytogenetic and cytoecological studies on
us 1. Chromosomes. Bot.
. & J. С. Packer. 1968. Chromosome num-
bers in the flora of Suniel Creek, N. W. Alaska. Bot.
Not. 121: PE
Jorgensen, C. A., Th. Sprenson & M. Westergaard. 1958
The flowering plants of cda A taxonomical and
cytological survey. Biol. Skr. 9: 1-172.
Kalkman, . The genus Rubus (Rosaceae) in Ma-
lesia. 2. The subgenus Malachobatus. Blumea 29: 319—
86.
. The genus Rubus pring in Malesia.
3. The e Micranthobatus. Blumea 3 јр“
Keep, E. 1958. Cytological notes. Pp. 26-28 n Annual
Report E. Malling Research Station for 1957, East
Malling, Engl
Kotimaki, M. & H. Hiirsalmi. 1979. Cytogenetic studies
n Rubus arcticus, Rubus stellatus and their hybrids.
Hereditas 91: 83-89,
Krogulevich, R. E. 1976. Rol poliploidii v genezise "id
Putorana. Flora Putorana. Nauka, Siberian Dept.,
vosibirsk.
978. Kariologicheskiy analiz vidov flory Vos-
tochnogo Sajana. V. Flora Pribajkal'ja. Nauka, Siberian
Dept., Novosibirs
& ¿ Rostovtseva (Editors). 1984. Khromosom-
nye Chisla Tsvetkovykh astenii i Sibiri Dal’nego Vos-
k.
toka. Nauka, Sibirian Dept., Novosibirs
Laane, M. M. 1969. Videre kromosomstudier i norske
karplanter. dog
Larsson, G
. Tet traploid Rubus arcticus, framställd
10: 151-159.
for breeding in n northe ern Rubi.
Lidforss, B. . Resumé seiner Arbeiten tiber Rubus.
Z. Indukt. Abstammungs-Vererbungsl. 12: 1—13.
Longley, A. E. 1924. Cytological studies in the genus
Rubus. Amer. J. Bot. 11: 249-282.
G. M. Darrow. 1924. Cytological studies of
diploid == "n em forms in raspberries. J. Agric.
Res.
Lóve, A. pony a evaluation of corre-
sponding taxa. Vegetatio 5-6: 212-224.
—— ——. 1987. Chromosome Reports XCVI. Taxon 36:
D. Lóve. e Aedes ir studies on
на plants. III. Some new chromosome numbers of
berries plants. Ark. d 31A (12): 1-22.
—— 1966. Cytotaxonomy of the alpine
aines plants of Mount Washington. Univ. Colorado
Press, Ser. Biol. 24: 1-74
& . 1982a. IOPB chromosome number
reports LXXV. Taxon 31:
&
1982b. IOPB сећање number
reports LXXVII. ‘Taxon 31: 7
—— & J. tchie. 1966. Chromosome numbers
from central iih Canada. Canad. ay Bot. 44: 429—
439.
Magulaev, A. 1976. “eng > ae tsvetkovykh
Е shch. IL) In € га
i osve she eniy
ology of some Indian species of
Rosaceae. Саба: 18: 139-149.
Malla, 5. В., 5. Bhattarai, М. Gorkhali, Н. Saiju € М.
у ~ IOPB epe number reports
ТУШ. Taxon 26: 561.
: & M. Shrestha. 1974. IOPB chromo-
some number reports. XLVI. Taxon
. Gorkhali & M. P. Singh. 1975. IOPB
chromosom me number reports XLIX. Taxon 24:
Markarian, D. & H. P. Olmo. 1959, Cytogenetics of Ru-
bus. 1. Reproductive behavior of R. procerus Muell. J.
Heredity 50: 131–
Markova, M. L. 1968. Chromosome number of four Bul-
garian taxa of the genus Rubus L. Izv. Bot. Inst. (Sofia)
18: 75-77.
1972. IOPB chromosome number reports
XXXVI. Taxon 21: 339.
Marks, G. E. 1952. Chromosome counts of species and
varieties of garden plants. Pp. 47-50 in Annual Report
John Innes Hort. Inst. for 1951, Norwich, Englan
Maude, P. F. 1939. The Merton catalogue. A list of the
Annals of the
Missouri Botanical Garden
chromosome “arg” s species of British flowering
tol. 38: 1-31.
Н. Dhawan. 1966. Cytological investi-
gations in the N. W. Himalayan Rosaceae. Proc. Indian
i. Congr. Assoc. 53: 276-277.
& 1971. IOPB chromosome number re-
ports XXXIV. Taxon 20: 792.
——, T. S. Sareen & А. S. Hans. 1973. Cytology of
some woody species of Rosaceae from the Himalayas.
Silvae Genet. 22: 188-190.
Мии. N. 1972. Taxonomic Study of the Genus Ки-
n Japan. Dr. Thesis, Kyoto University, Kyoto, Ja-
6. Taxonomical notes on the hybrid between
Rubus oni and К. hirsutus (1) аы. J. Geobot.
24: 26-34.
1989. Notes on Japanese Rubus (4). J. Phyto-
geogr. Taxon. 37: 14.
— & Y. Iwatsubo. 1993. Chromosome number of
Japanese Rubus. Acta Hort. 352: 429-431.
Nybom, H. 1980. Chromosome numbers in Rubus species
from Sri Lanka. Bot. Not. 133: 47-48
Chromosome numbers and reproduction
in Rubus subgen: Malachobatus. Pl. Syst. Evol. 152:
211-218.
Ono, M. pe parar studies on the flowering
plants endemic to the B Islands. Mem. Natl. Sci.
Mus. (Tokyo) 10: 63-80. "fi Japanese, with English
summary.
Packer, J. G. & R. Witkus.
number reports LXXV. Tax :
., R. Ingram, R. J. Abbott D. L. Jennings & P.
1981. Karyotype variation in Rubus with
particular reference to R. idaeus L. and R. coreanus
Мир, лу ар 8 125—132.
Ро, N. S., А. Р. Sokolovskaya 4 2. С. Rud
Chisla kromosom nekotorykh vidov sosudistykh
r, Kurilskie ostrova. Bot. Zurn
uns IOPB chromosome
Rohweder, H. 1937. Versuch zur Erfassung der mengen-
mássigen Bedeckung des Darss und Zingst mit poly-
ploiden Pflanzen. Planta 27: 501—549.
Rostovtseva, T. S. 1977. Chisla khromosom nekotorykh
vidov rasteniy yuga Sibiri. II. Bot. Zurn. SSSR 62:
1034—1042.
Rozanova, M. A. 1934. Puti formoobrazovaniya v rode
Rubus. Bot. Zurn. SSSR 19: 376-384.
. O proiskhozhdeniy sibirskoy maliny v ya-
blenem aviopoliplaiti: Compt. Rend. (Dokl.) Acad. Sci.
S 24: 58-60.
. On genotypic differences between races
of Rubus caesius L. Compt. Rend. (Dokl.) Acad. Sci.
URSS 27: 590—593.
Sarkar, A. K., M. Chakraverty, N. C. Saha, S. K. Das &
D. Hazra. 1977. IOPB chromosome number reports
LVII. Taxon 26: 450.
Scheerer, H. 1939. Chromosomenzahlen aus der Schles-
wig-Holsteinischen Flora. I. Planta 29: 636-642.
Sharma, . Sarkar. 1970. Chromosome number
reports of plan ts. Pu Report, 1967-1968. Res
Bull. Univ. CR A 1-50.
Sharpe, R. H. & J. п. 1958. Development of
temperate-climate fruite for Florida. Proc. Florida State
Hort. Soc. 71: 294—300.
Sherman, W. B. 1968. Blackberry breeding in Florida.
Sunshine State Agric. Res. Rep. 13: 30-32.
Shoemaker, J. S. € T. T. Sturrock. 1959. Chromosome
relations in blackberries. Proc. Florida State Hort. Soc.
72: 327
d V. K., B. S. Gill & M. S. Sidhu. 1990. Cytology
y nion bere of Rosaceae. Proc. Indian Acad. Sci.,
PL "Sei. 100: 17-21.
Skalinska, M., E. Pogan, R. Czapik et al. 1978. Further
studies in ا اد numbers of Polish Angiosperms.
XII. Acta Biol. Cracov., Ser. Bot. 21: 31-63.
Sokolovskaya, A. P., N. 5. Probatova € Z. С. Rudyka.
1985. Chisla khromosom vidov semeyst Asteraceae,
Poaceae, Rosaceae, iz Primorskogo Kraya, Kamchatki i
Sakhalina. Bot. Zurn. SSSR 70: 126-128.
. S. Strelkova. 1960. Geograficheskoye ras-
prostraneniye poliploidnykh vidov rasteniy v Yevraziat-
skoy arktike. Bot. Zurn. SSSR 45: 369—381.
rsa, V. 1963. Chromosomenzahlen Finnischer Kormo-
pue II. Ann. Acad. Sci. Fenn., Ser. A. IV. Biol. 68:
d 5 J. & H nt 1985. The genus Rubus in
South Africa. L Саса воно purohéra and geographical
distribution. — 15: ge
— € Н. Liebenberg. 1985. The genus Ru-
bus in South rue II. Meiotic chromosome behaviour.
Bothalia 15: 597-606.
Subramanian, D. 1987. Cytotaxonomic studies of South
Indian Rosaceae. Cytologia 52: 395—403.
Taylor, R. L. & С. A. Mulligan. 1968. Flora of the Queen
Charlotte Islands. Part 2. Cytological Aspects of the
"РВ ueen’s Printer, O
19 0а. йорисе versatility in Rubus.
The дааа and development. J. Genet. 40:
1940b. The origin of new forms in Rubus. Ш.
The chromosome pautas of R. lopanobaceus Bailey,
its parents and derivatives. J. Genet. 41-156.
Thompson, M. M. 961. Cytogenetics of Rubus. П. Cy-
tological studies of the varieties ‘Young’, ‘Boysen’, and
related forms. Amer. J. Bot. 48: 667-673.
995a. Chromosome numbers of Rubus species
at the National Clonal Germplasm Repository. Hortsci-
ence i 1447-
995b. Chromosome numbers of Rubus cultivars
at the ا Clonal Germplasm Repository. Hortsci-
ence 30: 1453-1456.
. M. Zhao. 1993. Chromosome numbers of
Rubus species in southwest China. Acta Hort. 352:
9
4934
Vaarama, А. 1 939. Cytological studies on some Ё innish
"e and hybrids of the genus Rubus L. J. Sci. Agr.
Soc. (Finland) 11: 72-85.
: . Om artkorsningsfórüdling inom släktet
Rubus. Nord. Jordbruksforskn. 31–32: 412-417.
. 1953. Cytology of hybrids between Pacific Rubi
ursini types and an eastern blackberry. Amer. J. Bot.
565—510.
1954. Chromosome numbers of some —
6
and hybrids of the genus Rubus. Arch. Soc. Zoo
"Е, 1958. Handbook of the Rubi of Great
Britain ar Ireland. Univ. Press, come
1 ки certain Ca-
chen tops: el ero A
1. Revision der Sektion Corpo (Gattung
Rubus, Rosaceae) in Skandinavien und im nórdlich
Volume 84, Number 1
1997
Thompson 139
Chromosome Numbers in Rubus
Mitteleuropa. Sonderb. Naturwiss. Vereins Hamburg 4:
1-229.
1991. ere Fu asa
r Gattun
W. Maurer.
der in Osterreich nachgeweisenen Arte
Rubus L. (Rosaceae). Phyton (Horn) 31: 6
Williams, C. Е, B. W. Smith & 6. M. 8 1949. A
pan-American blackberry hybrid. J. Heredity 40: 261-
65
2 3
Pee 9.6 1992. WR дү behavior as a factor in
reeding. Proc. Amer. Soc. Hort. Sci. 28: 114-117.
. 1936. быш behaviour in blackberry-
raspberry hybrids. J. Agric. Res. 52: 385-396.
Yeager, A. F. & E. M. Meader. 1958. Breeding better
fruits and nuts. New Hampshire Agric. Exp. Sta. Bull.
Yu, T. T (Editor). 1985. Flora Reipublicae Popularis Sin-
icae. Tomus 37; viii, 10—218. Science Press, Beijing.
Yurtsev, B. A. & P. G. Zhukova. 1982. Khromosomnye
chisla nekotorykh rasteniy severo-vostochnoy Yakutii.
Bot. Zurn. SSSR 67: 778-787.
Zandee, M. & C. Kalkman. 1981. The genus Rubus (Ro-
saceae) in Malesia. 1. Subgenera Chamaebatus and
Idaeobatus. Blumea 27: 75-113.
Zhukova, P. G. 1980. Khromosomnye chisla nekotorykh
vidov rasteniy yuzhniy Chukotskiy. Bot. Zurn. SSSR 65:
; 1982. Chisla khromosom nekotorykh vidov ras-
omnye chisla nekot ган vidoy rasteniy vostoka
асо И Yakutii. Bot. Zurn 62: 229-
Zielinski, Q. B
ers of certain trailing blackberry clones. Proc. Am
Soc. Hort. Sci. 57: 163-164.
Annals of the
140
Missouri Botanical Garden
20661 чоѕішоці, ‘Z661 !USPYNIBN 29
Oqnsjem] ‘CGI "үе 19 PÁBASAO[OAOS :8961 URSIT[N 29 3o[4e],
TSE а у rn
2961 ATEN
20661 поѕйшоц, ‘6661 човвјејепо)
22661 човашоцј,
GEGT УРИНА
+861 PAISIAOISOY 29 YOrasnsory 7 161 PAISIAO]ISOY
20661 uosduiou
2861 9191
2861 "еу
0861 vAoxnuz “8961 19xoeq AY uosuqof
20661 uosduiou] 2861 PA0%
-NYZ x APSUNA “1861 “SLOT AP{sutatdis $ eAoeápog “6L61
пшевлин 29 RUNON :8961 194984 9 uosuqof “0961 PA
-Ox[9MS 29 PÁBASAO[OAOS :6961 ‘LSGI 055167] ‘GEGT PUBIBBA
9961 uosiajeq Y [[9xSeH
2661 9947 Y
uosdwoy], :2661 !YSPYMIPN 79 oqnsyem] :8261 #1661 ouutf
22661 повашоцј,
0261 Tees Y иес
с861 лочәред $ PALI 861
BAASJAOISOY 2 цотләүпдогу :2861 eAoxnu7 #2861 9107
2۸91 “6L61 uoD[spodu :8261 ‘TF 19 exsurpexs “6961
ешюлерД eco] иоѕшең-дојѕәң *yp61 249] Y 2107 (6261
apne) moje] ‘££61 Хојаиој $ моме *yz61 4913uo7]
9x90, 8112074) snuodqng
8с
9x90, sisdospwo7y) snuasqng
2490] snypgapupy’) snuesqng
9S
9904 (ин) =п-ошәршту”) snuesqng
"ug snippod y
uoss1e'] йл $121]241)]]91$ ЭС о i| 5121]210 “Y
IQM ‘VM (XY) synova dsqns
уен (T) #217910 5020)
‘xor simpop
"Xqony simpop
9204 ("хцотур) simon *dsqns "T $n21240 "M
"1 $n21240 ^M
"1 $721524D “Y
‘ug s2p1023 “y
"WIXB Y зтргиноод "M
‘yooH хә звј пој sparnu “y
uo(p'(] хә TEM SNUDÁ]DO ^M
"т SNIOWIDUIDYO ^M
зоополојоц
uc
џохвј,
"uoreue[dxe лоцџпј 10у 641 9894 vag 'seroeds sngny jo siequinu ошовошолуо С] 9[qe],
141
Моште 84, Митбег 1
1997
Thompson
Chromosome Numbers in Rubus
0247 xy uosdwoy] ‘£661 OqnsieM] y ruseuniey :2661
е 19 0qnsjem] :7c6[ ЦН Y uonug :8061 *epcóq оишү
£661 oqnsie^[ x {SEMEN
8261 dəy
2661 9942, x uosduiou y,
2661 9847, x uosdwoy y
2661 !useunuey o OQNSIBM] :с7 61 Te 19 PITA
23204 зттодоцоојоју snuadqng
34904 snipdosooq42 “Y
Чапу snijofipnbo "у
"buy uaRıanq y
buy ә әп “y
iopqay (2204) wmapsnquipQ “IBA әдип Y ‘[SuOoH Muay “Y
9904 UNIDSNQUIDA Y
22661 uosdwoy y 9c ттеү-рчең umzoZups] "y
2661
овцу x uosduoq], :£661 !Ҷѕеҷтием $ oqnsm] :8с61 doox 8z 94904 407091 “Y
2661 99427, x uosdwoy y су soi хә 24204 sásoprydun `y
nq Y nA (2904) wnzsppapqipoq snuasiqns
ec661 чоѕішоц |. ср под зтово "у
ec661 uosduiou су puny snuaSiqnu "у
1861 wex3u] 9 eq су ypuəg па оолоош “y
90661 чозашоцј, су Читу ғп20490]9 "у
с661 uosduiou |, ср шод M2901109 "у
90661 uosduiou, ср ayooy зтухудоцтоор “y
ayooy sniDqo4() snuasqng
vz61 Aepsuoy y "qp4y (әчәәлсу) snonosoqns "у
6261
ru[esIHH $ MBunoy :6961 ‘LSGI Yossie] :усбү PUreJEPA I "uis snipjpots "У
1861 9191 8c әле (7]) sipas горко
S861 лоџәред ју ләшү 42861 2491
Y 2۸91 “6L61 чоГҳѕ[әЗиҷ :8/61 [e 19 PASUNEAS :8261
qor^o[n3ory ‘£961 #5105 “6961 ‘LEGT uossreT PEGI 9^0]
“PHOT 949] Y 9^0 ‘SGI “GEST PUBIBRA ‘GEGT 19199095 8б "1 зї!їюхї$ Y
2661 !USPUNIPN 2 oqnsjem] РТ тріо snoruodolopansd “y
1861 249] 7I IQIM "ум (JOY) зиговодпа 1047)
22661 чозашоцј,
‘q 8861 '9961 91071 Y 2491 PESGI FWEIBEA PEGI 2291 РТ ‘Jey suaosaqnd y
воополојоц uc џохвј,
penunuo) сү e[qup
O————————— адд:
Annals of the
Missouri Botanical Garden
142
9861 WoqAN 86 лаб snsodns y
2661 !SPYNIPN o очпвуем] 9G wig snsogna y
2861 uerreure3qns :c76[ "е 19 PIPA pI wg snsodns “Y
2661 9847, x повашоцј, Фр Jay SNxapfos "M
9961 ue^eqq X enjoy :с961 ew V1 US SID NIUDA Y
£661 'USPUNIEN 29 oqnsyem] 8z род oran Y
2661 9947, x човашоцј, 8с JOTUBA 39 AF] са У. |
9861 шод 8с тпркаморонроме up
2861 uerueurg1qngs VI Iupee спурјодолти “Y
2661 9947 $ човашоцј, 8c nq Y nA хә ny snofiojrmut y
8961 "SH 9c ejeÁey mow y
peg snsojnpupjd ва “Jag SNUDILAQUID) Y
90661 човашоцј, 8c ТешәҢ 129013 чел “Jag snuptioquin] Y
£661 9947 $ човашоцј, ‘Z661 ruseunaey o одпетему 8c 195 пили Y
2661 OFYZ x uosduoy y, Тр 34904 snapuan "M
9861 “0861 WOqAN 9s "qun, snarpur y
£661 9947, $ поѕішоц |, :2661 njseunaey y очпвуем] 8б әдипу Y “SUSY тиәЎирүз1 “Y
90661 uosduoy, 8с HN PeH TAS и
90661 човашоцј, cV пәп A 0Y Y
1161 ‘1F 19 теџеб 8c ‘qxoy snuádoxoy M
90661 човашоцј, 8c әдипу Y [suo Muay y
8961 "SH РТ ry] sn] 4ydovo0w тел вұвќең зарлошојро “Y
£661 9947, $ uosduioq], 666] ruseqnaev 2 Oqnsjem] 87 ‘rue N 1UNZpr09-DIDÁDY Y
£661 Oqnsiemy X njseqniey “8661 “1861 ouutf 9S 295 РЕН worry Y
2661 oqnsieA[ W ILYEN “9561 ouutf ср AUS Y че AN H
(00% '0№) p961 ччеш#лоң 86 eumg $n049w018 “y
0661 “TP 19 [euduts “PRET ‘IF 19 1119 96 azjuny snupt2upan ^y
1861 чегишет VI ozjunw SNUDILSUPIDE M
1861 uerueure1qng VI ayoog спајтј "M
2661 9847, x uosduoy], 87 ozjunw SISUISOULIOf “Y
£661 9947 ју човашоцј, 86 JorueA Y ‘491 tappof Y
9861 WOqAN 9G Joupiey snuvrujoynof “Y
1861 uetueueaqng vl 1oupae«) ститијоцтој “Y
2661 9942, $ човашоцј, 86 "ovy зәроитудә Y
£66[ 9842, $ човашоцј, 86 "zey-pueg тухцаоцоор 4
£661 9947, x uosdwoy y, 86 m] 29 nA snijofisspao y
воополојо Y uz похвј,
грепипио с 148],
143
Volume 84, Number 1
1997
Thompson
Chromosome Numbers in Rubus
22661 uosduiour
:2661 9847, ў човашоцј, :2661 ruseunrew A одпзјемј
2661 9947, x uosduoy y, :2661 !yseyney 9 Oqnsjem]
9961 чемец(] Y LIN ‘S961 THEN 6761 TE 19 sure,
(S9 `оМ) #961 uueus1og
S961 “Te 19 sardg “cg6] sisse[qnq y serdg
S86L ‘Te 19 59146 ‘eger sissajgng y seidg
2661 9947, № човашоцј, раб] моцед 9 Ko[guo'T
2861 9191
96661 чозашоцј, :8961
JI тјојџоцооо 7M
ng nfunp Y
inus хә "ureg-'qong sn4ojfiq "У
Á1194 Y “USA SNUDIP)OQYILD ^M
под snjojado "M
шод snypjadp “y
ajgoy таоцаоигрр "M
"qp&y (nny) штаојалра 122007
ова NA 29 10/48], ‘PEGI eurexeeA ‘себт Ko[pduo 5 ^oure(q VI "ny snaojfiaapd y
1861 эло] ial "аркы (7) штморо 4290qny
20661 uos
-Чшоцу, :/861 OSM 8661 19peopy Y 193294 *pco[ Aopsucy] VI "] snmaopo “Y
90661 uosduiou], VI Келе) “y snupoixauio2u "M
7261 fo[duoT Ic шо], ғпѕототәр “Y
90661 vosdwoy, РТ — PE H
33901 стдојаои у snuesqng
£66] 9947, x uosduour 8с s[euq хә 9904 SNANIUOYIUDX "У
22661 поѕйшоці, gz "2eW- pueg штыо8ит$] “y
906010 ЧЕРИН Ооо 8c "HS Нор Y
20661 uosduiou], 9c "zv|p-pueg (оед % ләт) snaojfijdum “ea 9oueg soporyda? y
E661 9847, x uosdwoy, ‘PEGI вшелеед 9c 9oueH sapouuda] y
22661 uosduiou r 9c 9oueg тәоуштѕ y
2661 !useu
-NIEN Y oqnsie^[ :2661 “Te 19 Oqnsyem] :8661 *epco[ ouutf 86 SURE зири 2
6961 SPIN 8с supp ae у
2661 9847, ў uosdwoy] ‘£661 IUSEJNIEPN у одпвјемј 9c `Цэиві x neoung sisuauanyojes "M
42661 uosduou], 86 ÁiI9qLI9M ‘A0 “WS snsodns "M
0861 WoqAN 86 000,1 118211001 ea “wg snsodns “Y
ваополојон uc џохвј,
‘ponunuoy `T [QEL
Missouri Botanical Garden
Annals of the
144
JOIUEA 29 'AY] SIsuaouay? "M
90661 човашоцј, ‘Z661 !Чѕеҷпзем X Oqnsiem] vt
9261 ловјпаеџј РТ €AOXUIS хә "USSOIC) (BAOUBZOY) пудтд "M
BAOULZOY MYosng “IBA "ү snappi ^M
62661 PUBIBBA РТ ‘Чиу SOUND IBA ERIP. Y
90661 чозашоцј, :с861 Aouapreq x PAI :1861 үе
ә [ood 6261 uoD[s[ou :8L61 [e 19 BYSUITEYS “6961 uos
Sue] “PSO Пела Y lesury “ES6T чоѕшең-доѕәң :40р61
ѕешоці, ‘LEGI 19pamuoy “OPEL “PEST PAouRZ0Y ‘ZEGT
Пәшед ‘L561 uojgurpre(] o опело ‘pz, моце( 7 Хојдио VI "| snappi Y
2661 9847, x uosdwoyy, al ayooy sisdoappi “Y
1861 IPH Y uonug VI ‘TM nsues uo ‘q snsojnounpod y
2261 Te 19 eya 86 mo ^) хә qua enam у
2561 ПАН Y uonug ‘S961 JEW 14! uoq '5 xe "еу snoatu "у
2561 IMH Y uonug 2261 SHEW VI 99904 nsues *XOY spond "y
BIH ^H споти чел 'мовр snaupdéay Y
90661 чоѕїшоці, РТ eyonog ӯ puny snupuaisrauffoy y
6661 uosduioq[, ‘E661 9947 x поѕішоці, ‘Z661
!YSPUNIPN 29 oqnsie^T “6861 njseunrey ‘BSGI ‘81661 ouutf VI ҷчо Mty Y
20661 човашоцј, 8261 deay PI Авдо “Y SISUADADY Y
£661 'SPUNIBN 9 одпвјемј PI "uie утиюХю1@ ^w
LLOL 18 19 PWN 8с uoq ‘q $nsojorpof ^M
2661 njseunaev X oqnsyem] ‘S961 ТРИ РТ uoq ‘q snsojorof y
90661 чоѕйшоці, :2661
овцу 39 повашоцј, :/ 261 [MH Y Uonug феб eureregA VI 9x20, зпзојповој Y
20661 uosduiou], РТ [әт] хә әҹ̧әо; sounydajsna "y
90661 повішоці, ‘£661 9947
Y човашоцј, :2661 mjseunaev y одпѕјему ¿0661 ‘TP 19
їеЧӣшс :/861 uerueure1qng “PRET ‘TF 19 1119 рт ‘TP 19 P|
“TP 6261 “TP 19 BIYPW ‘9961 ueeuq Y PUN :2961 үер VI "us snondijjo y
2661 !YSPUNIBN A) одпвјемј VI 'AY] smyuDoD22042 "M
8561 ‘91861 ouutf Vt Келсу "y 1y2un y
22661 uosdwioy y, :2661 9847, x повашоцј, :2661 n[seuna
-е№ 29 Oqnsiem] :6861 njseunaeN 8961 doow *pc6] ешелеед VI әдип snyofidopao "y
9666] човашоцј, :2661 oeuz x uosduioq] ‘1661 14589
“MPN $ oqnsie^| :ү861 “TP зә 1004 *766T IMH Y uonug
"2961 SHEN “OPEL “TP 19 surerprA *pg6T мое] y Хојдио] pI ‘bi гпироло “y
воополојом uz џохвј,
"ропипио сәер
145
Моште 84, Митбег 1
1997
Thompson
Chromosome Numbers in Rubus
2661 Tyseynsey oy одпајемј pI ‘Zploy sisuampulyo "Y
90661 чов оцу], ‘P561 woue( y Хојдио | усб Aa[3uo'] bl "1 $1191uapio00 “Y
2261 Te P Uyw gz аипуј, snəau ^M
9961 чемеца X PIYOW ‘SOGT rej 861 PWEIEEA vl "ug srdin90150] Y
2561 IMH Y uonug Vt Dp rue у
80661 uosduiou| :2661 oeuz x поѕашоці, :2661
mjseunaev A) oqnsje^[ :/861 uerueure1qno :996[ ue«euq
X VIIN ‘961 [FW ‘LS6T IMH Y uonug :qopogp зешоцј, Vt чин TS
2661 !Чѕецпеу 7p Ognsyem] “2261 CUD 8с En ssl. перини ү
p961 чивш8лод VI подом yg пшјоцпт-тиош су
92661 човашоц, :2661 'USTUNIPN o Oqnsjem] VI jotusA Y Ay] mque Ww
8261 ouutf I ‘zploy snsiou чел y] snjyjéudoaonu 7M
POOE ПРУ пр она VI 371 труди ^y
8S61 очш[ 5pgo] Mour(] y Ko|duoq I 34904 ѕпәрдоѕәш 'M
2с661 uosduiou J, bl UOJ9p[?7) 3 “pezy sNUDIUYINDADU Y
2961 TER vt SAA Y
2861
‘Te 19 59146 :861 sissopgnq y 59846 :/261 IMH Y uonug VI 'ЧАәў Y PPA 1810p] у
(gp 'oN) p961 uueuiz1ogq 971 a[[nq snun127u210] “Y
©861 ‘Te 19 59146 :с861 sisso[qnq 9 sordg се uns H'O (1509) snivjjootpodisuo] “y
C961 ‘TP 19 sards ‘eget sissapgng 29 sardg 87 "ung "HD (л=п) snipjjoorpadi2uo] y
S861 TF 19 sards :с861 sisse[qnq 29 seidg VI "uns “HD (1805) snipjjoorpadiduo] y
22661 чоѕашоц |, ‘EEG Buo 9 моме( РТ Аел “y 29 Коллој, xo se[ano(q sruuapoona] “y
2661 'YSPYNIBN % одпзјемј 9c "ury sndip300n3| y
90661 uosduou], vt пе 19 NA LiL sisuataqny чел 9x00, SNJÁISOISD] “Y
20661 чозЧшоц], :2661 9947,
№ uosduroq], ‘LG6T IMH $ uonug *pg6[ моле y Хојвиој VI әзәо,] snjÁiso1D] ^M
CLOT РЦ vt PER
£661 9942, 9 uosdwoy y, ¿2661 IMH X чопша *pG6[ FWEIEEA vt 2x20, (sie) зтодои “y
6061 T" PATA РТ PEH Г.)
Kopeg ^H] (SWH) зпирогјиту ea 9100]y "S snipuruouut у
92661
повашоцј, :2661 9847, x uosduiou] ‘£661 tuseuniev
m oqnsie^] :766[ IMH $ uonug *pz61 моле X Хој8иој VI 9100yy "S snipunuouu y
FG66T чозашоцј,
"8561 “91861 Ouurf ‘жабі моше(т X Хә[ӣиот 361 Ao[duo] VI 9x90 snso49900]]t “Y
8261 ouutf VI "unxejy snoruodpf ^y
sooua19JoM uc џохвј,
‘panunuoy `I APL
ИР,
146
Annals of the
Missouri Botanical Garden
96661 човашоцј, ‘Z661 tuseunaew $
oqnsie^[ :8961 UBB MA 2 1014e], 2261 Хој8ио тј oy моли v1 “Чела sipiqoioads "у
92661 човішоці, ‘£661 9947,
у uosdwoy] ‘E661 !YSPYNIBN у OGNSIBM] “рст PUIEIEEA VI 9320, xəduns “y
6661 PA0UBZOY 9c (BISy ам) “Woy 2111918 чел "T $пәрр1 “Y
6661 PA0UBZOY 9c (etsy ум) 93904 (A91) sisuouipyops “чел "J ѕпәррт Y
6661 PAOURZOY 9c (ersy N) 93904 snispjoupjaui "тел "J $пәрр1 “Y
90661 чоѕішоці, :2861
BAOYNYT Y ләѕыпд £276] үе 19 BaoynyT #861 84997401
“SOY y yotasmaory :8/61 “9/61 YrAe[nBory “6861 eAouezoy 8б 'AY] sisuautpuops “Y
20661 повашоцј, PL 9490] (suite) 21200100 “IRA "шс 8170/1801 “Y
20661 човашоцј, :2661 9947; W поѕйшоц PL "ug snyofisor y
8061 ouutf VI — POM
1861 uerueure1qno РТ "ход SNSOUIIDA "M
20661 повашоцј,
‘C661 9847, Y поѕішоц |, 2661 'YSBYUNIBN o» одпвјемј РТ ‘ssoquie’) suadund “y
2661 !'USTUNIEN јр oqneyemy] toco] ouutf РТ ouye 422Dopnosd y
20661 човашоцј, gz Аопва Wa мојјгти y
Kojeg ‘HT дола y
S861 ‘TP 19 sedg 5861 srssojqnq y serds 86 "PILA тиш y
S861 ‘TP 19 sard 4861 sissagng y 59146 I ‘PITA stapuuid "у
22661 човашоцј, РТ JOLUBA Y "лә" sisuapfuid Су
£661 9847, 9 uosduoy y,
:£661 !useunaey 29 одпејему *¿G6T [MH Y uontg 8561
‘qIS6T Ouurf :) 261 Ámstuoyo *pgop моме $ Хојвиој VI urxe snispjooruaotd "y
2661 "jseunrey $ oqnsie^p geo ouurf I urxe тога y
Собе SPORE Y Брин 16 q тодалра y
vz61 KopduoT vl quay, пуй y
92661
иоѕашоц І, :2661 9947, x uosduiou] :1661 tyseynsey
19 Oqnsiem] “6861 `Те 19 eAoreqoaq :49961 muureg :2261
[MH Y uonug “6p61 ‘Te 19 SWIM “8561 ‘1861 ouutf pI "1 smgofiavd у
20661 uosduiou j, pI azjuny (ABI *y) snjj&udojdoo ‘iea Áexinjyj хә 'qunyy, зтошура y
8S6T "41661 ouutf РТ әдипу sapiomuipd “y
8861 “41861 оишү VI JOJUBA 29 ‘A91 1оштој y
8261 ouurf VI Келе) “y sn fydordos y
ec661 uosduoqr ‘Z661 ЧЕЧЕ 5 Oqnsjem] I Águn]y хә "ип, spud су
ваополојоц uz џохвј,
:ponunuo) ср әде,
147
Thompson
Volume 84, Number 1
1997
Chromosome Numbers in Rubus
20661 човашоцј, gz uosja snsoonbs ^w
#0661 uosduio[, 9c "un PR Y
(202 `оМ) #961 uuew3iog 8c uewyey snpidsny "eA пвшхем 2202404 Y
2861 пен y Z1oquoznog 9c ueueyong snaupd y
92661 човашоцј, 8c Anm у eposo Y
(aued ола ayooy nsuas sn;pqodumr] =)
ueurx[ey (yosiy) зтрдоцтиоллуј зпиодапс
2661 Oqneyemy 29 тувецием Ic троє 10unjso& "у
9661 CRAP тиме VI "zproy 10uttsoK “y
8261 ouutf VI JOJUBA 29 "Ag'] 19994 "M
2261 uossjejsne) VI "y9UB1J x neoung sndapoowjupx "M
2061 EIEN опна РТ pm epa
92661 ‘
позва шоц, :2661 njseqnrev ә oqnsie^p :8061 “91561 ouurf VI Келти xo "qunq] spy y
92661 чоѕішоці, :2661 9947, $ uosduiou], РТ оа tS у
3261 ouurf РТ "ром STLIDUOLO) “IBA 9x90, souvydaisna Y
8561 “91861 оси PI под uou “uodef ‘10ne пиозоштиоз "Y
PR SENT у
2661 9947, 9 човашоцј, :/261 IMH $ uonug РТ поводу тирген Y
8861 оишү VI uoq ‘q хә "ем dsv “y
22661 uosduiou r
:2661 9847, 29 uosduioq] ‘Z661 !yseyney д) одпвјемј vt “bi smupamums су
22661 uosduioqu] РТ JorueA 29 “AY] smijofidapqpaoqns “eA 7y7] snyAydonu Y
8261 очи VI AE и U
JOIURA A ‘AQT (jorueA ду “AY]) стуојортолодп ^y
Lv61 19surg 1 “XI, 550115 y
PSOL Held ју јеви ‘ZEGT ПәШеД VI "wor snsoZ1is чел "T ѕпәррт ^y
9961 249] $ AQT VI (лошу сому) 99904 (ләт) sisauaunpyops чел "| snappt “Y
2861 SHIM 29 194984 “CTBT 91971 Y AQT 14! (ләшү сому) одоод ѕтғрјоирјәш чел “| ѕпәррт “Y
re61 Хој8иој РТ uospreqorj SISUZPDUDI лел "ү ѕпәррт ^M
20661 поѕішоцү ‘peg ешелевд *poo
пег4 Y PSU *Gpo[ TE 19 SWEM рт ‘TE 19 19yos
"M “6£61 PAOUBZOY ‘9661 [PUA “PZHT MOR 29 Хә[8иот РТ xor snsoZuis y
воополојон uz џохвј,
гропипиој `I әүвү,
Missouri Botanical Garden
Annals of the
148
0961 IPASEH
ECG] vosiuey-do¡soy
0861 YASNYJ
2061 uosrureH-do[soH
E61 "Ива
(6261 әрпЕд) чозше®ң
0661 uaisumq)
(6261 зрпвуј) uostueH
9061 x?uuofrog :6661 “EET uossjelsn?)
£€6] повишен -Чојвон (6261 орпеу) човшен
9961 OO $ muureg ‘eggs, twweg :1961 UNL X 119
-SEH :0961 IPASPH ‘6861 CULO Y UPLIPALE OSE ңәш
19199 ‘gs61 193199 :cgg[ SUBI) “EL6T /ә[8007] 9 MoE
ec661 чоѕішоці, :epo] UOSsyeIsNs)
c661 exsu&maog
061 SPW
2261 чоѕшең-Яојѕәң
(6661 орпеу) 931oqe, ‘SEGT 94610
2261 vostre y-do]soH
C661 "JB 19 одпзјемј
2261 чоѕшең-Яо[ѕәң
с661 [E 19 oqnsjem]
9561 зочмеПод
9661 доимоПоц
wexojg “Y хә See] SMIDAJDI ‘Y
(une1g °) uou (9661) uosieA, ‘HDA nsues 119/I9D.1Q "M se)
позмо “Y Y Seepy snsourunj]sia24Q "M
93904 хә “Way "y sisuaim3o4q "M
‘aguas спиргрлод "M
see (qeg) пшохојд Y
seyoing хопешојој ттиртирхојд Y
]s9A suo4fiq "M
ungig “9 11u04142Q. "M
wen UOJYSY '^9 појмом ^y 11u0140Q "M
“YOSIY SNNUDIS "M
‘fay sun pxo
лел oue] (oduvT) nuaya
a
snipuo42nui xo3d osnery THA $ndoumiuoum ^w
Y
Y
aduey] (әдив”ү) nuy
(әлрпс uou *uosjeA ‘HIM nsues) suo4figopnosd y
UO]MON “Y хә поме уд Suaodimsp ^M
(еќејешң “AD түп "fd. $742204d су se)
(вотоодв &uraqxoe[q ueədomy) "y sngny snuadqng
(оо ‘ds RT 'У se)
one snjpoppf Y
UO]M9N "у SUDIMPUDIO/FUD “Y
qur] хә ошто uou ‘oped oad “one mpppuazyoazyas ^M
sea] snipoyi]dum y
199294 “AH Nuas “Y
поте HOM зтјојигуо Y
1992M “TH хә APPM sniszodspp ^M
"IMA "fd srivusopo “y
ague”] от уолорш ‘dsqns “aguas спиоврр ^M
snqny uonoog
$99u919]9M
uc
uoxer
"penumnuo?)
т 9"
149
Thompson
Volume 84, Number 1
1997
Chromosome Numbers in Rubus
"pur] зтритуза “Y
ЄСЄТ uostueH-do[sog 9€ прије] (s19204 WM) s2pr01munjoo "M
2061 човшен-дојвон (6261 әрперү) иоѕшең gz "qeg $1ўиәтиоишпр ^M
2261 ови се ogduv] xo иәѕиәү 14224p "Y
#0661 uosduou], :6e6[ uosspejsne) 9c әЗив"] xo uosuof мобмр “Y
2261 чоѕшең-доѕәң 9c UOSIPA HIM пмотр ^M
9261 YuLPlieg *epg] позвјеузпо) Ic aued oid зоо Y AP snpimu ^w
2261 eneq QZ зоо Y әціәмҖ uou “yne при y
2261 човшлен-Фојвон TC S99N Y ош uou “yne snpınu ^M
TOW "га гтооиотр “y
9861 xouuefrog 8c PPM тәүшәш 7M
BUM "хә “IMA Га зторатр y
(6661 әрпер) повшлен 8c CIMA ‘fd sarvurun y se)
uosiA, ‘HOM (5888) smofimmop y
S661 ‘TP 19 Oqnsiem] :2р61 ‘661 uossjejsnz) 8c Teys "S73 (s1920 WA) sa 4ydásop “y
22661 чозашоцј, 9c "nf мАо y
себт әиел”) 97, (soo] snivam y se)
2061 uosruıeH-do[saH 9c uos "WA Snupdusidsaio y
2с661 uosduou] :с661 `8 19 Oqnsjem] 9c loqoA “YH SapiosiXyjouoa “y
6961 UNO 9c 94904 17347 "M
sequog nsnjo Y
6261 човцлен-дојвон gz (шлој pemp “ләпәсу стилраов “y se)
2261 vostuey-do¡soy 9c (Aguas, snutupdas “y sv)
ТРР Y uoueg “OM sisuounqssio “Y
6561 Voss aens 8с 9x90, snonquia ^M
(6261 әрпвуџ) uosureg gz s1930Y ‘WM (S193304 WM) чохои “Y
9S61 Punəfiog 9c 2x90 SNSLÁYI014074 “Y
(6£61 9pnejy) uosureg 8с (uneig “9 114:D.949q “y se)
aipng snyjkudoopo “y
9661 допџоПод gz зтрАудолоруо “IBA огупцоб ^A 29 141088 snppAydosavyo y
22661 uosduoy], 9c 9390, SNISDINDI "M
£S6[ човшлен-дојвон (6661 epnejy) позшен 8с TIN Га Y әлләјәт snypaudonpapo "у
8961 LAON FI Ipod 522019019 лел "мод =пѕојиәшо] “Y
1961 UNL Y [[?9€H :0961 [ISH :9661 xouuefrog
"£661 198199 ‘OGGI YPISHYD :2р61 ‘GEGT ‘61 uossjeisno) VI aud od "ippog $пзотиәшо] ^y
92661 uosduoy y, :6961 19 РТ Ја зижешо YY
ваополојо Uz џохвј,
“panupg uo”) т 91948],
150
Annals of the
Missouri Botanical Garden
2661 uostu]-do[soH се ‘pueg nupunav Y
ETOT ym) 87 И den aco E
95661 uosduiou], 8c "pues питшиту Y
2061 човшен-дојвон 87 левој WA ғпиотуту:9 y
9661
youuafteg ‘661 повшен-дојвон (6661 PPren) әйләде] 8c әҹәо 517019 Y
9661 xouuefrag 9c 1opuoy simpornpa ^M
POOL exsu&maog 8c [$914 '5 Y "f 81010019 y
OS6T uou) ‘усб Х9ј8и0] tz 238 wuy пәртозл{1 Y
6£61 uossjejsne) Ic uedry snuimngaaqns ^y
'qepurT snuzmjaagns ‘IEA DADO] SNYIUDSIÁNI ^M
€V6T “6£61 uosspeisnz) Ic DADOS snus? ^M
үе 19 JayUNy хә әцтәд\ 1135709015 ^M
966] дочџоћод 9c "IMA ‘fd меппт :dsqns одоше % booa] пиолрод су
v661 exsu£iaog 8c ‘qudg sisuamman "M
90661 човашоцј, 86 ipreqjog пзотриоја ^M
£66] чозшен-дојвон 8z UOSIBA HIM Ampun? y
9667 тю емиш 8с uneig ‘9 sosudynpunjs ^M
20661 човашоцј, 9c ayooy $n2124002 “Y
9261 xouuofrog 37 quayey snip nomad y
S661 “TP 19 одпвтем] “6861 uossjeisn?) 8с pu 11112128 y
(6£61 apnea) vostre 8c (ловом ‘NM ғ=пир/әр Y se)
£66] човилен-дојвон gz 9x90, гојолату y
6861 човшен-дојвон SE TOW ‘fd 29 924999] sipiqppruaof у
(6661 зрпеу) човшлен 8с "IMA Га Y 2148197] sipiqppruaof у
966] Ҹәишәйәя 82 "IMA ‘fd snsonxayf ‘dsqns əya snsoyof y
9661 [Ре офа 8с aU! snsoyof “Y
2061 повшлен-Чојвон “6861 uosspeisn?) 87 [pur] snssy “y
9661 xouuefrog Ic [pur] snssy "y
€S61 uosuueg-do[sog (6261 9pne]y) әйләде] 8с әціәм uou *uosjeA ‘HIM NSUIS #920804 "у
20661 човашоцј, 8с uoMaN “Y Y svepy заолу ла “y
ЄСЄ] чоѕшшең-до[ѕәң 8c uos]EA ‘HIM snpunqpaao ^M
C66] ‘TP 19 Oqnsjem] 9c Rq “AH (човшпцос *y) snsouidsimupojo y
€po] uossjeisne) IZ шәле) хә 3490] 1017072 "M
966] допџоћод *ecgq повшлен-Чојвон “6£61 uossjeisnz) 96 93904 5т19ә19ә "y
(6661 pne) човшең 8с IMA “Га stadsaosip y
воополојоц uz UOXB],
‘panunuoy '[ әү,
151
Thompson
Volume 84, Number 1
Chromosome Numbers in Rubus
1997
2961 човшлен-дојвон Ic (si330Y ‘WM тјојисттир rea 3490] sopp[2otsp] “Y вв)
2261 чоѕшен -дојѕәң 9c "us suyopisoono] “Y
"uis хә "цотојцос SÁYIDISOIMA] “Y
с661 '[? 19 oqnsjem] 9c 9x90, snupupomna] "M
2061 човштен-дојвон 8c 9x20, uou *(gc6[) поему HM nsues SMUPUDINA] "y
U0IMIN "V Y seepy smuaofupunong] y
2261 SABIA 9c “duweg титоз “y
pnearyy tasspuidsay "M
2561 SPW St soo] snsouiZi1u23] "M
(62 apne) әйләде 8z sae] т50и191иә] "M
€S6[ повшен-дојвон (6661 APNEA) uosuueg 8z UOSIEA HOM NND Y
6661 чозвјејепо) 87 19199) Y "ри; `y хә uəsuəf 122upj y
22661 човашоцј,
‘9961 OWO ју пише 20961 nuureg :0961 [oXseH *ecor
повишен -дојвон “gg6] SUBI) ‘L361 uoidurae x әиел) 8c ‘PITA 52120] У
6661 поверио) 56 Ten өлрипцоу ^w
(6£61 әрпеуџ) повилен 8с (‘yosery ^4 sunpnsu: "y se)
s1920y пол y
6261 uossjeisn^) 97 "qepur] snuifuoo y
"gosory 4 Саодт) smafuoo *dsqns 'цовату supynsur у
90661 uosduou] :2р61 “6£61 “ELST uossjersn) 8с ey sso Y
6361 повер) 86 Чери момент RI
(6261 9pnejy) әбләче 1 ev uueog xe оцтод стада ^y
2661 човилен-дојвон (6261 орпвуј) 9319q8 4 gz uus0g хә әцтәм smsəfur y
2061 човилен-дојвон 8c (шлој ремр) ови uou *(gcG[) чоѕјем ‘H7 nsuos SNSIÁYIOLIDU ^M
2261 човшлен-дојвон (6261 орпеуј) иовшлен 9c our] uou “(SEG T) uosjeA ‘HIM nsues SOSA YIOLIDUL Y
(6261 9pnej) uosuey I eue] uou 8261) UOSIEA ‘HIM nsues SOSIÁYIOLIDU “Y
појмом “Y 29 seepy snppdasysafin y
2061 иоѕшең-Яојѕәң :2261 ее gz "qeg зтрратои ^y
£S6[ uostueg-do[sag :2261 eneq 8с под тломфи Y
S661 ‘TF 19 oqnsjem] t9c6] xouuefrog 8с 93204 зпоррошодху “y
6961 uosrıreH-do[seH Ic PPH Y uoueg "уд 1077.0 Y
aguas) хә Áəjdry snjrydo¡4y y
6061 човилен-дојвон “(6£61 әрперү) uosureg 86 алроб pgmn, Y
рб] Seuou] ‘GEGT PULIN ‘L361 uoiZurpre(q 5 Quer) 9c qeg 4221514] "M
. UOSIEA WIM sruaofispuy ^y
sooua19]9M uc џохвј,
грепшпио) ү гјдвј,
Annals of the
152
Missouri Botanical Garden
TOA
2261 човшлен-ојвон [42 ‘PPH $ uoueg "уд sisuagpuiqnau "У
2661 поѕшең-дојѕәң 87 ‘PPH Y uoueg DA ѕ1иәдридтәи "У
(6661 9pneq) човшен 8c ‘qeg пруподтги y
0861 USO :6261 “ELST uosspeisna) 8Z uosiopuy °<) sng2aaqms `Y
с661 eYsuAeıog :с861 лопова A PADL TU
:9с61 ЯәчпәПәң 661 uosuueg-do[soH ‘Ep6 uosspersn?) 8с цен rm.
6261 човилен-дојвон 9c e1png ләруюшоәи ^w
6661 uostueg-do[san 86 uO) ‘HOM CqopurT) snjéudoaonu чел "пи “fd 51]юїошәи "у
6661 uossjejsn?) (6661 әрперү) 81949 4 8с "qopur] мәшјәѕ “y
v661 PAsuÁie1og ‘ESG, повмлен-Чојвон 8с TOR: Fd trmou у
2661 човшлен-Чојвон 8с пове WIA A № Г) 1724p]u "У
£S61 чоѕшең-до[ѕәң 9c попе SNIDULOPD "M
eco] човшлен-дојвон 9c aupng мошти "M
€b61 “6E6T uossjeisn?) 8G qepurg NAO ү
C661 '|? 19 Oqnsjem] 9c UOSSIBIA J, мерити су
966] »ouueoftog 8Z urexo[g SNIDUOJMU ^w
neaiog стотиоти "M
6961 119 ‘Ep61 “6E61 uossjeisnt) 1 quod fo, EOP M
с661 ‘TE 19 Oqnsjem] Ic far] хә "гү snupjuow су
(62661 apne) човшлен 9c s1930Y ‘WM тиште још ^M
22661 чоѕашоц [, 82 "znf 10gu2202$nu "M
2961 SHEW SẸ юне ASNO ar
(6661 орпеу) uosueH 86 поворци
gcp позвале -Чојвон 9c попе 4afimuids y
NOMON V te mw Y
2261 чоѕшең-до[ѕәң 9c "udeg sao "M
(6661 әрпер) әйләде 87 2x90, гишгроипјеш “Y
2061 чоѕшен -дојѕәң St s1920y ‘WM 29 9904 11]mysamui Y
Ep61 uossjeisn?) 9c APNG (IMN fd) отриудолош ва ѕәәм X APM sn 4ydosovw ^M
C661 ‘TF 19 oqnsje^] :с661 exsu&je1og
:9261 Youualieg 661 човилен-дојвон ‘Epp uossjeisnz) 9c вәә№ Y I три удолоош "M
(6661 9pnej) иовшен 87 UOSIEA "WA SYOfidUO] Y
uospA ‘HOM suo4figuo] y
9261 xyouuefrag ‘ZEGT eneq 9c $99] SNUDÁSpUY ^M
с661 71° 19 oqnsiem] ‘eco, uostueg-dopsag :6е61 uossjeisno 86 "IMA fd 222292pu] y
$99u919]9M uc UOXB],
‘panunjuoy с] әчү,
153
Thompson
Volume 84, Number 1
Chromosome Numbers in Rubus
1997
(6861 грпеџ) uosuaeg 8с "IMA “Cd Y әїлә}әт ansanb y
5961 поени -борән cv џојед) $ "pug snyofiaind чел ‘quey sippnupaid "у
2261 човшлен-дојвон ez uap y ‘pug snyofiasnd rea ‘quy syppruvsdd ^M
ec661 uosduiou]
"1961 UNL X позван :9eoq xouuefrog eco човшен-дојвон 8c quae y sypprunstd ^y
РӨТ uosspejsna) 8Z-9Z "queyew sypprupstd y
7661 PYsUAIBIOg 86 "1917, (apns) smuaofipopnua(d “y
6661 "2261 uossjejsn?) 8с snizans ^j әчтәдү snpyyod 7M
isng y xa sng snpyyodopnasd ^y
(6£61 грпеуј) повишен 9c (s1odoy "WA uojKxoskato "M se)
'q107) хә 31puaj9] y Авјпод зтодиојола су
S661 ‘E 19 oqnsjem] 9c iuo[ojrog xo22D4d "M
2261 чоѕшең-до[ѕәң gz (шлој pemp) әри” стпшоугирлјод `y
S661 E 19 oqnsjem] :9661 ңәш
aftog ‘g61 чозшен-дојвон *epop ‘GEGT 'egor човејезапе) 86 ‘qapury smuayyuvÁjod y
92661
позва шоцј, :с661 BYSUAIeIOg :9261 доишоПод feg; еш
“BIBRA :2061 чоѕшең -до[ѕәң “gp61 ‘6661 “EGET uossjersna 9c зао Y ош SIDO y
2261 SPW се "one туојплало ^M
966] Pung 82 IQM smjoftuidivs y
S061: p» 9 одани 9c AJOT Y "IMA “fd smymupoomjd `y
2261 чоѕшең-до[ѕәң 9c UOSIEA ‘HIM sndap2o2pud y
(6£61 зрпеу) повише 9c (TEN ‘fd =мојрхр ^y se)
UOSIEA ‘HIM ѕпритшләа су
2861 Ydezy ‘gG61 повилен-дојвон 87 ƏLƏM пргојед “Y
(6£61 PPren) әйләде 8 SOON Y "Чед пређе Y
9261 youuefteg :6661 “EEG uosspeisnz) S£ SOON Y MOM HPD Y
с661 ‘TP 19 Oqnsjem] ‘CEG, exsu&e1og 96 ша титиошграа y
6661 чоѕшең-до[ѕәң 9c a1png зишојутоа ^y
9S61
xouuefteg ‘Eco, повилен-дојвон *6g6[ “E£61 uossjeisne) 8с MISA snpyyod y
€S61 Uostue}]-dojsay 9c aipng SNYIUDÁXO су
(6661 9pnejy) vostre y ct ное MOM foptoptm ч
0961 TIPJSEH “ES6T човмлен
-Чојвон ‘POPOL ѕешоці, (6661 зрпеу) 93198. 4 ‘g6 оче) 96 поете Y IA OPIOPHIR E
eapng пишодрити y
$99u219]9M uc UOXB],
'penunuo с әү,
154
Annals of the
Missouri Botanical Garden
9c61 youuefrog 87 a1png snijofispaop ‘dsqns ти "pd 8101243) y
(6661 орпеу) uosieA, 9c иовте\ HOM spat "У
Ep61 uosspeisn2) 92x 1894 snjpoms Y
6961 19 6661 PVEN) 931oqe 4 9c 189A smiDo[ns ^M
£c61 човшен-дојвон 9c a2n1({ sapiouii2u1qns ^M
9261 хопџоћод :2261 иовилен-дојвон
{661 epnejy) човшен :6661 “E£6T uossjeisu?) ‘ZEGT Ned 85 әцтәдү пједиоа y
S661 ‘TP 19 Oqnsiem] ‘961 outrefrog ‘Ego, човшен-Чојвон 9c SOON 29 təm Monpas “Y
6661 uosspeisnz) 9c ‘qapury smujfuoo y
uOS]EA ‘HIM SDuoraquardos ^M
Ep61 uossjeisnz) 9c Buy snupjuow ^y
S661 TE 19 Oqnsjem] :9261 xouuefreg 9c 9qr9A хә лојцом пости "M
€c6] чоѕшең-до[ѕәң 9c "Qosery ^J SLDINIIID °F UOS МОМ SSS “Y
€V6T ‘661 uossjeisnz) 9c odueT snjnydotos y
2061 uosrureH-do[seH 8c пове “YOM (олрпб) #220195 y
1861 1998 9c 1999 гү штотрто Y
POOL BysuAieI0g 87 1999 "ан чә]рәиүэ< y
966] xouueftog 82 119919495 *dsqus VLIJ, хә IM 14949197498 "M
9861 youuefrag 9c MISA zppuenjoopyos Y
£61 чоѕшен-о[ѕәң (6£61 PEN) әтәче 87 a1129 220025 Y
996] PAOAAPIA РТ Al Snaumsups "Y
20661 uosdwoy J, VI 9912$ sni9UuDs Y
(6861 PPren) 9319qe 4 87 ‘qeg 1420058 y
2661 uostueH-do[soH 9c "IMA "га 29 2112397 suaosafns y
9661 xouuefrog :ecop 19319g GEGT UOSSJEISNO 87 IIM NPN Y
2661 uosrıreH-do[soH (6261 apne) 921908 4 87 aupng sauappidí4o ^y
uosjgpA WYA mumua ^y
£61 uosuueH-do[saH (6661 әрпеи) човилен 87 uexo[g “y (qeg) smjofipunios y
9261 PPunəfg gz IYIDA, SNIIDSOL "у
S661 ‘TP 19 Oqnsjem] (6661 орпеу) 9B19q8 4 8c 'uuaog хә әцтәдү snijofiquioua У
єсє] повшлен-Чојвон ec “әри” snjj&udo421ur “rea "uuoog хә IAM DImppa ^M
6261 2261 uossjejsn^?) 9c puny] snijofijsnZup ‘тел *uu20g хә IAM Dgmppa ^w
661 PASUÁTRIOY 5:276] PAOWLY :2061 uos
-шле | -Војвон “6€61 ‘EEGI поѕѕуеіѕпе) (6661 PEN) 92198 4 85 "uusog хә APM ојпрол Y
воопзлојоц uc uoxe[
'penunuo) °1 a]quL
155
Volume 84, Number 1
1997
Thompson
Chromosome Numbers in Rubus
2061 uosureg-do[sag 8c AJT Y “IMA "fd 4ofipidsno ^y se)
Áeqnog (2143j2 Y "IDA `Га) snaofipideno у
6661 uosspeisne) се S99N офа зтјодрло "у
(6261 Apnea) иоѕшең 8c "xo[g snyofipaoo y
6661 uosrueH-do[saH 8z fat aoe Y
2261 човшен-дојвон 8c DADO snssdyros0zyo “Y
6061 чозардепгу 82 ayioy, ғиәоѕәдпа ^y
uospeA, ‘HIM тројооојцо "у
Ep61 uossjejsn^) 8z (uoneuruuajop итеџмозип) 93204 11уәғу/рә су
(6861 грпејј) uosıureH 8c ("әр wou) "пи fd suo су
2261 чогшен-дојвон 8c TOW “fd Y 214919] srmpouadsp су
£€6] човшен-дојвон 9c (wmsneyur "шоп) UOSIPA HIDA 7102р ea шу snoudp ^y
8961 FAO :9261
youueftag :£c6 повшен-ојвон (6261 зрпеџ) uosureg 8c әчтәдү ѕтлојпоао "у
2261 човшен-дојвон 87 aipng suo4figsnup ^y
2061 човшен-дојвон 8c AJOT Y WOW Га Moiua "у
2261 човшен-дојвон 9c (pnu "шоп) uoswy ‘HOA sngojouapp ^y
(6£61 apne) поштен 9c (чең прутодтаи су se)
“IMA "га Y 214919] snjoradimoD y
(51815 oruouoxe) 10/pue виопеитилојор Inpqno() sngny чопәәс̧
2061 чоѕшең-дојѕәң 9c uosjeA, ‘HDA png) прор-Хођот ^y
(6261 apne) эзләде 8c 9x20, хә IMN "fq ana y
2261 ene 8c (33e лојцом synvoypa ^y sv)
UOIMIN ^V sruuofimpoipa u
с861 “TP 19 59146 :с861
sisso[qnq y sords :9e6] хопџоПоц ‘ggg, uosureg-do]
-SƏH (6661 зрпеу) 931998 4 ‘рөт ‘GEGT ‘ELGI uosspersne) 8z aued oid зао Y AM sr y
с661 “Te 19 oqnsjem] 9c BUM Y ти Га 578020212 су
9861 x?uuefteg 061 чәіѕицо) :epop ‘GEGT човајејепо) 8c эчтәдү sn71520 "M
Erol '6261 џоввјејапо) се ISNA) s15u221042]$20 "M
€£61 Ko[duoT 9 моли I SS3JUIOY |, Yueqing “AD "рү Surau ^M
GEGT SUBI) :/261 побише y оџел) РТ sTULIQUI “IBA лә1әлә Srupomsna ^M
#0661 чозашоцј, ‘1961 UN], Y [[oxseH VI 94904 (PILA) Nuur wea nouos snyofrum y
*Op6[ Seu y ‘GE6I UPI) ‘ZEGT eneq VI ләтәләрү smuponsna ^w
#0661 човашоцј, :[96[ un] Y [1958 H :0961
I[?X**H :9661 xouuefreg ‘ecg, повшен-дојвон рөт
BAOURZOY ‘GEGT “ELST UOSSJEISNO) :/261 uoidurpre(] A) oue) її nouos snijofnum y
вәәпәлә}ә uc uoxe[
~ ПУНИ И" "РИМИНИ
гропипио) `I ојдеј,
Annals of the
156
Missouri Botanical Garden
GEGT "2261 uosspeisn?)
€eoT чоѕшең-до[ѕәң
(6£61 OPTEN) човшен
(6661 орпеуј) човшлен
£c61 повишен -дојзон
(6661 орпеу) uosrueH
£C61 чоѕшең-до[ѕәң
GEOL "neq
2061 198198 “0561 siy)
(6661 Apne) 9319qe4
(6661 opney) әйләде
зррбт 5ешоцј, ‘SEGT 9181)
2061 чоѕшшең-дојѕәң ‘2861 SHEW
€b6[ uossjeisn)
(6661 орпеу) 9319qv4
£66] човмлен-дојвон
£c6 1 чоѕшең-Яојѕәң
6661 EEGI uossyeisn)
єсє] чоѕшең-Яојѕәң
€co] vostre y-do[soy
(6£61 opnew) әйләде
966] Pung
20661 uosdwoy, ‘C661 exsu&ra0g ‘gs61 uosrueg-dor
-SƏH (6261 әрпеу) uostueH :2р61 ‘661 “ESOT uossjeisn?)
ec61 uosııreH[-do[s2H]
6261 “££6T uossjeisn?)
9261 ХопџоПод :2261 SHEN
(6261 PPE) әйләде]
0261 меду y 1enbeg “3561 SPW
0961 MPASPH
2261 позвале y -do[soH
0961 ПӘХѕЕН
(z*euar) =) "IMA "fd snirudongrop “y
(xe uyun) ‘Agua smofimpod
"quaj[ew snunosqo
snunumjsipijjod ‘тел uos "Wr 2A (pns) 571551172590
snununjsipijod “IBA uosjeA "WA (олрпб) snunssunosqo
uosjEA ‘HIM (91png) smuissunosqo
Y
Y
(ə1png SISUIUDUOUII “Y se)
(edA101q [eoo] "ләцәс) 1 11919491
Y
(влодом ‘NM snuDÁa] “y se)
Зи Y TON ‘fd чојхоирјош
Y
"[PS-II9g машод ^M Se
(шлој порлед) "eS -[19g 142440Q “y se
(шлој порлед) "ләпәс) хә пи ‘fq поџидош “y
"IMA “fd $64201$042014
“ләпәсу sap10]¡4ydo1>0w
a1png sousporda]
әтәдү 1ounalo]
(xe шеџмооип) Yos 1у2рдиг роу
&enog хә “IMA ‘fd 81809499711
‘Bury хә “TDW "fd Dasu
TOW “fd Y OYT зтјојиави
ayra xus
uosjEA "WA (әрп) гпрраовтрллоу
Y
бе cd Бе ба nd à ee
Y
пуордио оу “eA YSW тугофиотру “dsqns "wy W SPPA MY Y
"98е IY SPPA питу
Y
"IMA ‘fd Y 2143197] тојтиола y
nealog 11210222 су
ayro snosnf Y
әчтәдү 12002 Y
әллә}ә JS TNW ‘fd трудошкар
(‘xe uyəun) sap10/£ydÁsop
Y
Y
воополојоц
џохвј,
penunuo) “| әј,
157
Volume 84, Number 1
1997
Thompson
Chromosome Numbers in Rubus
о
5661 TE 19 oqnsjem] 8с "РА пне и
2261 човшлен-дојвон Тр si9B0Y ‘AM тошиотад “Y
2261 човшеН-дојвон 9c UOSIEA Y'M smupruojZutqpQ ^w
6661 uossjejsn?) 9c puny] snsoynpupjdqns ea плу nsuos ѕлѕогошәи ^M
6€6[ uossjejsn^) 9c "jut ре таг оомодпу ‘тел злу nsuos snso40uigu ^M
6€61 '££6[ uosspejsne) 9c РИМ Y 9u&eg uou “yuy nsuas snsosowəu "M
Ep61 uossjejsno 8c puny "AY Y snsompupj2qns ва зәриеүү snjooan ^y
Ep61 човвјејепо) 9c пепео) па гооомодп °} зәрирүүү snjoaann су
Ep61 uosspejsn?) ‘Tp6T зәрирүүү 9c зәривүүү ѕлјоәто су
2061 uosuueg-do[sag SE qeu) snonajouapp "у
"purT 170/107) uonsag
2261 човшен-дојвон Ic S99N Y ALM S1770 ‘IEA SIN Y AP SUDÍ MA 'у
ЄСЄТ повшен-дојвон (6261 epneyy) uosuueg 8с aupng snsapdsijpoa “y
(6261 орпеу) повџлен 8с TOW га smivusun y
aıpng sruuofiuisun Y
2261 човшлен-дојвон 37 әлрпс (uedry) стуфошолојоу ‘rea Hoyo стјодшт су
ЄСЄТ човмлен-дојвон 56661 epnejy) әҷтәд\ 9c UM стао «у y
2261 човшеј-дојвон 9c 9x90, тутора M
2961 чоѕшең-до[ѕәң 8 TOW “fd Y gog smjomazás y
2961 uostuey-do[soy 986 SO понор Y
(£b61 џоввјејепе)) 19pue][y 86 SNUDUALD ^j stoumno?) 29 loq xo әцтәд\ suadaos “y
9261 хоиџоПод 9c “ә хә aytayy suadias “y
£S61 Uostuey-do¡soy се (xe итеџооип) олрпб sippprumakdnuos "у
2061 чоєшең-о[ѕәң Ge eipng snsopnpunp#isuo] "re^ o1png snuvdjapnuyos су
(6661 Apne) иоѕшең 8c "Wei хә ILM 11249121408 "M
6661 uosuueH-do[so Se "IMA “fd 575019028 "у
(0661 apne) човзшлен ср png snjjopunjoa ^M
8961 лову! :9261 xouusfrag 8c SUA P ти Га sn mau "у
Ep61 човвјејепо) Ic S99N Y YA snijofruumya у
(6661 9pnejy) позилен 8с (әчтәдү 1274204 "у se)
39904 =1=йоәрш9ха y
2061 човшен-дојвон 87 "IMA ‘faq $77/4ydopod y
(6661 opnejy) uosureg 9c (олрпб зтрио “y se)
'^9ua^) трАвоуиа ^w
$393U9.19J9M uc џохвј,
грепипиођ сәер
158
Annals of the
Missouri Botanical Garden
6661 чоѕеуе\впе) 82 sng зтшлоћотоја °} (палодуурт x ѕтѕәро Y)
EPG uosspejsn?) 87 ЛАН ^H хә sng sruaofionp8 “y
6261 uossJesn4) gz пешпә} хә "pu opruoyf y
2661 “Te 19 oqnsjem] 8с 1999 “YH 4029049f "У
србт Uossje1sno 66-86 182) “y (yosary) snyxruuad у
6661 uossjejsn?) 9c isng “y (yosary) snixruuod “y
6661 UOssfeisns) GE "IMA Та $mapfiqu» y
€V6T '6661 uosspeisn?) 87 IMA [a saunfiquo y
6661 uossjeisn?) cy TEN ‘fd $mapqmorospf y
2261 uossjejsn?) ST JPY] Snpinuqns *j ueumoy зпрохпјо "M
6261 uossjeisn?) 8с Jpr] snpinuqns *j иешпә\ $NIDANJA "Y
6661 uosspejsn^) 87 += виә8ләл snpipuqns pe UBUININ зтохтја "M
6261 uossjejsn) 9c әмва ола ‘puy sruaofiu20 "M
Ep61 uosspejsna) 9c ueumoN $NIDVANJA "M
V661 PYSUAIEIOg 9c ‘аид sısuəuzjop “Y
££6[ uossjejsno) Zp ueunəy (‘qapury) suənu y
6661 uossjejsn?) Gb "рыд холој “y
повцошој (ри) холој *j "әри" ѕирутштѕѕтр ^M
6261 uosspejsne) 9c папе) (ueumoN) sruu0fi2042qns лел "әри" SUD]NUNSSIP ^M
РӨТ ‘6261 uossjeisn?) ср 'puj $71]2ә]ә$ “IBA "әри ѕирутшпѕәр “Y
(2261
uossjejsne) UL Zp se пәлі Ајепоополо) EPG uosspelsnz) 9c ‘sn (zinoqog) sisuaismpq “IBA *qopurT supqnunssip ^M
C661 `үе 19 Oqnsjem] 92 USUAL Y 19q9A “YH sussnuop ^M
6261 чозвуе1впсу ev дәри" ст жудојоло “y
гәдәм “YH туфошојоко "M
с661 ‘Te 19 Oqnsjem] 8% 192 “YH Sadiuobaquos "M
£S61 vostre y-do]soH се UOSIEA ‘HIM simpor2ndamd y
£c6 1 човилен-дојвон се ѕләдоу ‘NM (qeg) suadunfuoo су
2261 чоѕшең-Яо[ѕәң 9c sio3oy "WM (qeg) suadunfuos y
(661
uossjeysn‘) ш Zp se џолта Ајвпоополло) epo] UOssje]sne) 32 “sng ("цоволу) snaopipu “IBA "qopurT S1101110 “Y
6561 UOssje1snt) 0€+ "qepurT snpipo “Y
6261 uossjejsn^) gz q9pur] snmp 'у
сс61 uossjejsn^) cv UBUININ SUITIIMIP ^M
uneig `©) SÁY9DISOIdUIDO ^w
s3DU919J9Y uc похве],
"ponunuo) сү QPL
159
Volume 84, Number 1
1997
Thompson
Chromosome Numbers in Rubus
ee А
2761 '6g6[ uossjeisnz) се “sng ^y хә sng (ившпә хә uəsuəf) rZuruupa “IRA плу snsoumud ^y
РӨТ uosspisn2) се pur] “AY y mundo ва чу пошта су
19924 “AH (ueumoy xo повиој) MFunusDM *y ‘Чиу snsoumid су
6261 uossjejsne) се “JUL ре ysna snonpapis “IBA ‘yry snsournid ^M
2261 поѕшең-ојѕәң се seo] sugsnjgns У
6261 uossjeisne) се snsourad °} ‘цілу sasoumad су
6261 човејејепе) 82 чеп) у (Puy) 010,30 44d y
P66L *suÁmaog 9c ипвд *£) SÁYIDISOYLIO "M
6261 uossyeqsn<) бъ "qapur] sndanoofo] лел *qopur] SNYIUDSOL “Y
Ep61 uosspeijsne) се "qapur] sndap2ofoj ва "qopur] snyrupsos "M
EPL “6E6T uossjeisnz) 86 "qapur] sndzn2ofo тел *qapur] snyruvsos y
чәвләрә Y 19qoA “YH snoFeasou су
2661 човшеН-дојвон SE qeg хә wexojg `y snuvumofjoq y
2961 чоѕшен-до[ѕәң 8с "qeg хә urexo[g “y зтиршпојјод y
€66] ‘TE 19 oqnsjem] 9c "PIA Y Ukey snsosowau y
6561 uossprisn?) ср HPD Y ‘PUY пигзиомош чел "pug sruaofiruoo "M
6661 uossjejsn?) 8с u9po Y PUA ттиә$иәмош чел "pu sruuofiruoo "y
"9189/0 пшоојдриу Y
6661 чозвјејапе) 9c 809 “y (19199) 15s20/pi] xea sruuofituao y
6861 uossjeisnz) SE-8Z 1509 “y (13099) пзмојрђ iea спшојпиго “y
oque] (19199) пзмојру y
6661 повары гу 8с PHA 1011p ^j "qepurg snipgmaaos су
£v61 Човвун1впгу 8с ‘ISN nsopmpunpjdo су *qapur] snpmuəs "у
с661 '[? 19 oqnsjem] 9c ипе ‘9 soympooadum] y
£P6L човвјејеп) 96 "qopur] 12120492] "y
6661 uossjejsn?) 9c пә ^y (pu) sisdoougséy y
2061 човшеН-дојвон SE поѕтв/ HIM (Kor) smuaofimppa y
повзе WIM зигрро роу “y
ер6т “6661 uosspyeisno 9c ueumoN ("цовагу xo лав) толриојоц y
p661 €xsuÁqe10g се 1992M “AH 5755049 "M
Ep61 uossjejsno) 9c Dso4quim `J osnery "Гил хә моје) Y "pup snonjo y
6261 uossjeisno 9c pyAydosnu "у asnery "pu^ xo моје Y "pup топлог су
6261 uossjejsn?) se 9snery “THA xe 19199 Y "pug myo? y
5261 uossjeisn?) 8с "аераг] saipunmumoo "у
6661 повертати) 8с+ 9snery “TH хә 19199) 29 “Puy 521209 y
ЄРӨТ “6£61 uosseisnz) 8с әѕпегу "На хә uepo?) Y "pug sono? y
воополојон uc џохвј,
"ponunuo) ү әре,
160
Annals of the
Missouri Botanical Garden
пдитшлрт SNSIBA “IBA "рид t1pjofuojso ^M
GEOL POSEA) су
GEGE пој 9c "рыд 4әриюрләтш лел "pi прјејново “y
(suoj [290] pow 1чәвәлфәл 'влел) "рың Mpjafuaiso "M
(cg = и se uaaa) p61 uossjersn?) ¿0L "pu (Чәвәгү) sno170q чел "qopur] п2лодл220] y
Ep61 uossjeisnz) Z/SE "pup ("Ҷәѕәлу) snonpoq чел "qopur] 1181291280] “y
egor UOssyeisns) 6b "цоволу 212709 ‘IA "qopurT 1121204927] “Y
GEDI "asen: 6v OSAT det sai ч
(adAj01q [890]-"S9][1 "шоп) поввлер J, SMUIADU uou ‘иешпәуу ('qosary xo "T) snunxow ^M
6261 uossjeisn^) DEF (edAjo1q [eoo] "цовогу sipaopna y
АН `н хә лепо snipuaoqput "M
Ep61 uossjeisnz) 9c ‘uey про Y
2961 SPIN SE equ whup Y
Z£61 "neq се "33e “wg 5120/1100 “Y
ve61 кој 8ио 82 "33e "ug smofi] 4109 Y
es61 човшен-дојвон се "Чеч попајоигро “y
6961 uossjeisne) Тр ¿C3pr]) supsauagoud xea *qopur] snm “Y
OEGI повернут су (Jpr]) sisuapana Tea *qopur] snmop Y
Ep61 човвјејепо) 9c UBUININ sruuofiso40uigu “IBA “әри” SNINID "M
Ep61 uosspeisna) 9c UBUIMaN зтшојотиој “IRA "qopur]T snnm ^M
6661 uossjejsn) ev 1oxonewag snjmop uou “3311 "шои *qopurT спор ^M
(snis огшопохр] 10/pue виопвипшлојор ipqnoq) пјојукло uon
€co[ човмлен-дојвон 9c e1png пигшот ^M
ЄБЄТ uossjeisn?) су (; и ходуцтем jo шлој [890]) зтишојмојпзит Y
GEOI мори SE pso1qum "упалу пглодууот Y
6261 човвјејепо) се jene) snd iea му 1181291400 7M
6861 uossje1sn<) c£ "pu snoyfiugpui ва "way 11312q400 Y
#0661 uosduou], ‘EVEL ‘GEGT uossjelsn?) SE "guy 11819q/400 Y
€co[ човилен-дојвон се one snyjupopudu Y
€co[ uosuaeg-do[soH ce 'qeg тојтолодт "M
5561 VOSTRA су a3ue”] snaopapn "M
ON “PH 240198 Y
Ep61 uossjeisnz) 9c лепе) snxifouaq “IBA "рыд amuoy y
uosspew Y зәдә "4H Mans "M
?có6[ uosduiou p су adue’] SISUINASAS "M
6£61 uosspejsn) се "'qapur] sndap20142 ва "qopurT SMYIUDSOL “Y
SODE Пе ефи 8с 1999 “AH snjyeudopriya 7M
воополојоц ug uOXE],
penunuo) сү әв,
161
Volume 84, Number 1
1997
—— ыдын
Thompson
Chromosome Numbers in Rubus
1961 T? 19 1990514
e61 &o[duo]
e61 Хој8ио]
e61 &o[duo
bz61 ^ә[йчот
261 surg
1761 surg
2761 PSU
bz61 Хој8ио]
$261 ој 8иој
2%61 sur
2761 PSU ‘p61 &o[duo]
bz61 4a[3uo]
9961 ПЕН
Y SISPIEY ребг Neg Y Sur “Lp6[ PSU *pzo[ Buo
(sorads Á119qx98]q ивоџошу цио) snqny snuasqng
әле Ape “AD “Ray, трмој у
&apteg "HT зпиштр y
(aued oad) proureag snaps ^w
лоџод ғіиәтиәуәрро “Y
sasuaruaoj]y чоцәәс
OT61 ‘PEGI PA0UEZOY раб Buo ec [odo snorupgsayan] ва *] створо “Y
20661 uosduoy ү :с661
eysu&ie10g “6261 uofxspo3u] ‘T1961 UNI, 9 цежеен :0961
TPASPH :9661 xouuefrog ‘ESEI 123104 ‘OSGI ueisuq)
‘GEGT FWEIBEA ‘LEGI лоромцон “OP6L “PEST PA0UBZOY
"EVOL ‘GEGT ‘EEGI uossjeisn?) ‘ZEGT ENE сб 4ә[8и0] 8c “Т snisono “y
stomog 29 "fo #5207) иопооб
6661 uosspeisnz) cv "pup таи "ea "pug snrpxoa y
(ad&yotq [eo01) ‘pug snipxoa ^y
2061 човшен-дојвон Er AAI] Y "IMA "Га sruofippaquen "у
££6I uossjeisn?) 9c sisuojmy y ‘pug snmma y
€S6[ човшен-дојвон ce 91201 snso1qnos “Y
(6661 зрпеу) uosuueg су алрпб snyjopungoa y
6861 uossjejsnz) FFE (әшви релш) ‘J ueumoy таро y
6661 uossjejsnz) bbs ngunun впвләл “IBA "pu прјејиово “y
воополојом uc џохеј,
гропипиој '[ әда,
Annals of the
162
Missouri Botanical Garden
9161 пеоре X uosee 9S "PILA 514077930) y
1661 19SULH 6b pareuouejg snpnjdoo2 "у
Vc61 Aepsuoy Se попу snso][la ^M
SB6L ‘TP 19 59146 :с861 stssejqnq 29 sordg 9c PILA $140]J020jf y
Vc6l &o[2uo'] 1 qouepg snurdsrusofiq “Y
vz61 9810] су qouepg snunpAyoal y
re61 fojduo] ср qoue[g smupjapunan "y
1961 uosduiou j, 9c soÁe]y unsny pue 'sso[urou], uysny ‘Məq urjsny “sao зарад umuiduoqo у
$210jj39»], | UONI
1%61 195014 9c Koqreg “H” smurissuos “y
C961 ‘TP 19 89146 :с861 sissopgnq 29 59145 8c Kajreg °Н”Т snnospd ^y
ego[ ‘Te 19 saidg :с861 sissajgng y seidg Ic Аорта ^H] 5772500 "y
c861 ‘TP 19 serdg $861 srssojqnq $ 59145 8c ysing snijofiouno ^y
C961 ‘TP 19 вә! ‘gol sissajqnq P 59145 IZ чета snijofiouno “y
c861 ‘TE 19 sardg ‘ger sisse[qnq y 59146 :8961 чешлоцб
#6661 YOUNIS Y гәувшәоцс̧ :9461 ALWAYS $P deys РТ ysing snijofiouno y
тојгипо uonoag
20661 чоѕішоці, ‘0961 FBI) / 61 198014 “PZOL 4919007] Ic "т чзигрриро y
vz61 &opduoT I "рМ прим y
20661 поѕішоці, *pz61 Ao[duoT VI "] чзигрриро "M
sasuappun;) попооб
2761 19814 Ic kojeg ^H] $2020] Y
Хоја ^H] smpian y
1%61 19801 87 kojeg ‘HT sanbiun y
раб Хә[#чот ev "joue[g $upa4moa4 "M
LV6T 198019 ЧЕ Аорта "и 2pupgom y
"цривј4 SUDAINIAL Y
LOT 19surg се (z'u&s) Kojeg "H"] snupipavo ^y
pz61 4918007 Ic "[oue[g SUDAMIAL “Y
Ly6T 195013 87 под snaruvapisuad y
vz61 Aepsuoy Ic qouepg 570040 "y
6561 ошо Y ueuexrepy V 198199 "V snUDISMO] “Y
vz61 91810] бї морәйт snsopuo4f "y
vz61 Kopguo] Ic Mojadig snsopuo "y
раб! 49[8uo] РТ moJo31g snsopuo4f "y
воополојом uc uoxe[
"penunuo) "ү әү,
163
Volume 84, Number 1
1997
Thompson
Chromosome Numbers in Rubus
bZ6L Хә[йчот IZ "qpáy supouZm ^y
bz61 4a2uo7 Ic moyodıg srsojas суу
te61 4a¡2uo7 tl моје ви snsojas "M
1%61 surg v1 әрең ^H] тиоцоод y
Kopeg ^H] тилои y
E61 Хој8ио Ic qoue[g supinoad y
әрең ит 21252 y
2761 surg 8% fopreg "ит пзгиоу Y
re61 Хој8ио tz qoue[g smporpupj2 су
LOL su Ic Aayieg и snsuassip ^y
2761 SU 8с Хота “Н” типог y
Kopreg ^H] спипзгриојо y
1502€ uonoag
vz61 Хојвиој Ic qoue]g сторло су
EVOL 1980: tz Kopeg HI Mus y
Zvó61 19809 tz Kopeg ‘H”] snjd y
2761 19814 87 Кәпе ‘HT пиопту y
vz61 Хә#чот 9G о етра у у
vz61 Хој8ио се 1 snpidsny y
90661 поѕішоці, :996[ Пен Y SIPPY vt "| snpidsiy y
ipidsigj чоцәәс
2761 PSU £9 ('wouerg стјојироуа -y se)
Хора ^H] sums y
1961 uosduioq, бӯ (enesony 'ло ‘ued ш “ирәЯоу y se)
qpÁy (&opeg HT) 2200101 y
рабт Хој8иој VI yourlg SNDNAMI "M
vz61 Aa13u07] [42 qoue[g стоглотиов “y
"joue[g sm70/i21]d y
vz61 fauo СЕ "youe[g стујојироца y
Уб PRI Se Хота 'H”] $72D4092a0u су
vz61 Ko[pguoq Ic "youe[g sruuofiynu y
2761 19814 б? Kopreg ^H] =пәрләш су
2761 198014 59 ТРИ 5407920 y
ваополоје uz
‘рәпициог) сү ejqe]
Missouri Botanical Garden
Annals of the
164
ec661 uosduou], ‘1861 шелвиј 9 ALA *6p6T “TP 19 SWEM
22661 позва шоцј,
20661 човашоцј,
1861 шелдиј y ова “6£61 UOSsyeisng
20661 uosduiou]
161 4I9 $ pisuerz ‘GEGT Хојаио | ју моле
q 86661
позва шоцј, ‘£661 "useunaey A) одпајемј “gp6] uossjeisn?)
tepo UMOIG ‘TOT 7° 19 19YOSIA ‘EEGI Kojduorq 9 моме(]
‘1961 човашоцј, ‘eGo, eure1eeA :1661 ÁP Y MSUNSIZ
*epo] UMOIG ‘TOT “TP 19 19u9st4 ‘EEGI Ko[duo'] 29 моллв(]
ссбј eureeA ‘q *egpo] seuoup 2261 A9[3u0'] 29 моме
ESOL вшелевдл :epgp UMOIG ‘TOT “TP 19 190814
CEST WEA
q “ec661 vosduoy :8961 ueuuaus :6661 4901
-1MIS Y лозешооцс geo 1әўешәоцс Y odreus :9261 PUA
¿vor 1994
87
puq4y оџеповапс-лоји] јеле pesodosg
(sarads Аллофуовја ueoureury ymos) тту snua3qng
v8
LL
0L
(¿ зтодогорј x snqny) "iuog snn? Y
под snyofinm "У
IN A |
Реис Ч
sejĝnoq snjp1ado42Dui ^M
RSs Y uq ones
PUPS Y wp шып
"Pues Y umo) muun о
"IPHYIS Y “Ryo snyofina ^M
при D Eq mue уу
Kojeg ^H] %122040и080] “Y
"Pues Y We) тилт Y
1111/7 uoroogs
"РАМ sniostuqna "У
"or пон Y
$2]D101410494 uotnjoag
Kopjreg "HI $nupjosouunu "У
Хоја "HT $1suauisuoosim ^M
sooua19JoH
uoxe[
penunuo) ст ARL
NOTICE
THE 1996 Jesse М. GREENMAN AWARD
The 1996 Jesse M. Greenman Award has been
won by Paul Kores for his publication “A system-
atic study of the genus Acianthus (Orchidaceae:
Diurideae),” published in Allertonia 7: 87-220
(1995). This study is based on a Ph.D. dissertation
from Tulane University under the direction of Dr.
Steven P. Darwin.
The Greenman Award, a certificate and a cash
prize of $1,000, is presented each year by the Mis-
souri Botanical Garden. It recognizes the paper
judged best in vascular plant or bryophyte system-
atics based on a doctoral dissertation published
during the previous year. Papers published during
1996 are now being accepted for the 29th annual
award, which will be presented in the summer of
1997. Reprints of such papers should be sent to
Dr. P. Mick Richardson, Greenman Award Com-
mittee, Missouri Botanical Garden, P.O. Box 299,
St. Louis, Missouri 63166-0299, U.S.A. In order to
be considered for the 1996 award, reprints must be
received by 1 June 1997.
Volume 84, Number 1, рр. i 166 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN
published on February 20, 1997.
165
poros and Molecular Approaches to Plant Biosystematics
The proceedings of the Fifth International Symposium of the International Organization of Plant
тање (IOPB)
Edited by Peter C. Hoch and A. G. Stephenson
Twenty-three original contributions that span the breadth of biosystematics, a dynamic field of study
that bridges the realms of systematics and population biology. The papers are arranged in four groups,
reflecting the original four symposia of the 1992 meeting. DNA and Plant Biosystematics presents
innovative work that uses the rapidly developing nucleic acid methods adapted from molecular biology.
Plant Growth Patterns and Biosystematics includes comparative and developmental analyses of plant
architecture and branching patterns. Plant Reproductive Strategies surveys new approaches in the anal-
ysis of plant reproductive biology, an area central to both systematic and population-level studies.
Phylogenetic Analysis and Population Biology emphasizes the application of the powerful new methods
of phylogenetic analysis to problems at the species and population levels. Monographs in Systematic
Botany from the Missouri Botanical dn ics ene. 53. ISBN: 0-915279-30-4. 416 pp. Illustrated.
1995. $60.00 U.S. $62.00 Non-U.S
Annals of the Missouri Botanical Galan: Volume 82, Number 2: Alternative Genes |
for Phylogenetic Reconstruction in Plants
A symposium cosponsored by the American Society of Plant Taxonomists and the Botanical Society of
America, organized by Pamela S. Soltis and Douglas E. Soltis, and presented at the 1993 AIBS meetings.
Although the chloroplast gene rbcL has been successfully used to reconstruct plant phylogeny, many
important questions of plant phylogeny and evolution cannot be addressed using it. The contributors to
this issue of the Annals explore the potential of eight alternative genes or DNA regions for phylogenetic
reconstruction at a variety of hierarchical levels. Both nuclear and chloroplast genes are evaluated. Three
regions of the nuclear ribosomal RNA cistron are explored: the 18S gene, the internal transcribed spacers
(ITS), and the 26S gene. Small multigene families from the nuclear genome may also carry phylogenetic
signal: the phytochrome gene family and the small heat shock gene family. Three genes from the chlo-
roplast genome are also considered: atpB, ndhF, and matK. Each paper describes the location, oe
structure, and rate of evolution of the chosen gene and discusses its potential for Pa ee study.
issue also contains: “The Comparative Pollination and Floral Biology of Baobabs (Ada bor га |
Te by David А. Baum and “In Memoriam: Peter G. Martin.” Annals о 1995. 174 | pages. $27 50.
U.S. $28. 00 Non-U.S.
To order, please indicate method of pani below. dala or or money rs should be- ye. U t funds, L5
payable through a U.S. bank, to Missouri Botanical Garden. Ord ers must be prepaid. $3. 00 handling ў
_ fee on all orders. Additional $2.00 handling fee will be added to
are made until payment is received. Phone: (314) 577-9534, fax: qu PEAK e-mail:
"дер Фтођогоге.
Send order to: ` | 5 Please pale _ copy(ies) of tn No. 53
e woe Missouri Botanical Dicke re Please send – SPs, -copy(ies) of Annals 829),
РО. Вох S
St. ecc Ma 63166-0299, U 3 А.
D Check/money order enclosed E Tad se Se sd zi |
O Send invoice ($2.00 fee will be added to total) e eee
O ae card number (MasterCard/Visa) |
Name |
Ennio dne | grece НЕ a 75 Address |
Name as it appears on Уа
Telephone number ( daytime) -
Postal Code Country
zi ы ARE SUBJECT TO CHANGE WITHOUT NOTICE — UL BAY:
CONTENTS
is, Pamela S. Soltis, Daniel L. Nickrent, Leigh A. Mum
William P EUR Sara B. Hoot, Jennifer А. Sweere, Robert K. Kuzoff,
Kathleen A. Kron, Mark W. Chase, Susan M. Swensen, Elizabeth A. Zimmer,
Shu-Miaw Chaw, Lynn J. Gillespie, W. John Kress & Kenneth J. Sytsma -
Tribal Relationships in the Gesneriaceae: Evidence from DNA Sequences of the ;
Chloroplast Gene ndhF ... M
ANE SEE Be E Smh ДС. Wolfram, K. D. Brown, C. L. Carroll & D. 5. Denton. 50 A
A йкы Conspectus of the Tribe Juanulloeae (Solanaceae) “Pek
andra Knapp, Viveca Persson & Stephen Blackmore wee id
A Hevision of. the Genus Xiphotheca (Fabaceae) —-- Anne Lise Schutte 90
A Review of the Genus Eccremocarpus (Bignoniaceae) -.................... William G. D'Arcy | 103
дейт "uiris. for Conservation of Plant Biodiversity in the Mediterranean Basin — — 3 ,
Frédéric Médail & Pierre Quésel 12.
“Survey о of Chromosome Numbers i in Rubus em: Rosoideae) В
Махіпе М. Јата ў,
Angiosperm Ман Inferred from 18S Ribosomal DNA Sequences
оив
ts ^A ters Notis: e
Annals
of the
Missouri
Botanical
Garden
1997 ҸҸ
Number 2
Volume 84, Number 2
Spring 1997
Annals of the
Missouri Botanical Garden
The Annals, published quarterly, contains papers, primarily in systematic botany,
contributed from the Missouri Botanical Garden, St. Louis. Papers originating out- |
side the Garden will also be accepted. Authors should write the Managing Editor -
_ for information concerning arrangements for publishing in the ANNALS. Instructions
to Authors are printed in the back of the last issue of each volume.
Editorial Committee
Henk van der Werff
and Michael H. Grayum
Co-Editors (this issue),
Missouri Botanical Garden
Amy Scheuler McPherson
Managing Editor,
Missouri Botanical Garden
Diana бош
Editorial Марне
Missouri Botanical Garden
Vicki Couture
Secretary
Teresa Johnson
Publications Order Processor
- Missouri Botanical Garden.
| Peter Goldblatt
Р. Mick Richardson
Missouri Botanical Gar den
Ihsan A. Al-Shehbaz
Missouri Botanical Garden
"Gerrit Davidse
Missouri Botanical Garden
Roy E. Gereau
Missouri Botanical Garden
Gordon McPherson |
Missouri Botanical Garden
For subscription information contact Depart- -
ment Eleven, P.O. Box 299, St. Louis, MO
63166-0299. Subscription price is $110 per
volume U.S., $115 Canada and Mexico, $135
all other countries. Four issues per volume. The
journal Novon is included i in the Каин |
рпсе of the ANNALS.
amcpherGadmin. mobot.org таш со
< . deptllGmobot.org (orders) |
—.. http://www.mobot.org
© Missouri Нови Garden 1997
` The mission of the Missouri Botanical Garden is to dud and vere knowledge а about oe |
their environment, in order to geste ium enrich m T
The ANNALS OF THE MISSOURI BOTANICAL —
- GARDEN (ISSN 0026-6493) is published quar-
Tower Grove Avenue, St. Louis, MO 63110. Pe _
t5 Hox 299, E Louis, MO e
terly by the Missouri Botanical Garden, 2345 —
riodicals postage paid at St. Louis, MO and ad-
ditional mailing offices. POSTMASTER: Send ad- 2x]
dress changes to ANNALS OF THE MISSOURI | |
BOTANICAL GARDEN, Department Eleven, РО. T
Volume 84
Number 2
1997
MISSOURI BOTANICAL
JUN2 7 1997
GARDEN LIBRARY
Missouri
Botanical
Garden
THE GENUS LYCIANTHES
(SOLANACEAE) IN
VENEZUELA!
Carmen Benítez de Rojas? and
William G. D'Arcy?
ABSTRACT
The 11 species of Lycianthes that occur in poten are described and distinguished with illustrations, a dichotomous
key, and notes on their appearance and ran,
he separation of Lyciant
nthes from Solanum and its closer relationship
to Capsicum are reviewed in light of paria ine and recent molecular evidence
The genus Lycianthes (Dunal) Hassl. includes
150-200 species of herbs and shrubs that range
throughout the Neotropics and in southeast Asia. A
few species have marketable fruits, and one species
is sometimes grown as an ornamental, but otherwise
the genus is largely unnoticed. This paper reviews
characteristics of the genus Lycianthes and revises
the species that occur in Venezuela. Photographs of
some of these plants are presented in Figure 1.
Salient features of the flowers of the Venezuelan
species are shown in Figure 2.
Lycianthes was long included in the large genus
Solanum,which is similar in its flowers and fruits
and also in its poricidal anthers. Solanum lycioides
L., a species with seeds enclosed in sclerenchyma,
was separated as the genus Lycianthes by Hassler
in 1917. Soon after, Lycianthes was monographed
by Georg Bitter (1920) who expanded the genus to
over 100 species, including many lacking scleren-
chyma in the fruit. Acceptance of the genus has
been hesitant, but most botanists now recognize it
as distinct from Solanum. D'Arcy (1986) studied
the calyx in Lycianthes, Capsicum, and other rela-
tives and concluded that the unusual calyx warrants
generic recognition. He supported Bitter's view that
Lycianthes is more closely related to Capsicum than
to Solanum. This was accepted by Barboza and
Hunziker (1992), Dean (1995), and others. More
recently, a classification of the Solanaceae based
on chloroplast DNA (Olmstead et al., in press) has
! This study was conducted with binational support, and we eer: певане үс funding from the Consejo Na-
cional de Investiga
(INT-9116039) of the United States.
Such herbaria in Venezuela were CAR, IRBR, MER, MERC, МЕКЕ
N, MA, MO, NY, P. US, W, and WIS. Thanks are - pta to Bruno
usly prepared our асно. м Richard e datos (MO) for photographic prepa
ewed the manuscript; p (B
other countries BM, BR, , CORD, F, G, K, LIN
Manara, who gracio
Nee (NY) entered into У helpful discussions and
en ете!
ciones Cientfficas Tecnólogicas (CONICIT Р1-56) of Venezu
Many herbaria made their facilities ratio or sent specimens
on loan for st tudy.
F, MY, PORT, TFAV, UCOB, VEN, VZU, and in
M) also зае и. ЕПеп
Dean (University of California-Davis) kindly biu an visit version ари" the manusc
? Facultad Agronomía, Universidad Central de Venezuela, Maracay, e ей 4597, Aragua, Venezuela.
3 Missouri Botanical Garden, РО. Box 299, St. Louis, Missouri 63166, U.S.A.
ANN. MISSOURI Bor. GARD. 84: 167-200. 1997.
168
Annals of the
Missouri Botanical Garden
Figure 1.
Selected Lycianthes species. A, B. Lycianthes pauciflora.—A. Fruits and calyx.—B. Fruiting calyx.—C.
Lycianthes asarifolia. Flowers and foliage.—D. Lycianthes pauciflora. Seeds.—E. Lycianthes lycioides. Pyrene
S. A, B.
After Benítez & D'Arcy 3271 (MO, MY). С. After D'Arcy 16261 (MO). D. After Benítez et al. 5148 (MY). E. After Nee
17760 (MO).
supported the separation of Lycianthes from Sola-
num and placed it near Capsicum.
STONE CELLS
Some solanaceous fruits contain sclerenchyma in
the form of stone cells or sclerocytes, which com-
monly do not enclose the seed. These are aggre-
gates of secondarily hardened parenchyma cells or
sclereids. Bitter (1911, 1914) examined many So-
lanaceae, and he found them in various berries of
Lycianthes, Solanum, and other genera of subfamily
Solanoideae. He hypothesized that they might be
relics from hard walls of ancestrally capsular fruits.
Danert (1969) used electron microscopy to examine
their genesis in Solanum. Stone cells have been
used in the Solanaceae as taxonomic clues (Schil-
ling, 1981), but there is still poor confirmation of
their constancy in taxa of different levels. They may
e diagnostic in some Lycianthes taxa, notably L.
lycioides, but are variable in others, such as L. pau-
ciflora. Their presence or absence was noted by
Bitter, Barboza and Hunziker (1992), and Dean
(1995) in their species descriptions.
THE САГУХ
In bud, the calyx in Lycianthes and Capsicum 1s
fused to the top (complete prefloration), and the
corolla, stamens, and other interior parts le
from it by a stretching of the calyx apex. This re-
sults in a truncate margin subtended by a thin
sleeve of tissue with reduced vasculature. Low in
the calyx, the five principal calyx traces each di-
verge into three traces, much as in a foliage leaf,
and the adjacent lateral traces are fused together,
resulting in a calyx with 10 nerves. Of these, ~
primary ones are produced by the continuation 0
the principal calyx traces, and 5 are produced sec-
ondarily by fusion of the adjacent lateral traces. In
Volume 84, Number 2 Benitez & D’Arcy 169
1997 Lycianthes in Venezuela
Figure 2. Closed and open flowers of Venezuelan species of Lycianthes. Left figure: olei fd bud ботов Mo re
Right figure: open flower showing corolla lobing and relative size of stamens.—A. Lycianthes f
cianthes pauciflora.—H. Lycianthes stenoloba.—1. Lycianthes ;
diata. aoe 6057 (VEN); к After Rutz-Terdn 12533 (MY); C. After Bunting 4433 (МҮ); D. pot
122959 (MO); E. After Liesner et al. 7803 (VEN); F. After Ru&-Terán 64 (МЕКЕ); С G. After Ferrari 733 (MY); H.
Mocquerys 978 (VEN) L After Davidse 18899 (VEN); J. After Steyermark 121528 (VEN); К. After 56628 (MY).
170
Annals of the
Missouri Botanical Garden
many cases, some or all of these 10 traces enate or
bend outward to form umbos or teeth laterally on
side of the calyx and below the apical sleeve area.
Thus, in Lycianthes calyces, all teeth or umbos are
lateral and arise below the calyx apex, and no pri-
mary teeth are found at the top of the calyx. These
structures are shown in Figure 2, and their vascu-
lature was illustrated by D’Arcy (1986).
The extent of lateral vein enation is variable, dif-
fering in different species, sometimes in different
flowers on the same plant, and sometimes in dif-
ferent veins on the same flower. In different spe-
cies, there may be no teeth at all or any number
from 1 to 10. When there are 10 teeth, they often
form two series, the 5 teeth arising from the primary
traces larger than those arising from the fused lat-
eral traces. In Solanum, calyces have as many teeth
as corolla lobes or stamens, usually 5, rarely 4 or
6. Having 4—6 calyx teeth is quite unusual in Ly-
cianthes.
The calyx in Lycianthes, thus, has three features,
thought to be specializations, that are not found in
the calyx of Solanum: (1) complete prefloration and
egress of interior parts by stretching rather than
bending out and tearing, which produces an entire,
thinned, apical sleeve; (2) fused adjacent lateral
traces of the ancestral lobes; and (3) enation of the
traces in many species to produce umbos or teeth.
However, because they are so alike in other ways,
Solanum and Lycianthes are assumed be derived
from a closely related ancestor.
Whether Solanum and Lycianthes arose indepen-
dently from a common ancestor or Lycianthes arose
from within Solanum is not revealed in morpholog-
ical studies of the calyx. The trees used by Olm-
stead et al. (in press) to illustrate their classification
of the Solanaceae depict one clade leading to two
phylads (Lycianthes/Capsicum and Solanum/Jalto-
mata), each with taxa displaying both porose and
longitudinal anther dehiscence. All the porose-de-
hiscent Solanaceae except the enigmatic Triguera
of Gibraltar and nearby areas are thus contained in
this single clade, and the question remains whether
porose dehiscence was derived twice within the
clade or was derived once and lost twice.
In Capsicum, although calyx prefloration is com-
plete and egress of the interior parts is by stretch-
ing as in Lycianthes, the sleeve is smaller and less
evident, and the lateral traces do not fuse and do
not enate to produce secondary umbos or teeth.
Thus, Capsicum has only two of the three special-
izations noted above for Lycianthes: the complete
prefloration and stretching egress of the floral parts,
and the enation of primary traces to produce teeth.
The fusion of adjacent lateral veins and enation of
a second series of teeth is hardly indicated or not
present at all. Capsicum calyces have 5 teeth; Ly-
cianthes calyces seldom have 5 teeth, they usually
develop either 10 teeth or fewer than 5.
In his morphological study, D’Arcy (1986) sug-
gested that floral parts in Witheringia have a basi-
cally similar way of egressing from the calyx and
that this genus is probably closely related to the
Lycianthes/Capsicum group, but the recent study of
Olmstead et al. (in press) gives a more distant
placement.
OTHER CHARACTERS
Other characters are also useful for recognizing
Lycianthes, although they are not diagnostic. Leaves
are always simple and entire, most anthers are yel-
low or orange, and, with the exception of Lycianthes
lycioides, seeds are discoid with the embryo coiled
around the edge of the seed and surrounding the
endosperm as is usual in seeds of the Solanoideae.
In most species (all those found in Venezuela), in-
florescences are fascicles or solitary flowers and
lack peduncles; anthers open by small terminal
pores, corollas are rotate, and the fruit is an orange
or red berry. Seeds are numerous, except in L. ly-
cioides. In most characters, plants of Lycianthes are
much like those of related genera such as Solanum,
Physalis, and Capsicum.
Diversity of corolla shape and androecium ap-
pearance is shown in Figure 2.
Hair TYPES
Several hair types, simple, branched, and stel-
late, are found in the indumentum of Venezuelan
species of Lycianthes, and these tend to have tax-
onomic correlation and diagnostic value at the sec-
tional level. In the Venezuelan species, all hairs are
uniseriate, and glandular hairs are not found. Sim-
ple hairs are usually several cells long with little
noticeable differentiation. In section Simplicipila,
coarse and tawny simple hairs give a strigose ap-
pearance. In Lycianthes acutifolia, a basal expan-
sion, not unlike that found in many Boraginaceae
(pers. obs.), gives the simple hair on the leaf lamina
the appearance of arising from a translucent foot.
Many species have only simple hairs. Species with
branched or stellate hairs often also have simple
hairs. Hairs with the radii all arising at one point
are termed stellate; when radii arise along the stalk
they are referred to as branched (Roe, 1971; Seithe,
1962; Haegi, 1991). On some parts, especially
abaxially on the corolla, hairs may be reduced to
simpler, difficult-to-interpret types.
Seithe (1962), in her study of hairs in Solanum,
Volume 84, Number 2
1997
Benitez & D’Arc
Lycianthes in Venezuela
171
examined 25 species of Lycianthes. Among these
were six that occur in Venezuela: amatitlanensis,
asarifolia, acutifolia (conicibaccata), lenta (vari-
ifolia), radiata (goudotii), and stenoloba (stephan-
ocalyx). She concluded that the elaboration of hair
types in Lycianthes evolved independently from that
in Solanum. She hypothesized that in Lycianthes
stellate hairs arose from branched hairs, unlike in
Solanum where they arose from gland-tipped hairs.
Unfortunately, she did not distinguish between the
simple, eglandular hairs (Fingerhaare) character-
istic of many species of Solanum and the coarse,
tawny strigose hairs of some Lycianthes, which seem
quite different in nature. These simple hairs in Ly-
cianthes may also be independent in origin from
those in Solanum.
UsEs
Fruits of some species, Lycianthes asarifolia in
Venezuela and elsewhere, are sometimes eaten, and
some, L. moziniana (Dunal) Bitter and others in
Mexico (Williams, 1993; Dean, 1995), even reach
fruit markets. Lycianthes rantonnei (Carriére) Bitter
is grown as an ornamental in upland tropical gar-
dens. No other uses are known.
CHEMISTRY
Lycianthes has a suite of alkaloids comparable to
other genera of Solanaceae (Roddick, 1986), but
few species have been examined (Bradley et al.,
1978; Evans & Somanabandhu, 1980; Lin et al.,
1987; Ripperger & Porzel, 1992), none of these
species occurring in Venezuela.
SYSTEMATICS OF LYCIANTHES
Lycianthes was divided into a hierarchy of sub-
genera, sections, and series by Bitter (1920), only
some of which are represented in Venezuela. The
subgenera and sections in Venezuela are amply dis-
tinct and not likely to be confused. Within the sec-
tions, however, distinctions are often difficult to
make, and species limits are sometimes poorly
known. The type species, Lycianthes lycioides, has
fruits with eight or fewer seeds that are enclosed in
sclerenchyma as pyrenes. It constitutes a perhaps
monospecific subgenus and section. In the other
four sections, seeds are numerous and not enclosed
in sclerenchyma. Section Asaropsis includes one or
perhaps three species of ground creepers. Section
Simplicipila includes perhaps a dozen species of
weak shrubs with strigose hairs. Section Polymeris
includes many species of night-blooming climbers.
Series Oligochondra and Virgatae are separated by
presence or absence of stone cells in the fruit.
Subgenus Lycianthes (L. lycioides): seeds enclosed
as pyrenes.
Section Lycianthes (L. lycioides): erect, woody
shrubs; flowers diurnal; stamens unequal (3+2).
Subgenus Polymeris (Dunal) Bitter: seeds numer-
] mec
ous, not enclosed in
with stone cells.
Section Asaropsis Bitter (L. asarifolia): plants
procumbent; leaves cordiform; flowers solitary,
diurnal; stamens equal; stone cells wanting.
Section Polymeris: shrubs or climbers; flowers
fasciculate, nocturnal; calyx teeth 10 in 2 un-
equal series; stamens unequal (4+ 1); stone cells
often present.
Series Oligochondra Bitter (L. ferruginea, L.
pauciflora, L. stenoloba): plants high climbing
or erect, small shrubs; flowers fasciculate, noc-
turnal; fruits dangling, stone cells often pres-
ent.
Series Virgatae Bitter (L. lenta, L. sanctaemar-
thae): plants low climbers; flowers fasciculate,
nocturnal; fruits dangling, stone cells wanting.
Section Simplicipila Bitter (L. acutifolia, L. ama-
titlanensis, L. inaequilatera, L. radiata); sub-
shrubs; flowers fasciculate, diurnal; stamens
equal; fruits held erect, stone cells present or
not.
GEOGRAPHY OF LYCIANTHES
Lycianthes is confined to the Neotropics and to
southeast Asia. Most of the species and most of the
diversity is in the New World, with distinctive
groups in Mexico and Central America. Section Ly-
cianthes (L. lycioides) is restricted to South Amer-
ican uplands. Section Asaropsis includes one (or
two) species that grow in lowlands in eastern South
America. Lycianthes asarifolia, of this section, was
recently reported as introduced into Texas (Darwin
& Feibelman, 1991). This section may also em-
brace L. lysimachioides (Wall.) Bitter, a wide-rang-
ing southeast Asian species that has similar growth
form and anthers, but which has well-developed
teeth on the calyx. Section Polymeris and its similar
series Oligochondra and Virgatae are widespread
at lower and middle elevations in South and Central
America and the Antilles. Section Simplicipila has
a similar range but is not known from the Antilles,
and it may not range north of Nicaragua.
SCOPE OF THE PRESENT STUDY
Both authors have studied plants of Lycianthes
in various countries for more than two decades, but
the present focus that led to this paper dates from
172
Annals of the
Missouri Botanical Garden
funding from binational grants in 1991. This per-
mitted three extensive field trips to cover almost all
parts of Venezuela thought to host Lycianthes spe-
cies, also study of a large suite of borrowed her-
barium specimens by both authors at the Missouri
Botanical Garden, St. Louis. Many other smaller
field trips were made in Venezuela during this pe-
riod, and other herbaria were visited. Many of the
species were grown for observation in the St. Louis
greenhouse.
Considering the large range sizes of some species
and frequent occurrence in disturbed habitats, the
number of specimens in herbaria is gga ком
low (D’Arcy, 1973: 632; Nee, 1981). This
meant few observations and scant data in some
cases, leading to hesitancy in assigning species
names and in documenting ranges outside of Ven-
ezuela. Despite our efforts, the genus remains se-
riously undercollected in Venezuela, and further
work should enhance knowledge of the genus.
LYCIANTHES in Venezuela
In Venezuela we record 11 species of Lycianthes.
They include procumbent herbs (L. т.
erect shrubs (1. lycioides), arching subshrubs
amatitlanensis, L. inaequilatera), and climbers ^s
remain in the understory (L. lenta) or climb to the
canopy (L. pauciflora). All are perennial. Some spe-
cies have diurnal flowers that generally lack scent,
while others have fragrant flowers that are open at
night. Night scented flowers have been observed to
open just before dusk and to close after dawn. Most
species are found at low elevations, but some spe-
cies (L. ferruginea, L. radiata, L. acutifolia) range
into cloud forests at upper elevations, and L. ly-
cioides is found up to 3500 m elevation. Lycianthes
asarifolia and L. lenta are often adventive in sec-
ondary habitats, and the latter, circum-Caribbean
in distribution, is often found near the sea.
KEY TO SPECIES OF LYCIANTHES in Venezuela
TAXONOMIC TREATMENT
Lycianthes (Dunal) Hassl, Annuaire Conserv.
J Bot. Сепеуе 20: 180. 1917. Based on
Solanum sect. Pachystemonum subsect. Ly-
cianthes Dunal, in DC., Prodr. 13(1): 29. 1852.
Solanum series Meiomeris Dunal, in DC.,
Prodr. 13(1): 29; 156. 1852. Solanum subg.
Lycianthes (Dunal) Bitter, Bot. Jahrb. Syst. 54:
424. 1917. Otilix Raf., Med. fl. 2: 87. 1830.
Solanum sect. Lycianthes (Dunal) Wettst., in A.
Engler & K. Prantl, Nat. Pflanzenfam. 4(3b):
22. 1891. TYPE: Solanum lycioides L.
Parascopolia Baill., Hist. pl. 9: 338. 1888. TYPE: Para-
scopolia acapulcensis Baill. — Lycianthes acapulcen-
sis (Baill.) D'Arcy.
The name Lycianthes is conserved against Otilix
Raf. and Parascopolia Baill.
Perennial herbs, shrubs, or vines; pubescence of
simple or branched hairs. Leaves often geminate,
simple, entire, membranous to coriaceous, petio-
late. Inflorescences mostly solitary flowers or fascic-
ulate at leaf axils. Flowers diurnal or nocturnal, of-
ten only a few opening at a time, sometimes showy,
mostly 5-merous; calyx cupular, apically truncate,
10-пегуед, the nerves often enated into lateral
teeth, pubescent or glabrous; corolla white, blue, or
purplish, mostly rotate or reflexed, apically suben-
tire or deeply divided; stamens equal or not, an-
thers yellow (the Venezuelan species), held together
in a cone, rarely connate, the pores mostly minute
and terminal, rarely elongate, adaxial slits; ovary
spherical or conical, style slender, mostly glabrous,
stigma capitate, sometimes bilobate; ovules many.
Fruit a fleshy or juicy berry, mostly red, orange, or
yellow, usually globose or nearly so, stone cells
(sclerocytes) present in some species; seeds many,
mostly discoid, light colored, testa reticulate; em-
bryo coiled around the periphery of the testa.
la. Leaves either basally cordate or vereri oblique, minor leaves often present; corollas white, mostly
hai
one opening at a time, ubere
га. Leaves basally co:
peduncles held erect abo
ve the leaves (Asaropsis)
simple; stamens all alike (Asaropsis, Simplicipila).
rdate, glabrous; vanis creeping or procumbent; flowers solitary, deflexed » erect
asarifolia
2b. Leaves conspicuously oblique, pubescent; plants erect; flowers fasciculate, erect on the ao gut
Hong beneath
the leaves (Simplicipi
Calyx teeth — (the 10 nerves slightly elevated and often conspicuous, often splitting pet
to appear tooth-
al
3b. Calyx teeth arb agin surpassing the margin of the cal
4a. Calyx 2 mm long or longer, calyx teeth 2-3 mm long; berry more than 6 mm lon
5a. Corolla more than 9 mm long; fruiting shoals mostly more than 15 mm Нед fruit
oblong to conical (often n 1200 m elevation in Venezuela)..............--------- L. acuti
5b. Corolla less than 9 mm
pr P nere ess than 15 mm long; fruit p (not
above 1200 m siimi in a ем
matitlanensis
4b. “n less than 2 mm long, calyx аі. e than 0.7 mm long; berry 5-6 mm bu ———
|
|
|
Моште 84, Митбег 2 Benitez & D’Arcy 173
1997 Lycianthes in Venezuela
lb. — basally narrowed or truncate, often — minor leaves seldom present; corollas white, blue, or violet,
veral opening at a time, often crepuscular or nocturnal; hairs mostly branched or stellate; stamens un
Coria, Polymeris).
Plants erect, branches rigid, sometimes with spinose branches; leaves small (less than
1 cm wide); stamens with 3 longer and 2 shorter filaments; fruit less than 6 mm
s fewe
(more than 1. pi cm wide); stamens with 1 longer and 4 shorter filaments; fruits more
than 6 mm across; seeds mostly more than 8, the testa entirely visible, not enclosed
in r Ea ма Reape
7a. Calyx lacking teeth L. sanctaemarthae
Tb. Calyx with conspicuous teeth.
. Pubescence of simple hairs; calyx teeth in one series; corolla lobed; fruit-
ing calyx teeth ascending or spreading, almost as long as the Ts ЖАО
L. че
8b. Pubescence of branched or stellate hairs; calyx teeth in two series; corolla
subentire or lobed; fruiting calyx teeth reflexed or spreading, much shorter
than the fruit.
9a. Leaves vera - Open overall with minute stellate hairs, leaf
bases mostly truncate or rounded, leaves near the inflorescences
mostly less than X cm long; fruits less than 9 mm across, lacking
stone cells.. L. lenta
Ob. n uh н. A каба bas I fi n!
dul ог сипеаје, pann leaves commonly more e than 5 cm long;
fruits more 10 mm across, mostly with stone cells.
10a. рем pre? Je жаши with yellowish or reddish brown
the pedicels Бе epa corolla lobes greenish and the
folds white; found between 1200 and 2900 m in Venezuela..
L
Ob. Plants glabrescent, pedicels glabrate, corolla mostly uniform
yellow; found between 600 and 1500 m in pue. vibe
CLAVE DE LAS ESPECIES DE LYCIANTHES DE VENEZUELA
la. Hojas basalmente cordadas u oblicuas; hojas menores generalmente presentes; corolas blancas, abriendo una flor
por inflorescencia, diurnas; estambres iguales; tricomas generalmente simples (Asaropsis, Simplicipü
2a. Plantas rastreras o procumbentes, glabris, flores solitarias, colgantes sobre pedúnculos erectos. y colocadas
arriba e las hojas (Asaropsis) ----------------- L. asarifolia
2b. Plantas erguidas no enraizando en los nudos, pubescentes, flores fasciculadas, erectas, colocadas debajo de
las WR implic ipila).
3a. Cáliz sin dientes, los 10 nervios conspícuos y un tanto elevados L radiata
3b. Cáliz con dientes conspícuos exediendo el margen del cáliz.
4a. Dientes del cáliz iguales o mayores de 2 mm de pame frutos mayores de 6 mm de longitud.
5a. Corola igual o mayor de 9 mm de longitud; fru os oblongos, cónicos u ovoides; pedicelos
= nua mayores de 15 mm de longitud; emi encima de 1500 m de akan en
уаш чо L. acutifolia
5b. Мает E 9 ad de Migkud Ku tos globosos; pedice a MR Hm
de largo; localizada debajo de 1200 m de elevación en Venezue
Ab. Dientes del cáliz menores de 0.7 mm de longitud; frutos 5-6 mm de longitud ........... Li
lb. Hojas basalmente truncas o angostas, con frecuencia — hojas menores гага vez метен corolas blancas,
nic o violeta, abriendo varias por
tricomas STA estrellados o о ramificados фоат, Polymeris).
L. amatitlanensis
se
а мисе frutos menores de 6 mm de ат, coi cn A wr
en pirenos ( (сі Ll
6b. nermes, pelos
mln» inp, he ni (us d ¡adri rbi
а mas que 8, no inclufdas еп pirenos (Polymeris).
Та. Cáliz edentado L. sanctaemarthae
7b. Cáliz con dientes subapicales conspícuos.
8a. Plantas con tricomas simples; dientes del cáliz en una serie; corola con el
borde lobado; en fruto, los dientes del caliz erguidos y casi tan — como
el fruto L. stenoloba
174
Annals of the
Missouri Botanical Garden
8b. Plantas con tri
comas estrelladas if d ; di
„үт. ја |
desiguales; согоја con el borde casi entero о › lobado; en fruto, los dientes
del caliz extendidos o reflejos y menores que el fruto.
Plantas en conjunto pubescentes con pelos estrellados diminutos, hojas
mente truncas o redondas, hojas cercanas de las inflorescencias
generalmente menor de 4 cm de largo; fruto menor de 9 mm de ancho,
sin células pétrea
o
>"
Plantas tas glabrescentes y con escasos pelos estrellados о tomentosas,
ojas basalmente obtusas o cuneadas, hojas de
5 ст de largo; fruto mayor de 10 mm де ancho, con pei con
células pétreas
10a. Plantas pubescen tes, pedicelos tomentosos; corola con los lobos
verdes y el pliegue blanco; localizadas en elevacionales entre
900 L ]
т
1200 је
10b. Plantas glabrescentes, pedicelos glabros; согоја generalmente
con coloración uniforme; localizadas en elevacionales entre 600
1. Lycianthes acutifolia (Ruiz & Pav.) Bitter,
Abh. Naturwiss. Vereine Bremen 24 (1): 453
1919 [1920]. Solanum acutifolium Ruiz &
Pav., Fl. peruv. 2: 33, t. 162, fig. b. 1799.
TYPE: Peru. Muña, Ruiz s.n. (B destroyed, =
F photo 2561).
ae умы (Dunal) Bitter, Abh. Naturwiss.
Vereine Bremen 24(1): Ms 1919 [1920]. i
ора Ан Dunal, іп DC., Prodr. 13(1): 1
1852. TYPE: "ти ш сааш Tovar, Moritz Ме
ре, С-ОС, = Е рћого 6765).
conicibaccata Bitter ex Seithe, Bot. Jahr
313. 1962. TYPE: Colombia. Chiquinquita, Воуаса
(Bogotá fide Seithe), blühend July 1909, Frère Félix
. (holotype, M not seen).
lycianthes xylopiifolia var. intermedia Bitter, Abh. Natur-
ereine Bremen 24(1): 455. 1919 [1920]. SYN-
TYPES: Venezuela. Colonia iie Moritz 345 (BM
en); Fendler 974 (G not se
ићи * lopiifolia var. maxima Bitte Abh. Natur-
wiss. Vereine Bremen 24(1): 455. "1919 [1920].
E: Venezuela. Colonia Tovar, Gollmer s.n. (ho-
lotype, B not seen).
Lyianthes bhoidtnii Bitter, Abh. Naturwiss. Vereine Bre-
n 24(1): 444. 1919 [1920]. TYPE: Colombia. Alto
de С. Vulkan Зогага, Lehmann (В not seen,
= F photo 2581).
Erect subshrub 0.5-2 m high, branches weak,
slender, drying brown, sometimes dark or reddish,
sparingly ascending-strigose; pubescence of coarse
tawny hairs, the bases often expanded and foot-like.
Leaves unequal-geminate, ovate, mostly narrow,
sometimes oblique, 5-7 X 1.5-6 cm, membranous,
above sometimes slightly bullate, paler beneath,
veins 6—8 on each side, plane or slightly impressed
above, elevated beneath, with scattered hairs
above, mostly sparingly glabrescent; petioles 3-15
mm long; minor leaves obovate, X 1-2 cm,
sessile or short-petiolate. Inflorescences solitary
flowers in leaf axils. Flowers downward-directed;
L. pauciflora
pedicels 1-2.3 cm long, pubescent; calyx 24 X
3-5 mm, the cup densely strigose, teeth 10 in one
series, 2-3(—6) mm long, weak, erect at anthesis,
pubescent; corolla white, 8-15 mm X 1-3 cm,
lobed Y%-4 way down, the lobes spreading or re-
curved, pubescent outside, especially distally; sta-
mens unequal, 4 filaments 1.5-2.5 mm long, 1 fil-
ament 2-3 mm long, anthers 2-3 mm long; ovary
subglobose to conical, 1-2 X 0.5-1 mm, style 7-
7.5 mm long, exserted 2.5 mm, stigma subglobose.
Berry yellow to orange, conical, ovoid, or oblong,
6-13 X 4—12 mm, stone cells lacking; fruiting ca-
lyx 4—5 mm long, applied to the berry, pubescent,
the slender teeth erect or spreading, to 5 mm long;
fruiting pedicels to 3 cm long; seeds ca. 65 per
fruit, yellow, flattened, the testa hardened 2 X 3
mm. Figures 2F,
Dentro de la sección Simplicipila esta especie presenta
pubescencia áspera, castafio-amarillenta y con los trico-
mas de base ligeramente expandida, flores solitarias, cor-
olas mayores de 9 mm de longitud; bayas oblongas, cón-
icas u ovoides, sin células pétreas.
The species has coarse yellowish brown pubes-
cence. Many of the hairs have slightly expanded
ases. The mature fruits are conical or ovoid.
Distribution. Venezuela, Colombia, and Peru.
Ravine sides and disturbed sites in cloud forests,
3300 m elevation. (Map, Fig. 4.) Flowering
and fruiting throughout the year.
Representative specimens examined. VENEZUELA.
eral: Alrededor de las Aguaditas, 2 km al
e de la Colonia Tovar, 1970-2000 m, Steyerma rmark &
Delascio 123500 (MO, > Aragua: Colonia Tovar,
Benítez et al. 4236 (MY). Lara: Parque Nacional Yacam-
bú, Fernández 3697 (MY). Mérida: Monte Zerpa, ed
2050 m, Benítez & eem 4609 (MY). Miranda: Silla de
Caracas, quebrada Los Palos Grandes, 1 1700-1800 "i
Morillo & Merced d (МҮ, хе. Tachira: Selva пи
lada, по Quinimarí, arriba d Copas, 2 2500-2800 m ^i
Steyermark et al. 100742 (MY, VEN). Trujillo: Selva nu
оса = (оваа осама а ак
Volume 84, Number 2 Benitez & D’Arcy 175
1997 Lycianthes in Venezuela
Ге.
Figure 3. Lycianthes acutifolia.—A. Flowering and fruiting branch.—B. Opened flower.—C. Fruiting calyx. After
Benitez 4236 (MY).
lada virgen de las montañas de Misisí, carretera Trujillo- Dist. Perijá, on international boundary, headwaters of río
Boconó, 2200-2400 m, Steyermark £ Manara 125326 ^ Guasare, 2700-3300 m, Wood & Berry 85 (VEN).
(VEN). Yaracuy: Sierra de Aroa, 9 km W de San Felipe,
1100-1500 m, iden & González 10038 (VEN). Zulia: 2. Lycianthes amatitlanensis (Coult. & J. D.
176
Annals of the
Missouri Botanical Garden
Figu Lycianthes acutifolia.—A. нне
НЬ a outside of Venezuela.—B. Geographic distri
bution in Venezuela.
Smith) Bitter, Abh. Naturwiss. Vereine Bremen
24(1): 441. 1919 [1920]. Solanum amatitla-
nense Coult. & J. D. Smith, Bot. Gaz. (Craw-
fordsville) 37: 420. 1904. SYNTYPES: Gua-
temala. Ата ап, Barranca de Eminencia,
А ee 1457 (F, = F photo 49339,
GH); a Verapaz: о Tuerckheim
7753 ey US), 8488 (C, NY.
Lycianthes ulei Bitter, Abh. Naturwiss. Vereine Bremen
— эу 1919 [1920]. TYPE: Brazil. Ule 9764 (В
= Е photo 2592, МУ).
Lycianthes ahs subsp. dolichodonta Bitter, Abh. Natur-
wiss. Vereine Bremen 24(1): 438. 1919 [1920].
TYPE: Ecuador. Balao, Eggers 14409 (A, M not
seen,
Lycianthes ulei var. strigulosa Bitter, Abh. Naturwiss. Ver-
eine Bremen 24(1): 438. 1919 [1920]. TYPE: Ec-
uador. S. Miguel, Sodiro 114/46 (B not seen).
Subshrub 1-2 mm long, erect or arching and wand-
like; stems terete, puberulent; pubescence of coarse,
simple, erect, or ascending hairs, often suffused with
aves un -geminate, ovate, 14-23 X 4—
8.5 cm, apically cuspidate or acuminate, basally nar-
rowed, strongly oblique and slightly decurrent on the
petiole, membranous, dark green above, bright green
beneath, glabrate on both sides, pubescent along the
nerves, more densely so beneath, veins 7-10 on each
side, arcuate-ascending, slightly impressed above,
slightly elevated beneath; petioles to 0.8-1.5 cm long,
mostly strigose; minor leaves subrotund, 1-1.5 X 0.5-
0.7 cm with petioles to 5 mm long, sometimes ca-
ducous before the major leaves. Inflorescences fasci-
cles of 2-5 flowers. Flowers diurnal, 1 or 2 open at a
time and held below the leaves; pedicels 7-12 mm
long, strigose; calyx 1.5-2 X 3-3.5 mm, strigose, with
10 subapical unequal linear teeth in one series, 2.5—
3 mm long, strigose, often hidden in the pubescence;
corolla white, rotate, 6-9 mm long, lobed halfway
down, sparingly pubescent outside; stamens equal, fil-
aments 1.5 mm long, anthers mm long, narrowed
apically; ovary subglobose or conical, 1–1.5 mm long,
style 5-6 mm long, exserted 1.5 mm, stigma capitate.
Berry globose, red, 6-8 mm across, lacking stone
cells; fruiting calyx 6–7 mm long, applied to the berry,
teeth erect or spreading, to 5 mm long; fruiting ped-
icel 13-15 mm long; seeds ca. 100 per fruit, dark
brown, 1 X 1.5 mm, the thickened margin less than
0.5 mm wide. Figures 2J, 5.
Especie con pubescencia estrigosa y densa en sus tallos
y hojas, los pedicelos florales entre 7 y 12 mm de longitud
y los dientes del cáliz de 2.5-3 mm de longitud, bayas
globosas sin células pétreas.
This species has conspicuously dense and strigose
pubescence on stems and leaves. Its stems are often
arching in forest understory. It is similar to L inaequi-
a m has shorter calyx En and floral pedem
“Solanum appeared as
nomen а in Enum. PL. вена (4: 110. 1899.
Distribution. Venezuela, Mexico, Guatemala,
Honduras, Nicaragua, Costa Rica, Panama, Colom-
bia, Ecuador, Bolivia. Wooded slopes and along га-
vines of cloud forests, 500—1200 m elevation. (Map,
Fig. 6.) Flowering and fruiting through the year.
Pollination of this species was studied in Panama
by de Nevers (1986), who reported vibratile pollen
extraction by two species of halictid bees. The flowers
have no odor and no nectar and are visited mainly in
the moming when they are illuminated by sunlight.
d specimens examined. Mauer
ucre: Camino a Mundo Nuevo-Manacal, 1 waa
of Irapa, 500-700 m, Morillo 2519 (VEN); Penn A
Paria, Knapp & Mallet 6766 (BH, МУ), 5 MAP rk
Liesner 120772 мо, NY, VEN), Steyerma 3
96125 (P, VEN), 5 121528 pe 12159
(VEN). Yaracuy: Sierra de Aroa, río Carabobo, 800-1200
m, Liesner & González 9773 (VEN).
teyermark et al.
»
Volume 84, Number 2
1997
Benítez & D'Arcy
Lycianthes in Venezuela
\
А
MÁ
AAN
Figure 5. Lycianthes amatitlanensis.—A. Flowering and fruiting branch.—B. Flower bud.—C. Opened flower. After
Steyermark 121528 (VEN).
Lycianthes asarifolia (Kunth & Bouché) Bit-
ter, Abh. Naturwiss. Vereine Bremen 24(1):
423. 1919 [1920]. Solanum asarifolium Kunth
& Bouché, Index Sem. Hort. Berol. p. 10.
1845. Solanum violifolium var. asarifolium
(Kunth & Bouché) Hassl., Repert. Spec. Nov.
Regni Veg. 15: 221. 1918. TYPE: cultivated
in Berlin from seed from Caracas (holotype, B
destroyed, = F photo 2562).
violaefolium var. majus Dunal, in DC., Prodr.
13(1): 164. 1852. TYPE: Bolivia. Santa Cruz, d’Or-
bigny 619 not seen.
Annals of the
Missouri Botanical Garden
178
ne” w’ s Су n’ Са so” CM
~ -
202 ON Р
у сч ү,
s я
TON A З
> Ф Pd
+ *
км"
2
A х
n’ т ° ut s sw
e 6. Lycia
SE пи на outside of Venezuela.—B. Geographic
distribution in Venezuela.
anthes amatitlanensis.—A. Represen-
Solanum chodatianum Huber, Bol. Mus. Paraense Hist.
at. 4(4): 602. 1906. TYPE: Brazil. Villa de Con-
= 15 Рес. ap =“ 1336 (holotype, MG
not s = F photo
Selena к рен. fo. cosa i Hassl., Trab. Mus.
rm. Fac. Cienc. Med. Buenos Aires 21: 106. 1909
TYPE: Paraguay. Pilcomayo, Rojas 605 not seen.
[Synonymy taken from Barboza & Hunziker (1992:
Solari violifolium var. majus fo. chacoense Hassl., Re-
Spec. Nov. Regni Veg. 15: 220. 1918. TYPE:
Paraguay. Lower Pilcomayo, Rojas 275 (MO).
Solanum violifolium var. asarifolium fo. decadontum
assl., Repert. Spec. Nov. Regni Veg. 15: 221. 1918.
SYNTYPES: Paraguay. Pilcomayo River, Morong
920 (G not seen, MO); Rojas 605 not seen.
Creeping herb, stems slender, sparingly pubes-
cent, rooting at the nodes, stoloniferous; pubes-
cence of weak, collapsing, mostly erect, simple
hairs. Leaves solitary, ovate, 3-10 X 4-10(-15) cm,
apically rounded or obtuse, basally deeply cordate,
dark green and sometimes shiny above, matte and
lighter beneath, membranous, glabrous, veins 3—4
on each side, strongly ascending, sparingly
branched; petioles with purplish tones, 3-14 cm
long, canaliculate, sometimes with scattered simple
hairs. /nflorescences solitary, the flowers held above
the leaves. Flowers diurnal, inodorous, reflexed on
the pedicels; pedicels often white or purplish, 4-9
cm long, 0.5-1 mm thick; calyx pale, 3-5 X 3-4
mm, angled, the margin undulate-denticulate, gla-
brate inside and out, sometimes with sparse hairs
on the costas, lateral teeth absent but the 10 nerves
evident on drying; corolla "n or pale-yellowish,
outspread or reflexed, 0.6-1.1 X 1.2-2.2 cm, the
margin almost entire; stamens ae. filaments 1-
3 mm long, glabrous, anthers held together in a
cone, 2.5-3 mm long, the pores extending slightly
toward the sides of the anther; ovary conical or
bose. Berry orange-red, compressed-globose, 8-25
mm X 0.7-2 cm, lacking stone cells; fruiting calyx
5-9 X 11-17 mm, not splitting at the sinuses;
seeds ca. 70 per fruit, grayish brown, 2 X 2.5 mm
long, the margin thickened. Figures 2E, 7; Saun-
ders, Refug. Bot. 4, t. 255. 1871, as Solanum asar-
ifolium; Martius, Fl. Brazil 10, t. 4, 12. 1846 (calyx
and stamens), as Solanum asarifolium; Benítez de
Rojas, 1974: fig. 12; Steyermark & Huber, 1978, t.
288; Huber, 1906: 603, fig. 7 (flower), as Solanum
chodatianum.
Unica especie de hábito regie y estolonifero de to-
zuela, aie: sus
hojas pe cordiformes, de flores solitarias
colgantes sobre pedúnculos erectos y los offices edenta-
dos, bayas comprimido-globosas sin células pétreas.
Common names and uses. — "Ajicillo," “Barba de
Tigre," “Childa,” *Huevo de Sapo,” “Nicua.” Fruits
are eaten out of hand and also made into jams. |
This is the only species with a prostrate habit
among A species of Lycianthes, presenting
its conspicuously cordiform leaves on erect petioles
and solitary flowers deflected on erect peduncles.
The calyces usually lack noticeable teeth.
Distribution. Venezuela, Bolivia, Peru, Para-
guay, southern Brazil, and northeastern Argentina.
Deciduous and semi-deciduous woods near moist
places, 50-1000 m. Also a component of adventive
and weedy vegetation in croplands. (Map, Fig. 8.)
Flowering and fruiting throughout the year
species is similar to рани repens
(Spreng.) Bitter of southeastern Brazil, which is
said to differ (Bitter, 1920: 426) in having smaller
leaves, shorter stalks, and distinctive hairs. We
have seen only two collections that were cited by
Bitter under this name. One of these, Dusén 11315
(F, MO) from Paraná, Ponta Grossa, has two un-
equal but well-developed leaves at each node. The
|
|
|
|
|
E
Volume 84, Number 2
1997
Benítez & D'Arcy 179
Lycianthes in Venezuela
II
Figure 7. Lycianthes asarifolia. Flowering and fruiting stems. After Licata et al. 32 (MY).
other specimen, Pohl 5407 (F), without locality, has
solitary major leaves and sessile minor leaves like
many plants from elsewhere. The Dusén specimen
is the only one we have seen with well-developed,
long-petiolate minor leaves. The original descrip-
tion of the basionym, Boldoa repens Spreng. (Syst.
Veg. 1: 179. 1825) and Bitter's description also note
unequally paired leaves. While the Dusén collec-
tion does have small leaves and peduncles, except
for the well-developed minor leaves it seems to be
conspecific with plants we have identified as L.
asarifolia. If L. asarifolia and L. repens are actually
the same, then L. repens is the prior and correct
name. We continue use of L. asarifolia for this spe-
cies, a name that has been used in many publica-
tions, rather than substituting the unfamiliar and
uncertainly equivalent name L. repens. Bitter cited
a specimen, “Otto ex hb. Kurt Sprengel,” which
Annals of the
Missouri Botanical Garden
de
Figure 8. Lycianthes asarifolia.—A. Representative
distribution outside of Venezuela.—B. Geographic distri-
bution in Venezuela.
would have served as a type. If L. repens is recog-
nized as the correct name, then Solanum violaefol-
ium Schott (in Sprengel, Syst. Veg. 4 (appendix):
403. 1827) may also be a synonym. Bitter placed
Solanum violaefolium into synonymy under L. re-
pens, but Schott’s description notes that the calyx
is “JOfidis,” seemingly contrary to the truncate,
nearly edentate calyces found in our species.
Representative specimens examined. VENEZUELA.
istrito Federal: 5 Naiguatá, vertiente N de Ја Cordil-
lera de la Costa. m, Morillo & Manara 2022
N). Apure: Boca del Uribante, Trujillo & Fernández
10597 (MY). Aragua: Río Hondo, km 32 carretera hacia
Choroní, 800 m, Benítez & Rojas 3993 (MY). Carabobo:
La Toma de Valle Seco, Dist. Puerto Cabello, 100-200 m,
Benítez £ Pons 4651 (MY). Cojedes: Las Tucuraguas,
Mpio. Angel Bravo, 700 m, Del
El Toro, 600-900 m, Liesner et al. 7803 (MY, VEN).
Lara: Barquisimeto, Saer 263 (V i
Guaire betwee
esa Cavacas-Biscucuy, Stergios & Aymard 4451 (MY,
PORT). Yaracuy: Finca Antonia, San Felipe, 250 m, Fer-
rari 790 (MY).
4. Lycianthes ferruginea Bitter, Abh. Naturwiss.
Vereine Bremen 24(1): 339. 1919 [1920]. So-
lanum ornatum Morton, Contr. U.S. Nat. Herb.
29(1): 59. 1944. SYNTYPES: Venezuela. Ar-
agua: Colonia Tovar, Moritz 1642 (B destroyed,
= F photo 2572, BM not seen, = MY photo;
HBG not seen, MO, US); Colonia Tovar, Gol-
mer s.n. (B not seen); Fendler 991 (BR, G not
seen, GOET not seen, NY-2). [Although we
have seen duplicates of specimens cited by
Bitter in the original protologue, we only saw
one specimen that he actually cited. This
specimen accords with our concept of the tax-
on and with all the duplicates noted, but we
chose not to designate a lectotype without op-
portunity to review the material again.]
Much-branched shrub or vine, often high-climb-
ing, branches often at right angles or zigzag, young
growth densely ferrugineous pubescent, stems gla-
brescent; pubescence of reddish or yellowish ses-
sile and stalked stellate hairs, mostly with 3-5 ra-
dii, and simple hairs. Leaves solitary, ovate or
narrowly ovate, 3.4-12 X 2-6 cm, basally obtuse
or rounded, sometimes slightly decurrent on the
petiole, apex acute, membranous, densely stellate-
pubescent on both sides, glabrescent and dark
above, brownish yellow beneath, veins 3—4 on each
side, mostly slightly impressed above, elevated and
conspicuous beneath, minor reticulate venation
mostly evident beneath; petioles 0.4—1.7 cm long.
Inflorescences fascicles of 2—4(—5) flowers, mostly
near the branch tips. Flowers diurnal, one or two
opening at a time; pedicels 0.8-1.7 cm long, to-
mentose; calyx 3-7 X 3-8 mm, tomentose, teeth
10 in 2 unequal series, erect or spreading, 0.54
mm long, sometimes appearing only as costas on
the calyx cup; corolla white, sometimes greenish on
the lobes, rotate, 0.9-1.5 cm long, 1.8-3 cm wide,
entire or subentire, the lobes conspicuously thicker,
glabrous outside overall, the lobes sometimes mi-
nutely ciliolate with reduced curved hairs, some-
times with small tufts of stellate hairs on the lobe
tips, glabrous within; stamens unequal, 4 filaments
1-2 mm long, the fifth 2-4.5 mm long, anthers 2.5-
3 mm long; ovary ovoid, 2-4 X 1-3 mm, glabrous,
style 6-7 mm long, glabrous, exserted 1 mm, stigma
subclavate. Berry orange-red, subglobose, 10-20
mm across, with 2-4 stone cells; fruiting calyx
slightly accrescent, the cup 4—6 mm long, applied
to the berry, sometimes splitting irregularly, the
teeth slightly reflexed, not accrescent; seeds са. 36
Volume 84, Number 2 Benitez & D’Arcy 181
1997 Lycianthes in Venezuela
\ NN A
OSLO
SECTAS
Figure 9. Lycianthes ferruginea.—A. Flowering and fruiting branch.—B. Opened flower.—C. Fruiting calyx. After
Benítez 5148 (MY).
Annals of the
Missouri Botanical Garden
182
mw w Ld Су те wv Ld СМ
EE. v
М >. © =ч},
ve
a wv
*
% y
=:
[ d
гё
А х"
Figure 10. Lycianthes ferruginea.—A. Representative
distribution outside of Venezuela.—B. Geographic distri-
bution in Venezuela.
per fruit, yellowish brown, 3-3.5 X 3.54 mm, the
thickened margin ca. 0.7 mm wide. Figures 2A, 9.
En esta especie el cáliz presenta 10 dientes en dos
cuamente verdosos y engrosados y el pliegue blanco; las
‚ bayas son subglobosas, sin células pétreas o desde 1 hasta
4
This species is much like Lycianthes pauciflora,
differing in its conspicuously dense pubescence
and corolla color. The calyx lobes of the flowers of
this species vary greatly in length and appearance,
sometimes appearing as short points spreading from
the calyx apex and other times as slender tomentose
teeth arising from low on the sides of the calyx. A
difference between this species and L. pauciflora
that was not noted in the key to species is in the
coloration of the corollas when seen from outside;
in L. ferruginea the lobes are green and the folds
white, while in L. pauciflora, the entire surface is
white.
This species is also similar to L. armentalis J. L.
Gentry from Central America, but that species has
pubescent corollas, and so far as is known, equal
stamens. It also resembles L. jelskii (Zahlbr.) Bitter
from Peru, but we cannot judge well from the photo
of the type of that species.
Distribution. Colombia. Cloud forests, 1200-
2900 m elevation. (Map, Fig. 10.) Flowering and
fruiting throughout the year but most flowering from
April to August.
Representative specimens examined. VENEZUELA.
Distrito Federal: El Junquito to Colonia Tovar, 1770 m,
Davidse 4035 (VEN). Aragua: Colonia Tovar to E] Agua-
catal, Benitez et al. 4235 (MY). Falcón: Sierra de San
Luis, 1500 m, Demey (MY-86695, CORO). Lara: between
Cubiro & La Escalera, 1600-2000 m, Steyermark et al.
110242 (MY, VEN). Mérida: Caserío El Portachuelo, NW
of Guaraque, 2 т, Marcano-Berti & López-Palacios
1758 (MER, MY). Miranda: Cortada del Guayabo, Ta-
mayo 397 (VEN). Táchira: above Betania, below Páramo
de Tamá, 2530 m, Steyermark 57433 (MY, VEN). Truji-
llo: El Paramito, near Escuque, Lasser 1203 (VEN). Yar-
acuy: Los Quinquines, road to La Candelaria, Diederichs
173 (VEN).
5. Lycianthes inaequilatera (Rusby) Bitter, Abh.
Naturwiss. Vereine Bremen 24 (1): 439. 1919
[1920]. Bassovia inaequilatera Rusby, Mem.
Torrey Bot. Club 6: 90. 1896. Brachistus in-
aequilaterus (Rusby) Rusby, Bull. New York
Bot. Gard. 4: 470. 1907. SYNTYPES: Bolivia.
Between Tipuani and Guanai, Bang 1708 (B
not seen, G, M not seen, MO).
Erect or wandlike subshrub 1-2 m tall, branches
flexuous, arching, strigose, adult branches somewhat
glabrescent, ridged, often drying dark brown; pubes-
cence of coarse simple hairs. Leaves unequal-gem!-
nate, narrowly ovate, 16.5-20 X 46.5 cm, apically
acuminate, sometimes abruptly so, basally strongly
oblique, one side obtuse or rounded, the other de-
current on the petiole, membranous, lighter beneath,
glabrate or with scattered hairs on both sides, es-
pecially along the nerves, more so beneath, ciliate,
veins 6-8 on each side, major veins plane or slightly
impressed above, slightly elevated beneath; petioles
0.5-0.8 cm long, compressed; minor leaves ovate,
2.3-3.6 X 1.3-1.6 ст, subsessile. Inflorescences fas-
cicles of (1—)4 flowers. Flowers with pedicels 2-3 cm
long, strigose; calyx 2.5 X 2 mm, pilose, teeth 10,
slightly unequal in one series, subulate, 0.5-0.7 mm
long, strigose; corolla white, rotate, 8-9 X 9-10 mm,
lobed %—М way down, pilose outside; stamens equal,
filaments 3.5 mm long, anthers 2.5-3 mm long; ovary
ovoid, 1 X 0.5-0.6 mm, style 6.5 mm long, €
2 mm, stigma capitate. Berry red, subglobose, 5%
mm across, without stone cells; fruiting calyx ассгеѕ-
cent, the сир 3—4.5 тт long, the teeth slightly ac-
|
|
|
|
Volume 84, Number 2
1997
Benitez & D’Arcy 183
Lycianthes in Venezuela
Figure 11. Lycianthes inaequilater
After Stergios et al. 6360 (MY); B, C. After Davidse 18899
crescent, fruiting pedicels 2 mm long; seeds ca. 70
per fruit, yellowish brown, 1.5-1 mm across, the con-
spicuously thickened margin 0.5 mm wide. Figures
21, 11.
Se caracteriza por ser pubescente, con hojas menores
с cuas, los dientes del cáliz en número de 10, liger-
amente desiguales, en una serie y subulados, los cuales
ilatera.—A. Flowering and fruiting branch.—B. Flower bud.—C. Opened flower. A.
(VEN).
son generalmente poco conspícuos en fruto, bayas sub-
globosas, sin células pétreas. :
This species closely resembles L. amatitlanensis,
from which it differs in its lesser pubescence and
shorter calyx teeth, which are usually less conspic-
uous in fruit, and longer pedicels.
Distribution. Venezuela and Bolivia. Cloud for-
Annals of the
Missouri Botanical Garden
Se
n “ s 0°
“Fos Р
“ @ fy.
ч e t
У 10°
e
10°
*
2
x
ге 12. Lycianthes inaequilatera.—A. Representative distribution outside of Venezuela.—B. Geographic distri-
la.
gu
bution in Venezuela
ests and along ravines, 1500-1900 m elevation.
(Map, Fig. 12.) The six collections seen from Ven-
ezuela are all in fruit, made scattered throughout
the year. Further collecting should reveal the pres-
ence of this species in other countries.
Representative specimens examined. VENEZUELA.
Ме
Paramitos, SE of Biscucuy, 1000-1500 m, Stergios et
al. 6360 (MY, PORT).
6. Lycianthes lenta (Сау.) Bitter, Abh. Naturwiss.
Vereine Bremen 24(1): 364. 1919. Solanum
lentum Cav., Icon. 4: 4, plate 308. 1797. So-
lanum virgatum Lam. var. lentum (Cav.) O. E.
Schulz, in Urban, Symb. Antill. 6: 189. 1909.
TYPE: cultivated Madrid, seed from Mexico
(holotype, MA)
Solanum cumanense Roem. & Schultes, Syst. Veg. 4: 662.
1819. TYPE: Venezuela. Cumaná, Humboldt 71 (ho-
lotype, P, = IDC microfiche 4316, = F photo,
002896).
Solanum lentum var. echinatum Dunal, in DC., Prodr.
13(1): 173. 1852. TYPE: Mexico. Without collector
(G not seen, = F photo 34120). [The typification of
this p was interpreted by Nee (1986: 97) to be
ed on specimens of Sesse & Mocino now at F,
MA, and G, including the one of which we have seen
oto.
-Solanum s sylvaticum sensu Schlecht., Linnaea 5: 112.
830, non Dunal (1852), or Bitte: Abh. Naturwiss.
Vereine Bremen 24(1): 364. 1919 [1920]. Based on
material from e ы
?Solanum quadriflorum М. s & Galeotti, jer
Acad. Roy. Sci. Brexellne 120) 139. 1845. TYPE
наг Galeotti 1231 (holotype, ВК not seen; iso-
not seen, = F photo ene
SA declinatum. Sessé & Мос., x. ed us
1893 [1894]. SYNTYPES: vat тека
7) cited as ee
een, — photo 48235).
parer oe Fernald, Bot. Gaz.
895. TYPE: Mexico. Sinaloa
i 20:
Villa Union,
poste i“ (holotype, GH not seen; i sotype,
Solanum nocturnum Fernald, Proc. Amer. Acad. Arts 35:
Volume 84, Number 2
1997
Benitez & D’Arcy
Lycianthes in Venezuela
A
Figure y
inserted on corolla. After Bunting 4433 (M
570: 1900. SYNTYPES:
533 (GH not seen); Tehuantepec, Seler 1625 (GH not
seen).
Lycianthes lenta var. endopsila Bitter, Abh. Naturwiss. Ver-
eine Bremen 24(1): 367. 1919 [1920]. SYNTYPES:
Venezuela. Caracas, Н 748 (B not seen); Var-
gas s.n. (lectotype, designated here, US-601441).
Solanum virgatum Lam. var. caracasanum O. E. Schulz,
Mexico. Oaxaca: Palmer
13. Lycianthes lenta.—A. Flowering and fruiting branch.—B. Opened flower.—C. Flower bud.—D. Stamens
у).
in Urban, Symb. Апш. 6: 190. 1909. TYPE: Ven-
ezuela. Near Cura, Humboldt 748 (holotype, P).
Lycianthes lenta var. scotinophila Bitter, Abh. Naturwiss.
Vereine Bremen 24(1): 367. 1919 [1920]. TYPE:
Venezuela. Valle del Aragua bei San Mateo, Otto 788
(holotype, B destroyed).
Lycianthes pauciflora (Vahl) Bitter subsp. tobagoensis Bit-
ter, Abh. Naturwiss. Vereine Bremen 24(1): 343.
186
Annals of the
Missouri Botanical Garden
1919 [1920]. SYNTYPES: Tobago. Broadway 4523
(holotype, B destroyed); Trinidad, Crueger 148
(GOET not seen).
Lycianthes variifolia Standl., Field Mus. Bot. 4: 259. 1929.
TYPE: Belize. Tower Hill, Karling 13 (holotype, F).
[Taken from synonymy of Gentry & Standley (1974).]
Shrub to 3 m, erect, climbing or sprawling, stems
slender, drying nish, young branches pubescent;
pubescence of stalked, stellate hairs with 4–6 rays.
Leaves solitary or geminate, ovate, sometimes narrowly
or broadly so, 4.4—10 X 2.6–5 cm, apically rounded,
obtuse, or short-acuminate, basally rounded, truncate
or slightly cordate, sometimes short-decurrent on the
поје, membranous, softly pubescent, veins 3—4 on
each side, slightly elevated; petioles 1-2 cm long.
Inflorescences fascicles of 5 or fewer flowers, some-
times leaving a cicatrix from fallen pedicels. Flowers
nocturnal, strongly sweet-scented, only a few open at
a time; pedicels 1–2.5 cm long, slender, pubescent;
calyx 3-4 X 3–5 mm, sparingly pubescent, with 10
subapical teeth in two unequal series, 1-2(-3) mm
long, erect or spreading; corolla white or bluish, 1.5—
3 X 3-5 cm, lobed less than % way down, the lobes
conspicuously thicker, puberulent in bud with stellate
and reduced hairs, glabrescent except at the tips; sta-
mens unequal, 4 filaments 1.5-2 mm long and an-
thers 3-4 mm long, the fifth filament 3.5-8 mm long
and anther 5-6 mm long; ovary ovoid, 1-1.5 X 1 mm,
glabrous, style 8-12 mm long, exserted 2 mm, stigma
subemarginate or clavate. Berry red or orange-red,
subglobose, 6-9 mm across, stone cells lacking; fruit-
ing calyx 6-8 mm long, generally applied to the berry,
teeth spreading, hardly accrescent; seeds ca. 35 per
fruit, pale yellow, 2 X 2.5 mm, the thickened margin
ca. 0.5 mm wide. Figures 1C, 2C, 13; Nee, 1986: p.
101. fig. 10.
The name Solanum nocturnum is placed into
synonymy here based on Fernald's description.
Es de amplia distribución en áreas de alturas bajas y
se caracteriza por sus tallos castaño-claros y ramificación
densa, los dientes del cáliz en dos series desiguales y los
frutos carecen de células pétreas.
The species may be recognized by its bright
brown stems, dense branching, and large corolla
diameters.
Distribution. Mainly Caribbean in distribution:
Mexico, Belize, Guatemala, El Salvador, Nicaragua,
Trinidad and Tobago, Cuba; Venezuela. Deciduous
woods and thickets, sea level to 500 m. Often near
the sea, but actually of wide distribution in medium
and low elevations. (Map, Fig. 14.) The species
flowers and fruits throughout the year.
Representative specimens examined. VENEZUELA.
arque Nacional Santos Luzardo, isla El Vapor,
40 m, Duno et al. 184 (MY). Aragua: Ocumare de La
Costa, 0—400 m, Badillo 1816 (MY); Carretera Cagua-La
w € x 0 nt st и СП
~ :
• aie С P
y 0 "ey
H
^ HE A
~ e
e:
v
a
A x
n 70) [^d “e 62° Су
Figure 14. Lycianthes lenta.—A. Representative dis-
tribution outside of Venezuela.—B. Geographic distribu-
tion in Venezuela.
Villa de Cura, 4 km al S de Cagua, Bunting 4433 (MY).
i Río Caparo, E del Cantón, 100 m, Steyermark
uebrada María Te-
resa, Dist. Valencia, 500 m, Benítez & Rojas 3214 Mn
Falcón: Dist. Silva, NE of La Soledad, 5 m, Steyermark
& Manara 110995 (MO, NY, VEN). Lara: Santa Rosa,
Pittier 13088 (MO, NY, VEN). Miranda: Рараго, río Chi-
co, 10 m, Aristeguieta 3994 (MO, VEN). Nueva Esparta:
Cerro Los Cedros, SW of San Francisco, Península de
Macanao, 100—300 m, Benítez 2614 (MY). Suere: Parque
Nacional Mochima, El Tacal, Cumana 1850 (IRBR, MY).
Yaracuy: Finca Los Apamates, La Llanada, between Ur-
ama & San Felipe, Romero 486 (MY). Zulia: Carretera
Caja Seca-Bobures, Bunting 5808 (MO, VEN, VZU).
7. Lycianthes lycioides (L.) Hassl., Annuaire
Conserv. Jard. Bot. Geneve 20: 181. 1917. So-
lanum lycioides L., Syst. Nat. ed. 12, 2: 174.
1767; Mantissa Pl. 1: 46. 1767. TYPE: Peru.
LINN 248.48 (lectotype, designated by Knapp
& Jarvis (1990)).
Solanum lycioides var. tomentosa Dunal, Hist. Nat. et
num 174. 1813. Solanum candicans Dunal, Sol. Syn-
Volume 84, Number 2
1997
Benitez & D’Arcy 187
Lycianthes in Venezuela
Fi 15. Lycianthes lycioides.—A. Flowering and fruiting branch.—B. Flower bud.—C. Opened flower.—D. Py-
).
23. 1816. Lycianthes candicans (Dunal) н, Ап-
Jard. Bot. Ge 1917.
& Tarmae (С not seen, = F photo 34114, MPU not
seen).
gure 5 ;
renes (seeds enclosed in sclerenchyma). After Benttez 5373 (MY
Solanum phillyreoides Dunal, Sol. Syn. 24. 1816. TYPE:
Colombia. Fluvium Magdalenae, (holotype, P, = IDC
microfiche 4375
Solanum pseudolycioides Rusby, Bull. Torrey Bot. Mes 26:
193. 1899, SYNTYPES: Bolivia. La Paz, 1 E
Rusby tm 12,000 ft, ec 835 (both NY neither
seen); Bang 32 (NY no
Solanum deos din Dammer, Eng. и. 37: 168. 1905.
Annals of the
Missouri Botanical Garden
188
no 100° E [d т “ s “*
«зы. x
M a ex у.
Y
A 35
.
.
> Q ”
+
wi w
EJ
LE
ы •
• 2°
°
A x?
з ке «° ut e “
Ld
v
Figure 16. Lycianthes lycioides.—A. Representative
distribution outside of Venezuela.—B. Geographic distri-
bution in Venezuela.
SYNTYPES: Argentina. Jujuy: Santa Catalina, Clar-
en 11550; Yavi, Fries 985 (B not seen, CORD not
seen). [Synonymy of Bitter, 1919 [1920]; Morton et
al., 1976: 33; Barboza & Hunziker, 1992.]
Erect shrubs 0.5-1 m tall, much branched, young
branches green, tomentose, adult branches strongly
lignified, grayish, glabrescent, conspicuously lined,
the shoots sometimes becoming spinose; pubes-
cence of erect, sparingly branched hairs. Leaves
solitary, often with short internodes and appeari
dense or even fasciculate, elliptical, smaller leaves
sometimes obovate, 0.6-2.5 X 0.3-1.1 cm, mem-
branous or papery, both sides densely fine-pubes-
cent, glabrescent, remaining apically tufted, veins
3—5 on each side, slightly elevated; petioles 2—4
mm long. Inflorescences fascicles of 3 or fewer flow-
ers. Flowers diurnal, showy; pedicels 0.5-1.6 cm
long; calyx 2-3 X 0.3-0.5 mm, pubescent to gla-
brate, teeth 10 in 2 unequal series, the upper series
1 mm long, the lower teeth 0.5 mm long, erect;
corolla blue-violet with a yellow eye, rotate, the
margin almost entire, the lobes strongly contrasting,
1 X 2-2.5 cm, mostly glabrate; stamens unequal,
3 filaments 1.5-2.5 mm long, 2 filaments smaller,
0.5-1.2 mm long; anthers yellow to orange, 1.8-2.7
mm long; ovary ovoid, 1-2 X 1-1.2 mm, style 4–
5 mm long, exserted 1 mm and curved, stigma bi-
lobate. Berry orange, depressed-globose, 2-6 mm
across, with 7—8 stone cells (pyrenes); fruiting calyx
hardly accrescent, applied to the berry, the teeth
spreading; fruiting pedicels slender, 8-20 mm long;
seeds 1-2 per pyrene, each enclosed in scleren-
chyma in the form of a pyrene. Figures 1E, 2B, 15;
Jacquin, Icones Plantarum Rariorum 1, t. 46 (as
Solanum lycioides); Ruiz € Pavon, Fl. peruv. 2: t.
177 (as Solanum lycioides); Edward's Bot. Register
32, t. 25. 1846 (as Solanum lycioides); Weddel,
Chloris Andina 2, t. 55 (as Solanum lycioides); Bol.
Mus. Paraense Hist. Nat. 4: 603. 1905-1906 (flow-
er, as Solanum lycioides). [J. J. Jacquin, 1782. Ico-
nes Plantarum Rariorum. 1: t. 46, cites Jacq. Misc.
3.]
Especie con ramas espinescentes y corolas de un azul-
T RN dala id zl 3. IIS һа
anaranjado de las anteras, bayas deprimido-globosas con
6 pirenos. Su distribución geográfica corresponde exclu-
sivamente al estado Mérida en elevaciones entre 1900 y
Common пате. “Cuchuva de Perro."
This species is distinct with its spinescent
branches, irregular branching, and small leaves.
The corollas are intense blue-violet with a contrast-
ing yellow or brownish eye.
The filaments in our material of this species are
not gibbous as recorded by Barboza and Hunziker
(1992). Solanum lycioides subsp. parvifolium
(Wedd.) Bitter and Solanum pseudolycioides Rusby
were considered to be synonyms by Bitter, and we
hesitantly consider them to be the taxonomically
typical Solanum lycioides. Material we have seen
has slender, flexuous branching, apparently reflect-
ing young growth rather than a distinct taxon.
Distribution. Uplands of Colombia, Ecuador,
Peru, Bolivia, northwestern Argentina, Paraguay,
and southern Brazil; in Venezuela in the Andean
region. Semi-arid woody formations and remnants,
1900-3000 m elevation. (Map, Fig. 16.) Almost all
collections are in flower, and most were made from
April to July.
m, Nee & Whalen 17053 (VEN); Caserío El Vergel g^
tween Mucurubá & Mucuchíes, 2900 m, Ruiz-Terán
López-Palacios 12533 (MERF, MY).
Volume 84, Number 2 Benitez & D’A
1997 Lycianthes in Venezuela
v.e
AN
се
A
ХЫ
\
|
EE
Figure 17. Lycianthes pauciflora.—A. Flowering and fruiting branch.—B. Opened flower.—C. Flower bud. After
Steyermark 123694 (MY) and Trujillo 13947 (MY).
А А : - Solanum geminatum Vahl, Eclog. Amer. 1: 21. 1797. Ly-
y: Lycianthes рапейога (Vahi) RAN d cianthes geminata (Vahl) Bitter, Abh. Naturwiss. Ver-
Ба Baer ees 24(1): на eine Bremen 241): 392. 1919 [1920]. TYPE: French
. Not Sendtn. . Solanum iana. von s.n. (С, = oto 22887).
rum Vahl, Eclog. Amer. 1: 20. 1796. TYPE: Solanum neglectum Dunal, Hist. Sol. 177. 1813. Based
Martinique (holotype, C-hb Vahl). on Solanum arborescens, solani hortensis folio, fructu
Annals of th
Missouri Lede Garden
n? 100° Са Су n" “ so” СЫ
| 8
a „
Т و a
E •
Fe 5
У e
+:
6
О
”
E
ES
A x”
n хе P P а «
dw
ГА
P
e
Je
4
e 18. Lycianthes pauciflora.—A. Representative
Р ыыы outside of Venezuela.—B. Geographic distri-
bution in Venezuela.
corallino. Maius, Plumier, Pl. Amer. ed. Burmann
242, t. 245, fig. 4. 1760; Plumier, Cat. 4: 34 ined.
mss., fide Lourteig (1987). Lycianthes neglecta (Dun-
al) Lourteig, Phytologia 62: TYPE: Plu-
mier plate based on a plant from — i 5 (lecto-
type, designated by Lourteig (198
Solanum speciosum Dunal, Hist. Sol. n 1813. Based on
г.
1760; Plumier, Cat. 4: 35 ined.
, fide Lourtei eig ` 1987). Lycianthes speciosa (Dun-
al) Tae eig, Phytologia 62: 442. . TYPE: Plu-
mier plate based on a plant from Santo Domingo
(lectotype, designated by Lourteig (1987).
Solanum sylvaticum Dunal, Solan. Syn. 24. 1816. TYPE:
Venezuela. ees Humboldt & Bonpland, s.n. (ho-
lotype, P-LA, = IDC 427 microfiche, = F photo,
39015).
Solanum glandulosum Sendtn., in Mart., Fl. Brazil 10: 52.
1846. SYNTYPES: " azil. Amazonas : Rio Japura
ана s.n. (В destroyed, М not extant, neither
en); southern Brazil, Sellow (B
Solo auge Dunal, in DC., Prodr. 13(1): 174. 1852.
me for Solanum каганы Sendtn., in
Mart., Fl. Brazil 10: 52. 1846, non Pe & Pavon
(1799). Lycianthes japurensis Bitter, Abh. Naturwiss.
Vereine Bremen 24(1): 350. 1919 че SYNTY-
~
PES: Brazil, Rio Negro, Rio Japurá, Martius s.n. not
seen; southern Brazil, Sellow s.n. (B not seen).
Solanum guianense Dunal, in DC., Prodr 13(1): 166.
1852. Lycianthes guianensis (Dunal) Bitter, Abh. Na-
turwiss. Vereine Bremen 24(1): 346. 1919 [1920].
TYPE: French Guiana. Cayenne, Aublet? (BM not
een, G-DC not seen, = IDC microfiche).
Solanum urbanum var. ovatifolium Chodat, Bull. Soc. Bot.
Сепеуе 2(8): 152. 1916. TYPE: Paraguay. Grand for-
ét de Caaguazu, Balansa 2080 not seen. [Synonymy
of Barbosa & Hunziker (1992
e гене Вишег,
Abh. Naturwiss. Vereine
: za
4): 17-34. 1992. Illegitimate combination. TYPE:
Paraguay. Arroyo Mocoy, Hassler 4912 (G not seen,
— F photo, 23070, MO, P not seen, — US, photo,
var. glabrescens C. V. Morton, Contr.
U.S. Natl. Herb. 29(1): 62. 1944. TYPE: Colombia.
Archer 2132 (holotype, US).
High-climbing shrub 5-6 m tall, upper branches
often dangling and interlocking with other vegetation;
branches ferrugineous tomentose, often glabrescent;
pubescence of stalked, pauciradiate stellate or den-
dritic hairs. Leaves solitary, ovate, 7-11 X 2-5 cm,
apically obtuse or acuminate, basally obtuse or round-
slightly impressed above, elevated beneath, glabres-
cent above, softly tomentulose beneath; petioles 0.7—
1.6 cm long, tomentose. Inflorescences fascicles of 2-
5 flowers. Flowers crepuscular and nocturnal; pedicels
7-16 mm long; calyx 2-7 Х 3-6 mm, sparsely pu-
bescent, teeth 10 in 2 unequal series, sometimes dark
purple, later white, 2-6 mm long, reflexed or not;
corolla white, rotate, 0.9-2.2 X 1.3-3.5 cm, the mar-
gin almost entire, the lobes conspicuously thicker, gla-
brate; stamens unequal, 4 with filaments 1-2 mm
long, the fifth filament 3.3 mm long, anthers 2-5 mm
long; ovary ellipsoidal, 1.5-2 x 1-1.5 mm, style 8-
9 mm long, exserted 5 mm, stigma capitate. Berry
bright orange-red, depressed globose or ovoid, 15-20
mm across, stone cells 2-3 or wanting; fruiting сајух
accrescent and thickening, 10-14 mm across, the
margin sometimes reflexed, the teeth usually reflexed,
3-7 mm long; fruiting pedicels 15 mm long, glabrate;
seeds 40-80 per fruit, brown, 2.5—4 X 3.54 mm, the
ngeye margin ca. 0.7 mm wide. Figures 1A, B,
D, 2G, 17; Plumier, Pl. Amer. t. 245, figs. 4, 5. 1760;
D’Arcy, 1973: 640, fig 13. [Plumier, Pl. Amer. ed
Burm. t. 245, fig. 4. 1760. as Solanum peduncularis
alaribus. |
Es la especie de más amplia ——— geográfica
el país, muy variable en cuanto a grado ii iet ae
14
Volume 84, Number 2 Benítez & D'Arcy 191
1997 Lycianthes in Venezuela
|
|
|
|
SUM
vel a M
5 cm
Figure 19. Lycianthes radiata.—A. Flowering and fruiting branch.—B. Opened flower.—C. Fruiting calyx. After
Schwartzkoff 12 (MY)
siendo ésta densa y ferrugínea en las ramas jóvenes y pueden estar ausentes o en número de 1, 2 6 4, localizadas
floríferas y casi ausente en las ramas adultas y fructíferas, еп la parte apical interior del fruto
еза
| así mismo los dientes del cáliz varían en tamafio e Conimon ‘name: © “Coronilla.”
| ta 6 mm де longitud, siendo reflejos о по еп flor y/o This species is often seen climbing high i
en fruto, y el borde puede presentarse reflejo o aplicado;
| la corola presenta medidas entre 0.9-2.2 cm de 2o shrubs or low in the TET and displaying large
| у el borde casi entero. Respecto а células pétreas, és clusters of showy red fruits. The flowers, usually
192
Annals of the
Missouri Botanical Garden
Figure 20. Lycianthes
ا онна of Venezuela. 8. Geographic distri-
bution in Venezuel
radiata.—A. Representative
during the day, are less conspicuous. The
yx has conspicuous, porrect, recurved or re-
Pul sometimes almost woody teeth.
Distribution. Collections have been seen from
Costa Rica, Panama, the Greater and Lesser Antil-
les, and all tropical countries in South America ex-
cept Chile and the Guianas, and it probably occurs
in the latter region. Lycianthes pauciflora is the
most wide-ranging species of the genus within Ven-
ezuela, and it is variable as to its degree of pubes-
cence and the size and form of its calyx teeth.
Cloud forests and gallery forests 600-1500 m ele-
bes PM. Fig. 18.) The species appears to flow-
fruit throughout the year, but most flowering
specimens were collected from April to July and in
November and December.
The concept employed here unites concepts of зеу-
eral regional treatments (D'Arcy, 1973; Barboza &
Hunziker, 1992) under the oldest name for this wide-
species. The Panamanian plants for which
D'Arcy used the name L. guianensis Dunal have larg-
er fruits and longer, thinner calyx lobes than those
from the Antilles, Venezuela, and other eastern parts
of the continent, and they may represent a distinct
taxon. However, in most particulars, particularly in
having fruits with 2 stone cells, the Venezuelan plants
agree with the other plants discussed here. Plants of
the species from the Antilles, particularly Dominica
and Martinique, the type locality of L. pauciflora, tend
to have slightly smaller calyces and flowers and more
rotund leaves than those of Venezuela and the Guian-
as, but they are otherwise similar. Fruits examined
from the Lesser Antilles had varying numbers of stone
cells (Martinique, Duss 364 (US), no stone cells, Duss
4430 (US), 1 stone cell, Dominica, Ernst 1942 (US),
2 stone cells). Plants from lowland Paraguay, Bolivia,
and Peru have dimensions like those of plants from
the Antilles.
Representative specimens examined. VENEZUELA.
Distrito Federal: between el topo Macanillal & El Pico
Izcaragua, 7-12 km E de los tanques de la Electricidad
de Caracas, 700—800 m, Morillo et a 20 (VEN). Ama-
: Reserva forestal Sierra Im , 1988 m, Sanoja
. Amazonas: 5 to 7 7 km rede river E of Cerro La
Neblina, 140 m, Liesner & Funk 15838 (MO, MY, VEN).
Apure: Reserva Forestal de San Camilo, Chiricoa, 200
m, Steyermark et al. 101704 (MO, NY). Aragua: Carre-
us Maracay-Choroní, 1200 m, Benítez et al. 4911 (MY).
Barinas: Dist. Bolívar, near feldspar mine, between La
Soledad and Santo Domingo, 1300 m, van der Werff &
Ortega 6124 (MO, NY). Bolívar: El Dorado-La Gran Sa-
mw 2908 (M arabobo: Colinas
Fortaleza, Pittier 8803 (VEN). Fal-
abla, 1450 m, So
B
pa
~
(УЕМ). 5 erro Patao, М of Ри
ermark "y Аай 91324 (VEN). Tachira: between Que-
brada Grande and El Nula, border with Apure, 250 m,
Gentry € Puig-Ross 14293 (MO). Trujillo: Vía
Escuque-El Socorro, Benitez 1952 (MY). ا Ser-
ranía Santa María-Cerro La Chapa, 6 km N de Nirgua,
1200-1350 m, Meier et al. 3903 (MY, VEN).
9. Lycianthes radiata (Sendtn.) neo зак Na-
turwiss. Vereine Bremen 24(1): 433. 1919
[1920]. egi radiatum Sendtn., in Martius,
Fl. Brazil. 10: 53. 1846. TYPE: Hartweg 1293
[129 in publication] (BREM not seen, W not
seen, B destroyed, — F photo 2586).
Solanum goudoti Dunal, in DC., Prodr 13(1): 158. 1852.
Lycianthes goudoti (Dunal) Bitter, Abh. Naturwiss.
Vereine Bremen 24(1): 435. 1919 [1920]. TYPE: Co-
n Vin 13 (holotype, G-DC not seen, = IDC
rofiche, = F photo, 006772, W not seen
Icones гар Bitter, Abh. Naturwiss. Vereine Bre-
n 24(1): 459. 1919 [1920]. TYPE: Ecuador. Prov-
incia Santo Domingo, Sodiro 114/38 (B destroyed, =
F photo 2577)
Volume 84, Number 2
1997
Benitez 4 D’Ar
Lycianthes in Venezuela
193
KZ
LX A
Se \
Va
um
NIIS =
SITAS
SALÓ DAD
OSTIAS
A
4
NS
Q
We
INS
~
XS
N
\
y
Y
Ч
zh
Figure 21. Lycianthes sanctaemarthae.—A. Flowering branch.—B. Flower bud.—C. Opened flower.—D. Infruc
tescence.—E. . A, B, C. After Steyermark et al. 122959 (VEN); D, E. After Bunting 10185 (MO).
Erect or wandlike shrub 1-2 m tall, young stems beneath, sometimes glabrescent above, veins 7-9
densely pubescent, sometimes glabrescent; pubes-
cence of weak, simple, ascending hairs, those on
the internodes to 5.5 mm long. Leaves unequal-
geminate, major leaves elliptical, sometimes
oblique, basally obtuse, the reduced side cuneate,
apicaily acuminate, 10-19 Х 3-7 cm, membra-
on each side, elevated beneath; minor leaves de-
current on the petiole, 5-9 X 2-3 cm; petioles 0.8—
2 cm long, plane above. Inflorescence 5—16-flow-
ered. Flowers diurnal, a few opening at a time;
pedicels 0.6-1.8 cm long, slender, dispersed or
densely pubescent with ascending hairs; calyx 2—
nous, paler beneath, pilose on both sides, more so 3.2 mm long and wide, densely pubescent outside,
Annals of the
Missouri Botanical Garden
Figure 22. Lycianthes sanctaemarthae.—A. Represen-
tative S аы outside of Venezuela.—B. Geographic
distribution in Venezuela.
inside with minute, dispersed glands, teeth want-
ing, the 10 nerves evident, sometimes somewhat
elevated; corolla white, rotate, 6-8 mm long, deeply
lobed, puberulent outside; stamens equal, filaments
2-2.9 mm long, anthers 2-2.5 mm long, drying with
a dark dorsal connective; ovary ovoid, 1.2-1.5 mm
long X 0.8-1.3 mm across, style 6 mm long, ex-
serted 3 mm, stigma capitate. Berry globose, 5.5—8
mm across, stone cells absent; fruiting calyx slight-
ly accrescent, 2 mm long, mostly strigose; seeds
70 per fruit, dark brown, 0.5-1 x 0.75-1.5
mm. Figures 2K, 19.
Esta especie es la única dentro de la Sección Simpli-
cipila en Venezuela que no presenta dientes en el cáliz,
mostrando 10 nervios conspícuos у un tanto elevados.
This is the only species in section Simplicipila
in Venezuela that has no teeth on the calyx.
The foliage and pubescence of this species is
extremely variable, and the material cited may rep-
resent more than one taxon. Dorr et al. 5104 has
smaller, more symmetrical leaves with much less
pubescence than the other specimens.
Distribution. Venezuela, Colombia, Ecuador,
and Peru. Evergreen forests, 1900-2800 m eleva-
tion. (Map, Fig. 20.) The species flowers and fruits
throughout the year.
> ET a examined. VENEZUELA.
Mérida: 1900 m, Benítez et al. 4185 (F, MO,
MY, NY, VEN): E dia of Prado Verde-Las Cuadras, 2150
m, D’Arcy & Benitez 18257 (MO, MY). Táchira: 1 km
SE of alcabala Páramo El Zumbador, 2750 m, Pietrangeli
388 (MY). Trujillo: between La Playa SW of Сагасће and
Potreritos de Cendé, 2200 m, Dorr et al. 5104 (MY, NY).
10. Lycianthes sanctaemarthae Bitter, Abh.
Naturwiss. Vereine Bremen 24 (1): 377. 1919
[1920]. TYPE: Colombia. Sierra Nevada de
Santa Marta, Smith 1189 (holotype, B not
seen; isotypes, CM, F-2, MO).
Shrub climbing to 8 m, young growth tomentose,
glabrescent; pubescence of stellate hairs. Leaves
mostly solitary, ovate, 8-10.5 X 4-8 cm, membra-
nous, paler beneath, veins 3—5 on each side, major
veins elevated beneath, both sides with sparse
hairs; petioles 1-3 cm long, basally compressed,
rulent; minor leaves ovate, 1.5-2.8 X 1.5-2.
cm. Inflorescences in fascicles of 6-13(-16) flowers;
peduncles obsolete or to 10 mm long. Flowers with
pedicels 11-14 mm long, sparsely pubescent; calyx
3-3.5 X 4-5 mm, sparingly evenly pubescent, dry-
ing dark, the margin translucid, teeth wanting; co-
rolla white with a light violet line, the margin sin-
uate-lobed, the lobes less than % the length of the
corolla, 1.1-1.8 X 2-3.5 cm, evenly dispersed pu-
bescent with simple or stellate hairs; stamens un-
equal, 4 filaments 1.5-2.2 mm long, the fifth 4-5
mm long, anthers equal, 5-6 mm long, the tips dry-
ing discolorous; ovary subglobose, 1.5-2 X 1. 5-3
mm, style 10-12 mm long, exserted 2 mm, stigma
subcapitate. Berry orange, subglobose, 9-12 X 8
mm, stone cells 2, apical in the fruit; fruiting calyx
drying uniformly dark brown, 6-8 mm wide, some-
times developing umbos on the sides; fruiting ped-
icels to 2 cm long; seeds ca. 55 per fruit, yellowish
brown, 2-2.5 mm, the thickened margin ca. 0.5 mm
wide. Figures 2D, 21.
Presenta el cályx edentado y la pubescencia está re-
presentada por tricomas estrellados. Los frutos presentan
células pétreas, a diferencia de lo sefialado en la literatura
en donde en la ción Virgatae a la que pertenece sta
especie, no se mencionan células pétreas
The calyx of this species lacks teeth and the pu-
bescence is stellate.
Distribution. Restricted to the Sierra Nevada
t
Volume 84, Number 2
1997
Benítez & D'Arcy 195
Lycianthes in Venezuela
Figure 23. Lycianthes stenoloba.—A. Flowering and fruiting branch.—B. Flower bud.—C. Opened flower. After
i
Mocquerys 978 (MY).
de Santa Marta in Colombia and in the Río Guasare
watershed in Zulia State in Venezuela. Riverine for-
ests from 500 to 600 m elevation. (Map, Fig. 22.)
We have seen flowering specimens from May and
June and fruiting specimens from August.
Although Bitter reported an absence of stone
cells in the fruits of this species, we found two in
the fruit we examined (Bunting & Kauffman
10257).
Representative specimens examined. VENEZUELA.
Zulia: Dist. Mara, Cuenca de los ríos Socuy-Guasare, en
la Paloma, 600 m, Bunting 10185 (MO), Bunting & Kauff-
man 10257 (MO); E of río Guasare, 600 m, Steyermark et
al. 122959 (NY, VEN); Cerro Los Manantiales, E of río
Guasare, W of Hacienda Los Manantiales, 600 m, Stey-
ermark et al. 123281 (NY, VEN).
11. Lycianthes stenoloba (van Heurck &
Muell.-Arg.) Bitter, Abh. Naturwiss. Vereine
Bremen 24(1) 358. 1919 [1920]. Solanum
stenolobum van Heurck & Muell.-Arg., Observ.
Annals of the
Missouri Botanical Garden
196 í
ме" 100° s к n’ Са so” LM
Tna „
-o CT
t
SY w
4 ГА
ө
•
10
x
A w
n* те Су u s Си
4 1°
v
Figure 24. Lycianthes stenoloba.—A. Representative
Pih anis didis of Venezuela.—B. Geographic distri-
bution in Venezuela.
Bot. 69. 1870. Bassovia stenoloba (van Heurck
& Muell.-Arg.) Britton, Mem. Torrey Bot. Club
4(3): 232. 1895. TYPE: Peru. Prope Tarapoto,
Spruce 4210 (G-DC not seen, = IDC micro-
fiche). [A duplicate (syntype) of this collection
was cited as being in van Heurck's personal
herbarium, but its whereabouts are unknown.]
Lycianthes acutangula Bitter, Abh. Naturwiss. Tene
Bremen 24(1): 357. 1919 [1920]. Solanum
gulum (Bitter) MacBride, сым Mus. Nat Hist.
_ 1962. . San
compressibaccata Bitter, Abh. Naturwiss.
Vereine гане, 24(1): 358. 1919 [1920]. ТҮРЕ:
Реги. (Ле 6800 (В not seen, = Е photo 2560 MO,
МҮ
Lycianthes longidentata Bitter, Abh. Naturwiss. Vereine
Bremen 24(1): 356. 1919 [1920]. Solanum longiden-
tatum (Bitter) C. V. Morton, Contr. U.S. Natl. Herb.
29(1): . TYPE: Brazil. Rio Acre, Estella,
Seringal, s. Francisco, Ule 9733 ee B de-
stroyed, = F photo 2582; isotype, US).
Sprawling subshrub to 4 m tall, internodes 3-6
cm long; pubescence of simple hairs. Leaves soli-
tary or geminate, ovate, 9-14 X 3-6.5 cm, apically
acuminate, basally cuneate, veins 4-6 on each
side, both sides with sparse simple hairs on the
main veins, glabrescent, ciliate; petiole 0.5-2 cm
long, slightly pilose. Inflorescences 1(—4)-flowered
fascicles. Flowers ?diurnal; pedicels slender, very
gradually expanding upwards, 1.5—4.5 cm long; ca-
lyx 2.5-3 X 1-3 mm, pilose, with 10 subapical,
erect, unequal teeth in one series, 2.5-5 mm long;
corolla white, rotate or reflexed, lobed Y%-4 way
down, 1-1.5 X 1-2.5 cm, glabrous; stamens mostly
unequal, 4 with filaments 0.2-1 mm long, the fifth
0.1–2.5 mm long, glabrous, anthers 4.5—6 mm long;
ovary ovoid-conical, style 5-8 mm long, glabrous,
exserted 1.5 mm, stigma bilobed. Berry orange,
compressed-ovoid, 9 X 8 mm, sometimes with 2
stone cells; fruiting calyx 8.5 mm long; seeds ca.
24 per fruit, yellowish brown, 3 X 2 mm. Figures
ZH x.
ta especie es conspícua por la longitud de sus ped-
icelos florales cuyas medidas oscilan entre 1.5-4.5 cm де
longitud y la corola presenta el borde lobado, con lobos
de longitud %—% de la longitud de la corola.
This species can be recognized by its slender,
often wide-flaring calyx teeth, its usually elongate,
slender usually arching pedicels, and its anthers
held tightly together in a cylinder. The corolla is
conspicuously lobed.
“Solanum acutangulum Griseb." appeared as a
nomen nudum in Lechler, Berberid. Amer. Austral.
(58. 1857).
Distribution. Venezuela, Brazil, and Peru. De-
ciduous woods and semi-deciduous riversides,
200—700 m. (Map, Fig. 24.) Collected in flower and
fruit from June to September.
Lycianthes stenoloba is much like the following
species from Central America: Lycianthes n
malense Bitter, Lycianthes stephanocalyx, (L5
Brandeg.) Bitter, and Lycianthes solitarium (Blake)
Standl. The Central American species were treated
in part by Gentry and Standley (1974) and Dean
(1995), and Nee (1986) treated Lycianthes stephan-
ocalyx for Veracruz, Mexico. The name L. stenoloba
is employed for this species with hesitation. In the
original description and in the type photos and oth-
er material seen for the synonyms cited, the calyx
is mostly longer—4-9 mm as opposed to 2-3
mm—than in the Venezuelan речь) Bitter re-
ported finding two stone cells in fruits of this spe-
cies, but we found none in the fruits we examined.
Representative specimens examined. VENEZUELA.
Volume 84, Number 2
199
Benitez 4 D’Arcy 197
Lycianthes in Vente
Carabobo: Valle Seco, camino hacia La Toma, 100-200
m, Benítez & Pons 4654 (MY). Falcón: ima de San
re, , van der
MY). Mérida: El Vigia,
Miti 978 (MY, P). uod Finca Antonia, San Fe-
lipe, Ferrari 791 (MY).
Literature Cited
Barboza, G. E. & A. T. Hunziker. 1992, Estudios iid
nire XXXIII. El iere Lycianthes en La Arg
a. Darwiniana dup
вафа de Rojas, С. Е. an géneros de las Sola-
naceae de Venezuela. и Fac. Agron. (Maracay)
7(3): 25-108.
Bitter, G. 1911. Steinzellkonkretionen im Fruchtfleisch
beerentragender жул сееп und deren systematische
Bedeutung. Bot. Jahrb. Syst. 45: 483-507.
— 1 19 Weitere dada! эрсә iiber das Vor-
kommen von Steinzellkonkretionen um Fruchtfleisch
beerentragender Solanaceen. Abh. Naturwiss. Vereine
vind 23: 114-163.
0. Die E о Abh. Naturwiss.
"ran Proosa 24(1): 292
E D. J. Collins, P. 5: Crabbe. F. W. Eastwood,
Wind. ОМ Suah & D. E. Symon. 1978. A
survey 4 Australian Solanum plants for potentially use-
asodine. Austral. J. Bot. 26: 723-754
9. Uber die Entwicklung der Steínzellkon-
kretionen in Пи Gattung Solanum. Kulturpflanze 17:
299-311.
pie W. С. 1973. Family 170. Solanaceae. Jn Flora of
a. Ann. Missouri Bot. Gard. 60: 573—780.
Я The calyx in Lycianthes and some other
Мата Ann. Missouri Bot. Gard. 73: 117-127.
arwin, S. P. & T. Feibelman. 1991. Ly cianthes asarifolia
о d to North America. "Sida 14
par . А. 1995. Systematics and e ohotad of Ly-
ianthes series Meizonodontae. Ph.D. sai Uni-
Wem of California, Berkeley.
Silii e. k A. De Candolle, Prod-
56-183.
Evans, W. C. & A. Somanabandhu. 1980. gps
taining паран secondary
Cyphomandra, Lycianthes and rito a ark
chemistry 19(11): 2351-2356
Gentry, J. L., Jr. & P. С. Standley. 1974. Solanaceae. In
Flora of oe Fieldiana, Bot. 24 (pt. 10, nos. 1
: 1-1
I Trichomes of Solanaceae Tribe
N
onomy-Chemistry-Evolution. Royal Botanical Gardens,
Kew
Hassler, E. 1917. Solanaceae Austro-Americanae. An-
. Bot. Genéve “a 180-189.
ra Amazonica. VI.
Ucayali е no Pampa del Sacramento, п
bre a dezembro de 1898. Bol. Mus. “Goeldi 4: 510-
Kaai S. & C. E. Jarvis. 1990. The typification of »
nam
W
. J. Linn. Soc. 104: 325—367.
¿Mi L Chong & S. Y. Lin. 1987. Steroidal
alkaloids from Solanum capsicastrum. Phytochemistry
26: 305-307.
ен А. 1987. Mirrors ingen Americana-
V. Solanaceae. Phytologi 448.
т С. У. A. Т Hunziker & Е В. Зтић. 1976. А
i
Nee, M. 1981. Tips for salvo егетин Solanaceae
Newslett. 2:
1
‚Бйз Fasc. 49: 1—191. Jn Flora
de Veneta: yes Nac. Inv. Recurs. Biot., pa.
Nevers, G. de. 1986. Pollination of Lycianthes amatitla-
nensis in 5 Panama. Solanaceae Newslett. 2: 36—
Olmstead, R. G., J. A. Sweere, R. E. Spangler, L. Bohs &
J. Palmer. (In press.) derum and provisional
DRM of the Solanaceae based on chloroplast
. Symposium volume, Ath ا e
on the Solanaceae. Royal Botanic Gardens
Ripperger, H. & A. Porzel. 1992. 2 паста
ulcidine from Lycianthes biflora. Phytochemistry 31:
26.
Roddick, J. G. 1986. Steroidal alkaloids of the Solana-
ceae. Pp. 201-222 in W. С. D'Arcy (editor), Solanaceae
eed and Systematics. Colombia Univ. Press, New
Ros: K. 1971. анат d of hairs in the genus Sola-
Solanum pseudolycioides in an enu-
meration of South American plants. Bull. Torrey Bot.
Clu
Schilling, E. E. 198 1. Systematics of Solanum sect. So-
lanum (Solanac eas) in North America. Syst. Bot. 6:
172-185.
Seithe, A. 1962. Die Haararten der Gattung Solanum
und ihre taxonomische Verwertung. Bot. Jahrb. Syst. 81:
–336.
Williams, D. E. 1993. Lycianthes moziniana (Solanaceae):
An underutilized Mexican food plant with “new” crop
potential. Econ. Bot. 47: 387-400
List OF SPE
Lycianthes acutifolia (Ruiz & cn Bitter
Lyci s amatitlanensis (Coult. & J. D. Smith) Bitter
Lycianthes asarifolia (Kunth & Bouché) Bitter
| Lycianthes
iL Lycianthes stenoloba (van ели) & Muell.-Arg.) Ви-
COLLECTIONS OF LYCIANTHES FROM VENEZUELA STUDIED
Specimens are listed alphabetically by нын fol-
eor b Рона or number and herbarium of deposit; th
"s is ar y a number in n paren с
to the number in the text and in the List of
(MO, NY, VEN) (1); 7122 (NY) (1). Aristeguieta & Foldats
1406 (VEN) (3). Aristeguieta & Pannier 1973 (VEN) (3).
198
Annals of the
Missouri Botanical Garden
Aymard & Flores 216 (MY, PORT, VEN) (7). Aymard et
al. 1643 (MY, PORT) (8); 2839 (MY, PORT) (3).
Badillo 1816 (MY) (6); 1914 (MY) (8); 4426 (MY) (8);
5672 (MY) (4); 6584 (MY) (7); 6636 (MY) (8); 6687 (MY)
(4). Badillo & Holmquist 6217 (MY) (8). Badillo et al.
7841 (MY) (8). Benftez 318 (MY) (6); 517 (MY) (3); 683
(MY) (7); 1181 (MY) (8); 1335 (MY) (8); 1353 (MY) (8);
1408 (MY) (4); 1418 (MY) (1); 1446 (MY) (4); 1551 (MY)
(1); 1554 (MY) (6); 1562 (MY) (8); 1952 (MY) (8); 2068
(MY) (4); 2243 (MY) (8); 2614 (MY) (6); 3214 (MY) (6);
3621 (MY, NY) (3); 3868 (MY) (1); 3869 (MY) (4); 3898
MY) (1). Benítez & Otero 4609 (МУ) (1). Benítez & Pons
MY) (11). Benítez € Rojas 308
M
4261 ничу (8); 4611 (MY) (9);
(MER, VEN) (1). Bunting 2908 (MY) (8); 3020 (MY) (8);
4433 (MY) (6); 5808 (VZU, MO, VEN) (6). Bunting 10185
(MO) (10). Bunting & Kauffmann 10257 (MO) (10); Bur-
kart 16324 (VEN) (6).
Cárdenas 4038 (MY) (8). Castillo 1930 (MY) (9). Car-
nevali et al. 614 (MY, VEN) (8). Cesari (VEN-249120)
(је Colonello 930 (CAR) (8). Croat 54841 (VEN) (5). Cu-
a 1850 (IRBR, MY) (6).
Arcy & Benitez 18257 (MO, MY) (9). Davidse 4035
tede (4). Davidse & Steyermark 18168 (VEN) (7). Dav-
idse & González 18899 (NY, 6 (5); 21914 (МУ) (8).
Davidse & Miller 27476 (MO, MY) (8). Delascio 51 (CAR)
o р: (САВ) (1); 981 (САВ) (9 т (CAR) (4). De-
о & de Delascio 2692 (CAR, VEN) (3). Delascio &
ud 12838 (MO, VEN) (3). Denie же МҮ-86695
(4). Diederichs 173 (VEN) (4); 177 (VEN) (8); 270 Lp
(8); 279 (VEN) (8). Dorr & Barnett 71
Dorr et al. 4749 (MY, NY) (8); 5104 (NY, VEN) o, a
(MY) (6).
(M
girde 974 (MO, P) (1); 991 (G, MO, NY) (4); 1065
(3). Fernández, A. 379 (MY) (3); 497 (MY) (6); 606
Му (8); 620 MY) 3) (8; 1112 (MY) (8); 3697 (MY) (b
3762 (MY) (8); 3 MY) (8); Fernández, F. 98 (VEN
(8). Ferrari 733 a (8); 790 (MY) (3); 791 (MY) (11);
859 мо (7).
Puig-Ross 14293 (MO) (8). Gentry & Stein
47275 (MO, МЕ VEN
Humbert 26108 (MER, P) (1); 26156 (MER, P) (1).
Humboldt 748 (P) (6).
Ijjasz & Madriz 164 (MY, VEN) (7).
Jahn 1211 (VEN) (6); 1248 (VEN) (6). Jeffrey & Trujillo
2510 (MY) (8).
ME & Mallet 6766 (BH, MY) (2); 6807 (BH, MY,
VEN) (1
Lasser 1068 (VEN) e gd dra xd ~ (VEN)
(8); 2212 (VEN) (8). Lic t al. 32 (MY, PORT) (3).
Liesner pes а, (1); 12888 on VEN) (5). Liesner
unk 15838 (MO, MY, VEN) (8). Liesner & González
Mes E e Kul (VEN) (2); 9969 (VEN) (8). Liesner
& Guariglia 11631 (MO, NY, VEN) (8). Liesner et al.
7803 (MY, em m (3). Linden 437 (C) (4); 478 (P) (3).
pez-Figueiras & Rodriguez 9080 (ME 7). López-
Ló
Palacios 1457 (МЕКЕ, МО, MY) (7); 1539 (МО) (4); 2206
(МЕКЕ, MY, VEN) (8). López-Palacios € Bautista 3485
(MER) (8).
Miplimd (MY-9313) (6). Manara (MY-83285) (11);
pen 71770) (3); (VEN-176544) (3). Marcano-Berti 1409
ER) (8). Marcano-Berti & López-Palacios 1758 (MER,
E (4). Marcano-Berti & Carrillo 29-4-78 (MER) (8).
Mocquerys 880 (P, VEN) (6); 978 (MY, P) (11); s.n. (P)
(6). Маја 3756 (MY) (8). Morillo 2519 (VEN) (2);
3384 (VEN) (1); 11148 (MERF, MY) (7). Morillo & Сагсја
11472 (MERF, MY) (1). Morillo & Manara 1600 (MY,
VEN) (i 2022 (VEN) (8) 2070 (MY) (3). Morillo & Seres
8614 (VEN) (1). Morillo & Smith 6057 (MY, VEN) (4).
Morillo E al. Pp (VEN) (8); 3272 (VEN) (8). Moritz
164: se
==
Oats (и Y, NY, VEN) (8). Nee € Whalen
16899 (МО, NY) (8); 17053 (VEN) (7); 17146 (NY, VEN)
E 00 60 (МО, NY) (8). Nilsson & Steyermark 221
8).
че
NAS 892 (VEN) (3).
Pannier 199 (МЕКЕ, VEN) (5). Pefour 6 (MERC) (7).
Pefialoza 205 (CAR) (1). Pietrangeli 338 (MY) (9); 1331
(MY) (9). Pittier 5972 (NY) (3); 8803 (VEN) (8); 9129
(NY, US, VEN) (3); 9378 (NY, VEN) (1); 10036 (VEN)
1); 10057 (NY, VEN) (1); 11179 (VEN) (6); 11867 (MO,
NY, VEN) (8); 13088 (MO, NY, VEN) (6); 13237 (MO,
NY, VEN) (7); 13514 (MO, VEN) (4). Pittier & Nakichen-
ovich 15543 (VEN) (8). Plowman 7766 (P, MO, NY) (1).
Plowman et al. 13445 (NY) (8). Poelt & Oberwinkler
14991 (VEN) (7).
Quintero 2173 (MER) (9). Quintero & Hernández 246
(M
~
Reales 2071 (MY) (4). Ricardi 1093 (MERC) (1).
G. 770 (MY, VEN) (9); Rodrí ат Н. 74 (МҮ)
.R &
(3). ard
1712 (MY) (1). Romero 486 (MY) (6); 854 4 (MY) (8). Ro-
sales 11 (MY, VEN) (4). Ruiz-Terán fin (MER) (8); 2128
(МЕКЕ) (1); 8802 (MERF) (7). Ruiz-Terán & López-Fi-
gueiras 64 (MERF, MY) (1); 245 E HERA MY) (7).
Ruiz-Terán & López-Palacios 6708 (МЕКЕ) (7); 11470
(MERF, MY) (8); 12533 (MERF, MY) (7).
Saer 118 (VEN) (6); 263 (NY, YEN) (3). Sanoja, E.
. Schwarzkopf 12
; V4464 (MY, UCOB) (8); V4465 (VEN) (4); V4466
VEN; (4); 7557 (мт, VEN) (4). Smith V9384 (МУ) (8). |
bel et al. 2062 (NY) (3). Stein $ Laa 1512 (MO,
MY, VEN) (8). de & Aymard 4451 (MY, PORT) (3).
Stergios et al. 6360 (MY, PORT) (5). Steyermark 55405
(MY, VEN) (1); 55945 (MY, VEN) (4); xs am M
56262 (VEN) (7); 56494 (NY, VEN) (4); 56628 (F, M
VE
а (м0. NY ) 1
(МО, VEN) (8); 1
121593 (VEN) (2); aeons ухо (l); 126813 (VEN) 68;
126823 (VEN) (8). Steyermark & Agostini 91324 (VEN)
Steyermark & Liesner 118330 (VEN) (9); 1 8386 (MO,
VEN) (9); 119186 (VEN) (8); 120743 ся [^ 8); 120772
(MO, NY, VEN) (2). Steyermark & Manara 110995 oa
NY, VEN) (6); 125326 (VEN) (1). rl & M
123694 (MY, VEN) (8). Steyermark & Stoddart 118046
а аса а ~ EE EN E T чый Са Нн T а A a РАВНИ
Volume 84, Number 2
1997
Benitez & D’Ar
Lycianthes in Venezuela
199
(VEN) (8). Steyermark & Steyermark 95433 (VEN) (3).
Steyermark & Rabe 96125 (P, VEN) (2); agis за "1
Steyermark & Wessels-Boer 100481 (MO, N (8).
sit et al. 92969 (VEN) (8); 98795 = ven, (4);
(VEN) (8); 127216 (MY, VEN) (3). Suarez & Gil 53 (MER)
Tamayo 283 (VEN) (9): at (VEN) (4); 1249 OE (1).
Trujillo 1260 (MY) (4); 2917 (MY) (1); 3325 (MY) (3);
; 15829
à Y)
(1). Trujillo 4 де! vee 8265 (MY) (1). Trujillo & Fer-
597
16574 (MY) (6); 1
van der Werff d (VEN) (8); 4811 (VEN) (8); 5453
(VEN) (8); 6124 (MO, NY) (8). van der Werff & Vera 864
(CORO, MY) (11). Velasquez 110 (CAR) (8); 310 (CAR)
1
(1).
Whetzel & Muller 485 (VEN) (1). Williams 10269
(VEN) (8); 10789 (VEN) (8); 11084 (MY, VEN) (3). Wing-
field 12989 CORO, n (11). Wood 448 (VEN) (3). Wood
& Berry 85 (VE
Xena 626 at (4 dg
INDEX TO SCIENTIFIC NAMES
Asaropsis
lenta 182
Section 171
Bassovia 182
inaequilatera 182
stenoloba 196
о 179
repens coma by!
Brachistus 182
inaequilat 182
Capsicum 167, 170
Jaltomata MID
Lycianthes
enta 171
acapulcensis 172
acutangula
acutangula subsp. compressibaccata............---------
acutifolia 171, 174
amabtlanenals с, м а, 171, 172, 175
armentalia о 182
asarifolia 171, 172; 177, 179
australe 190
ind CT а сна 187
йир o E Li ue 171
(етика. ао 171, 172, 180, 182
EE. ш а Ре
iude AAA A 192
O ен eens 171
iti ver ee и етен 174
CUAL еее 196
БШ Aa AER ыен ота 190, 192
hola .— -___-_-______._ ___-_--
bei! ПРАВНА he 171, 172, 176, 182
PEA з SNORE Нар deu dd 190
КЫШ еј os 182
lehmannii 174
lenta
lenta var endopsila
171, 172, 184
185
185
lenta var. scotinophila
"prom tata
196
168, 170-172, 186
187
oido subsp. tomentosa
171
“
то
niana 171
pauciflora... 168, 171, 172, 182, 185, 189, 190, 192
radiata 171-173, 192
rantonnei 171
repens 178, 179
t 171, 194
Section 171
awe Asaropsis 171
solitariu 196
кис; 171, 195, 196
stephanocalyx 171, 196
Subgenu 171
subsect 172
ulei 176
ulei subsp. dolichodonta 176
ulei var. strigulosa 176
variifolia 171, 186
xylopiifolia 174
xylopiifolia var. intermedia 174
series 172
Oligochondra
ries 171
til 172
Pachystemonum
172
Physalis 170
Polymeris
Section 171
Subgenus 171
Simplicipila
Section 171, 194
lani
hortensis folio, fructu corallino..................... 189, 190
lanum
acutangulum 196
acutifolium 174
amatitlanense 176
asarifoliu 177, 178
australe 190
candican 186
caucense var. glabrescens 190
chodatianum 178
ib 196
n 184
declinatum 184
geminatu 189
goudoti 192
guianense 190
japurense 190
lambii 184
lentum 184
lentum var. echinatum 184
longidentatum 196
lyciifo 187
lycioides 167, 172, 186
lycioides subsp. parvifolium ................................... 188
lycioides var. tomentosa ODT 186
neglectum 189
200 Annals of the
Missouri Botanical Garden
nocturnum 184 A EII bc Lo EE 180
MEN Lieu Р ОВОЧЕВА CIEN DR diis a 189 violaefolium fo. lilaciniflora ı.2 178
phillyreoides 18 violaefolium var. maj 177
OOO CIES ERA 187, 188, 190 violiifolium var. asarifolium cc 177
quadriflorum 1 em var. asarifolium fo. decadon-
radiatu 192 178
sect. Lycianthes 171 violiifolium var. majus fo. chacoense .................. 178
sect. cena subsect. Lycianthes............ 172 virgatum 184
series Meiomeri 172 virgatum var. lentum 184
Ире бит A A ON SEE 190 ylopiaefoli 174
stenolobum 195 Triguera 170
subgenus Lycianthes 172 Virgatae
sylvaticum 184, 190 Series 171
urbanum var. ovatifolium 190 Witheringia 170
DIALYPETALANTHUS
FUSCESCENS KUHLM.
(DIALYPETALANTHACEAE):
THE PROBLEMATIC
TAXONOMIC POSITION OF
AN AMAZONIAN ENDEMIC!
Frederic Piesschaert,?
Elmar Robbrecht? and Erik Smets?
ABSTRACT
al and anato
ers vid calcium oxalat
postu Mae pde calicis In the past, Dia
Myrtales or ‘Contes es
ac
elucidation of gu relationships of Dialypet
nd corrections de erroneous rune observations, e.g., on su
nced gynoecial сет hos such a features as free
ead. It is concluded that ontogenetic and mic
talanthus.
е neotropical rainforest tree iiem prieto fuscescens Kuhlm. is the only species of the етан.
cal documentation is presented, including newly reco features
e packages, silica bodies in
such as inde-
the wood, U-shaped ere peer
pposed stipular dimorphism. The genus
petals, dimery, stipules,
ost frequently а to be related to
However, because of the lack of of e " ne bundles, pa e of well-de veloped
ial origin of q
Ru e in particu
and macromolecular data are much needed for a "уты
The monospecific Amazonian genus Dialypeta-
lanthus is enigmatic as regards its systematic po-
sition. Characters such as the opposite, entire
leaves with sheathing stipules and the dry frui
with winged seeds obviously point to a position in
(or near) the Rubiaceae. However, a close relation-
ship with the Rubiaceae must be questioned be-
cause of the presence of certain deviant floral char-
acters (e.g., free petals and numerous stamens in
two whorls with their filaments fused into a basal
ring). As for the classification of Dialypetalanthus,
all present authors accept its familial status, but
they do not agree about the relatives of the Dialy-
petalanthaceae. While some prefer a position in the
Myrtales or Rosales, others assign the family to
Gentianales or Rubiales. In the present paper we
aim (1) to document knowledge of Dialypetalanthus
fuscescens Kuhlm. as broadly as possible and (2) to
clarify its systematic position.
HISTORICAL SURVEY
In October 1874, Ferreira was the first to collect
some flowering branches of a yet unknown tree from
the Amazonian region. The specimen (Ferreira 438)
was filed in the herbaria of K and LISU as an un-
identified taxon of the Rubiaceae. In 1925 the tree
was re-collected and described by Kuhlmann as the
new genus and species Dialypetalanthus fuscescens
Kuhlm
According to Kuhlmann (1925, 1942), the as-
cending imbricate ovules, winged seeds, and em-
bryo “formando núcleo central” (1942: 25) clearly
indicate a relationship between Dial nthus
and Rubiaceae, more specifically with the tribe
Cinchoneae of the subfamily Cinchonoideae sensu
Schumann (1891). Kuhlmann established a mono-
generic tribe Dialypetalantheae in the Rubiaceae,
characterized by free petals and anisomery.
After a reinvestigation, mainly of wood and leaf
anatomical characters, Rizzini and Occhioni (1949)
established the monotypic family Dialypetalantha-
ceae. They concluded that the Dialypetalanthaceae
are to be placed in the Myrtales, somewhere near
the Myrtaceae (Eugenia), Melastomataceae (Hub-
епа), or Нудгосагуасеае (= Trapaceae). Note that
We are grateful to Steven Jansen for preparing
1 the wood sections, to L. Macias and Р. Delprete for providing
additional distribution data, to the directors of the herbaria of K, NY, L, and US for providing herbarium spec
i mens,
to Marcel Verhaegen for the tiem qut of SE-micrographs of pollen, and to Johan Buelens for making photographs of
hw
e um specimens. Thi
projet G.0143.95
supported by the Fund for Scientific Research-Flanders (F.W.O., Belgium,
).
* Laboratory of Plant Systematics, Botanical Institute, K.U.Leuven, Kardinaal Mercierlaan 92, B-3001 Heverlee,
Belgium
з National Botanic Garden of Belgium, Domein van Bouchout, B-1860 Meise, Belgium.
ANN. Missouri Bor. Garb. 84: 201-223. 1997.
202
Annals of the
Missouri Botanical Garden
Kuhlmann (1925) had already mentioned that the
copious flowering of Dialypetalanthus reminded
him of certain Myrtaceae. In addition, Rizzini and
Occhioni sent samples of Dialypetalanthus to some
“eminent botanists” (Burkart, Erdtman, Lam, Jans-
sonius, and Bremekamp), asking for their opinion.
In 1952, Occhioni and Rizzini assembled their an-
swers into a publication. All the taxonomists con-
sulted accepted the establishment of a new family,
which probably explains why it is generally ac-
cepted today. They disagreed, however, about its
possible relatives. Burkart stated that relatives of
Dialypetalanthus should be looked for somewhere
in his Myrtiflorae; Erdtman concluded that the pol-
len of Dialypetalanthus resembles that of certain
Rhizophoraceae (see also Erdtman, 1971); Lam,
while admitting some resemblances to the Rubi-
aceae, suspected a relationship with Lythraceae or
Melastomataceae; Janssonius found similarities be-
tween the wood of Dialypetalanthus and that of the
genus Psychotria of the Rubiaceae; finally, Bre-
mekamp agreed with Rizzini and Occhioni (1949)
that relatives of Dialypetalanthus should be sought
near the Myrtaceae and Melastomataceae, notwith-
standing the large leaf-like stipules and lack of in-
traxylary phloem (see also Bremekamp 1966: 3, 8).
Emberger (1960), Melchior (1964), Cronquist
(1968), Stebbins (1974), and Dahlgren (1975) fol-
lowed Rizzini and Occhioni (1949) and placed the
Dialypetalanthaceae in the Myrtales. Cronquist
(1981), however, reconsidered his opinion, placing
Dialypetalanthus in “the rather amorphous Ro-
sales” (p. 551), where it was kept in his system of
1988. He refused to associate the family either with
Myrtaceae (because of the stipules and the lack of
internal phloem) or with Rubiaceae (because of the
free petals and numerous stamens on top of the
ovary).
Dahlgren also changed his view on the system-
atic position of Dialypetalanthus. At first, he trans-
ferred the Dialypetalanthaceae from Myrtales to
Cornales (Dahlgren, 1980); later he moved the fam-
ily to Gentianales (Dahlgren, 1983; no arguments
were provided for this transfer, but are presumably
the same as given in Dahlgren & Thorne, 1984;
discussed further).
According to Hutchinson (1959), free petals and
polyandry occur in Dialypetalanthaceae, as well as
in some genera of the Rubiaceae, and should not
be considered as distinctive characters (but his ar-
gument does not hold; see Discussion). He placed
the Rubiaceae and Dialypetalanthaceae in a sepa-
rate order Rubiales, closely related to the Logani-
aceae (Gentianales).
In a study of the Myrtales, Dahlgren and Thorne
(1984) concluded that Dialypetalanthus should be
excluded from the order. Stressing inter alia the
paracytic stomata, they considered the Dialypeta-
lanthaceae as an “early off-shoot of the Rubiaceae
or a relict family closely related to the Rubiaceae
in the Gentianales” (p. 690).
Robbrecht (1994), in a review of the delimitation
of the Rubiaceae, added a few new characters that
are in common between Dialypetalanthaceae and
Rubiaceae, namely the thickening pattern of the
inner tangential walls of the exotestal cells and the
occurrence of colleters on the adaxial surface of the
stipules. He concluded that a profound study of the
Dialypetalanthaceae is needed to shed more light
on the possible relationships.
In their consensus classification of the Gentian-
ales, Nicholas and Baijnath (1994) suggested that
Dialypetalanthus represents an archaic group with-
out any close living relatives. They excluded the
genus from the Gentianales and placed it near the
order Cornales and its allies, concluding that it
needs much closer analysis.
MATERIAL AND METHODS
This study is based on herbarium material from
NY, US, and K (herbarium abbreviations following
Holmgren et al., 1990). Numerous attempts to ob-
tain fixed material were unsuccessful. The distri-
bution map was completed with locality data from
specimens from the Museu Goeldi (MG) (data pro-
vided by L. Macias, Belém).
Both for scanning electron microscopy (SEM)
and light microscopy (LM) observations, the her-
barium material was prepared by simple boiling or
by the method described by Peterson et al. (1978),
using a mixture (6:1) of 10% aqueous di-(2-ethyl-
hexyl) sodium sulfosuccinate, and 98% acetone for
24 hours (see also Erbar, 1995). For LM, the ma-
terial was dehydrated in an alcohol series and em-
bedded in paraffin, using a Histokinette 2000. Sec-
tions (10-14 jum) were made with an MIR Shandon
rotation microtome, and subsequently colored with
safranin (10 min) and fastgreen (4 min) in a Var-
istain 24-3, Shandon. After staining, the sections
were mounted in Eukitt. Observations were made
with a Dialux 20 Leitz microscope.
For SEM, the material was dehydrated in an al-
cohol series and FDA (formaldehydedimethylace-
tal). After critical-point drying (CPD 030 Critical
Point Dryer, Balzers), the material was mounted on
stubs with Leit-C Conductive Carbon Cement and
sputter-coated with gold (+ 180 nm) in a Spi-mod-
ule® Sputter Coater (Spi Supplies). Observations
were made with a JEOL JSM-6400 electron micro-
Volume 84, Number 2
1997
Piesschaert e
poor ie fuscescens from Amazonia
Figure 1;
vein; bar = 0.25 mm.—B. Dendritic hair on the
= 50 р
scope. In order to show the thickenings of the inner
tangential walls of the exotestal cells, the outer tan-
gential walls had to be removed. Therefore, boiled
seeds were transferred to a 3:1 mixture of alcohol
96% and acetic acid 99% for 24 hours. Afterward
they were put in a 3% aqueous mixture of sulphuric
acid at 50°C for at least 10 hours (modified from
Braune et al., 1967). Remaining specks of dirt were
removed by ultrasonication.
e methods used for acetolysis and breaking
pollen grains are discussed in detail by Huysmans
et al. (1994). Wood anatomical sections were pre-
pared as described by Jansen et al. (in prep.).
RESULTS
VEGETATIVE STRUCTURES
Нађи. Dialypetalanthus fuscescens is a large,
slender rainforest tree reaching up to 30 m (but
usually smaller). It has a soft, fibrous (because of
the formation of several concentric phellogen lay-
Th
ers), red to cinnamon-colored bark. The trunk is
main vein; bar =
m.—C. Tran
m.—D. Paracytic stoma in pera view; bar = 10 jum. fs ае Ferreira 9018 (A-C) and Lobo et al. 328 (D).
Leaf structure of diving tes fuscescens. (A-C Ae E SEM. pud Transverse section of n ma
verse section of the lamina; "e
)
fluted toward the base. The wood is light but ex-
tremely hard. It is used locally for house construc-
tion (Rizzini & Occhioni, 1949), but as far as we
know it has little economic value.
Leaves. The simple leaves are decussately ar-
ranged and have blades with entire margins. The
petiole is well developed (0.6–3.5 cm) and shallow-
ly sulcate above. The blades are broadly elliptic
(L/W-ratio 1.5:1), elliptic (2:1), suborbiculate (1.2:
1), to narrowly (2:1) or widely obovate (1.2:1)
(sometimes slightly asymmetric), with a shortly acu-
minate to rounded tip, and an obtuse, acute or cu-
neate to slightly decurrent base (terminology follow-
ing Hickey, 1988); they are (2-)6-17(-20) x (1–)
4—11(-14) cm. The leaf venation is pinnate and
camptodromous. The veins, especially the midvein
(Fig. 1A), are prominent on the abaxial side of the
blade. The divergence angle between primary and
secondary veins is about 35° in the center of the
blade and gradually rises toward the base of the
leaf. The veins are detectable up to the fifth order.
204
Annals of the
Missouri Botanical Garden
The areoles have a more or less constant shape and
size. The leaves become gradually smaller and have
a lower L/W-ratio toward the inflorescence region,
and are eventually replaced by bracts. The petiole
and veins are densely covered with unicellular to
uniseriate hairs on the abaxial side. Dendritic hairs
occur as well very sporadically (Fig. 1B). The ad-
axial side of the leaf is almost completely glabrous
or bears few hairs along the main veins. Domatia
are absent. The upper epidermis consists of flat,
rectangular cells (Fig. 1C) that occasionally contain
crystals of an unknown nature. The strongly devel-
oped cuticle runs over the leaf margin and gradu-
ally becomes thinner on the lower epidermis. The
outer tangential wall of the epidermal cells is
straight, except above the veins, where it is round-
ed. Beneath the upper epidermis, a well-developed
hypodermis occurs with clearly larger cells. The
palisade tissue is well developed and consists of a
maximum of 6 cell layers (often less). It is inter-
rupted only by the main veins. At the leaf margin,
the palisade tissue is replaced by angular collen-
chyma. According to Rizzini and Occhioni (1949),
the spongy mesophyll and palisade tissue are sep-
arated by a layer of isodiametric cells (not seen by
us). The lower epidermis (inclusive of the hypoder-
mis) is much thinner and more irregular than the
upper epidermis. The difference in cell size be-
tween epi- and hypodermis is much less рго-
nounced. Stomata are paracytic, randomly oriented,
more or less sunken, and found only in the lower
epidermis (Fig. 1D). The vascular configuration of
the petiole and midvein is complex and rather vari-
able in successive sections (Fig. 1A). In the center,
short radial vessel rows are arranged in an arc,
open toward the adaxial side. The main leaf ana-
tomical characters were described by Rizzini and
Occhioni (1949). Most of their observations were
confirmed in our preparations.
Stipules. At each node, two large leaf-like stip-
ular lobes—the most conspicuous vegetative char-
acter of Dialypetalanthus—occur on either side be-
tween the two petioles (Fig. 2). Rizzini and
Occhioni (1949), Hutchinson (1959), and Cronquist
(1981) considered them to be intrapetiolar, while
Dahlgren and Thorne (1984) described them as in-
terpetiolar. We observed that the four stipular lobes
are connected at their base to form a low sheath
around the stem, i.e., they are intra- as well as
interpetiolar. This sheath is covered adaxially with
a basal row of long silvery hairs and colleters of
the standard Rubiaceae type (that is, consisting of
an axis of elongated cells, covered by a palisade-
like epidermis; Robbrecht, 1988). The abaxial sur-
Figure 2. Stipular dimorphism at the base of an inflo-
rescence; lb = leaf base, s = stipule, la = lateral inflo-
rescence axis, ma = main inflorescence axis.
face is densely covered with unicellular to unise-
riate trichomes. Rizzini and Occhioni (1949)
claimed that Dialypetalanthus has two types of stip-
ules, namely normal and “bud stem covering” ones
(“bud stipules” as we will call them in the following
discussion). They considered this so-called “stipu-
lar duality” to be unique in the dicotyledons. The
bud stipules would bear a “remarkable pilose scale
at the base” and fall off when the “gemma” (apical
bud) increases. We have dissected several intact
terminal buds (Fig. 3A), looking for possible scars
of former structures, but could not confirm the find-
ings of Rizzini and Occhioni. After we had removed
the outer stipules (these are the bud stipules of
Rizzini and Occhioni) of a terminal bud, two struc-
tures, densely covered with hairs, indeed appeared
(Fig. 3B, C). However, their position (alternating
with the last normal pair of leaves and alternating
with the hairy structures of the next node; Fig. 3D),
and their clearly pinnate venation pattern, proves
that these “pilose scales” are reduced leaves.
Therefore, there is no essential difference between
bud stipules and the other stipules. The only gen-
uine stipular dimorphism we noticed is in the tran-
sition zone between vegetative and generative parts,
i.e., at the base of the inflorescence. Here consec-
utive nodes bear stipules that differ strongly, mainly
Volume 84, Number 2
1997
Piesschaert et al. 205
Dialypetalanthus fuscescens from Amazonia
If: leaf (sy, scar of leaf; py, petiole); rl,, rl: reduced leaves; ss,, 55,, SS}: stipular sheaths
Figure Dissection of a terminal vegetative bud of Dialypetalanthus fuscescens.—A. Intact bud.—B. Half of the
outer em sheath removed, revealing a reduced hairy leaf and the stipular sheath of the next node.—C. Outer
stipular sheath removed completely.—D. Schematic representation of a vegetative bud. After Nee 34472.
in size (upper ones much larger than the short, tri-
angular lower ones; Fig. 2). Anatomically, the stip-
ules have a characteristic adaxial epidermis, con-
sisting of thickened cells. Scattered through the
homogeneous chlorenchyma are numerous small
vascular traces that are surrounded by fibers. Riz-
zini and Occhioni (1949) recorded occasional par-
acytic stomata in the abaxial epidermis.
Wood anatomy. The wood of Dialypetalanthus
shows indistinct growth rings (probably reflecting
the precipitation cycle), marked by a transition
from thin- to thick-walled fibers. The fibers are sep-
tate (Fig. 4C) and have simple pits in vertical rows
(Fig. 5D) (libriform fibers sensu Baas, 1986). The
wood is diffuse-porous. The vessels are solitary or
arranged in short radial rows (2-8 cells) (Fig. 4D,
E). In young wood, close to the pith, the radial
vessel rows are often longer. The outline of the sol-
itary vessels is rounded (Fig. 4E), although the
smaller vessels are often compressed between the
larger ones. Axial parenchyma is present and oc-
curs as scanty, paratracheal strands (Fig. 4B). The
width of the rays varies from (1—)2 to 6 cells (Fig.
4A); their height may exceed 100 cells. In radial
sections, the procumbent body-ray cells have a
margin of one or often several layers of square cells
(Fig. 4F). Sometimes a mixture of procumbent and
more or less square cells occurs. The rays are vis-
ible to the naked eye as clear, narrow, parallel
lines. The pith and the rays contain very small cu-
bic to prismatic crystals; navicular crystals occur
as well. The septate fibers as well as the rays con-
tain silica-bodies (Fig. 4G, H). The perforation
plates of the vessels are simple, bearing a single
elliptical to almost circular opening (Fig. 5A). The
vessel-ray pits are simple (Fig. 5B). The intervessel
pits are alternately arranged and vestured (Fig. 5C).
Internal phloem is absent.
REPRODUCTIVE STRUCTURES
Inflorescence. The bloom of Dialypetalanthus is
copious, with numerous white, fragant flowers in
large inflorescences. The inflorescence of Dialype-
talanthus (Figs. 6—8) was described by Kuhlmann
(1925) as paniculate and racemose. Rizzini and Oc-
chioni (1949) and Hutchinson (1959) also indicated
it was a panicle. According to Weberling (1992), a
panicle is characterized by terminal flowers on its
main axis and side branches. Thus, a panicle is a
determinate inflorescence. Dialypetalanthus, how-
ever, does not have a terminal flower but a terminal
bud that at first sight may be confused with a single
terminal flower. Dissection of the terminal bud re-
veals a floral meristem where acropetal inception
of lateral flowers occurs (the flowers in the most
terminal zone seem to be poorly developed) (Fig.
7). The inflorescence of Dialypetalanthus is thus
indeterminate, contrary to a panicle. Dahlgren and
Thorne (1984) provided the first correct illustration
of the inflorescence but did not describe it. Cron-
quist (1981) was the first to correctly call the Di-
alypetalanthus inflorescence a thyrse. More specif-
ically, it is a frondobracteose, heterothetic,
indeterminate thyrse with opposite branches (Fig.
Annals of th
Missouri تن Garden
^+ ow „ж а
є
|
E
r
1,
142
ea
A
T CY
4. Wood anatomy of Bab vidis lire ios Es —A. lor eine section; bar = 0.5 mm.— B. Radial
section showing paratracheal parenchyma (arrows); bar — 0.35 mm.— C. Tangential section рум sept fibers; bar = 50
pm.—D, E. Transverse sections with short radial pr rows en 4-cellular "E bars: D = 0.5 m = 0.2 mm
F. Radial section of a ray with нии layers of square marginal cells; bar = 0.5 тт. —С. Radial section showing
silica bodies in a ray; bar = 0.2 mm.—H. Radial эзен ү detail i silica bodies; bar = 50 pm
8) (all terms sensu Weberling, 1992). The inflores- else). Bracteoles are situated somewhere near the
cences are (usually) terminal. Normally, two side middle of the pedicel. The bracts, bracteoles, and
branches occur at each node (three branches in the true leaves of the inflorescence are all deciduous-
type specimen, Kuhlmann 1514; observed nowhere As mentioned earlier, the transition zone from
Volume 84, Number 2
97
Piesschaert et al.
Dialypetalanthus fuscescens from Amazonia
а а | Le
igure 5. Wood anatomy of Dialypetalanthus fuscescens (SEM).—A. Simple rod Я а vessel.—B. Simple pits
between a vessel and underlying ray parenchyma.—C. Vestured pits.—D. Simple fiber pits
woody to herbaceous parts at the base of the inflo-
rescence is characterized by a short stem portion
with different-sized stipules.
Calyx. The calyx of Dialypetalanthus consists
of four green, semicircular lobes. These lobes arise
as two decussate whorls of two primordia (Fig. 9G).
In early ontogenetic stages, the sepals cover the
other floral structures completely (cryptopetaly sen-
su Sprague, 1940). Later, they are pushed apart by
the developing petals, stamens, and style. The ab-
axial side of the sepals as well as that of the petals
is covered with a typical, velvet-like indumentum,
consisting of long uniseriate hairs, with the excep-
tion of a rather broad (1 to 2 mm) hairless border.
The margins are fringed. Apart from the base, the
adaxial side of the sepals is glabrous. As in the
other floral structures, the sepal lobes are thick and
fleshy. In the center, they usually consist of about
20 cell layers. The cells of the epidermis are clearly
smaller than the cells of the underlying parenchy-
ma. Numerous vascular traces occur in the center
of the parenchyma. The sepal lobes are persistent.
During maturation and dehiscence of the fruit, they
are split in two (Fig. 15A
rolla. The corolla is composed of four free,
white petals. Just like the calyx lobes, they arise
as two decussate whorls of two primordia each
(Figs. 9G, 10). This dimerous origin of calyx and
corolla (sometimes noted in the literature: Cron-
quist, 1988; Maas & Westra, 1993) should be rep-
resented on a floral diagram with petals opposite to
sepals (Fig. 10), instead of alternating with them.
In very young buds, all petals are globular and соу-
er almost completely the interior floral parts. In old-
er stages, petals enclose the interior floral parts in
pairs. During anthesis, the petals unfold into broad-
ly elliptic (2 to 2.5 cm long and 1.5 cm wide),
shortly ungulate structures. The innermost petal is
distinguished from the other petals because of an
apical prolongation with transparent hairs at the
margin in the bud stage. The inner petals are often
more or less wrinkled, thus allowing rapid expan-
sion during anthesis. Aestivation is imbricate, but
with variations. Sometimes a petal will overlap an-
208
Annals of the
Missouri Botanical Garden
New T Botan піса Garden
Fernandes Cas
Y PLANTAE BOLIVIANAE
Javier FERNÁNDEZ CASAS
PANDO, Nicolás Suárez: entr obija y
Porvenir, malezas y restos "Ae ү eod
Fdez. Casas n & Susanna, 12
Arbolito m, flores Prancis, .
e б. Flowering gene of Dialypetalanthus fuscescens with flowers in late bud stage (Fernández Casas &
Paises. 8188, NY); bar =
other on both sides, as is always the case with the
inner petal whorl (and the sepals); sometimes the
overlapping will occur only on one side (this has
only been observed in the outer petal whorl). As
mentioned above, the abaxial indumentum of the
petals is comparable to that of the calyx lobes. The
adaxial side is glabrous. The corolla is shed shortly
after anthesis (Fig. 9D). It is not clear whether pol-
lination has already occurred at that stage or if the
yellow-colored stamens take over the attractive
function of the corolla. Anatomically, the corolla is
very similar to the calyx. The fleshy petals have a
distinct epidermis with relatively small cells. Be-
neath the epidermis there is a poorly differentiated,
many-layered parenchyma enclosing numerous,
parallel vascular bundles. Crystal sand occurs scat-
tered through the parenchyma.
Androecium. The androecium of Dialypetalan-
thus consists of 16 to 25 (usually less than 20; com-
monly 16 or 17) stamens that are united at their
base into a short androecial ring atop the ovary, free
from the corolla. Two whorls can be distinguished
within the androecium, the outer whorl always hav-
ing more stamens than the inner. A slight dimor-
phism can be seen between stamens of the two
whorls: the inner ones have slightly longer anthers
and a shorter filament than the outer ones (Fig. 9А).
The stamens are closely packed together in bud
stage and tightly enclose the developing style. They
Моште 84, Митбег 2
1997
Piesschaert et al
Dialypetalanthus fuscescens from Amazonia
Їз
>
b2 | ЈЕ: (а) tie bi
ba
D
Figure 7. Dissection of a terminal bud of the inflorescence of Marmein pa fuscescens.—A. Right bract removed,
gu
revealing the right floral bud surrounded by two bracteoles
И ;
are yellow-green and remain after the corolla has
fallen off. Later, the androecium falls off as a whole,
leaving only the style
Each locule consists of two clearly separated pol-
len sacs (Fig. 9B). The locules are large, elongate
(7 to 8 mm long and 2 mm wide), and pointed to
the lateral-adaxial side. The short filament (Fig.
9A) is basifixed and continues in a massive, essen-
tially abaxial connective (Fig. 9B). A peculiarity of
7
ne
n
E
аа
Ал
7 3;
ы
Figure 8. Schematic ренина of the inflores-
cence рч Dialypetalanthus fusces
(f1).—B. The same dissection showing a frontal view of the
ud of the — whor
mm. ag te Ducke 228
ud. ame view as A; ka bract (b2), Hur floral bud КП), and bract of the
b4) also removed, showing left floral bud of the first whorl (f2) -— (f4).—
. Schematic representation of a terminal inflorescence bud; a — axis
; bar =
the stamens of Dialypetalanthus is the (very) spo-
radic occurrence of stellate trichomes on the an-
thers (Fig. 9E). In most cases, however, the stamens
are completely glabrous. The locules possess two
horn-like appendages at their apex (Fig. 9C). These
locular appendages are curved toward each other
and have two basal branches, one to each pollen
sac (Fig. 11A). Kuhlmann (1925) did not mention
these structures at first, but later he called them,
rather unfortunately, “little valves" (Kuhlmann,
1942). Neither Rizzini and Occhioni (1949),
Hutchinson (1959), nor Cronquist (1981) paid any
attention to these appendages; n and Thorne
(1984) illustrated them but did not refer to them.
The locular appendages are clearly visible in bud
stage (Fig. 11A). Just before anthesis the anthers
tear open between the basal branches of the ap-
pendages (Fig. 11B), leaving small, more or less
round pores at the top of the locules (Fig. 110)
through which the pollen is released. Anther de-
hiscence therefore is porate (as stated by Kuhl-
mann, 1925; Rizzini & Occhioni, 1949; Cronquist,
1981; Dahlgren & Thorne, 1984). It has to be men-
tioned, however, that in some cases the pollen sacs
are split open over their entire length, probably ex-
plaining why Hutchinson (1959) stated that dehis-
cence is longitudinal. The locular appendages con-
sist of parenchymatic tissue filled with crystal sand,
surrounded by a more or less sclerified zone (Fig.
9C). A similar kind of calcium oxalate accumula-
tion in anthers was described for Solanaceae and
some other families by D’Arcy et al. (1996). They
reported a special type of hypodermal tissue in the
stamens, the so-called resorption tissue, filled with
calcium oxalate crystals and situated at the stomium,
running lengthwise around the anther. During mat-
uration of the stamens, the tissue surrounding the
Annals
ibd а Garden
flower after the petals have fallen
bar = 0.25
off; S = , A = androecium, С = calyx = 1 ст E Stellate hair on stamen:
mm.—F. Endothecial thickenings; bar 1 mm.—G. Young ise showing the dimerous origin of cal
and corolla; В = whorl of bracts, 51 = first sepalous who second sepalous whorl, Pl =
After Ducks 22816 (B, C, E, F, С), Ducke 21684 (A), and Ne ee 34472 (D).
first petalous whorl.
Volume 84, Number 2
1997
Piesschaert et al. 211
Dialypetalanthus fuscescens from Amazonia
Figure 10. Floral diagram of Dialypetalanthus fusces-
cens. Note the alternation of dimerous whorls. The stamens
are united at their base into a short androecial ring.
resorption tissue disintegrates and a mass of crys-
tals, the “oxalate packages” (O.P.’s), is set free. It
appears that the situation in Dialypetalanthus is
very similar, although the location of the crystals
(at the top of the anthers instead of the stomium)
is somewhat different (see also the section on Pol-
lination Biology). Various possible functions (e.g.,
in anther dehiscence, discouragement of herbi-
vores, as pollinator reward) for the O.P.’s were dis-
cussed by D’Arcy et al. (1996), but the exact role
of these structures remains unclear.
The inside of the pollen sacs is covered with
small (< 1 um), globular, smooth orbicules. Huys-
mans et al. (1997) have recently drawn attention to
these easily overlooked structures in Rubiaceae.
Between the pollen sacs of each locule, a clear epi-
dermal protuberance with remarkably enlarged epi-
dermal cells occurs (Fig. 9B). The same epidermal
configuration is found in between the locules, at the
adaxial side of the connective. In transverse sec-
tions, the cortex cells of the connective are isodi-
ametric with large intercellular cavities. Longitu-
dinally, the cells are elongated and clearly smaller
than the epidermal cells. They become smaller in
the proximity of the pollen sacs. The cortex zone
beneath the epidermal protuberances is strongly
lignified, often with off-shoots into the endothecium
(Fig. 9B). A one-layered endothecium occurs, ex-
cept at the dorsal side of the pollen sacs. The en-
dothecial thickenings are mostly spiral, seldom cir-
cular or U-shaped, or may even be absent (Fig. 9F).
In longitudinal sections the endothecial cells have
the polygonal appearance of cambial cells, with
their longitudinal axis perpendicular to that of the
cortical cells. The well-developed connective is tri-
angular in cross section (Fig. 9B). It contains one
central vascular bundle that seldom splits into two
smaller bundles at the apex of the stamens. In the
lament, at the adaxial side of the vascular bundle,
one can find a half circle of dark-staining cells,
possibly with high metabolic activity. A transverse
section of the androecial ring shows a ring of dark-
colored cells connecting the vascular bundles of all
stamens. The function of these cells is unclear.
It is not yet clear how the androecium is initi-
ated. Most probably it is not an example of primi-
tive polyandry (that is, each stamen arising as a
separate primordium; Ronse-Decraene & Smets,
1992), but a case of complex polyandry. In view of
the dimerous origin of calyx and corolla, we expect
the existence of two alternating whorls of two com-
plex primordia each, giving rise to numerous sta-
mens by “dédoublement.” This would also be in
Figure 11.
sis.—B. Stage ee
appendages are completely torn open. ће де Мега et 4 1295 (А), Ducke 22816 (B), and Anderson 11865
(C).
between A an pendage tears open
A odii a of the anthers of рим pones (SEM).—A. Locular appendage before anthe-
d C:
n its basal branches.—C. Locules after anthesis; the
212
Annals of the
Missouri Botanical Garden
SPT а A ES. XR. €
Ic UPS
Fatt 12, Gneci structure a Dialypetalanthus gear (A, B LM; C-E SEM.)—A. Transverse section а! the
top of a young gynoecium; P — ental zone; bar —
polarized light; note ca 'uplighting oxalate crystals in der placeita (arrow); 0 = 0.5 m
sk at the base of the style (S); A
papillose stigmatic surface.—
hairy di
and Thomas et al. 5205 (E
placenta in fruit stage. After Vilhena et al. 1003 (A), Nee 34518 (B), de Med: et al. 1295 (О), Ducke 21684 (D).
).
accordance with the alternation pattern of dimerous
whorls in the flower as a whole.
Pollen. The pollen of Dialypetalanthus is tri-
zonocolporate (Fig. 13A). The grains are oblate-
spheroidal in equatorial view (P/E-ratio 0.88-1)
and rounded to triangular in polar view. The polar
diameter (P) varies from 17 to 21 jum, the equato-
rial diameter (E) from 19 to 24 jum. The margins
of the ectocolpus are irregular, but form sharp an-
gles at the end (Fig. 13B). The mesoaperture is a
porus or colpus (2 to 3 jum long and mostly less
than 1 jum wide), often provided with a clear margo
or annulus (Fig. 13B). The endocolpus (Fig. 13E)
is oriented perpendicular to the exocolpus; it has
the same dimensions as the exocolpus but is more
regularly bordered. The ends of the endocolpi are
sometimes bifurcate (Fig. 13F). The apocolpium is
well-developed (apocolpium-index 0.35—0.45; Fig.
13A, C). The sexine is tectate perforate (perfora-
tions < 1 jum; muri larger than lumina but still <
1 pm; Fig. 13C, D). Muri are smooth to мапу. The
inside surface of the pollen grains is granular be-
tween the margins of the endocolpi (Fig. 13F, white
star). At the top of the endocolpi, the endexine 15
SOE GR a weak-looking zone with many
gular grooves that connects the endocolpi (Fig.
13F > black de. Sexine is as thick as or slightly
thinner than the nexine (Erdtman, 1971). The col-
umellae are short and thic
Gynoecium. The inferior ovary of Dialypetalan-
thus is composed of two fused carpels. It is more
or less flattened, grooved, and densely covered with
uniseriate hairs (white in the living state; Rizzin
& Occhioni, 1949). The septum is oriented perpen-
dicular to the widest part of the ovary, parallel to
the outer sepal whorl (wrongly illustrated by Rizzini
Volume 84, Number 2 Piesschaert et al. 213
1997 Dialypetalanthus fuscescens from Amazonia
Figure 13. Pollen morphology of Dialpetalanius fiscescens (SEM). A-D. Entire pollen. E, F. Broken pollen.—A.
Polar view with well-developed apocolpium.—B. Equatorial view of ecto- and mesoaperture.—C. Detail of the apocol-
pium.—D. Detail of the mesocolpium.—E. Endoc ни. —F. Endexine ornamentation; note the difference between the
warty (white star) and weaker looking zone at the top of the endocolpus (black star). After de Albuquerque et al. 1295.
214
Annals of the
Missouri Botanical Garden
& Occhioni, 1949). In very young buds, the two
halves of the septum‘ are fused in the lower part
of the ovary. In the upper part, however, they are
distinct (Fig. 12A). In mature flowers, the septum
is continuous throughout the ovary (Fig. 12B). As
a consequence of this septal development, the pla-
centa is U-shaped. This can be seen mainly in
young flowers, where septal development is still in-
complete, but even in mature fruits the apical split
of the placenta is often clearly visible (Fig. 12E).
Placentation is therefore axile in the lower part of
the ovary and parietal in the upper part. This is
often obscured by the fact that the ovules and seeds
are closely packed together. Anatomically, the pla-
centa is characterized by the occurrence of cells
that are filled with crystal sand (Fig. 12B). Each
locule contains numerous ascending, imbricate
ovules. Even before anthesis they have the sigmoid
shape that is characteristic of the seeds. Most au-
thors (Kuhlmann, 1925; Rizzini & Occhioni, 1949;
oT 1981) have described the ovules as bi-
tegmic. However, we believe that the exotestal
seed-coat structure (described later) indicates that
the ovules are in fact unitegmic, as was already
mentioned by Dahlgren and Thorne (1984). The
style is long (up to 12 mm) and glabrous. The pa-
pillose stigma is shortly bilobed (Fig. 12C). In
stained sections, the papillae are very dark, indi-
cating that the stigma probably secretes a sticky
substance in vivo. In bud stage the style is tightly
enclosed by the stamens. After anthesis it elongates
and rises above the androecium. The style persists
even after the other floral parts (with the exception
of the calyx lobes) have fallen off. In mature fruits,
it falls off as well. The epidermal cells of the style
are somewhat smaller than the cortical cells. The
latter are more or less arranged in radial rows. The
center of the cortex is formed by the transmission
tissue. On each side of the transmission tissue there
is a small vascular bundle. Cells containing crystal
sand occur scattered through the style. Very occa-
sionally, prismatic crystals occur as well. At the
base of the style, on top of the ovary, there is a low
annular disk, covered with long unicellular tri-
chomes (Fig. 12D). Based on its position (inside the
androecial whorls), the origin of the disk is most
probably gynoecial and not receptacular. The na-
ture of the secretion, if any, is not known. Just as
the style, the disk is characterized by the presence
of cells containing crystal sand. The ovary wall
shows few striking characters. In the subepidermal
* Each septal half is formed by fusion and subsequent
inward growth of the two carpels; finally, the septal halves
meet in the center of the ovary.
zone, numerous tannin idioblasts occur. A clear
zone of flattened cells forms the wall of the locules.
This zone transforms into a sclerified endocarp in
the fruit stage.
Fruit and seeds. Dialypetalanthus bears fruits
starting from the end of March or beginning of April
(immature fruits are green). The fruit (Figs. 14, 15)
is an elongated to more or less round (1.2-1.8 X
0.5–0.9 cm), bilocular capsule with persistent calyx
lobes. In old capsules these calyx lobes may break
off. The bracts and bracteoles are deciduous. The
top of the fruit often protrudes above the calyx
lobes. The hairy disk as well as the scars of the
other floral structures remain visible (Fig. 15A).
The fruit is sparsely covered with hairs (more
densely so at the base of the calyx lobes), becoming
almost completely glabrous when the fruit grows
older. After maturation, the fruit splits at first along
the septum (from the top downward), by which the
outer calyx lobes are split in two. Afterward the
locules are split (equally from the top downward)
and the seeds set free (Fig. 15A, B). The four parts
of the fruit wall remain attached together at the
base.
Anatomically, the fruit consists of a thin exocarp,
a “fleshy,” fibrous mesocarp (due to the vascular
bundles that are situated here), and a strongly
sclerified endocarp. The exocarp and mesocarp can
be easily removed, leaving two coffee-bean-like
halves (i.e., endocarp and seeds). When a locule is
opened, the numerous seeds and well-developed,
U-shaped placenta are visible (Fig. 15B).
The seeds are rather small (up to 7 mm long, but
usually smaller and about 1 mm wide at the widest
part). Their sigmoid shape (Fig. 16A) causes a ro-
tation when they fall out of the fruit and probably
allows further dispersal by wind. The seeds have a
wing that is well developed basally and apically,
and extremely narrow in the central endosperm-
bearing part (Fig. 16E). They are attached to the
placenta with the basal part of the wing. The apical
part of the wing differs from the basal in that it is
wider and not twisted. The endosperm and embryo,
visible as a distinct swelling in the center of the
seed, are present in only a minority of the seeds.
The majority of the seeds are devoid of embryos;
some of the examined fruiting see ge seemed
to lack embryos almost completely. The seeds are
exotestal (sensu Corner, 1976). The endotesta con-
sists of a few layers of crushed cells. Only in the
wings is a clear cellular structure visible (Fig.
160). The inner tangential walls of the exotestal
cells have a reticulate thickening pattern (lumina
mostly about 10 рт, wider than the muri and often
in two rows); their radial walls are also thickened
Volume 84, Number 2
1997
Piesschaert et al.
Dialypetalanthus fuscescens from Amazonia
CNPq - INPA
MUSEU PARAENSE EMILIO GOELDI
The New York Botanical Garden
Plantas da Amazó:
Medios & Jo Moria 3230 00-65-1977
Trabalhos de campo em convénio CNPq (Brasil! - NSF (USA)
Figure 14. Fruiting specimen of Dialypetalanthus fuscescens (Silva & Morio 3238, NY); bar = 5 ст.
(Fig. 16B, C). The seeds have a thin but very ob-
vious, oil-containing endosperm (Fig. 16E). The
embryo is large, straight, and has two short coty-
ledons; in cross section it is round.
POLLINATION BIOLOGY
Rizzini and Occhioni (1949) are the only authors
who postulated a pollination strategy for Dialype-
talanthus. They claimed that Dialypetalanthus
shows “obvious adaptations for anemogamy,” name-
ly many light, dry and small pollen grains. How-
ever, Juncosa and Tomlinson (1987: 1315) pointed
out that “the existence of light powdery pollen in
the taxa with animal pollination is an example of
the way in which pollen characteristics may be mis-
leading about the method of pollination.” In our
opinion, Rizzini and Occhioni’s (1949) hypothesis
is wrong. The pollen characteristics mentioned as
well as the exposed stamens and stigma of Dialy-
petalanthus are indeed often found in wind-polli-
nated taxa, but the following facts are hardly
brought in line with anemogamy: bisexual flowers,
unexposed inflorescences, a large attractive corolla
(although it falls off soon after anthesis), fragrant
flowers (described by Rizzini and Occhioni as “jas-
mine”), a disk that may secrete nectar (or oil), loc-
ular appendages that may have a reward function
for pollinators, numerous ovules (indicating the
need of successful pollination, contrary to wind-
pollinated species, which mostly have few ovules
216
Annals of the
Missouri Botanical Garden
É + f Dialy Ри | &
Y .
У...
+. ..
st
"s
. cre
. B
Sete 2 . pat
~. + „== e. AS ae ee io Pie ba
N SS с ы” а e ta ean eo, Ud,
ара E "Ж" . . mrss of
OE BEA ^з? а е бета се * ...
Figure 15.
and more silos oli Slits. С-В. win) view of a half fruit in the same
septum, se — seeds, pl — placenta. After Silva & Morio 3238 (A) and Silva 740 B.
calyx remnants and sc
per style) and pollen sacs closely associated with
the filament instead of the versatile anthers that are
typical of wind-pollinated species. Moreover, one
should bear in mind that Dialypetalanthus is a rain-
forest tree. The dense vegetation of mostly ever-
green species in rainforests, the large amount of
precipitation, and the low turbulence due to the
closed canopy, by which pollen cannot stay in the
air long enough to cause successful pollination,
make wind pollination unlikely. All of these mor-
phological and ecological data strongly suggest an-
imal pollination. А plausible type of pollination is
cantharophily. Beetles are especially attracted to
white, fragrant flowers with abundant and easily at-
tainable pollen (Willemstein, 1987). All of this is
offered by Dialypetalanthus. Grant (1950) stated
that an inferior ovary may be an adaptation to de-
structive pollinators (in this case beetles) as a pro-
tection for the vulnerable ovules. Cantharophily
would be a good explanation for the simultaneous
occurrence of numerous stamens and an inferior
ovary in Dialypetalanthus. The rough flower treat-
ment that is so typical for beetles would also ex-
plain the earlier-mentioned lignified zones in the
E . 1 pti £211
A. Fruit in late st f |
stage after artificial separat ion;
stamens and the overall firm, fleshy structure of the
flower.
On the other hand, pollination by bees seems to
be an acceptable alternative, because the poricidal
anthers may point to buzz-pollination. Moreover,
D’Arcy et al. (1996) stated that the occurrence of
oxalate packages is rather typical for bee-pollinated
species. The pollen becomes mixed with the cal-
cium oxalate crystals when the anthers are vibrated
by the insect, although it is still unclear what the
crystals are used for. Field observations are need
to establish the pollination strategy of Dialypeta-
lanthus.
DISTRIBUTION AND ECOLOGY
In the e it was assumed that Dialypetalanthus
occurs m in the eastern part of Brazil, around
Belém први 1994; Nicholas & Baijnath,
1994). This was probably due to the fact that the
first collections (Kuhlmann 1514; Ducke 17921,
23660, and 21684) were from this region. The
specimens presently available reveal, however, that
the center of distribution is in the northern border
Volume 84, Number 2
Piesschaert et al. 217
Dialypetalanthus fuscescens from Amazonia
Figure 16. Seed morphology and anatomy of Dialypetalanthus sapere Lr SEM; D EI
with typical sigmoid shape (base of seed is at the right); bar
— С. Detail of the thickening patt
coat showing the thickened exotestal cells (X), crushed endotestal cells
note the cotyledons (C) and well-developed endosperm
)
the exotestal cells; bar = 10pm.
Transverse section of seed;
et al. 5205 (А, D, E) and Silva 740 (B, С
region between Bolivia and Brazil (Guajará-Mirim,
Costa Marques, Pórto Velho, Rondónia, Rio Branco)
(Fig. 17). The distribution area of Dialypetalanthus
is situated in Takhtajan's (1986) Amazonian Prov-
ince, i.e., the southern half of his Amazonian re-
gion. In the phytogeographic division of Andersson
(1992), Dialypetalanthus occurs in the (i) western
Amazon of Colombia, Peru, and Brazil; (ii) central
Amazon of Brazil; and (iii) eastern Amazon of Bra-
zil.
Although the tree inhabits the terra firme forest
(information derived from herbarium labels), it seems
M.)—A. Entire Ed
ed thic gen inner —— walls o
ern ak i i .—D. gential section of the o^
(N), and endosperm (ES); bar = 0.1 mm.—E.
(ES); = 0.1 mm. After Thomas
to be linked to the rivers in one way or another. This
could of course be due to the fact that rivers are
avenues for collectors to reach the collecting locali-
ties. Dialypetalanthus mostly grows on sandy or grav-
elly soils, schist outcrops (where it occurs together
with such taxa as Orbignya phalerata Mart., Hevea,
and Bertholletia), or granite outcrops. Dialypetalan-
thus has been found only south of the Amazon River.
No collections are known from the region between the
Amazon and the Orinoco.
According to Prance (pers. comm.), Dialypeta-
lanthus is a rather common tree in Brazil, occurring
218
Annals of the
Missouri Botanical Garden
[ 1979 by the University of Utrecht ?
\
[3
Figure 17. Distribution map of Dialypetalanthus fus-
escens.
in primary forest as well as in partly disturbed,
more open regions. However, as the forest becomes
more and more devastated due to agricultural ex-
pansion and the construction of dams, it is not im-
possible that D. fuscescens will become a threatened
species in the near future.
pecimens examined (square brackets indicate coordi-
Rio Yata, [1175, 66°W], Anderson 11865 (NY). Pando:
Nicolás Suárez, between Cobija and Porvenir, "me
68°30'W), Fernández Casas & Susanna 8188 (NY); Puettó
Oro, 74 km SW of Cobija, 11%25'S, 69°05’ W, Pennington
et al. 143 (K). gr" Acre: Mun. Sena Madureira,
basin of Rio Purus, Rio Macauá, Colocacáo Apui,
09%48'S, 6911'W, Daly et al. 8068 (NY not seen); Mun.
Rio Branco, road Rio Branco-Pórto Velho, km
22, Quinoá, [10°S, 67730'W], de Albuquerque et al. 1295
(NY, US); Mun. Sena Madureira, Rio Масаџа, Seringal
pod Luzia, 09*39'S, 69*01"W, de Lima et al. 575 (NY
not seen). Amazonas: Rio Madeira, Humaitá, [07°30’S,
63 W]. Ducke 35428 (K, US). Goiás: Xambioa, = *
ка а W, Edison Mileski 397 (MG not seen); Rio Ara
egiáo de Araguatins, Ilha Santa Isabel, pa. 9, 49W],
Oliveira 1572 (NY). Maranhão: Mun. a Luzia, Fa-
zenda Agripec da Varig, Rio Pindaré, [04*S, 45*30'W],
Lobo et al. 328 (K, NY, US); Funai, 40 km from Arame,
Vilhena et al. 1003 (NY). Mato Grosso: Dardanelos, Sta.
Elena, Cordeiro 175 (NY); Fazenda Cachimbo, [09°30" s.
55°W], Cordeiro 1085 (NY); Rio Madeira, Salto Theo
Aripuanã, Fontanilha, [10*30'S, 58°W], Silva & Morio
3238 (NY). Pará: Rio Tapajós, ae ie 55°30'W],
Ducke 17921 (K, US) SYNTYPE; a de Santarém,
[02°30'%, 55°W], Ducke 21 је (K, NY): piss de Santa-
rém, [02*30'S, 55°W], Ducke 23660 (US); Igarapé Fer-
nando do Noronha, Rio Sao Manuel, Caldeirao, i
64°30’ W], Pires 3853 (NY, US); Rio Itacaiuna-Rio Toc
, 50715", Pires et al. 12531 (MG not Wes
io Xi ^38'S, 52?00' W, Silva 740
(NY); Cabeca, 09^45'8, 51720", Silva 740 (NY); Tucuruí,
Breu е [03%45'S, 49°45'W], Silva 5457 (MG not
seen); Rio Tocantins, Tucuruf, Breu Branco, [04°S, 49*W],
Silva & "Bahia 3514 (NY); Rio Itacaiuna, 2 km down-
crossing to AMZA camp3-alfa, 05°53’ 5,
ikini et al. 57941 (K : Mun. de Presidente
Medici, BR364, road eae и km 300,
m S, 61°62'W, Cid et al. 4801 (NY); Mun. Costa Mar-
ques, BR429, near Rio Cautarinho at 5 km o
12%10'S, 63?25'W, Cid Ferreira viti (NY); Mun. Pórto
Velho, Estrada da Serra do Balateiro a 3 km da Vila Cam-
po Novo, 10°35’S, 63°39’ W, Cid F. аы 8895 (NY); Mun.
Alvorada do Oeste, BR429, estrada para Costa Marques,
Serra da Опса a 35 km da cidade de Alvorada do Oeste,
11°30’S, 62°30’ W, Cid Ferreira 9018 (NY); Guajará-Mir-
im, border region with Bolivia, [11°S, 65°W], Cordeiro
1007 (MG not seen); Pôrto Velho, [10°S, 63°W], Maciel et
al. 1674 (MG not seen); 2 km of Guajará-Mirim, [11°30’S,
65°W], Nascimento 316 (MG not seen); 15 km N of Ari-
quemes, on hwy. BR364 and 1 km E on “Linea 75,”
09%47'S, def W, Nee 34357 (NY, T 21 km SE of
Ariquemes, on hwy. BR364 and 1 km E on “Linea 45,”
10%07'S, 62*56' W. Nee 34426 (NY); Mun. Costa Marques.
2 km W of Rio Cautarinho, along hwy. BR429, 12*04'5,
63°28'W, Nee 34472 (N Y); Mun. Costa Marques, road
Costa Marques-Forte Príncipe da Beira, 9.5 km E of Forte
Príncipe da Beira, 12?26'S, 64?20' W, Nee 34518 (NY);
road Abuña-Guajará-Mirim, 1 km N of Riberáo, [10°30’S,
Rosa 485 (MG not seen); Mun qu
Mibrasa, Setor Alto Candeias, ы 128, 10°35'S, 63735 W,
Teixeira et a ); e Velho, Represa Samuel,
©, у 5'W, Thomas et al. 5205 (NY, US). PERU.
Madre s: Tambopata, "1249'S, 69°18'
al. 45057 (NY) LOCALITY UNCERTAIN. Ferreira 438
K).
DISCUSSION
Because of its polyandrous androecium, most au-
thors (Cronquist, 1968; Dahlgren, 1975; Emberger,
1960; Melchior, 1964; Rizzini & Occhioni, 1949;
Stebbins, 1974) have placed Dialypetalanthus neat
petalanthus and the Myrtales is unlikely because
Myrtales have indi phloem, leaves typically
covered with glandular dots, and seldom have par-
acytic stomata, unlike Dialypetalanthus. Moreover,
Myrtaceae lack septate fibers and their wood is of-
Volume 84, Number 2
1997
Piesschaert et al.
Dialypetalanthus fuscescens from Amazonia
ten characterized by well-developed axial paren-
chyma. Melastomataceae have an acrodromous ve-
nation pattern, a synapomorphy for this family
according to Renner (1993); this venation pattern
does not occur in the Dialypetalanthaceae. The
only vegetative resemblances between Dialypeta-
lantus and Myrtales are the simple, opposite, entire
leaves and the vestured intervessel pits of the wood,
but these characters are common in the Gentianales
as well and are by themselves insufficient to sup-
port a close relationship. The stipules of Dialype-
talanthus give no more indication of relationship.
Myrtales are exstipulate (e.g., in Melastomataceae)
or possess rudimentary stipules (e.g., Myrtaceae,
Trapaceae; Dahlgren & Thorne, 1984). Gentianales
s.l. often have well-developed (Rubiaceae, Loga-
niaceae) or rudimentary stipules (Apocynaceae).
In vegetative (and also fruiting) stage, Dialype-
talanthus is hard to distinguish from certain Cin-
chonoideae (Rubiaceae). Robbrecht (1994: 23) not-
ed that “the habitual resemblance of vegetative or
fruiting material to Rubiaceae is impressive, and
almost all Amazonian field workers assign the tree
to the Rubiaceae.” Indeed, the simple, opposite,
entire leaves and the well-developed stipules are
characteristic of Rubiaceae as well as Dialypeta-
lanthus. The paracytic stomata, colleters of the
standard Rubiaceae-type at the inside of the stip-
ules, the lack of internal phloem, the vestured pits,
and septate libriform fibers are shared as well.
The following features are in our opinion critical
to assess the relationship of Dialypetalanthus, and
need to be discussed in detail:
Dendritic hairs. Dendritic hairs have a rather
restricted taxonomic distribution in the angio-
sperms (Theobald et al., 1988). Our observation of
the occasional occurrence of dendritic and stellate
hairs on the petioles and stamens of Dialypetalan-
thus is new. Rubiaceae generally have simple or
uniseriate trichomes (Robbrecht, 1988), although
Theohald et al. (1988) mentioned the occurrence of
non-glandular stellate hairs in this family (they did
not specify species or genera, however). Stellate
hairs were not mentioned in Verdcourt’s (1958) and
Robbrecht’s (1988) survey of Rubiaceae trichomes.
Since stellate and dendritic hairs each were ob-
served only once in Dialypetalanthus, it remains
unclear whether any systematic value should be at-
tached to it.
Choripetaly. Dialypetalanthus is characterized
by the occurrence of four free petals. According to
Hutchinson (1959), the free petals of Dialypetalan-
thus are no reason to exclude it from the Rubi-
aceae, because “the corolla is divided to the base
in at least three genera” (p. 386) [Molopanthera
(Rondeletieae; Delprete, 1996), Synaptanthera
(Hedyotideae; note that the correct name is Syn-
aptantha; Robbrecht, 1994), and Aulacodiscus (=
Pleiocarpidia; Urophylleae; Robbrecht, 1994)].
However, the three genera cited by Hutchinson are
characterized by a deeply to very deeply divided
corolla tube instead of free petals. Schumann
(1891) described the corolla of Molopanthera as
“sehr tief geteilt” (p. 54) and that of Synaptantha
as “fast bis zum Griinde geteilt” (p. 24). Breme-
kamp (1940a) paid no special attention to the co-
rolla in his monograph of Pleiocarpidia. He de-
scribed it as “breviter hyperocrateriformis, tubo
suburceolato.” If Pleiocarpidia possessed free pet-
als, this most probably would not have escaped
Bremekamp’s attention.
Hedstromia (Psychotrieae) has also been de-
scribed as “having five essentially free petals”
(Darwin, 1979) or “polypetalous” (Smith € Darwin,
1988), but again we assume that the utmost base
of the corolla lobes is fused, as the stamens are
said to be “attached at base of the corolla” (Smith
& Darwin, 1988). The same authors (p. 239) de-
scribed Mastixiodendron as having “semisuperior
ovaries and polypetalous corollas, characteristics
that occur in a number of rubiaceous tribes.” The
latter statement apparently only refers to semisu-
perior ovaries.
Altogether, Mastixiodendron seems to be the only
genus in Rubiaceae that may possibly have free
petals. Note that this genus has previously been
placed in the Cornaceae because of its “choripe-
taly” (Melchior, 1925, fide Darwin, 1977). Howev-
er, the only modern documentation of Mastixiod-
endron (Darwin, 1977) contains insufficient
information to judge its choripetaly. It states that
“at anthesis, the corolla lobes are always entirely
free,” but gives no information on early ontogenetic
stages or the state of immature corollas. We have
looked at flowers of Mastixiodendron (M. pachycla-
dos (Schum.) Melch., Schodde & Craven 4245, L)
in order to verify the true nature of the corolla. At
anthesis, the corolla lobes are indeed entirely free.
In bud stage, however, they are fused; when the
flower opens they split down to the base along the
fusion margins. The mature stamens are completely
free from the corolla (and alternating with the lobes)
and are inserted below the intrastaminal disk.
Thus, all reports of true choripetaly within the Ru-
biaceae seem to be based on inadequate observa-
tions.
Free petals are, however, not the major problem
in postulating a relationship between Rubiaceae
and Dialypetalanthus, since both taxa could have
и
220
Annals of the
Missouri Botanical Garden
developed from a common ancestor with free petals.
In this context, it is interesting to recall that the
Gentianales are included in the Cornidae in some
modern angiosperm classifications (Frohne & Jen-
sen, 1992; Smets, 1988 unpublished Ph.D. thesis).
Here, the basal family Cornaceae (Cornales) does
have free petals. Whereas Lee and Fairbrothers
(1978) demonstrated serological links between Ru-
biaceae (a derived family of the Cornidae) and Cor-
naceae, Dialypetalanthus seems to add a morpho-
logical dimension to this connection.
Polyandry. The major problem in postulating a
close relationship between Dialypetalanthus and
Rubiaceae, and at the same time the strongest ar-
gument of Rizzini and Occhioni (1949) for the in-
clusion of Dialypetalanthus in Myrtales, is the poly-
androus androecium. Rizzini and Occhioni stressed
the fact that Rubiaceae are isomerous: “Ora, sa-
bemos serem as Rubiaceae notáveis pela fixidéz de
seu número estaminal: este 6 sempro igual ao de
segmentos da corolla, a tal ponto de, mesmo em
formas anómalas por cultivo, persistir essa carac-
terística” (p. 249). As an extreme example, they
mentioned Gardenia thunbergii L.f., which some-
times has up to 11 petals and 11 corresponding
stamens. Note that pleiomery is a rather common
feature within the Rubiaceae—Gardenieae, and that
it is not restricted to cultivated species (Robbrecht
& Puff, 1986).
Polyandrous androecia are, according to Hutch-
inson (1959), also known in the genus Praravinia
(Rubiaceae-Urophylleae). Bremekamp (1940b) at-
tributed the notion of polyandry in Praravinia to
Korthals's protologue of the genus, but doubted its
correctness, because all the specimens he exam-
ined were isomerous.
Whereas the polyandry of Praravinia is probably
based upon wrong observations, this cannot be said
of Coprosma (Anthospermeae); some species have
a secondarily increased stamen number [e.g., С. ne-
phelephila Florence, corolla 5—6-merous, stamens
5-8(-12); Florence, 1986]. Theligonum, now gen-
erally accepted as being a rubiaceous genus
(Robbrecht, 1994; Rutishauser et al., 1997), is
polyandrous as well. In Coprosma and Theligonum,
polyandry seems to be an adaptation to anemophily.
Oxalate packages. Our study revealed the oc-
currence of oxalate packages (O.P/s) in the anthers
of Dialypetalanthus at a moment when these pe-
culiar structures were receiving renewed attention
(D'Arcy et al., 1996). According to D’Arcy et al.,
the only positive reports of O.P's are from the So-
lanaceae, Ericaceae, Theophrastaceae, Balsamina-
ceae, Bromeliaceae, Araceae, Lemnaceae, Areca-
ceae, Liliaceae, Onagraceae, and Tiliaceae. In the
Dialypetalanthaceae, the O.P.’s are different from
the common or Solanaceae type described by these
authors, since they are not located in the stomium
but in apical, sterile appendages of the anthers.
O.P’s are not recorded from the candidate rela-
tives Myrtaceae, Melastomataceae, or Rubiaceae.
We have observed, however, very obvious O.P.’s in
Cinchona pubescens M. Vahl. (unpublished); here
they occur as crystal sand and belong to the type
described in Solanaceae (D’Arcy et al., 1996), i.e.,
situated at the stomium and running the length of
the anther. Since the Rubiaceae are characterized
by the common occurrence of several types of cal-
cium oxalate crystals, one may expect that O.P’s
occur much more generally in this family but re-
main overlooked.
Rubiaceae and Dialypetalanthaceae both have
О.Р, but these belong to a different morphological
type. It seems that the taxonomic distribution of
these anther structures is insufficiently known at
present to draw further systematic conclusions.
Dimery. The calyx and corolla of Dialypetalan-
thus are initiated as dimerous whorls. Dimery has
a rather restricted occurrence within the dicotyle-
dons. As far as we know, it is only reported for the
Begoniaceae, Berberidaceae, Cecropiaceae, Bras-
sicaceae, Capparidaceae, Ebenaceae, Elatinaceae,
doni e Gunneraceae, Lauraceae, Nepentha-
e, Nymphaeaceae, Oleaceae, Onagraceae, Phy-
олды igor: Portulacaceae (Ronse
Decraene & Smets, 1991: 95), Ranunculaceae, and
Winteraceae (Watson & Dallwitz, 1991; Ronse-De-
craene, 1992 unpublished Ph.D. thesis). Almost all
Myrtales and Gentianales having four sepals and
petals are truly tetramerous instead of dimerous, as
can be derived from their floral diagram. Therefore,
the dimery of Dialypetalanthus more or less isolates
the genus. It should be noted, however, that The-
ligonum (Rubiaceae) may be dimerous as well (Ru-
tishauser et al., 1997).
Anther dehiscence and monadelphy. Rizzini and
Occhioni (1949) considered the united filaments
and porate dehiscence of the anthers as strong ar-
guments to exclude Dialypetalanthus from the Ru-
biaceae. However, porate anthers occur in the ru-
biaceous genera Rustia (apically porate; Delprete,
1995), Tresanthera (laterally porate; Delprete, p
comm.), and Argostemma (Robbrecht, 1988). Th
character is definitely without value to wed
or exclude a possible rubiaceous relationship.
Monadelphy is not especially rare in Rubiaceae.
It occurs, for example, in Capirona, Bikkia, Cou-
tarea, and Exostema (Robbrecht, 1988) and is even
Volume 84, Number 2
1997
Piesschaert et al. 221
Dialypetalanthus fuscescens from Amazonia
considered to be a synapomorphy of the Chiococ-
ceae (Bremer & Struwe, 1992). In all these cases,
however, the ring formed by the filament bases is
attached to the corolla tube. In this feature Rubi-
aceae differ from Dialypetalanthus. However, in a
few genera of the Chiococceae sensu Delprete the
stamens are not attached to the corolla tube, but on
top of the ovary (Delprete, pers. comm.).
Gynoecial structure. Whereas the corolla and
androecium of Dialypetalanthus and Rubiaceae
show fundamental differences, their gynoecial
structure is completely similar. Both Dialypetalan-
thus and many Rubiaceae are characterized by an
inferior, bilocular gynoecium with numerous ovules
in each locule and a disk on top of the ovary, sur-
rounding the base of the style. The supposedly
primitive (Leinfellner, 1951), U-shaped placentas
of Dialypetalanthus occur in numerous genera of
the Rubiaceae as well (De Block & Robbrecht,
1997).
е numerous, ascending, imbricate ovules of
Dialypetalanthus offer a strong argument for exclu-
sion of this genus from the Myrtales. As Dahlgren
and Thorne (1984) already concluded from obser-
vations of the seed-coat structure, these ovules are
unitegmic. This character is typical for Gentianales
and Cornales, but is lacking in Myrtales (except for
Syzygium; Dahlgren & Thorne, 1984).
With the exception of Melastomataceae and Me-
mecylaceae, both Myrtales and Gentianales are
provided with a more or less well-developed disk.
Smets (1988) stated, however, that these disks may
not be homologous, because the disk of Myrtales
has a receptacular origin, whereas that of Gentian-
ales is gynoecial. The disk of Dialypetalanthus aris-
es late in the floral ontogeny as a swelling, situated
at the inside of the androecial ring, surrounding the
base of the style. Its origin is therefore most prob-
ably gynoecial, as in Gentianales.
Pollen. Erdtman (1971) considered the pollen
of Dialypetalanthus to be more similar to pollen of
Rhizophoraceae than to that of Myrtaceae, Melas-
tomataceae, and Rubiaceae. We do not deny the
resemblances between the pollen of Dialypetalan-
thus and certain Rhizophoraceae, nor the differ-
ences from that of Myrtaceae (often synaperturate
pollen) and Melastomataceae (usually with pseu-
docolpi); however, the pollen fits perfectly within
the pollen morphological variation of the Rubi-
aceae. At the time Erdtman wrote his book, this
variation was insufficiently known, as is demon-
strated by his limited treatment (only + 5 % of the
genera) of this huge family. The tricolporate, tec-
tate-perforate pollen of Dialypetalanthus is now
known to be the most common type encountered in
Rubiaceae (Robbrecht, 1988).
Fruit and seeds. The capsules with numerous
winged seeds of Dialypetalanthus are hardly distin-
guishable from those of certain Rubiaceae—Cin-
chonoideae. For example, capsules that split into
four parts and seeds with basally attached seed
wings (as in Dialypetalanthus) are very common in
several genera of the Rondeletieae s.l. (Delprete,
1996). Winged seeds occur in many Rubiaceae
tribes. Seed wings are extremely diverse; the con-
dition found in Dialypetalanthus (wing mainly re-
stricted to base and apex) occurs, e.g., in Mitragyna
(Coptosapelteae; Stoffelen et al., 1996: fig. 21). Al-
though Rizzini and Occhioni (1949) thought that
winged seeds are not very useful for classification
purposes, they add another resemblance between
the fruit and the seeds of Dialypetalanthus and the
Cinchonoideae.
The seed-coat structure of Dialypetalanthus can-
not be classified as one of the three seed-coat types
that Corner (1976) distinguished for the Myrtales,
and therefore presents another argument for ex-
cluding Dialypetalanthus from this order. On the
contrary, the pitted thickenings of the inner tan-
gential walls of the exotestal cells and the crushed
endotestal cells of the seed coat of Dialypetalanthus
are considered to be typical of the Cinchonoideae
(Bremekamp, 1966; Robbrecht, 1988).
Myrtales (with the exception of some rare cases
in Myrtaceae, where scanty endosperm may be
formed) are characterized by the lack of endosperm
(Dahlgren, 1991). This again argues against inclu-
sion of Dialypetalanthus in the Myrtales. With its
well-developed endosperm, it fits far better in the
Gentianales (and Rubiaceae).
CONCLUSION
Summarizing the evidence obtained from floral
and fruit structure, a close relationship between Di-
alypetalanthus and Myrtales is unlikely because of
the following character states of Dialypetalanthus:
unitegmic ovules, gynoecial disk covered by uni-
cellular trichomes, exotestal seed coat, and well-
Rubiaceae (more specifically the subfamily Cin-
chonoideae), whereas pollen morphology does not
exclude a relationship with that family. The dim-
erous origin of calyx and corolla, the free corolla
lobes, and the supposed cantharophily are unique,
possibly primitive, features of Dialypetalanthus,
giving it a somewhat isolated position. The inter-
pretation of the polyandrous androecium remains
Annals of the
Missouri Botanical Garden
problematic as long as its origin is not elucidated
by ontogenetic studies. We consider Dialypetalan-
thus to be a textbook case of heterobathmy, com-
bining primitive features (the presence of stipules
and placentas may be added to the char-
acters enumerated above) with derived gynoecial
features. Our observations consequently corrobo-
rate a Dialypetalanthus—Rubiaceae relationship,
but the genus can clearly not be included in that
family. The translation of this conclusion into a pro-
posal for a classification above the family level (1.е.,
should the Dialypetalanthaceae be placed in the
order Gentianales s.l. or in the Rubiales) is not
easy. Since the delimitation of the Gentianales, an
more specifically the position of the Rubiaceae, is
still problematic and subject to frequent change
(Nicholas & Baijnath, 1994; De Laet € Smets,
1996), it seems best to maintain at present a widely
conceived order Gentianales, including the Dialy-
petalanthaceae and Rubiaceae and arising from a
cornalean stock. In order to gain a better under-
standing of the position of the Dialypetalanthaceae
within this gentianalean complex (e.g., should Di-
alypetalanthaceae and Rubiaceae be put together
in the Rubiales s.l., closely related to the Gentian-
ales s. str.?), a cladistic analysis including Gen-
tianales, Cornales, and other members of the Cor-
nidae alliance most certainly would be very useful.
In the latter context, however, it is regrettable that
phytochemical (Darnley-Gibbs, 1974; Hegnauer,
1990), floral ontogenetic, and macromolecular data
of Dialypetalanthus are still very poorly or not at
all known. Consequently, future research should
mainly concentrate on these and other missing
characters of Dialypetalanthus to further elucidate
its systematic position.
Literature Cited
Andersson, L. A cecilia of neotrop-
ical Md Seria Bot. Be
Baas, P. 1986. Terminology of malos tracheary el-
ements—In defence of libriform fibres with minutely
. A. Bull., N.S. 7: 82-86.
‚ Та ubert. 1967. Pflanzenan-
ені Praktikum: zur Einfiihrung in die Anatomie
der Vegetationsorgane der hóheren Pflanzen (Sperma-
tophyta). VEB Gustav Fischer Verlag, Jena.
Bremekamp, C. E. B. 1940a. A monograph of the genus
ptio n za dis Recueil Trav. Bot.
Néerl. 3
nus Praravinia Korth. (Rubi-
асад) іп Borneo and Celia Recueil Trav. Bot. Néerl.
37: 237-278.
1966. Remarks on the position, the delimitation
and the subdivision of the Rubiaceae. Acta Bot. Neerl.
-33.
Bremec, B. & L. Struwe. 1992. Phylogeny of the Rubi-
aceae and the Loganiaceae: Congruence or conflict be-
tween morphological and molecular data? Amer. J. Bot.
79: 1171-1184.
Corner, E. J. H. 1976. The Seeds of No Vol.
1. Cambridge Univ. Press, Cambri
e Evolution San earne of
An Integrated System of Classification of
Flowering Plants. Columbia Univ. Press, New York.
——. 1988. The Evolution and Classification of Flow-
ering Plants, 2nd ed. The New York Botanical Garden,
New York.
Dahlgren 1991. Steps toward a natural system of the
dicotyledons: Embryological characters. Aliso 13: 107-
165.
Dahlgren, R. 1975. A system of classification of the an-
giosperms to be used to demonstrate the distribution of
characters. Bot. Not. 128: 119-
80. A revised system of classification of the
angiosperms. Bot. J. Linn. Soc. 80: 91-124,
—— ———. 1983. General aspects of angiosperm evolution
and macrosystematics. Nordic J. Bot. 3: 119-149.
Thorne. 1984. The order Myrtales: Cir-
cumscription, variation and кк меме а Ann. Missou-
ri Bot. Gard. 71: 63
D'Arcy же ЊЕ Kenting & S. L. Buchmann. 1996.
The caleiim "ali Ace or so-called resorption tis-
he
R. C. ыд (editors), The Anther.
Function and Phylogeny. Cambridge Univ. Press, Cam-
ridge.
Darnley-Gibbs, R. 1974. Chemotaxonomy of Flowering
Plants. Vol. m McGill. Queens Univ. Press, Montreal.
Darwin, S. P. 1977. The en. Mastixiodendron (Rubi-
aceae). J. лим Arbor. 58: 3
A synopsis of de фи genera of
Pacific еам Allertonia 2: 1—44.
De Block, Р. & Е. Robbrecht. 1997. On the ovary struc-
ture of Dictyandra and Leptactina compared with other
Pavetteae (Rubiaceae-Ixoroideae), or evolution from
multiovulate to uniovulate placentas. Bot. Jahrb. Syst.
De Laet, J. & E. Smets. 1996. commentary on the
circumscription and evolution of the order Gentianales,
with special emphasis on the position of the Rubiaceae.
Opera Bot. ia 11-18.
Delprete, P. G. Three new species of Rustia (Ru-
biaceae, ал from Panama and Ecuador.
Novon 5: 133-139.
Evaluation of the tribes Chiococceae,
morphological characters. Opera Bot. Belg. е 165-192.
mberger, L. 1960. Les végétaux vasculaire . 154
1539 in M. Chadefaud & L. Emberger (editori Traité
de smi d (systématique), tome II. Masson et Cie
éditeurs,
Erbar, C. 1995. On the floral pss of Sphenoclea
zeylanica (Sphenocleaceae, Campanulales)—SEM in-
vestigations on herbarium dere Bot. Jahrb. Syst
117: 46
m G. = Pollen Morphology and Plant Taxon-
my—Angi (An introduction to palyno ology),
Vol L Hafner P Publishing, New York.
Florence, J. 1986. Sertum Polynesicum II. Rubiaceae
nouvelles des îles Marquises (Polynésie Franca
ull. Mus. Nat. Hist. Nat., 4e sér., B, Adansonia 8: 3-
11.
Frohne, D. & U. Jensen. 1992. Systematik des Pflanzen-
Volume 84, Number 2
Piesschaert et al.
Dialypetalanthus fuscescens from Amazonia
reichs: unter besonderer Beriicksichtigung chemischer
Merkmale und pflanzlicher Drogen, 4., Aufl. Gustav Fi-
scher Verlag, Stuttgart.
Grant, У. . The protection of ovules in flowering
plants. Evolution 4: 179-201
Hegnauer, R. 1990. Chemotixonentie der Pflanzen 9.
Birkhauser Verlag, Basel.
Hickey, C. J. 1988. A revised classification of the archi-
tecture of dicotyledonous leaves. Pp. 25-39 in C. R.
Metcalfe & L. Chalk (editors), Anatomy of the Dick
ledons, 2nd ed., Vol. 1, Systematic Anatomy of the Leaf
and Stem. Сава Press, Oxford.
Holen, E к> . Н. Holmgren & L. С. Barnett (edi-
ors). 1990. ен Herbariorum. Part 1: The Herbaria
o ^s World, ed. 8. New York Botanical Garden, New
Hie J. 1959. The Families of Flowering devis
Arranged According to a New System Based on
Probable Phylogeny, 2nd ed., Vol. 1, euet riii
Clarendon Press, Oxford.
Huysmans, S., E. Robbrecht м E, arts 1994. Are the
genera Hallea and Mitragyna (Rubiaceae—Coptosapel-
> pollen omita dino? Быны 39: 321-
340
©; ee 5. mes & E. Smets. 1997.
Омей; A prelim and their пуча
value in the Слаба го Выпей Canad. J. Bot
n press.
Juncosa, A. M. & P. B. Tomlinson. 1987. Floral devel-
opment in seaman ty Rhizophoraceae. Amer. J. Bot. 74:
1263-1279
Kuhlmann, J. с. ib o conhecimento
de algumas чере, поуаз, зрела tambem um tra-
balho d de critica e novas combinagés. Arch. Jard. Bot.
Rio de d 4: 347- 365.
. Rubiaceae. Dialypetalanthus, Kuhlmann.
сити 6: oe
Lee, Y. S. & D. airbrothers. 1978. Serological ap-
proaches to An ا of the Rubiaceae and re-
lated families. Taxon 27: 159-185.
Leinfellner, W. 1951. Die U-fórmiche Plazenta als der Pla-
zentationtypus der Angiospermen. Oesterr. Bot. Z. 98:
38-358
Maas, P. J. M. & L. Y. T. Westra. 1993. Neotropical Plant
Families. A Concise Guide to Families of Vascular
Plants in the Neotropics. Koeltz Scientific Books, Ko-
enigstein.
Melchior, H. 1964. Englers Syllabus der Pflanzenfami-
lien, ed. 12, Gebr. Borntraeger, Berlin.
Nicholas, A. €: H. Baijnath. 1994. A consensus classi-
T for the order Gentianales with additional details
n the suborder Apocynineae. Bot. Re : 440-482.
Occhioni, P. & C. T. Rizzini. 1952. Dialypetalanthaceae.
Rodriguésia 15, 27: 181-183.
Peterson, К, L, . E. Bani: & J. D. т. dd
for histological i Stain rer І. А 1- à
Renner, S. S. 1993. Phylogeny and classification of the
Melastomataceae and Memecylaceae. Nordic J. Bot. 13:
Rizzini, C. T a E t idm 1949. *Dialypetalanthaceae.”
Lilloa 17: 2
Robbrecht, E. “ioes, “Tropical woody Rubiaceae. Opera
Bot. Belg. 1: 1-271.
1994. “Advances in Rubiaceae macrosystemat-
ics. Opera Bot. E
&
Je rvey of the Gardenieae and
туна tribes (Rubiaceos), she Jahrb. Syst. 108: 63-
ide Decraene, L.-P. & E. F. Smets. 1991. The floral
ontogeny of some members of the Phytolaccaceae (sub-
igni bait cues with a discussion on the evo n
of t roecium in the Rivinoideae. Biol. Jaarb. 5
77-99
& . 1992. Complex polyandry in the Mag-
noliatae: Definition, distribution and systematic value.
Nordic J. Bot.
Rutishauser, "ay у Ronse Decraene, E. Smets & I.
-Неџег. 1997. T r cynocrambe—The
a peculiar rubiaceous
Schumann, K. 189 Rubiaceae. Pp. 1-156 in A. Engler
& К. Prantl 72 Die Natiirlichen gore
4 (4). Verlag von pesos познао Leipz
Smets, E. 1988. e des * ria de stentia"
chez les rp r te r e arme Candollea
43: 709-716.
Smith, A. С. & S. P. Darwin. 1988. Rubiaceae. Pp. 143—
376 in A. C. Smith т Flora og xy Nova. A
New Flora of Fiji (spermatophytes only 2 ue
Tropical Botanical Garden, Lawai, Kauai;
1940. Taxonomic botany with BRER ref-
s. Pp. 435—454 in J. Huxley
(editor), The New a Д Oxford Univ. Press, Ox-
ford.
о
Stebbins, С. L. 1974. Flowering Plants. Evolution Above
the bay cies айн I elknap Press of Harvard Univ.
hus
Pre ТИ, вас
Stoffelen, D. Robbrecht & E. Smets. 1996. A revision
of Corynanthe and Pausinystalia (African PM.
Coptosapelteae). Bot. J. Linn. 0: 2
Takhtajan, A. 1986. оне Regions of the World, Univ.
California Press, Californ
beard Ж Lo, 3. La Krahulik & R. C. Rollins. 1988.
40—53 in С. В. Metcalfe & L. age (editors), Anat-
td of the Dicotyledons, 2nd ed., Vol. 1, Systematic
pane of the Leaf and Stem. Clarendon Press, Ox-
Ve er rt, В. 1958. Remarks on the classification of the
Rubiaceae tg Jard. Bot. Natl. Belgique 28: 210—290.
po» son, L M. J. —— 1991. The families of an-
aci n tom escriptions, with interactive
identification and Мина retrieval. Austral. Syst.
ot. 4: 681-695
Weberling, F. 1992. Morphology of а "qm Inflores-
ences. Cambridge Univ. Press,
Willemstein, 5. С. фо леш ‘Basis for Pol-
lination Ecology. Leiden Botanical Series 10. Leiden
Univ ом , Leiden
CONSPECTUS OF THE GENUS
PALICOUREA (RUBIACEAE:
PSYCHOTRIEAE) WITH THE
DESCRIPTION OF SOME NEW
SPECIES FROM ECUADOR
AND COLOMBIA!
Charlotte M. Taylor?
ABSTRACT
Palicourea Aublet comprises about 200 species of shrubs and small trees found throughout the moist and wet
Neotropics. This genus is eripe onim маг кин of the corolla that apparently represent adaptations for pollination
by hummingbirds and seems to be a me
of a hummingbird-pollinated group closely rela
ted to Psychotria subg.
Heteropsychotria, which is она check pollinated. зар recor geese classification + is proposed here for 188 species,
fcatinn
based largely on Characters of the stipules an
centered in Amazonia, and — 52 iun in ur sections, and the other generally middle to high ‘elevation,
pollination, together with habitat diversity, distyly, and variation among loc
isolation. ollowing new combinations
and comprising 136 species in five sections. Several
in some cases different su
istinctive features shared by
ttern
bgenera are noted. s suggest that
ocal populations leading to their reproductive
are made: Palicourea sect. Corymbiferae (МИП. Arg.) С. M. Taylor, Palicourea
subtomentosa (Ruiz. & Pav.) C. M. Taylor and P. subser. Subcymosae (Müll. Arg.) C. M. Taylor; the following new taxa
d
are described: P. subg. Montanae.
, P. sect. Cephaeloides, P. sect. Grandiflo
rae, P. sect. Montanae, P. sect. Obovoideae
ect. Реиповњиаћу лде, md P. sect. Psychotrioides; and the ei new MA are described, illustrated, and
Р g
classified: P. anderssoniana, P. anianguana, P. asplu
P. corniculata, P. cornigera, P. детае.
ndii, P. azurea
, P. gentryi, P. harlingii, P. premeg P те P. luteonivea, Р. prodiga, P.
P. cana candida, P. chignul, P. condorica,
subalatoides, P. subtomentosa subsp. lojana, P. ulloana, and P. vulcanalis.
Palicourea Aublet comprises about 200 species
of shrubs and small trees distributed from central
Mexico and the Antilles to southern Peru, Bolivia,
Paraguay, and northern Argentina (Fig. 1). This ge-
nus has not been surveyed as a whole since Schu-
mann’s (1891) review of the family and has never
been studied in any detail in its entirety, although
several recent regional treatments are available
(Steyermark, 1972, 1974; Bacigalupo, 1952; Taylor,
1989, 1993), and monographic work is currently in
progress. Palicourea is notable for the large number
of apparently closely related species it specs
overall, and also for the frequent sympatric oc
rence of several (to numerous) different species at
a given site (e.g., Kappelle & Zamora, 1995; So-
brevila et al., 1983; pers. obs.).
Species of Palicourea are typically members of
the understory and subcanopy of moist to wet forest,
and are found from low elevations to the upper lim-
its of wet montane forest. A few species grow in
savanna habitats and show pyrrhophytic adapta-
tions. Both widespread weedy species and locally
endemic, apparently more specialized species are
found in this genus. Palicourea flowers have согој-
las with relatively well-developed tubes and typi-
cally are odorless, brightly colored, and pollinated
by hummingbirds; the carnose, usually blue to pur-
ple fruits are dispersed by birds. Nearly all Pali-
originally suggested Palicourea as a subject for study.
? Mi
. Andersson,
. Gunter, and in particular R. E. Gereau, and also R. С Wilbur, who
issouri Botanical Garden, Р.О. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
ANN. Missour! Bor. GARD. 84: 224—262. 1997.
Volume 84, Number 2
1997
Taylor
Conspectus of Palicourea
courea species are distylous; this appears to be the
ancestral condition for the genus (Taylor, 1993).
Palicourea is closely related to the neotropical
subgenus Heteropsychotria Steyerm. of Psychotria
L. (Taylor, 1996). These groups are separated only
by characters of the corolla, as discussed below.
Schumann (1891) presented an infrageneric
classification for Palicourea as a whole, based pri-
marily on the regional treatments by Mueller (1881,
who included it in Psychotria) and Grisebach
(1861); no subsequent author has made reference
to this classification, including Steyermark (1972,
1974), even though he treated 56 and 47 species,
respectively, and presented a detailed infrageneric
classification for the closely related genus Psycho-
tria. No infrageneric classification has been pro-
posed more recently for this genus.
In this article I outline the circumscription of
Palicourea, present an infrageneric classification,
discuss some of the character patterns now evident
within the genus, and describe several new taxa.
CIRCUMSCRIPTION OF PALICOUREA
Schumann (1891) was one of the first authors to
distinguish Palicourea explicitly from Psychotria.
In his key to genera, he separated Palicourea by
its bent corolla that is gibbous at the base. He nev-
ertheless included in Palicourea species such as
Psychotria domingensis Jacq. (Schumann, 1891:
115), with white corollas that are curved in the tube
but never gibbous at the base (Taylor, 1987), and
excluded from it species such as Palicourea auran-
паса Miq. (Schumann, 1891: 114), a name here
considered synonymous with Palicourea fastigiata
Kunth and based on a set of specimens with the
corollas bright orange, gibbous, and straight to
slightly bent at the base, but generally straight in
the tube. Thus, his actual See of the
genus was not entirely cons ^
Standley (1936, 1938) erica Palicourea
by its “corolla tube elongate, more or less curved,
gibbous at the base; branches of the inflorescence
usually pale or red or yellow,” and this concept was
followed closely by Dwyer (1980). Standley placed
in Palicourea several species with these distin-
guishing features but with green inflorescence
branches and white or nearly white corollas, e.g.,
Palicourea andrei Standl., P. andaluciana Standl.,
and P. garciae Standl.; simultaneously he placed in
Psychotria several similar species that have white
or yellow corollas that are swollen at the base and
green to brightly colored inflorescence branches,
e.g., Psychotria bella Standl. [= Palicourea bella
(Standley) Dwyer], Psychotria copeyana Standl. &
L. O. Williams [= Palicourea garciae], and Psy-
chotria brenesii Standl. [= Palicourea standleyana
С. M. Taylor]. Standley's circumscription of Pali-
courea seems to have been based on the aggregate
classification of individual species rather than char-
acterization of the genus as a whole.
Steyermark's (1972, 1974) generic concept of
Palicourea was more consistent and based more
strongly on corolla features. He distinguished Pal-
icourea from Psychotria in his key as follows (Stey-
ermark, 1974, my translation from the Spanish):
"[b]ase of the corolla tube with a slight swelling ог
a curvature on one side; interior of the corolla tube
with a ring of trichomes usually near the base; co-
rolla usually blue, purple, yellow, orange, red, or
combinations of these colors; rachis and axes of the
inflorescence with similar colors." In practice, spe-
cies included by Steyermark in Palicourea usually
also have a relatively well-developed corolla tube.
As a basis for distinguishing Palicourea species,
I (Taylor, 1989, 1993, 1996) have used corollas
with a swollen base closed internally by a ring of
pubescence (as emphasized by Steyermark), and
the general tendency to have brightly colored floral
displays and relatively well-developed corolla
tubes. These features appear to represent adapta-
tion for pollination by hummingbirds: nectar ac-
cumulates in the enlarged basal part of the corolla
and is protected by the ring of pubescence from
weaker or smaller flower visitors, and the pollina-
tors are often attracted by the colorful inflorescence
displays.
Palicourea appears in general to represent a lin-
eage closely related to or perhaps even derived
from Psychotria subg. Heteropsychotria through ad-
aptation for hummingbird pollination (Taylor, 1989,
1996). This genus has so far been assumed to com-
prise a monophyletic group (Taylor, 1989, 1996)
that can be recognized by the characteristic corolla
morphology. Additionally, the pollen of the very few
species of Palicourea that have been studied lacks
or nearly lacks exine (Erdtman, 1952; Taylor,
1996). No evidence has been found to date to sug-
gest that this genus is polyphyletic, although its
unusual corolla morphology could conceivably have
been derived more than once. A few species that
show this characteristic corolla morphology and are
similar to other Palicourea species, but do not ap-
pear to be solely hummingbird pollinated, may rep-
resent secondary pollinator shifts. For example,
Palicourea andrei is very similar to P. boyacana
Standl., which has tubular orange flowers, but P.
andrei has sweetly fragrant, tubular white corollas
that could also be adapted for insect pollination.
As discussed below, hummingbird pollination ap-
226
Annals of the
Missouri Botanical Garden
Figure 1
Approximate number of species of Palicourea by country, for the entire range of the genus. Numbers
е
followed by “+” represent conservative estimates for country floras not yet surveyed in detail
pears to be an important factor in the diversification
of this genus. Adaptations for hummingbird polli-
nation are generally considered to include brightly
colored flowers, but these birds frequently do visit,
and even actively defend, Palicourea plants with
green inflorescences and relatively short white co-
rollas (e.g., P. calophlebia Standl., Colombia, pers.
obs.).
PHYTOGEOGRAPHY AND ECOLOGY OF PALICOUREA
Species of Palicourea are concentrated in tropi-
cal South America; they are also well represented
in Central America, particularly in montane Costa
Rica and Panama (Fig. 1). Eight species are found
in the Antilles, most of them in the Greater Antil-
les, and more than 32 species are now known from
Central America (Taylor, 1989, 1990). In South
America, Palicourea is widespread in low-elevation
wet tropical forests and also shows notable centers
of species diversity in wet tropical montane areas.
Palicourea is more species-rich in montane than
lowland habitats. When montane species are de-
fined as those predominantly distributed at or above
1000 m elevation (Gentry, 1988), 22 of the 26 spe-
cies known from Costa Rica can be considered
montane (Burger & Taylor, 1993), as can ca. 50 0
the ca. 80 species known from Ecuador (Taylor, in
prep.. The two subgenera recognized below ap-
proximate this habitat distribution: of the 188 spe-
cies of Palicourea classified below, 52 are placed
in subgenus Palicourea, which is primarily distrib-
uted at low elevations, while 136 are placed in sub-
genus Montanae, which is primarily montane. P al-
icourea is usually well represented locally ™
montane forests in both number of individuals and
number of species (e.g., Kappelle & Zamora, 1995;
Silverstone-Sopkin & Ramos-Pérez, 1995). The dis-
a
Volume 84, Number 2
1997
Taylor
Conspectus of Palicourea
tributions of most species of subgenus Montanae
correspond to Takhtajan's (1986) regions, and in
particular support his circumscription of the Cen-
tral American region, which extends south along
coastal Colombia into Ecuador, rather than Gentry's
(1982) circumscription of this region, which ends
in northwestern Colombia. Several groups of ap-
parently closely related species of Palicourea are
restricted to individual phytogeographic provinces
(sensu Takhtajan, 19.
The relatively Dé species diversity of Pali-
courea in montane habitats is very likely due in
part to the larger number of different habitats, or
microhabitats, in montane regions. In general, low-
elevation species of Palicourea usually have wider
geographic distributions than do montane species,
and fewer of the low-elevation species are local en-
demics. For example, none of the four lowland spe-
cies known from Costa Rica is endemic and all
extend at least as far south as Peru; in contrast, 6
of the 22 montane species from that country are
endemic to it and the rest are otherwise found only
in Panama or in a few cases in western Colombia.
Montane areas have been considered by a number
of authors to comprise a greater diversity of habitats
than lowland areas, due to both physical and his-
torical factors (Gentry, 1982, 1995; Young, 1995).
Additionally, apparently related species of Palicou-
rea frequently show distinct elevational zonation
(see discussion of P. ovalis and related species, be-
low), further partitioning montane habitats, as noted
by Webster (1995).
The relative youth of these areas may also be a
factor contributing to high species diversity. Al-
though mountains have existed in the region of the
Andes since at least the Eocene (Taylor, 1995), the
great height and perhaps also the pluvial climate
of the northern Andes are apparently more recent,
with the principal uplift occurring during the last
5 million years (van der Hammen, 1974). Signifi-
cant alteration in local conditions and therefore
plant communities has been noted during at least
the last 3.5 million years (Hooghiemstra & Cleef,
1995), which suggests that many populations of
Palicourea have long occupied a changing environ-
ment. Palicourea has not been reported from fossil
assemblages and probably will not be found be-
cause of its limited pollen exine, so its historical
occurrence is unknown. Modern species of Pali-
courea show variation among populations in flower
size (Sobrevila et al., 1983), inflorescence color,
and local flowering time (pers. obs., Puerto Rico,
Costa Rica), which may favor selection for repro-
ductive isolation through differing phenology and
perhaps pollinators. Fragmentation and isolation of
montane populations of Palicourea species would
reinforce this reproductive isolation, while distyly
would continue to promote outcrossing among in-
dividuals within small populations. Most species of
Palicourea, including many of those known only
from a few widely separated sites, are represented
on herbarium specimens by both long-styled and
short-styled plants. Only a very few cases of loss of
this feature have been demonstrated, mostly in
plants of the Antilles (Taylor, 1989), and in popu-
lations peripheral to the principal range of the gen-
erally distylous species (Sobrevila et al., 1983).
Widespread maintenance of distyly in the genus
suggests that this feature is adaptive and main-
tained through selection, and that pollen transfer
among plants is effective in most species at least
through the principal part of their range.
Many species of Palicourea are notoriously sim-
ilar vegetatively and in fruit, sometimes to the de-
gree that they can be separated only by their ma-
ture flowers. Sympatric assemblages of Palicourea
are common, with the sympatric species usually
separated reproductively by phenology and perhaps
subtle to marked differences in morphology (e.g.,
Sobrevila et al., 1983). For example, Palicourea
crocea (Sw.) Roem. & Schult. and P. croceoides
Ham. frequently grow sympatrically in Puerto Rico,
with plants of both species often found side by side
in mixed populations. These species have been
treated by a number of authors as conspecific (e.g.,
Liogier & Martorell, 1982). In one site in Puerto
Rico where these species are sympatric, plants of
P. crocea are usually less than 1 m tall, bear dark
red corollas that are shorter than those of P. cro-
ceoides on yellow inflorescence branches, flower for
a period of 1-2 weeks once or twice a year, and
are diploid; plants of P. croceoides are taller, bear
longer yellow corollas on red inflorescence branch-
es, produce flowers during a period of a month or
more approximately every 3—4 months, and are
polyploid (Taylor, pers. obs.; Kiehn, 1986). Individ-
ual plants are difficult to identify to species when
not in flower, but the two populations are repro-
ductively isolated and clearly separable by repro-
ductive features, and are considered separate spe-
cies here. Such reproductive isolation among
sympatric Palicourea species is probably common;
little work has been done in this area.
The relatively greater diversification of Palicou-
rea in montane habitats is likely due also in part
to its exploitation of hummingbirds as pollinators.
Ап increase in importance of species pollinated by
hummingbirds at higher elevations has been noted
in surveys covering various families (Webster,
1995), which suggests that this mode of pollination
228
Annals of the
Missouri Botanical Garden
is more effective than insect pollination at higher
elevations. This is probably partly why with in-
creasing elevation, Palicourea becomes relatively
more important than, or even replaces, Psychotria
subg. Heteropsychotria, the group most closely re-
lated to it, which is generally pollinated by small
insects.
CHARACTER PATTERNS IN PALICOUREA
Diagnostic morphological characters used here to
distinguish species of Palicourea include the fol-
lowing features. Stipule form may be sheathing,
i.e., united around the stem in a continuous, usu-
ally truncate sheath, or laminar, 1.е., with the in-
terpetiolar portion well developed and the intra-
petiolar portion reduced or absent; the stipule lobes
may be broadly rounded or acute to obtusely an-
gled, and straight at the base or rounded to lobed
or subauriculate on one or both sides. Leaf ar-
rangement may be paired or occasionally verti-
cillate. Inflorescence shape ranges from pyrami-
dal, in most species, to elongated and cylindrical
or spiciform, or to rounded-corymbiform with the
basal branches ascending and nearly as well as to
better developed than the central axis. Inflores-
cence and corolla colors may be pale green to
white, or dull to bright yellow, orange, red, purple,
violet, or blue; corolla color may be similar to or
contrast with the inflorescence branches. In gen-
eral, most species with brightly colored inflores-
cences or corollas vary from yellow to red, or purple
to blue; relatively few species combine colors from
these two groups. Calyx limb length varies from
relatively short, 0.2-1.2 mm long, as in most spe-
cies of Psychotria subg. Heteropsychotria, to rela-
tively well developed, to 20 mm long in Palicourea
bella and related species. Corolla shape varies
from somewhat swollen and a little gibbous at the
base and generally straight there and in the tube,
to strongly gibbous at the base, bent to 90° just
above this swelling, and curved in the tube; the
tube may be slender to relatively stout. Corolla
length varies from relatively short in the tube, 4—
5 mm long, to ca. 40 mm long in species of several
different groups; the length of the lobes varies, but
generally it is similar in proportion to the length of
the tube. Linear corolla lobe appendages and
colored multicellular corolla trichomes are
found in a few species. Fruit shapes described
here are evident mainly on dried specimens and
are due to the aggregate shape of the pyrenes.
Dried fruits range from subglobose to ellipsoid or
obovoid and may sometimes be laterally flattened.
Pyrene number is typically 2, but ranges to 4-5
in a few species; this number may be constant or
variable within a species. The longitudinal pyrene
ridges may be rounded to nearly flat or raised and
n
=
arp.
Characters that may vary within an individual
species include general shape and apex (ie.,
rounded vs. obtuse or acute) of the calyx lobes;
density of the pubescence on vegetative and some-
times reproductive structures (e.g., see discussion
of Palicourea alpina in Taylor, 1993); color of the
inflorescences and corollas, within the general color
groups noted above; length of the stipules and ped-
‘icels (at least to some extent); and external pubes-
cence of the corolla, including presence or absence
of the multicellular colored trichomes. Some mor-
phological structures appear to vary in correlated
fashion on individual plants, in particular the pro-
portional widths of leaves, stipule lobes, inflores-
cence bracts, and calyx lobes are usually similar.
Inflorescence and corolla color appear to vary
among closely related species and perhaps are cor-
related with reproductive isolating mechanisms, as
in the example of P. crocea and P. croceoides, above.
Among closely related species, for example, three
that are morphologically similar have very different
inflorescence and corolla colors: P. ovalis Standl.,
found at 1200-2000 m in Colombia and Ecuador,
has blue to purple inflorescences and corollas; P
chimboracensis Standl., found at 150—1400 m in a
similar range, has orange to red inflorescence
branches and yellow corollas; and Р. heilbornii
Standl., found at 1400-2200 m only in Pichincha
Province, Ecuador, has yellow to orange inflores-
cence branches and deep red corollas. Species of
subgenus Palicourea sect. Grandiflorae present a
similar example: of four similar species, two (Р.
grandiflora (Kunth) Standl., P. macarthurorum C.
M. Taylor) have red-orange to yellow inflorescences
and flowers, while two others (P. nigricans K. Krause,
P. amapaensis Steyerm.) have violet to purple inflo-
rescences and flowers; one yellow-flowered species
and one purple-flowered species are sympatric
through most of Amazonian South America, in var-
ious combinations, while both of the similarly col-
ored species are allopatric.
Several unusual or distinctive features found in
one or several species of Palicourea are also found
in other genera of Rubiaceae, often from other
tribes or subfamilies, and thus appear to have been
derived more than once in the family. One such
feature is a spathaceous calyx, which is completely
fused in bud and split irregularly by the elongating
calyx; this feature is found in P. spathacea
Taylor and is also known in other genera, e.g., Pen-
tagonia Benth., Hippotis Karst. (both Hippotideae),
Volume 84, Number 2
1997
Taylor 229
Conspectus of Palicourea
and Phellocalyx Bridson (Gardenieae, African). The
calyx lobes of several species of Palicourea are
somewhat to strongly unequal in length on an in-
dividual flower; this feature is notable in Palicourea
mexiae Standl., P. gomezii C. M. Taylor, P. gibbosa
Dwyer, P. discolor K. Krause (all subg. Montanae),
P. fastigiata, P. cymosa (Ruiz & Pav.) DC. (both
subg. Palicourea), and other species, and also in
some species of Rondeletia L. (Rondeletieae), Pen-
tas Benth. (Hedyotideae), and many other genera.
Verticillate leaves are found in several species of
subgenus Palicourea, including P. triphylla DC., P.
officinalis Mart., and the species of section Cor-
ymbiferae, and also in species of Duroia L.f. (Gar-
denieae), Remijia DC. (Cinchoneae), Declieuxia
Kunth (Psychotrieae), Bathysa C. Presl, Rondeletia
(both Rondeletieae), and other genera. Parallel (or
convergent) derivation of distinctive conditions can
be outlined for additional characters including cap-
itate inflorescences with well-developed bracts,
well-developed slender white corollas, and relative-
ly large pyrenes.
Within Palicourea, the diagnostic characteristics
noted above as used to distinguish species are dis-
tributed among the individual species in various—
for some features, nearly all possible—combina-
tions. This distribution of character states suggests
that at least some of these features have been de-
rived more than once, in some cases apparently
several times. Within the classification presented
here, characters that have apparently originated
more than once include the following:
* Multicellular, elongated, colored trichomes
on the exterior of the corolla are found in a number
of species with inflorescences and flowers of both
color groups, including Palicourea lachnantha
Standl. and P mansoana (Müll. Arg.) Standl. of
subgenus Palicourea, and P. thermydri J. H.
Kirkbr, P rigidifolia (Dwyer & M. V. Hayden)
Dwyer, P. eriantha DC., P. dorantha Wernham, P.
Justicioides Standl., P. calycina Benth., and P. cor-
nigera C. M. Taylor, of several different series in
subgenus Montanae sect. Montanae.
% Distinctive obovoid, laterally somewhat flat-
tened fruits are found in a number of species of
subgenus Montanae, including the species of sec-
tion Obovoideae series 5, which have stipules with
truncate continuous sheaths separating the lobes,
and the species of section Obovoideae series 6,
which have laminar stipules with the lobes closely
set on each side of a narrow, acute to concave sinus.
Thus, either fruit shape is a shared character and
stipule form variable among these species, or the
distinctive fruit shape has arisen at least twice in
these groups while the stipule form has been de-
rived only once.
4 The form of the longitudinal pyrene ridges is
distinctive for several species and useful in sepa-
rating some species groups. The pyrenes of most
species have rounded, usually rather low ridges,
but relatively sharp, pronounced ridges seem to
have arisen independently in such species as Pal-
icourea lachnantha and P. calophylla DC., which
belong to two different sections of subgenus Pali-
courea, and in the species of subgenus Montanae
sect. Cephaeloides series 10 and section Montanae
series 4 subseries g.
¢ Calyx length shows marked variation
throughout the genus, and to some degree within
each species group. It is most variable among spe-
cies of subgenus Montanae. Relatively long calyx
limbs seem to have arisen several times in the ge-
nus, in P. cymosa of subgenus Palicourea, and in
subgenus Montanae in species such as P. orosiana
C. M. Taylor, P. chignul C. M. Taylor (both section
Obovoideae), and the species of section Cephaelo-
~.
5.
¢ Perhaps the most striking example of а fea-
ture that has arisen more than once is the linear
corolla lobe appendages found in P. rigidifolia, P.
denslowiae J. Н. Kirkbr, P. lehmannii (Rusby)
Standl., P. corniculata C. M. Taylor, and P. corni-
gera C. M. Taylor, which are all purple- to blue-
flowered species of the western Andean cordillera
of Colombia and Ecuador. These appendages differ
morphologically: the first four species bear a single
appendage originating from the middle of the ab-
axial surface of the corolla lobe (Fig. 9D, E), which
is developed to various degrees in the different spe-
cies and which is found in other species of Psy-
chotrieae and other tribes; whereas Р. cornigera
bears two appendages per lobe, one originating
from each side of the base of the lobe (Fig. 9B), an
apparently unique feature in the family.
These combinations of morphological characters
indicate that at least some of them have been de-
rived more than once. They may suggest also the
possibility that the classification proposed here
should be reordered; however, any reordering will
indicate the repeated derivation of other features,
which suggests that the repeated origin of at least
some of them is a real occurrence. This in turn
suggests that different species may be responding
in similar ways to similar patterns of selection in
the environment. Particularly suggestive of this last
possibility is the repeated similar variation in inflo-
rescence and corolla colors (green and white, yel-
low-orange-red group, and purple-blue group) with-
230
Annals of the
Missouri Botanical Garden
in many species groups; the repeated derivation of
relatively long, colored calyx limbs; and the unusu-
al, prominent linear corolla lobe appendages de-
scribed above. In this last case, convergent occur-
rence of unusual structures that are morpho-
logically distinct suggests that some aspect of the
environment of this particular region favors these
structures.
AN INFRAGENERIC CLASSIFICATION OF PALICOUREA
The scheme presented below is based on a sur-
vey of the entire genus and monographic work on
part of it. It classifies 188 species. This scheme
does not include all species in this genus: those not
included either have not been studied in adequate
detail or are as yet too poorly known for classifi-
cation. Synonyms at the species level are not in-
cluded here, but can be found on the World Wide
Web at http://www.mobot.org/ MOBOT/Staff/Re-
search/taylor/palihome.html. Species included in
this classification that are described in this article
are indicated with an asterisk. Geographic distri-
butions presented for the species are approximate.
* Amazonia" is used here in the sense of Takhtajan's
(1986) “Amazonian Region” (ie., including the
Orinoco and other non-Amazonian drainages of
lowland, moist to wet, eastern to central South
America). Previously published infrageneric taxa of
Palicourea are included here. Most of the infrage-
neric taxa subordinate to section that are proposed
here are informal; formal description of these and
phylogenetic analyses will depend on the comple-
tion of monographic work.
This scheme suffers the limitations inherent in
translating what is clearly a branched pattern to a
linear list. Species are listed in an order that at-
tempts to place similar and apparently closely re-
lated species together.
Palicourea Aubl. Hist. pl. Guiane 172, t. 66.
1775. TYPE: Palicourea guianensis Aubl.
Subgenus I. Palicourea. Leaves paired to ver-
ticillate, usually membranaceous to papyraceous, or
subcoriaceous in species of savanna habitats, usu-
ally minutely pustulose abaxially; stipules united
around the stem into a continuous sheath, this often
reduced, the lobes reduced to well developed or
obsolete; inflorescences and flowers variously col-
ored; corollas tubular to somewhat funnelform,
somewhat to strongly swollen and gibbous at base,
generally straight to somewhat bent here and in
tube, externally glabrous or pubescent, sometimes
with colored multicellular trichomes; pyrenes 2-5
per fruit. Generally found at lower elevations, 0—
1200(1500) m, throughout the range of the genus.
Section А. Palicourea.
Psychotria sect. Oribasia Müll. Arg., in Mart., Fl. bras.
6(5): 223. 1881. Palicourea sect. Oribasia (Müll.
to base his name on Oribasia Schreb., Gen. pl. 124.
1789, nom. illeg., nom. superfl. for гера Aubl.
t. W. I. 346
Palicourea sect. Stephanium Griseb., Fl. B :
861. : Palico ourea guianensis. ree : Author-
ship of section Stephan ит is here ascribed solely to
atelia Aubl. and therefore nomenclaturally not syn-
onymous with Palicourea, although as originally cir-
cumscribed it included P. guianensis as we
Palicourea ser. Suberosae (Miill. Arg.) K. Schum., У Engl.
antl, Nat. Pflanzenfam. 4(4): 115. 1891. E
chotria ser. Suberosae Müll. Arg., in Mart., Fl.
6(5): 227. 1881. TYPE: Palicourea rigida Kui
Psychotria ser. Paniculatae Müll. Arg., in Mart., Fl. bras.
(5): 228. 1881. TYPE: Psychotria sellowiana DC.,
lectotype, here designated, — Palicourea guianensis.
Leaves paired; stipule lobes ligulate, usually obtuse
to rounded, relatively well-developed; inflorescences
pyramidal to broadly so; pyrenes 2-5. One wide-
spread species, Mexico and Antilles to Bolivia and
southern Brazil, the remaining species Amazonian.
P. guianensis Aubl. Lowlands of Mexico and
Antilles to southern Brazil and Bolivia; pyrenes
2-5 per fruit.
P. grandifolia Kunth. Amazonia; similar to Р
guianensis and perhaps not actually distinct; py-
renes 4—5 per fruit.
P. buntingii eiie n енген perhaps not
distinct from Р guia
P. tepuicola Seem. Venen perhaps not
distinct from P. guiane
P. rigida Kunth. йаш зауаппаз
throughout South America.
P. semirasa Standl. Middle elevations, Venezue-
la and Colombia.
P. flavifolia (Rusby) Standl. Middle elevations,
Bolivia. ;
P. attenuata Rusby. Middle elevations, Bolivia.
P. mansoana (Müll. Arg.) Standl. Western Ama-
zonia.
P. lasiantha K. Krause. Western Amazonia.
P. расћусајух Standl. Western Amazonia; sim-
ilar to P. lasiantha, perhaps not distinct from it.
P. lachnantha Standl. Western Amazonia; char-
acteristically drying turquoise-black, perhaps an
uminum accumulator.
P. anianguana C. M. Taylor*. Local in easte™
Volume 84, Number 2
1997
Taylor 231
Conspectus of Palicourea
Ecuador; combines characters of P. lachnantha
and P. lasiantha.
Section B. Grandiflorae, sect. nov. TYPE: Pali-
courea grandiflora Kunth.
Folia lobis stipularum reductis. Fructus magnitudinem
mediam generis aliquantum ad valde excedens.
Leaves paired; stipule lobes relatively reduced,
triangular to deltoid or shortly ligulate; inflores-
cences broadly pyramidal to corymbiform-rounded;
fruit somewhat to markedly larger than average, 4—
15 mm long; pyrenes 2. Amazonia.
P. grandiflora Kunth. Amazonia.
P. amapaensis Steyerm. Northeastern Amazo-
nia.
P. nigricans K. Krause. Western Amazonia; sim-
ilar to and perhaps a western sister species to P.
amapaensis.
P. macarthurorum C. M. Taylor. Western Ama-
zonia, low to middle elevations.
Section C. Crocothyrsae Griseb., Fl. Brit. W. I.
345. 1861. TYPE: Palicourea crocea (Sw.)
Roem. & Schult., lectotype, here designated.
Leaves paired or verticillate; stipule lobes del-
toid to narrowly triangular or lanceolate, acute,
rather short to well developed; inflorescences py-
ramidal to broadly so, corymbiform-rounded to fas-
tigiate or spiciform (series 1) or to narrowly pyra-
midal (series 2); pyrenes 2. Mexico and Antilles to
Paraguay and Bolivia.
Series 1. Croceae (Müll. Arg.) K. Schum., in
Engl. & Prantl, Nat. Pflanzenfam. 4(4): 115.
1891. Psychotria ser. Croceae Müll. Arg., in
Mart., Fl. bras. 6(5): 228. 1881. TYPE: Pali-
courea crocea, lectotype, here designated.
Inflorescences pyramidal to broadly so, corym-
biform-rounded, fastigiate, or spiciform.
Subseries a. Subeymosae (Müll. Arg.) С. M. Tay-
lor, stat. et comb. nov. Basionym: Psychotria
ser. Subcymosae Müll. Arg., in Mart., Fl. bras.
6(5): 229. 1881. TYPE: Palicourea barraensis
Müll. Arg., lectotype, here designated, —
longiflora (Aubl.) Rich.
Stipule lobes triangular to narrowly so. Mexico
and Antilles to Paraguay and Bolivia.
P. croceoides Ham. Antilles, and perhaps
throughout lowland moist South America; this
and the next two species comprise a poorly un-
derstood, variable, widespread complex of forms
that are most diverse in Amazonia.
P. сгосеа (Sw.) Roem. & Schult. Mexico to Par-
aguay and Bolivia.
P. fastigiata Kunth. Swamps and e of black-
water rivers, lowland South Ameri
P. charianthema Standl. Жыйын اند
P. subspicata Huber. Western Amazonia.
P. herzogii Standl. Middle elevations, Bolivia,
perhaps to Peru or Ecuador.
P. huberi Steyerm. Guayana Highlands.
P. lancigera Steyerm. Guayana Highlands.
P. longiflora (Aubl.) Rich. Northern and eastern
Amazonia.
P. maregravii A. St.-Hil. Eastern Amazonia.
P. coriacea Mart. Savannas of eastern South
America, pyrrophytic.
P. mello-barretoi Standl. Savannas, Brazil;
perhaps not distinct from P. coriacea.
P. officinalis Mart. Savannas; Brazil.
P. longistipulata (Müll. Arg.) Standl. Amazonia,
with marked clinal variation from east to west.
P. bracteosa Standl. Western Amazonia.
P. lasiophylla Standl. Western Amazonia, to
middle elevations.
P. nitidella (Müll. Arg.) Standl. Eastern Ama-
zonia.
Subseries b. Stipule lobes lanceolate. Western
Amazonia.
P. conferta (Benth.) Sandw. Western Amazonia.
F. тоге (Ruiz & Рау.) DC. Western Amazo-
P. punicea (Ruiz & Pav.) DC. Southwestern
Amazonia.
P. iquitoensis K. Krause. Local in northeastern
Peru.
P. plowmanii C. M. Taylor. Local in southeast-
ern Peru.
Р. jatun-sachensis С. M. Taylor. Northwestern
Amazonia.
Series 2. Verticillatae (Miill. Arg.) K. Schum., in
Engl. & Prantl, Nat. Pflanzenfam. 4(4): 115.
1891. Psychotria ser. Verticillatae Müll. Arg.,
in Mart., Fl. bras. 6(5): 227. 1881. TYPE: Pal-
icourea triphylla DC., lectotype, here desig-
nated.
Psychotria ser. Cylindricae Müll. Arg., in Mart., Fl. bras.
6(5): а 1881. TYPE: Psychotria tabacifolia Müll.
Arg., lectotype, here designated, = Palicourea ma-
crobotrys (Reiz & Pav.) DC.
Inflorescences pyramidal to elongated and nar-
rowly so. One widespread species, Mexico to Bo-
livia and southern Brazil, the remainder in Ama-
zonia.
Annals of the
Missouri Botanical Garden
P. macrobotrys (Ruiz & Pav.) DC. Southern
mazonia.
P. tetraphylla Cham. & Schltdl. Southeastern
razil.
P. longepedunculata Gardner. Southern Brazil.
P. calophylla DC. Northern Amazonia.
P. affinis Standl. Southern Amazonia.
P. anisoloba (Miill. Arg.) Boom € M. T. Cam-
pos. Central Amazonia.
P. triphylla DC. Mexico and Cuba to Bolivia
and southern Brazil; leaves usually verticillate,
though consistently paired in Cuba.
P. melheana Jung-Mend. Brazil; very similar to
P. triphylla.
Section D. Corymbiferae (МШ. Arg.) С. M. Tay-
lor, stat. et comb. nov. Basionym: Psychotria
ser. Corymbiferae Miill. Arg., in Мап., Fl. bras.
6(5): 229. 1881. TYPE: Palicourea corymbi-
fera (Müll. Arg.) Standl., lectotype, here des-
ignated.
Psychotria ser. Bracteosae Müll. Arg., in Mart., Fl. bras.
6(5) . 1881. TYPE: Palicourea virens (Müll.
Arg.) Standl., lectotype, here designated.
Leaves verticillate or occasionally paired; stipule
sheaths subtruncate, lobes reduced or obsolete; in-
florescences pyramidal to corymbiform-rounded,
ith floral bracts often well developed; pyrenes 2—
5 per fruit. Amazonia.
P. corymbifera (Müll. Arg.) Standl. Amazonia;
pyrenes 2—5 per fruit.
Р. virens (Müll. po Standl. Eastern Amazonia;
pyrenes 4-5 per
P. irwinii ola па Amazonia; py-
renes 2 per fru
E ma (Кидре) DC. Amazonia, with
marked clinal variation from east to west; py-
renes 2 per fruit.
Subgenus II. Montanae, subg. nov. TYPE: Pali-
courea thyrsiflora (Ruiz & Pav.) DC.
Folia binatim jugata, papyracea chartacea subcori-
aceave, abaxialiter plerumque epustulata: stipulis lami-
naribus vel circum caulem connatis vaginam continuam
formantibus.
Leaves paired, usually papyraceous to charta-
ceous or subcoriaceous, usually not pustulose abax-
ially; stipules united around the stem in a usually
well-developed, continuous sheath with two short to
well-developed lobes, or laminar (1.е., interpetiolar)
and bilobed to varying degrees; inflorescences and
flowers variously colored; corollas tubular to fun-
nelform, somewhat swollen and gibbous at base and
generally straight there and in tube, to very strongly
gibbous and swollen at base, constricted and
strongly bent, to as much as 90°, just above this,
and straight to curved in tube, externally glabrous
or pubescent with trichomes of various types, in-
cluding short to long, multicellular, colored tri-
chomes; pyrenes 2(4) per fruit. Generally found at
higher elevations, (150)1000-3500 m (to limit of
woody vegetation). Greater Antilles, Mexico and
Central America to Venezuela and Bolivia.
Section E. Montanae, sect. nov. TYPE: Palicou-
rea thyrsiflora (Ruiz & Pav.) DC.
Folia stipulis circum caulem connatis vaginam contin-
uam formantibus. Fructus pyrenis ellipsoideis ovoideis
subglobosisve
Stipules united around the stem in a continuous,
usually well-developed and truncate sheath, with
lobes triangular to narrowly so or rarely lanceolate;
inflorescences pyramidal to narrowly so or some-
times relatively short and few-flowered in species
of higher elevations; infructescences typically be-
coming purple in all species; fruits ellipsoid, ovoid,
or subglobose, sometimes laterally flattened. Great-
er Antilles, Mexico and Central America to Vene-
zuela and Bolivia.
Series 3. Fruits generally ellipsoid to ovoid and
laterally flattened. Mexico and Antilles to Ven-
ezuela and Bolivia.
Subseries e. Stipules with sheath moderately to
well developed or sometimes reduced in spe-
cies of higher elevations, with lobes triangular
to narrowly so, generally persisting with the
stipules. Greater Antilles, Mexico and Central
America to Venezuela and Bolivia.
P. thyrsiflora (Ruiz & Pav.) DC. Andes of Co-
lombia and Venezuela to Bolivia.
Р. padifolia (Willd. ex Roem. & Schult.) С. М.
Taylor & Lorence. Mexico to Panama; sim-
ilar to P. thyrsiflora.
P. alpina (Sw.) DC. Greater Antilles; similar to
P. thyrsiflora, flowers monomorphic (i.e., по!
distylous).
P. eriantha DC. Hispaniola; similar to P. alpina,
flowers monomorphic.
P. angustifolia Kunth. Costa Rica to Venezuela
and Peru
P. rigidifolia (Dwyer & M. V. Hayden) Dwyer
Panama to western Colombia; ес _
placed in Palicourea, similar in several fea-
tures to P. angustifolia and perhaps P. densa
Standl. (Series 4, Subseries f)
P. weberbaueri K. Krause. Colombia to Peru;
perhaps a color variant of P. thyrsiflora.
Volume 84, Number 2
1997
aylor 233
Conspectus of Palicourea
leucantha Donn. Sm. Guatemala.
acetosoides Wernham. Western Colombia.
lutulenta Standl. Colombia.
heterantha Standl. Colombia; perhaps not
distinct from P. lutulenta.
anarina C. M. Taylor.* Ecuador; similar to
"s thyrsiflora.
subalata Standl. ex Steyerm. Colombia to
eru.
gentryi C. M. Taylor.* Local in east-central
Ecuador; similar to P. subalata.
subalatoides C. M. Taylor.* Ecuador; similar
to P. subalata.
. danielis Standl. Local in northwestern Colom-
bia.
. luteonivea C. M. Taylor.* Southern Colombia
to Ecuador; similar to P. thyrsiflora.
holmgrenii Standl. Southern Colombia to Ec-
uador.
vulcanalis Standl. ex C. M. Taylor.* East-
central Ecuador.
ulloana C. M. Taylor.* Ecuador to northern
Peru; pyrenes 4.
calothyrsus K. Schum. Ecuador; perhaps a
high-elevation form of P. thyrsiflora.
pennellii Standl. Colombia; perhaps not dis-
tinct from P. calothyrsus.
calantha Standl. Ecuador.
ponasae K. Krause. Peru.
psittacorum Standl. Colombia to Ecuador.
ionantha Standl. Colombia.
herrerae Standl. Peru to Bolivia.
lobbii Standl. Ecuador.
subtomentosa (Ruiz & Pav.) C. M. Taylor.*
Ecuador to Bolivia, with marked variation
from north to south.
. buchtienii Standl. Bolivia; provisionally
placed here.
P. corniculata C. M. Taylor.* Northern Ecua-
or.
P. anderssoniana C. M. Taylor.* Ecuador.
а Менде | x TN
о о ш G
a-
a-
as eee a
4
Subseries d. Stipules with sheaths well developed,
with lobes narrowly triangular to setaceous and
usually deciduous to caducous, or obsolete.
Ecuador and perhaps Peru.
P. flavescens Kunth. Ecuador and perhaps
Peru
P. tectoneura K. Schum. & K. Krause. Ecua-
d
or.
P. prodiga Standl. ex C. M. Taylor.* Central Ec-
uador
Series 4. Fruits generally subglobose to ovoid, not
laterally flattened. Costa Rica to Venezuela
and Bolivia
Subseries e. cala tubular to somewhat funnel-
form, externally glabrous or pubescent with
relatively short, usually colorless trichomes.
Costa Rica and Venezuela to Peru.
P. amethystina (Ruiz & Pav.) DC. Colombia to
P, сай Standl. Costa Rica; similar іо Р.
amethystina.
P. purpurea С. M. Taylor. Costa Rica and Pan-
P. demissa Standl. Venezuela and Colombia to
Ecuador.
P. ovalis Standl. Western Colombia to Ecuador;
similar to P. chimboracensis and P. heilbor-
nii.
P. chimboracensis Standl. Western Colombia to
Ecuador; similar to P. ovalis and P. heilbor-
nii.
heilbornii Standl. Local in Ecuador; similar
to P. ovalis and P. chimboracensis.
latifolia K. Krause. Colombia to Peru; similar
to P. amethystina.
, discolor К. Krause. Costa Rica and Panama;
similar to P. latifolia.
deviae C. M. Taylor.* Ecuador to western Co-
lombia.
perquadrangularis Wernham. Venezuela to
Colombia.
stipularis Benth. Western Colombia to Peru.
albert-smithii Standl. Eastern Colombia to
Venezuela; similar to P. stipularis.
lehmannii (Rusby) Standl. Local in western
Colombia; provisionally placed here.
a-
R-
ve ем T N
Subseries f. Calyx relatively well developed; co-
rollas tubular, often relatively well developed,
externally glabrous to pubescent with short
colorless trichomes or sometimes with multi-
cellular, colored, showy trichomes. Central
America to northern Ecuador.
P. heterochroma K. Schum. & K. Krause.
Western Colombia to Ecuador.
P. albocaerulea C. M. Taylor. Local in Costa
Rica.
P. lancifera Standl. & L. O. Williams. Honduras
to Costa Rica.
P. macrocalyx Standl. Costa Rica to Panama.
P. standleyana C. M. Taylor. Costa Rica to Co-
lombia.
P. antioquiana Standl. Northwestern Colombia.
P. tunjaensis C. M. Taylor. Colombia.
234
Annals of the
Missouri Botanical Garden
P. calophlebia Standl. Western Colombia to Ec-
uador.
P. toroi Standl. Western Colombia.
P. dorantha Wernham. Western Colombia.
P. densa Standl. Western Colombia.
P. justicioides Standl. Western Colombia.
P. calycina Benth. Local in Ecuador.
P. azurea C. M. Taylor.* Local in Ecuador.
P. jaramilloi C. M. Taylor.* Southern Ecuador.
P. cornigera C. M. Taylor.* Southern Ecuador.
Subseries g. Inflorescences pyramidal to broadly
so or corymbiform-rounded, usually with floral
bracts relatively well developed and broad; co-
rollas tubular, with tubes relatively stout; py-
renes generally triangular in cross section.
Western Colombia to Peru.
P. quadrilateralis C. M. Taylor. Northwestern
Colombia (in press, Novon).
P. thermydri J. H. Kirkbr. Northwestern Colom-
bia
P. ни J. H. Kirkbr. Local in northwest-
ern Colombia.
Section F. Obovoideae, sect. nov. TYPE: Pali-
courea hospitalis Standl.
Folia stipulis laminaribus vel circum caulem connatis
vaginam continuam formantibus. Aras ok а ue obo-
voideus etiam lateraliter subcomplanatu
Stipules united around the stem in a continuous
sheath or laminar; corollas generally funnelform,
somewhat to strongly gibbous at base, slightly to
strongly constricted and bent there, externally gla-
brous or pubescent with usually colorless tri-
chomes; fruit generally obovoid and somewhat flat-
tened laterally. Costa Rica to Venezuela and
Ecuador.
Series 5. Stipules united around the stem in a con-
tinuous truncate sheath; inflorescences gener-
ally yellow, or sometimes flushed with purple
or blue, generally remaining or becoming yel-
low in fruit. Costa Rica to northwestern Colom-
bia or perhaps Ecuador.
P. lasiorrhachis Oerst. Costa Rica to north-
western Colombia.
P. pendula C. M. Taylor. Local in Panama.
P. vestita Stand]. Local in Costa Rica.
P. obtusata K. Krause. Venezuela.
P. orosiana C. M. Taylor. Local in Costa Rica
and western Panama.
P. chiriquina Standl. Local in western Panama.
P. montivaga Standl. Costa Rica, perhaps to
western Panama.
P. adusta Standl. Costa Rica, perhaps to western
Series 6. Stipules laminar; inflorescences yellow
to orange or red, often becoming purple in
fruit. Panama to Ecuador.
P. hospitalis Standl. Western Colombia to Ec-
uador.
Р. diet Standl. Local in Ecuador; perhaps not
distinct from P. hospitalis.
P. chignul C. M. Taylor.* Northwestern Ecuador.
P. kalbreyeri Standl. Western Colombia to Ec-
uador.
P. lugoana C. M. Taylor.* Western Colombia to
Ecuador.
P. asplundii C. M. Taylor.* Ecuador.
P. gibbosa Dwyer. Panama to Ecuador.
P. lyristipula Wernham. Colombia to Ecuador.
P. myrtifolia K. Schum. & K. Krause. Ecuador.
Series 7. Stipules laminar; inflorescences pyra-
midal to corymbiform-rounded, variously col-
ored in flower and fruit; pyrenes with ridges
rounded and often becoming planar to nearly
smooth at maturity. Panama to Venezuela and
Ecuador.
P. apicata Kunth. Venezuela and Colombia to
cuador.
р, пенни Standl. Western Colombia to
Ecuad
F а (Ruiz & Pav.) DC. Peru, perhaps
north to Colombia.
P. vaginata Benth. Colombia and Venezuela;
identity not clear, provisionally recognized
ere.
P. tubuliflora Dwyer. Panama.
P. tumidonodosa Dwyer. Panama, perhaps to
northwestern Colombia.
P. salicifolia Standl. Costa Rica to western Pan-
ama.
P. pauciflora Standl. Local in Costa Rica and
western Panama; provisionally placed here,
fruit unknown.
P. nubigena Standl. Western Colombia, perhaps
to Peru
P. mexiae Standl. Costa Rica to Peru
P. gomezii C. M. Taylor. Costa Rica to | Ecuador.
P. amplissima (Standl. ex Steyerm.) С. M. Тау-
lor. Western Colombia.
Section G. Pseudoamethystinae, sect. nov.
TYPE: Palicourea obovata (Ruiz & Pav.) DC.
Folia stipulis laminaribus. Toia ex subgloboso ovoi-
deus, lateraliter nec complan
Stipules laminar; inflorescences pyramidal; inflo-
Моште 84, Митбег 2
1997
Taylor 235
Conspectus of Palicourea
rescences and flowers blue to purple; corollas with
relatively well developed tubes; fruits subglobose
to ovoid, not laterally flattened. Western Colombia
to Peru.
P. lineata Benth. Colombia to Peru; similar to
P. stipularis but with stipules laminar.
P. obovata (Ruiz & Pav.) DC. Colombia to Peru;
similar to P. latifolia but with stipules laminar.
P. subscandens Standl. ex Steyerm. Western
Colombia to Peru; similar to P. perquadrangularis
but with stipules laminar.
Section H. Psychotrioides, sect. nov. TYPE:
Palicourea petiolaris Kunth.
Folia stipulis laminaribus. M plerumque ellipso-
ideus etiam lateraliter complanatu
Stipules laminar; inflorescences pyramidal to
broadly so or rounded-corymbiform, frequently yel-
low or sometimes pink; corollas yellow to white or
pink; fruit generally ellipsoid and distinctly flat-
tened laterally. Mexico and Costa Rica to Venezuela
and Ecuador.
Series 8. Corollas with tubes relatively short. Cos-
ta Rica and Venezuela to Ecuador and in Ja-
maica.
P. petiolaris Kunth. Venezuela to eastern Co-
lombia.
P. leuconeura Standl. Venezuela to eastern Co-
ombia.
steyermarkii C. M. Taylor. Venezuela.
. tilaranensis С. M. Taylor. Costa Rica.
garciae Standl. Costa Rica to Ecuador.
jahnii Standl. Venezuela, perhaps to Colom-
bia.
caloneura Rusby. Local in northeastern Co-
lombia.
abbreviata Rusby. Local in northeastern Co-
lombia
ч
sarnechii C. M. Taylor. Northwestern Colom-
bia (in press, Novon).
ers Standl. Venezuela and eastern Colom-
uud Griseb. Jamaica; provisionally
placed here.
wilesii C. D. Adams. Jamaica; similar to P
pulchra.
aschersonianoides (Wernham) Steyerm.
Venezuela to eastern Colombia.
Series 9. Inflorescences corymbiform-rounded; со-
rollas with relatively long tubes, these often
elongating shortly before anthesis from rela-
tively short buds. Southern Mexico and Vene-
zuela to Ecuador.
я 39 Cw SD SD OO Ww т
P. lineariflora Wernham. Venezuela to eastern
Colombia.
P. macrantha Loes. Southern Mexico; provi-
sionally placed here.
P. boyacana Standl. Eastern Colombia.
P. andrei Standl. Colombia to northern Ecuador.
P. candida C. M. Taylor.* Northern Ecuador.
Section I. Cephaeloides, sect. nov. TYPE: Pali-
courea bella (Standl.) Dwyer.
Folia stipulis laminaribus. Inflorescentia bracteis et
limbo calycino modice vel bene evolutis.
Stipules laminar; inflorescences pyramidal to
broadly so or corymbiform-rounded, or congested to
subcapitate, usually pink to red-purple; usually
with relatively well-developed and showy bracts;
calyx limb moderately to very well developed; co-
rollas tubular to somewhat funnelform, usually
white to purple, often with tubes relatively well de-
veloped. Costa Rica to Ecuador.
Series 10. Inflorescences pyramidal to corymbi-
form-rounded; calyx moderately to well devel-
oped, usually colored and showy; pyrenes gen-
erally relatively large, with ridges relatively
sharp and well developed. Costa Rica to west-
ern Colombia.
P. bellula C. M. Taylor. Costa Rica.
P. stenosepala Standl. Western Colombia to Ec-
uador.
P. bella (Standl.) Dwyer. Costa Rica to Panama.
P. spathacea C. M. Taylor. Local in Costa Rica.
P. hammelii C. M. Taylor. Costa Rica to Pana-
ma.
P. killipii Standl. Local in western Colombia.
P. ochnoides Dwyer. Western Panama.
Series 11. Stipules often relatively large; inflores-
cences congested to subcapitate, with bracts
usually well developed and showy; pyrenes
generally not unusually large, with ridges
rounded and frequently low. Panama and Ven-
ezuela to Peru.
P. grandistipula (Standl. ex Steyerm.) C. M.
Taylor. Western Colombia.
P. macbridei Standl. Peru.
P. condorica C. M. Taylor.* Southern Ecuador.
deca C. M. Taylor.* Northeastern Ecua-
Р. سو (Standl. & Steyerm.) Steyerm. Ven-
ezuela and Colombia.
P. skotakii C. M. Taylor. Local in Costa Rica.
P. sopkinii C. M. Taylor. Western Colombia (in
press, Novon).
236
Annals of the
Missouri Botanical Garden
Figure 2.
subsp. lojana, flowering bran
&
P. dimorphandrioides (Dwyer) C. M. Taylor.
Eastern Panama to western Colombia.
P. acanthacea C. M. Taylor. Western Panama to
northwestern Ecuador.
P. frontinensis Cogollo & C. M. eae Western
Colombia ae press, Cal
P. orquidea C. M. Taylor. Western Colombia (in
press, Nobb).
DESCRIPTIONS OF New TAXA
The following newly described taxa are included
in the infrageneric classification presented above,
which indicates their relationships within Palicou-
rea. They are treated here in alphabetical order; the
position of each species in the infrageneric classi-
fication is indicated at the end of the morphological
description. Descriptions and diagnostic characters
—A. Paia anderssoniana, flowering branch, based on Asplund 17430.—B. Palicourea subtomentosa
sed on Harling & Andersson 21367. C,
pd ieu 21825.—C. Fiosan branch.—D. Flower in bud. A-C to 5-ст scale.
D, Palicourea canarina, based on Harling
of the genus Palicourea have been presented by
Taylor (1989) and Burger and Taylor (1993).
Palicourea anderssoniana C. M. Taylor, sp. по“.
TYPE: E cuador. Pichincha: Reserva Florísti-
Colorados, a 3.5 km al NE de la carretera,
estribaciones occidentales del Volcán Pichin-
cha, 0013'53"S, 78°48'10"W, 11 Aug. 1985,
2 pores & V Zak 7992 (holotype,
-4990804; isotype, AAU). Figure 2A.
Haec species a congeneris inflorescentiis sessilis rams
imis reflexis, pedicellis sat longis atque corolla lutea gib-
bosa glabra distinguitur.
Flowering at 1.5 m tall, to 5 m tall; stems gla-
brous. Leaves paired; blades elliptic to rather broad-
наб
Volume 84, Number 2
1997
Taylor 237
Conspectus of Palicourea
ly so or oblanceolate, 7-15 Х 2.5-8 cm, at apex
rather abruptly acuminate with tips 5-15 mm long
and usually falcate, at base cuneate to acute, pa-
pyraceous to chartaceous, adaxially glabrous, abax-
ially sparsely to moderately pilosulous, usually
markedly more densely so on costa and secondary
veins; secondary veins 8-15 pairs, generally ex-
tending to margins, with 1-3 weak intersecondary
veins usually present between pairs of secondary
veins, adaxially venation plane or costa sometimes
thickened, abaxially costa and secondary veins
prominulous and lesser venation thickened; mar-
gins thinly cartilaginous, entire; petioles glabrous,
6-25 mm long; stipules glabrous, united around the
stem in a continuous truncate sheath 2.5-3 mm
long, lobes narrowly triangular, 4-5 mm long,
acute, ciliolate. Inflorescences deflexed, sessile;
panicles broadly pyramidal to corymbiform, 6-15
X 6-15 cm excluding corollas, with secondary axes
24 pairs, the basalmost pair reflexed, with flowers
pedicellate in cymules of 3—7; bracts narrowly tri-
angular, 3-9 mm long, acute, ciliolate, those sub-
tending secondary axes 5-9 mm long, those sub-
tending pedicels 3-5 mm long; pedicels 5-17 mm
long; axes, bracts, and pedicels glabrous, yellow to
red; flowers with hypanthium glabrous, cylindrical,
1-1.3 mm long, calyx limb glabrous, 1.5-2 mm
long, divided for % to nearly completely to base,
lobes triangular to ovate, rounded to acute or slight-
ly acuminate, equal or somewhat unequal on an
individual flower, entire; corolla tubular to some-
what funnelform, yellow, at base swollen and gen-
erally strongly gibbous, generally straight there,
straight or slightly curved in tube, externally gla-
brous or sometimes pilosulous on lobes in bud, in-
ternally glabrous except for a densely villous ring
ca. 4 mm above the base, tubes 17-19 mm long,
de diam. near middle, lobes 3-6 mm long,
slightly thickened adaxially at apex; anthers in ap-
parent short-styled form partially exserted, ca. 4
mm long; stigmas not seen; disk ca. 1 mm high.
Infructescences similar to reno fruit ellip-
soid to subglobose, ca. 5 pyrenes with
3–5 low, rather sharp а даре. Cloud
and wet montane forest at 1325-1930 m, north-
central Ecuador. [Subg. Montanae, Sect. Montanae,
Ser. 3, Subser. с
This species is distinguished by its sessile inflo-
rescences with the basalmost pair of secondary axes
reflexed, relatively long pedicels, and glabrous, gib-
bous, yellow corollas. This inflorescence arrange-
ment is unique in Palicourea. Vegetatively P. anders-
soniana resembles P. thyrsiflora. This handsome
plant is named in honor of Swedish botanist Len-
nart Andersson, whose work has contributed sig-
nificantly to our understanding of the Ecuadorian
flora and its biogeography.
Paratypes. ECUADOR. Pichincha: El Volante on
road from Chiriboga to Santo Domingo de los Colorados,
Asplund 17430 (S); along road between Tandayapa and
Mindo, 19 km from Tandayapa, about 5.5 km from Mindo,
Croat 49391 (MO); cantón Quito, parroquia Mindo,
“Bosque Protector Mindo,” cumbre de la montaña entre
río Mindo y río Bagasal, 00°02'N, 78°48'W, Delprete et al.
6098 (MO, TEX); Guajalito, carretera antigua Quito-Santo
Domingo, 00°10'S, 78%55'W, Palacios 525 (MO); along
new road Nanegal-Mindo, van der Werff 13397 (MO); Río
Guajalito, near Chiriboga, along old Quito-Santo Domingo
road, van der Werff et al. 12214 (MO)
Palicourea anianguana C. M. Taylor, sp. nov.
PE: Ec , : Afiangu, Rfo Napo,
‘oe 76°23'W, 16-27 Apr. 1983, J. E.
Lawesson, T. Læssøe & P. M. Jørgensen 39442
ере MO-4279338; isotypes, AAU, ОСА
3.
not seen). Figure
Haec species a congeneris lobis stipularibus bene ev-
olutis ex obtusis rotundatis, inflorescentia purpurea dense
pubescente, corolla trichomatibus purpureis longis vestita
atque pyrenis acute porcatis distinguitur.
Flowering at 5 m tall, to 8 m tall; stems glabrous
to densely puberulous. Leaves paired; blades elliptic
to broadly so, 15-32 X 7-19 cm, at apex shortly
acuminate with deltoid tips 5—10 mm long, at base
obtuse to truncate or somewhat rounded, membra-
naceous to papyraceous, adaxially minutely puber-
ulous to glabrous, abaxially moderately to densely
short-pilosulous throughout; secondary veins 12-18
pairs, occasionally looping to interconnect in distal
part of leaf, without or with 1(2) weak intersecon-
dary veins usually present between pairs of sec-
ondary veins, adaxially costa prominulous to prom-
inent, secondary veins thickened to prominulous,
and minor venation plane, abaxially costa promi-
nent, secondary veins prominulous, and minor ve-
nation plane and not particularly evident; margins
thinly cartilaginous, entire; petioles puberulous to
glabrous, 13-38 mm long; stipules glabrous to pu-
berulous, united around the stem in a continuous
truncate to concave sheath 1-1.5 mm long, lobes
ligulate, 9-13 mm long, obtuse to rounded, cilio-
late. Inflorescences erect; peduncles 4.5-9 cm long;
panicles pyramidal, 4-10 X 4–10 cm excluding co-
rollas, with secondary axes 4-9 pairs, with flowers
pedicellate in cymules of 3-7; bracts narrowly tri-
angular to lanceolate or linear, acute, those sub-
tending secondary axes displaced distally for Y to
% of length of first axis internode, 5-8 mm long,
those subtending pedicels 1-3 mm long; pedicels
1-6 mm long; peduncle, axes, bracts, and pedicels
238
Annals of the
Missouri Botanical Garden
Figure 3. Palicourea anianguana.—A. Flowering branch, based on Balslev et al. 62299.—B. Flower, based on
44.2.
Lawesson et al. 39.
densely tomentellous or hirsutulous, violet to red;
flowers distylous, with hypanthium densely lanate,
turbinate to cupuliform, 1.5-2 mm long; calyx limb
1-1.2 mm long, subtruncate to shallowly lobed, in-
ternally glabrous, externally densely lanate; corolla
tubular to somewhat funnelform, white to violet or
pink, somewhat swollen at base, generally straight
there and in tube, externally densely pubescent
with purple to violet trichomes 1-3 mm long, in-
ternally glabrous except for a densely villous ring
ca. 2 mm wide at ca. % of length of tube above
base, tube 15-20 mm long, 3-4 mm diam. near
middle, lobes triangular to deltoid, 2.5-3 mm long,
acute; anthers in short-styled form partially exsert-
ed, ca. 5 mm long, in long-styled form included,
positioned in upper % of tube, ca. 5 mm long; stig-
mas in short form ca. 3 mm long, positioned near
middle of tube, in long form ca. 1.5 mm long, ex-
serted; disk 1.5-2 mm long. Infructescences red-vi-
olet; fruit ellipsoid, 8-9 X 7-8 mm, lanate; py" enes
with sharp longitudinal ridges. Rainforest in tierra
firme at 260—350 m, northeastern Ecuador. [Subg.
Palicourea, Sect. Palicourea.]
This species is distinguished by its stipules with
well-developed obtuse to rounded lobes and short,
truncate to concave sheaths, thin-textured leaves.
well-developed pyramidal purple panicles, densely
tomentellous to hirsutulous inflorescence axes, la-
nate calyces, corollas that are externally pubescent
with long purple trichomes, fruit not at all flattened,
and pyrenes with sharp longitudinal ridges. It 18
similar to Palicourea lasiantha, which has corollas
externally pubescent with trichomes 0.5 mm long
Volume 84, Number 2
1997
Taylor 239
Conspectus of Palicourea
gure 4. A, B, Palicourea subalatoides, based on Asplund 18928.—A. Flowering branch.—B. Calyx and -e
based о
E ipea. C, D, Palicourea asplundii.—C. Flowering branch,
Tipaz et al. 280.—E. Palicourea lugoana, flower, based o
Neill 7532. res С to 5-cm scale; B, D-F to 5-mm scale
or shorter and inflorescence branches, calyces, and
fruits glabrous to puberulous; and to P. lachnantha,
which has acute stipule lobes 2-8 mm long, calyx
limb 1.8-2 mm long, and pyrenes with rounded
ridges. Palicourea anianguana combines distinc-
tive characteristics of both P. lasiantha and P. lach-
nantha, but it is not intermediate in these features
and is consequently here considered a separate, lo-
cal species. The specific epithet refers to the area
where this new species has been collected, Afiangu.
Paratypes. ECUADOR. Napo: Añangu, 5 bank of Río
Napo 95 km downstream from Соса, 00°32'S, 76°23’ W,
n Asplund 8672.—D. Flower, based o
n Lugo 14.—F. Palicourea gentryi, flower in bud, based on
Balslev et al. 62299 (AAU); Añangu, NW corner of Parque
Nacional Yasunf, ani - 7 digi 23. ^ Korning & Thom-
sen 47044 (AAU); Afiangu, in Parq acional Yasunf,
00°31-32'S, 76°23’ Ww. > et al. 39247 (AAU, MO).
Palicourea asplundii C. M. Taylor, sp. nov. TYPE:
Ecuador. Carchi: cantón Tulcán, Reserva In-
Tipaz, D. Rubio
MO-4990803; isotype, ОСМЕ not seen). Fig-
ure 4C, D
Haec species a congeneris stipularum interpetiolarium
240
Annals of the
Missouri Botanical Garden
lobis rotundatis, limbo calycino sat bene evoluto atque
corolla flava valde gibbosa curvataque distinguitur.
Flowering at 3 m tall; stems glabrous to hirsu-
tulous. Leaves paired; blades elliptic, 9.5-23.5 X
4.5-12.5 cm, at apex shortly acuminate with del-
toid to slender tips 3-8 mm long, at base cuneate
to obtuse, papyraceous, glabrous throughout; sec-
ondary veins 13-18 pairs, sometimes uniting with
margins in distal part of blade, with 1(3) interse-
condary veins usually present between pairs of sec-
ondary veins, adaxially and abaxially costa promi-
nulous, secondary veins plane to thickened, and
minor venation plane to slightly thickened; margins
thinly cartilaginous; petioles glabrous to pilosulous,
6—40 mm long; stipules glabrous to hirsutulous,
laminar, ovate in outline, 7-10 mm long, bilobed
for ca. %, lobes rounded to elliptic, often overlap-
ping laterally, ciliolate, with sinus concave to nar-
rowly so. Inflorescences erect; peduncles 2.5—4.5 cm
long; panicles pyramidal, 9-18 X 8-16 cm exclud-
ing corollas, with secondary axes 5-12 pairs, with
flowers pedicellate in cymules of 5—9; bracts nar-
rowly elliptic to ligulate, acute to obtuse, entire to
ciliolate, those subtending secondary axes 3-11
mm long, those subtending pedicels 0.8-3 mm
long; pedicels 1.5-6.5 mm long; peduncle, axes,
bracts, and pedicels moderately to densely short-
pilosulous, yellow; flowers with hypanthium turbi-
nate, glabrous, ca. 1 mm long; calyx limb glabrous,
2-2.8 mm long, divided nearly or completely to
base, lobes elliptic to lanceolate, triplinerved, mi-
nutely ciliolate, obtuse to rounded; corolla funnel-
form, yellow, strongly swollen and gibbous at base,
constricted above the swelling, bent there to ca.
90°, generally straight in tube, externally glabrous,
internally glabrous except for a narrow pilose ring
at the constriction, tube ca. 9 mm long, 1-1.5 mm
diam. near middle, lobes deltoid, 1.5-1.8 mm long,
acute; anthers in long-styled form ca. 1.8 mm long,
included, positioned in upper % of tube; mature
stigmas not seen; disk ca. 0.5 mm high. Infructes-
cences similar to inflorescences; fruit ellipsoid to
usually obovoid, laterally somewhat flattened, ca. 5
X 6 mm; mature pyrenes not seen, perhaps with 3-
5 somewhat sharp longitudinal ridges. Wet forest at
1 1800 m, northwestern Ecuador and adjacent
Colombia. [Subg. Montanae, Sect. Obovoideae, Ser.
6.]
This species is distinguished by its laminar stip-
ules with rounded relatively short lobes, yellow in-
florescences and flowers, relatively well-developed
and deeply divided calyx limb, strongly gibbous
and bent corollas, and usually obovoid, laterally
somewhat flattened fruits. It is similar to Palicourea
gibbosa Dwyer, which has stipules divided for half
or more of their length with the lobes acute and
calyx limb 0.8-1 mm long; to P. lyristipula, which
has stipules usually divided for half or more of their
length with the lobes acute and calyx limbs 1–1.8
mm long; and to P. lugoana, which has calyx limbs
0.3-0.9 mm long. The specific epithet honors
Swedish botanist Erik Asplund (1888-1974), who
first collected this species and who documented the
Ecuadorian flora with excellent specimens.
Paratypes. COLOMBIA. Nariño: municipio Barba-
coas, corregimiento El Divise, de Benavides 1577 (РЗО);
municipio Ricaurte, camino los Cruces-Curcuel, Ramírez
et al. 8709 (PSO). ECUADOR. Pichincha: Chiriboga, on
road from Quito to Santo Domingo de los Colorados, As-
plund 8672 (MO, S).
Palicourea azurea C. M. Taylor, sp. nov. TYPE:
Ecuador. Zamora-Chinchipe: Loja-Zamora
road at pass, 12 Feb. 1985, G. Harling « L.
Andersson 21999 (holotype, СВ; isotype,
MO-4278731). Figure 5D, E.
Haec species a congeneris inflorescentia floribusque
azureis, floribus pedicellatis sessilibusque admixtis atque
tubo corollino 15-17 mm longo extus trichomatibus azur-
eis multicellularibus vestito distinguitur.
Flowering at 1 m tall, to 4 m tall; stems mod-
erately to densely hirtellous becoming glabrescent
with age. Leaves paired; blades elliptic, 4.5-11.5 X
1.8—6.2 cm, at apex acute to shortly acuminate with
deltoid tips 5-8 mm long, at base cuneate to ob-
tuse, chartaceous to subcoriaceous, adaxially gla-
brous except hirtellous along costa, abaxially mod-
erately to densely hirtellous on costa and secondary
veins, glabrous to hirtellous on lamina; secondary
veins 9-13 pairs, generally extending to unite with
margins, with 1(2) distinct intersecondary veins
usually present between pairs of secondary veins,
adaxially venation plane, abaxially costa promi-
nent, secondary veins prominulous, and reticulated
minor venation plane to thickened; margins thinly
cartilaginous; petioles 5-12 mm long, hirtellous;
stipules glabrous to hirtellous, united around the
stem in a continuous truncate sheath 3—4 mm long,
lobes narrowly triangular, 4-5 mm long, acute, en
tire to ciliolate. Inflorescences erect, sometimes tri-
partite and appearing sessile; peduncles 2-3 cm
long; panicles pyramidal, 5-7.5 33.00
excluding corollas, with secondary axes 44 pairs,
with flowers sessile and shortly pedicellate together
in cymules of 3-7; bracts subtending secondary
axes 4-8 mm long, triangular to ligulate or lanceo-
late, acute to obtuse, ciliolate, those subtending
icels 2-4 mm long, triangular to narrowly 50,
acute, ciliolate; pedicels 0—4 mm long; peduncle,
Volume 84, Number 2
1997
Taylor 241
Conspectus of Palicourea
шш с
Figure 5.
27144.—C. ата based оп Barbour 4161. D, E, Palicourea azurea,
h.—E. Calyx and corolla, partially dissected. F, G, Palicourea jaramilloi.—F. Flowe
brane
==
Е:
А
X
LN
5 cm
ka Winnerskjold 5784.—G. Corolla at anthesis, based on Jorgensen et al. 459. A, D to ^i scale; B, C, E-G to
cale.
axes, bracts, and pedicels blue, moderately to
densely hirtellous; flowers distylous, frequently
pendulous, with hypanthium turbinate, glabrous,
са. 1 mm long; calyx limb glabrous to hirtellous,
1-2 mm long, lobed generally to base, lobes lan-
ceolate to triangular, acute, equal to unequal on an
individual flower; corolla tubular to somewhat fun-
nelform, blue, a little swollen at base, generally
straight there and in tube, externally sparsely to
densely pubescent with blue multicellular tri-
chomes 0.3-1 mm long, internally glabrous except
for a densely pilose ring ca. 2 mm wide at ca. Y
of length of tube above base, tube 15-17 mm long,
3-5 mm diam. near middle, lobes triangular, 2.5—
3.5 mm long, slightly thickened and hooked adax-
ially; anthers in long-styled form ca. 3 mm long,
positioned ca. 25 of length of tube above base, in
short-styled form ca. 3 mm long, partially exserted;
242
Annals of the
Missouri Botanical Garden
stigmas in long-styled form exserted, ca. 1 mm
long, in short-styled form ca. 2 mm long, positioned
ca. % of length of tube above base; disk ca. 0.8
mm high. Infructescences and fruit not seen. Wet,
cloud and elfin forest, to the edge of forest vege-
tation, at 2550-2950 m, southern Ecuador. [Subg.
ontanae, Sect. Montanae, Ser. 4, Subser. f.]
This species is distinguished by its truncate
sheathing stipules, blue inflorescences and flowers,
bracts subtending pedicels and flowers 2—4 mm
long, and corollas with tubes 15-17 mm long and
externally pubescent with multicellular blue tri-
chomes. The specific epithet refers to the strong
blue color of the flowers, even on dried specimens.
Palicourea azurea is similar to P. amethystina,
which has bracts subtending flowers and pedicels
1-2 mm long and corollas with tubes 9-15 mm long
and externally glabrous or hirtellous with colorless,
uniseriate trichomes; and to P. stipularis, which has
bracts subtending pedicels and flowers 0.5-1.5 mm
long, calyx limb 0.8-1.2 mm long, and corolla
tubes 2-2.5 mm in diameter and externally gla-
brous or pubescent with colorless, uniseriate tri-
chomes.
Paratypes. ECUADOR. Loja: Parque Nacional Po-
docarpus, new road Loja-Zamora, E of Cerro Yanococha,
03^59'S, 79°07’ W, Madsen 75572 (AAU); summit of Loja-
acotos road on Nudo de Cajanuma, 6 mi. S of Loja,
ins 10879 (F). Zamora-Chinchipe: Nudo de Saban-
illa, just E of the pass on the road to Valladolid, Harling
& Andersson 21517 (GB); limit of Parque Nacional Po-
ocarpus, around pass on road Loja-Zamora, 03^58'S,
79°07'W, Madsen & Ellemann 75983 (AAU).
Palicourea canarina C. M. Taylor, sp. nov. TYPE:
Ecuador. Zamora-Chinchipe: above Valladolid
on road to Yangana, 1 Feb. 1985, G. Harling
& L. Andersson 21411 (holotype, GB; isotype,
MO-4278929). Figure 2C, D.
Haec species a congeneris stipularum vaginis continuis
truncatis ac lobis latis brevibusque, inflorescentia pyr-
amidali atque corollae luteae tubulari-infundibuliformis
sinibus (praesertim in alabastro) basi in alas breves ali-
quam vel bene evolutis distinguitur.
Flowering at 1 m tall, to 5 m tall; stems glabrous.
Leaves paired; blades elliptic, 5.5-20.5 x 2-11 cm,
at apex acute to shortly acuminate with tips to 5
mm long, at base acute to usually cuneate or ob-
tuse, papyraceous to chartaceous, adaxially gla-
brous, abaxially glabrous to sparsely hirtellous or
sometimes moderately to densely so along costa;
secondary veins 8-12 pairs, usually extending to
unite with margins, with 1(3) sometimes weak in-
tersecondary veins usually present between pairs of
secondary veins, adaxially costa plane to slightly
canaliculate and remaining venation plane, abaxi-
ally costa prominulous to prominent, secondary
veins prominulous, and reticulated minor venation
plane; margins thinly to distinctly cartilaginous;
petioles 0.5-2 cm long, glabrous; stipules glabrous,
united around the stem in a continuous truncate
sheath 1.5—4 mm long, lobes ligulate to deltoid,
0.5-2 mm long, obtuse to rounded, entire to sparse-
ly ciliolate. Inflorescences erect, sometimes tripartite
and appearing subsessile; peduncles 1.5-7 cm
long; panicles pyramidal, 2-23 X 3-12 cm exclud-
ing corollas, with secondary axes 4-8 pairs, with
flowers pedicellate in cymules of 3—7; bracts entire
to ciliolate, those subtending secondary axes
mm long, lanceolate to elliptic or narrowly so, acute
to obtuse or rounded, entire, those subtending ped-
icels 1-3 mm long, narrowly to broadly triangular,
ovate, or elliptic, acute to obtuse or rounded; ped-
icels 1-7 mm long; peduncle, axes, bracts, and
pedicels glabrous or sparsely to densely pilosulous,
green to yellow; flowers distylous, with hypanthium
glabrous, ellipsoid, 1.2-1.5 mm long; calyx limb
glabrous, 1-1.5 mm long, divided nearly to base,
lobes triangular to deltoid, acute to obtuse, entire;
corolla tubular-funnelform, yellow, a little swollen
at base, straight there and in tube, externally gla-
brous, internally glabrous except for a pilose ring
ca. 2 mm wide at ca. 2 mm above base, tube 16-
17 mm long, 4—6 mm diam. near middle, lobes tri-
angular, ca. 3 mm long, acute, not much thickened
adaxially, in bud with sinuses at base prolonged
into wings 0.3-1.5 mm long; anthers in short-styled
orm ca. 4 mm long, partially exserted, in long-
styled form са. 3 mm long, positioned ca. % of
length of tube above base; stigmas in short-styled
form ca. 4 mm long, positioned ca. % of length of
tube above base, in long-styled form ca. 1 mm long,
exserted; disk ca. 1 mm high. Infructescences and
fruit not seen. Wet forest at 1650-2600 m, south-
eastern Ecuador. [Subg. Montanae, Sect. Montanae,
Ser. 3, Subser. c.]
This species is distinguished by its stipules with
truncate continuous sheaths and short broad lobes.
pyramidal inflorescences, and tubular-funnelform
yellow corollas with the sinuses in bud shortly to
well expanded into short wings. The specific epithet
refers to the yellow color of the flowers. This new
species is similar to Palicourea thyrsiflora, which
has usually narrower and longer stipule lobes, Ш"
florescence branches yellow to usually red ог or-
ange, and corollas usually tubular and yellow be-
coming orange or red with age, with the sinuses
between the lobes sometimes a little saccate but
not developed into wings, and the buds generally
truncate rather than rounded on the top.
Volume 84, Number 2 aylor
1997 Conspectus of Palicourea
243
Figu —A. Palicourea candida, коч branch, based on Molau & Eriksen 2149. В, С, Palicourea vulcanalis,
ка оп TENE 2699.—B. Flower.—C. Stipule. D, E, Palicourea luteonivea.—D. Flowering branch, based on Zaruma
776.—E. Calyx and corolla, partially phe peo on Neill et al. 7401. F, 6, Seige prodiga.—F. Stipule, based
on Holm-Nielsen et al. 27163.—G. Flower, based on Holm-Nielsen et al. 27250. A, D, F to 5-ст scale; С to 1-ст
scale; B, E, G to 5-mm scale.
Paratypes. ECUADOR. Loja: Nudo de Sabanilla, W Parque Nacional Podocarpus, 04°13'S, 7856" W, Madsen
slope on road to Yangana, Harling & paar] 21713 & Bloch 75879 (AAU).
(GB, MO); Nudo de Sabanilla, W slope n road to Yan- Palicourea candida C. M. Taylor, sp. nov. TYPE:
gana-Valladolid, Harling & Andersson 21825 (GB, MO).
Zam
Ecuador. Napo: E upper slopes of Cordillera
ora-Chinchipe: Nudo de Sabanilla-Valladolid, horse
trail to Caserfo ا Honda, Harling & Stáhl 26313 de Guacamayos, 11-13 km S of Cosanga on
(GB), Harling 27166 (GB); Romerillo, trail at limit of the Baeza-Tena road, 00°40’S, 77°52'W,
244
Annals of the
Missouri Botanical Garden
2100-2200 m, 22 Dec. 1987, U. Molau & B.
Eriksen 2149 (holotype, GB; isotype, AAU).
Figure 6A.
Haec species a Palicourea andrei corolla et limbo ca-
lycino elongatis distinguitur.
Flowering at 2 m tall, to 15 m tall; stems some-
what succulent, glabrous. Leaves paired; blades el-
liptic to somewhat obovate, 8-18 X 4-10 cm, at
apex acuminate with deltoid tips 3-5 mm long, at
base cuneate to somewhat obtuse, papyraceous,
adaxially glabrous or costa often puberulous, abax-
ially apparently paler, sparsely to moderately ap-
pressed-puberulous, more densely so on costa and
secondary veins; secondary veins 12-16 pairs, gen-
erally extending to unite with margins, usually with
1 well-developed and often 2—3 weak intersecon-
dary veins present between pairs of secondary
veins, adaxially costa plane to prominulous and re-
maining venation plane, abaxially costa prominent,
secondary veins plane to prominulous, and reticu-
lated minor venation inconspicuous, plane to a little
thickened; margins ciliolate; petioles glabrous, 5—
25 mm long; stipules glabrous, laminar, ovate in
outline, 12-18 mm long, overlapping laterally, bi-
lobed for ca. У, lobes ligulate to triangular, obtuse
to rounded, ciliolate, with sinus acute to narrowly
concave. /nflorescences apparently erect; peduncles
7.5 ст long; panicles pyramidal, 3-3.5 X 3.5-
5 em excluding corollas, with secondary axes 1—
2(3) pairs, with flowers sessile in glomerules of 2—
4; bracts entire, glabrous, acute to obtuse, those
subtending secondary axes 8-15 mm long, lanceo-
late to oblanceolate, those subtending glomerules
and individual flowers 3-7 mm long, lanceolate to
ovate or suborbicular; peduncle, axes, and bracts
puberulous, apparently green; flowers with hypan-
thium turbinate, 1-3 mm long, glabrescent to pu-
berulous; calyx limb glabrous, 4-6 mm long, divid-
ed nearly to base, lobes elliptic, overlapping
laterally, obtuse to rounded, entire to minutely cil-
iolate; corolla salverform to narrowly funnelform,
white, a little swollen at base, generally straight
there and perhaps also in tube, externally sparsely
to moderately puberulous, internally glabrous ex-
cept for a pilosulous to villosulous zone ca. 18 mm
long in upper part of tube and ending below stamen
attachment, tube 32-42 mm long, 2-2.5 mm diam.
near middle, lobes narrowly triangular to ligulate,
8-9 mm long, acute; anthers in apparent short-
styled form exserted, ca. 7 mm long; stigmas in
apparent short-styled form positioned ca. % of
length of tube above base, ca. 3 mm long; disk not
seen. Inflorescences and fruits not seen. Wet forest
at 1700-2200 m, east-central Ecuador. [Subg.
Montanae, Sect. Psychotrioides, Ser. 9.]
This species is distinguished by its relatively
large, laminar, bilobed stipules, flowers sessile in
glomerules, relatively long calyx limb, and relative-
ly long slender white corollas with a rather long
pubescent portion internally. The species epithet
refers to the white color of the corollas. The flowers
appear to be adapted for pollination by nocturnal
visitors such as hawkmoths, based on their white
color and relatively long tubes. Palicourea candida
is similar to P andrei, which has sweetly fragrant
flowers, calyx limbs 1-2 mm long, and corollas with
tubes 20-25 mm long and lobes 6-8 mm long. The
corolla tube appears to elongate markedly shortly
before anthesis.
Paratypes. ECUADOR. Napo: Baeza-Tena road, south-
ern slope of Cordillera de Guacamayos above Jondachi,
Harling & Andersson 16364 (GB). Pastaza: Colonia Al-
varez Miño, ca. 6 km from Mera, Lugo 828 (GB). Tun-
gurahua: Cashurco, near Río Negro, northern side of Río
Pastaza, Lugo 800 (GB).
Palicourea chignul C. M. Taylor, sp. nov. TYPE:
Ecuador. Carchi: Maldonado, Parroquia Tobar
Donoso, Reserva Etnica Awá, Sabalera,
00%55'N, 78*32'W, 900 m, 22 Nov. 1992, C.
Aulestia, E. Aulestia & M. Guanga 852 (ho-
lotype, МО-4990807; isotype, QCNE not
seen). Figure 7A.
Haec species a congeneris stipularum laminarium lob-
ulis acutis, inflorescentia | iped lata y dicellis satis
longis ac bracteis sat grandibus latisque praedita, lobulis
limbi calycini satis bene evoluti lobulis lateraliter imbri-
catis, corolla sat longa atque fructu obovoideo distinguitur.
Flowering at 2 m tall, to 5 m tall; stems glabrous.
Leaves paired; blades elliptic to broadly so or ellip-
tic-oblong to somewhat oblanceolate, 14-29.5 X 7-
18 cm, at apex shortly acuminate with deltoid to
triangular tips 3-8 mm long, at base cuneate to
obtuse, papyraceous, glabrous throughout or pu-
berulous abaxially along costa; secondary veins 10-
19 pairs, often looping to interconnect, without or
usually with 1-2 weak intersecondary veins present
between pairs of secondary veins, adaxially costa
prominulous, secondary veins plane to prominu-
lous, and minor venation plane, abaxially costa and
secondary veins prominulous and minor venation
plane to thickened; margins entire, thinly cartilag-
inous; petioles glabrous, 1.5—6.5 cm long; stipules
glabrous or pubescent to minutely sericeous €s-
pecially on distal parts, laminar, ovate in outline,
6-12 mm long, bilobed for ca. %, lobes triangular,
acute, entire, with sinus acute to concave and ob-
tuse. Inflorescences generally erect; peduncles 0.5-
Volume 84, Number 2
1997
Taylor 245
Conspectus of Palicourea
gure 7. —A. Palicourea chignul, flowering branch, based on Hoover et al. 2494. B, C, Palicourea deviae, based
on — 12337.—B. Flowering branch.—C. Flower. A, B to 5-cm scale.
2 cm long; panicles pyramidal to narrowly so, 5—
14 X 5—7 cm excluding corollas, with secondary
axes 5—10 pairs, the basalmost pair often strongly
reflexed, with flowers pedicellate in cymules of 3—
7; bracts ovate to lanceolate, elliptic or oblanceo-
late to ligulate, acute to obtuse or rounded, entire,
those subtending secondary axes 6-15 mm long,
those, subtending pedicels 4—10 mm long; pedicels
4—14 mm long; peduncle, axes, bracts, and pedicels
glabrous to puberulous, orange, red-orange, purple,
or pink; flowers with hypanthium glabrous, turbi-
nate to usually cupuliform, 2.5—4 mm long; calyx
limb glabrous, pink, orange, or purple, 3—5 mm
long, divided nearly to completely to base, lobes
subequal to somewhat unequal on an individual
flower, elliptic to suborbicular, reticulate-nerved,
usually overlapping laterally, entire, obtuse to
rounded; corolla tubular, orange to red or yellow,
often grading in color along its length, swollen and
a little gibbous at base, generally straight there and
in tube, externally glabrous to usually minutely pu-
berulous, internally glabrous except pilosulous in a
ring ca. 1.5 mm wide at ca. % of length of tube
above base, tube 20-25 mm long, ca. 2 mm diam.
near middle, lobes triangular, ca. 2 mm long, acute;
disk shorter than calyx. Infructescences similar to
inflorescences except purple to rose; fruit obovoid,
somewhat laterally flattened, 8-10 X 6-7 mm, gla-
brous, rose perhaps becoming black; pyrenes with
3-5 rather sharp longitudinal ridges. Wet forest at
900-1450 m, northwestern Ecuador to adjacent Co-
lombia. [Subg. Montanae, Sect. Obovoideae, Ser. 6.]
This species is distinguished by its laminar
stipules with acute lobes, shortly pedunculate in-
florescences with relatively long pedicels and rel-
atively large and broad bracts, well-developed
calyx limb with laterally overlapping lobes, rel-
atively long corolla, obovoid fruits, pyrenes with
sharp longitudinal ridges, and combination of or-
ange to purple or pink inflorescences with red to
yellow corollas. It is similar to Palicourea hos-
pitalis, which has longer inflorescences and
shorter calyx limbs. The specific epithet is re-
ported by several collectors to be the vernacular
local name for this species.
One collection, Drew & Wiggins 41 (F), is pro-
visionally referred here, though the material is in-
adequate for conclusive identification. It conforms
to the characters of this species except that it was
collected in Imbabura Province, Ecuador, and it
has secondary inflorescence axes, pedicels, and ca-
lyx limbs that are sparsely to moderately hirtellous
and corollas that are densely pubescent externally
246
Annals of the
Missouri Botanical Garden
with multicellular stout trichomes 0.1-0.2 mm long.
No mature corollas are present on this collection,
which may represent another population of Pali-
courea chignul with a different pubescence pattern,
or an undescribed related species.
Paratypes. COLOMBIA. Nariño: Altaquer, margen
derecha de la carretera hacia Tumaco, G. López-J. 253
(PSO). ECUADOR. Carchi: cantón Tulcán, parroquia Ch-
ical, Reserva Etnica Awá, centro Gualpí Medio, 01%02'N,
78716", Aulestia & Grijalva 1152 (MO), Праг et al. 1972
(MO); trail from Pailón to Gualpí Chico area of Awá Res-
ervation, 00%51'N, 78°16'W, Hoover et al. 2394 (MO),
2419 (MO); Gualpí Chico area, 00°58'N, 78716", Hoover
et al. 2494 (MO), 2778 (MO), 2781 (MO), 3392 (MO);
plateau above San Marcos de los Coaiqueres, on trail to-
ward Gualpí Bajo, 01%06'N, 78^17'W, @ligaard et al.
57365 (AAU), 57445 (AAU); near Maldonado, van der
Werff & Gudiño 10714 (MO).
Palicourea condorica C. M. Taylor, sp. nov.
TYPE: Ecuador. Zamora-Chinchipe: cantón
Nangaritza, río Nangaritza, Pachicutza, faldas
inferiores de la Cordillera del Cóndor,
04%07'S, 78*37'W, 5 Dec. 1990, W. Palacios
& D. Neill 6556 (holotype, MO-4990806; iso-
type. QCNE not seen). Figure 8D, E.
Haec species a congeneris foliis sat amplis, stipularum
bilobarum sat grandium lobulis lanceolatis, inflorescentia
subsessili ex pyramidali rotundata, limbo calycino brevi
atque corolla lutea tubulari distinguitur.
Flowering at 3 m tall, to 5 m tall; stems appar-
ently somewhat succulent, glabrous to puberulous
or hirtellous often becoming glabrescent with age.
Leaves paired; blades elliptic to broadly so, 8.5-
36(50) X 5.5—18(26) cm, at apex acuminate with
deltoid tips 3-10 mm long, at base cuneate to ob-
tuse and sometimes attenuate, papyraceous to char-
taceous, adaxially glabrous, abaxially puberulous,
more densely so on costa and secondary veins; sec-
ondary veins 9-17 pairs, usually extending to unite
with margins or occasionally looping to intercon-
nect in most distal part of blade, without or some-
times with 1(2) weak intersecondary veins present
between pairs of secondary veins, adaxially costa
prominulous and canaliculate and secondary veins
and reticulated minor venation plane to slightly
thickened, abaxially costa prominent, secondary
veins thinly prominulous, and minor venation plane
to thickened; margins thinly cartilaginous; petioles
2-9(14) cm long, puberulous; stipules moderately to
densely puberulous, united around the stem in a
continuous truncate sheath 2—4 mm long, lobes lan-
ceolate to ovate, 15-20 mm long, acute, rounded
laterally, margins entire, with sinus concave to
truncate. Inflorescences erect, subsessile; panicles
pyramidal, 6(20) x 9(22) cm excluding corollas,
with secondary axes 3-5 pairs, the basal pair rel-
atively long and usually reflexed, with flowers ped-
icellate in cymules of 3-7; bracts subtending sec-
ondary axes triangular to lanceolate, 5-12 mm long,
acute, those subtending pedicels 0.5-1 mm long,
triangular, acute; pedicels 2-6 mm long; axes,
branches, bracts, and pedicels glabrous or sparsely
to moderately hirtellous, yellow to red-orange; flow-
ers distylous, with hypanthium turbinate to cylin-
drical, 1-1.5 mm long, glabrous; calyx limb gla-
brous, 0.5—1 mm long, divided nearly to completely
to base, lobes triangular, entire, subequal, acute;
corolla tubular, yellow, a little swollen at base, gen-
erally straight there and in tube, externally gla-
brous, internally glabrous except for a pilose ring
mm wide at ca. 2 mm above the base, tube
ca. 12 mm long, ca. 3 mm diam. near middle, lobes
triangular, 2-4 mm long, acute, adaxially thickened
and hooked, abaxially thickened to usually some-
what appendaged, appendages to 0.5 mm long; an-
thers in long-styled form ca. 4 mm long, positioned
at ca. % of length of tube above base, in short-
styled form not seen in good condition; stigmas in
long-styled form exserted, not seen in good condi-
tion, in short-styled form ca. 4 mm long, positioned
ca. % of length of tube above base; disk ca. 1 mm
high. Infructescences and fruit not seen. Wet pre-
montane forest at 900-1700 m, southern Ecuador.
[Subg. Montanae, Sect. Cephaeloides, Ser. 11.]
This species is distinguished by its relatively
large leaves, relatively large stipules with lanceo-
late lobes, subsessile pyramidal to rounded inflo-
rescences, relatively short calyx limb, and tubular
yellow corollas. It is similar to Palicourea harlingú,
which has rounded-corymbiform inflorescences and
calyx limbs 1.5-2 mm long; and to P. macbridet of
Peru, which has purple flowers and calyx limbs 2-
3 mm long with the lobes unequal on an individual
flower. The specific epithet refers to the Cordillera
del Céndor, in the border region of Ecuador and
Peru, where the type specimen was collected.
Paratypes. ECUADOR. Morona-Santiago: banks of
the Río Ontza, Cordillera Cutucú, 02%40'S, 78°W, Camp
1182 (5); cantón Gualaquiza, Cordillera del Cóndor, Cuan-
gos, m E of Gualaquiza, 03°29’S, 78°14’ W, Gentry
80116 (MO); Plan del Milagro at cross-road between Li-
món and Indanza, Harling & Andersson 24522 (GB, MO);
30-35, van der We
& Gudiño 11261 (МО). Zamora-Chinchipe: cantón Nan-
garitza, Pachicutza, camino al Hito, 04%07'S, 78" 37W,
Palacios et al. 8287 (MO).
Palicourea corniculata C. M. Taylor, sp. nov
TYPE: Ecuador. Napo: Baeza-Tena road, Cos-
anga, 4 Feb. 1980, G. Harling & L. Andersson
16227 (holotype, GB; isotype, мо-4278911).
Figure 9C, D, Е.
Volume 84, Number 2
1997
aylor
Conspectus of Palicourea
Figure 8. A-C, Palicourea harlingii, based on На
partially dissected. D, E, Palicourea condorica,
sected. A, B, D to 5-cm scale; C, E to 5-mm scale.
rling & Andersson 16917. —А. Habit.—B. Stipule.—C. Flower,
based on Palacios et al. 8287.—D. Habit.—E. Flower, partially dis-
Annals of the
Missouri Botanical Garden
Figure 9.
Palicourea corniculata, based on Balslev & Madsen 1033
partially dissected. A, C to 5-cm scale; B, D, E to 5-mm scale.
Haec species a congeneris stipulis truncatis vaginanti-
bus, hypanthio ex hemisphaerico ovoideo subglobosove
buliformi typice inferne azurea et superne flava lobulos
icibus ben Таң 1+ hah 2 се vi
r ©
Flowering at (0.6)2 ш tall, to 8 т tall; stems
glabrous. Leaves paired; blades elliptic, 6.5-20 х
2—7 cm, at apex acute to usually acuminate with
tips 5-10 mm long and deltoid, at base acute to
cuneate, chartaceous, glabrous throughout or often
sparsely to moderately hirsute along costa on ab-
axial surface; secondary veins 8-16 pairs, usually
extending at least weakly to margins in basal part
d looping to interconnect in distal part, with 1—
2(3) intersecondary veins usually present between
pairs of secondary veins, adaxially costa slenderly
prominulous and remaining venation plane, abaxi-
ally costa prominulous, secondary veins thickened
to prominulous, and reticulated minor venation
plane to sometimes a little thickened; margins thin-
ly cartilaginous; petioles glabrous, 0.8-2 cm long;
stipules glabrous, united around the stem in a con-
tinuous truncate sheath 3-5 mm long, lobes deltoid
to narrowly triangular, 1-2 mm long, acute to ob-
tuse, entire. Inflorescences erect; peduncles 1-3 cm
long; panicles pyramidal, 6.5-13 х 9-12.5 ст ex-
cluding corollas, with secondary axes 4—8 pairs,
A, B, Palicourea cornigera, based on Harling & Stáhl 26319.—A. Flowering branch.—B. Flower. С-Е,
9.—C. Flowering branch.—D. Flower іп bud.—E. Flower,
with flowers pedicellate in lax cymules of 3-7;
bracts lanceolate to ovate or triangular, acute, еп-
tire to ciliolate, those subtending secondary axes
mm long, those subtending pedicels or borne
along their length 0.5-2 mm long; pedicels 2-7 mm
long; peduncle, axes, bracts, and pedicels red to
violet or purple, glabrous; flowers distylous, with
hypanthium hemispherical to ovoid or subglobose,
glabrous, green to yellow, 2-3 mm diam., usually
with pedicel attached obliquely; calyx limb gla-
brous, green to yellow, 1—1.5(2) mm long, divided
partially to completely to base, lobes ligulate, sub-
equal to somewhat unequal on an individual flower,
obtuse to rounded, entire to ciliolate; corolla tu:
bular to somewhat funnelform, yellow, rarely white,
or usually blue to violet in tube and yellow on lobes
and sometimes also upper part of tube, somewhat
swollen at base, straight to bent there, generally
straight in tube, externally glabrous, internally gla-
brous except for a densely pilose ring ca. 1.5 mm
wide at ca. % length of tube above base, tube 10-15
mm long, 2.5—4 mm diam. near middle, lobes tni-
angular to somewhat deltoid, 2.5-3 mm long, acute,
at apex with abaxial, carnose, deltoid appendage
1.5-2.5 mm long; anthers in long-styled form ca.
2 mm long, included, in short-styled form ca. 4 mm
Volume 84, Number 2
1997
Taylor
Conspectus of Palicourea
long, partially exserted; stigmas in long-styled form
ca. 0.8 mm long, well exserted, in short-styled form
ca. 2.5 mm long, positioned just below anthers; disk
ca. 1 mm high. Infructescences similar to inflores-
cences; fruit subglobose, 5-7 mm diam.; pyrenes
with low, rather sharp longitudinal ridges. Wet for-
est at 1800-2600 m, eastern slopes of Andean Cor-
dillera in northern Ecuador. [Subg. Montanae, Sect.
Montanae, Ser. 3, Subser. c.]
This species is distinguished by its stipules with
truncate sheaths and short lobes, pyramidal inflo-
rescences, hypanthia hemispherical to ovoid or
subglobose and relatively large, and tubular to
somewhat funnelform corollas that are typically
blue in the lower part and yellow in the upper and
have lobes with well-developed appendages. The
specific epithet refers to these appendages, which
develop on the young flower buds, giving them a
distinctive profile.
Paratypes. ECUADOR. Napo: cantón Quijos, Sierra
Azul (Agrícola Industrial Río Aragón), campamento Estero
°56'W, Alvarez et al. 530 (МО, ОСМЕ),
n
Alvarez et al. 855 (MO, QCNE), 1017 (MO, QCNE); Cor-
dillera Guacamayo, Asplund 8824 (S); Cosanga, Asplu
9565 (5); road Baeza-Napo, Cosanga, 20 km 5 of Baeza,
00°37'5, 11*52' W, Balslev & Madsen 10339 (AAU, МО);
Baeza-Tena road on southern slopes of Cordillera de Gua-
peg he Harling & пуат. 16415 (GB); slopes of Gua-
rcu, on the lom ve Río Bretania, 00°
77:45, " Holm- Nielsen r al. 26779 (AAU), 26887
(AAU); parroquia Cosanga, entre a à — 3 el río Cos-
anga, Jaramillo et al. 12048 osanga, a
km de la carretera Cosanga-El "Aliso. Jaramillo et al.
12107 (GB); ar Baeza- КО. Little & Сатригапо 158
(MO); along а River at Cosanga, 5 of focus on trail
to Tena, боде. 693 (МО); сапібп Quijos, bad n en-
N .
leading W 2 km N of Cosanga, 00°33" S, 7753 W, Persson
et al. 3
Palicourea cornigera C. M. Taylor, sp. nov. TYPE:
Ecuador. Zamora-Chinchipe: above Valladolid on
road to Yangana, 2700 m, 2 Feb. 1985, G. Har-
ling & Andersson 21465 (holotype,
MO-4278930; isotype, GB). Figure 9A, B.
Haec species a congeneri ll
1-3 mm longis externe vestita ac sinibus lobulorum ad
margines appendicibus corniformibus bene evolutis prae-
ditis distinguenda.
Flowering at 0.5 m tall, to 2 m tall; stems oc-
casionally clambering, densely pilosulous or hirtel-
lous to hirsute. Leaves paired; blades elliptic, 4—
15.5 X 1.44.5 ст, at apex acute to usually acu-
minate with tips slender, sometimes falcate, 1–1.5
cm long, at base acute to cuneate or obtuse, mem-
branaceous to papyraceous, adaxially sparsely to
moderately pilosulous or hirtellous except densely
so on costa, abaxially densely pilosulous to hirtel-
lous; secondary veins 9-17 pairs, frequently ex-
tending to unite with margins, usually with 1-2
weak intersecondary veins present between pairs of
secondary veins, adaxially venation plane, abaxi-
ally costa prominulous, secondary veins plane or
usually a little thickened, and minor venation
plane; margins thinly cartilaginous, entire to cilio-
late; petioles moderately to densely hirtellous or pi-
losulous to hirsute, 3-12 mm long; stipules
moderately to densely pilosulous or hirtellous, unit-
ed around the stem in a continuous truncate sheath
2.5-3 mm long, lobes linear, 2-5 mm long, spread-
ing-pilose to hirsute. Inflorescences perhaps nod-
ding, sometimes tripartite and apparently sessile;
peduncles 1-3 cm long; panicles subcapitate or
branched once, 1-2 X 2-2.5 cm excluding corol-
las, with flowers sessile to subsessile in glomerules
of 3—5, bracts linear, those subtending glomerules
7-9 mm long, those subtending flowers 3-8 mm
long; peduncle, axes, and bracts densely hirtellous
to hirsute with blue to purple trichomes; flowers
distylous, with hypanthium ca. 1 mm long, turbi-
nate, densely pilose with linear blue trichomes 1—
3 mm long; calyx limb green, 3.5-6 mm long, di-
vided nearly to base, lobes narrowly lanceolate, un-
equal by ca. 2596 or so on an individual flower,
acute to acuminate, glabrous to moderately pubes-
cent with blue linear trichomes 1-3 mm long; co-
rolla funnelform, white to blue or sometimes yellow
distally, a little swollen and gibbous at base, gen-
erally straight there and in tube, externally mod-
erately to densely pubescent with linear blue tri-
chomes 1-3 mm long, internally glabrous except for
a densely pilose ring ca. % of length of tube above
the base, tube 15-20 mm long, 3.5-5 mm diam.
near middle, lobes deltoid, 2-2.5 mm long, slightly
thickened at apex adaxially, with 2 linear pubes-
cent appendages 3-5 mm long near base, one on
each side; anthers in short-styled form ca. 3 mm
long, partially exserted, in long-styled form ca. 2
mm long, positioned ca. % of length of tube above
base; stigmas in long-styled form 0.5-1 mm long,
exserted, in short-styled form ca. 3 mm long, po-
sitioned near or just above tube middle; disk not
seen. Infructescences and fruit not seen. Wet forest
at 1700-2700 m, southern Ecuador. [Subg. Mon-
tanae, Sect. Montanae, Ser. 4, Subser. f.
This species is distinguished by its corollas that
are externally pubescent with blue trichomes 1–3
mm long and bear two well developed horn-like
appendages from the base on each side of the ab-
250
Annals of the
Missouri Botanical Garden
axial part of the lobes, hirtellous or pilosulous pu-
bescence throughout, linear stipule lobes and
bracts, sessile or subsessile flowers borne in one or
a few glomerules, and well-developed calyx limb
with narrow lobes. The paired appendages attached
near the base on each side of each corolla lobe are
unique in the genus and perhaps the family. The
specific epithet refers to these appendages. When
the flower is in bud, these appendages are spread-
ing and can be mistaken for narrow corolla lobes,
but the corolla lobes are broad and much shorter.
Palicourea cornigera is similar in aspect to P. ca-
lycina, which lacks corolla lobe appendages.
Paratypes. ECUADOR. Loja: Nudo de Sabanilla, north
part, Harling et al. 20520 (GB, MO). Morona-Santiago:
road General Proaño-Aschlí, Km 22, 02%15'S, 78?13'W,
: inchipe: Loja-
Zamora road, E of pass, Harling & Andersson 22108 (GB,
MO); along road Valladolid-Nudo de Sabanilla, Harling &
Madsen 25255 (GB); Nudo de Sabanilla-Valladolid, horse
trail to Caserío Quebrada Honda, Harling & Stáhl 26319
(AAU, GB)
Palicourea deviae C. M. Taylor, sp. nov. TYPE:
Ecuador. Sucumbíos: Cartagena, Km 25 from
El Carmelo on road towards La Bonita,
00*37'N, 77°30'W, 13 Apr. 1979, B. Løjtnant,
U. Molau & M. Madison 12337 (holotype,
AAU; isotype, GB). Figure 7B, C.
Haec species a congeneris lobulis stipularibus lanceo-
latis asymmetricis bene evolutis, limbo calycino 1.5-1.8
mm longo atque corolla ex azurea purpurea distinguitur.
Flowering at 3 m tall; stems moderately to dense-
ly hirsutulous becoming glabrescent with age.
Leaves paired; blades elliptic to oblanceolate, 7-17
X 2.5-5.8 cm, at apex acuminate with tips 1-2 cm
long, at base cuneate to obtuse, chartaceous to sub-
coriaceous, adaxially glabrous except hirtellous
along costa, abaxially moderately to densely hirtel-
lous throughout; secondary veins 13-14 pairs, usu-
ally extending to unite with margins at least in dis-
tal part, with 1(3) weak intersecondary veins
usually present between pairs of secondary veins,
adaxially with costa plane to slightly sulcate and
secondary veins and reticulated minor venation im-
pressed, abaxially with costa prominent, secondary
veins prominent to prominulous, and minor vena-
tion prominulous to thickened; margins thinly car-
tilaginous, often somewhat revolute; petioles mod-
erately to densely hirsutulous, 0.8-2 cm long;
stipules moderately to densely hirsutulous or pu-
berulous, united around the stem in a continuous
sheath 5-7 mm long, with interpetiolar free portion
5-14 mm long, divided for ca. % to completely,
lobes lanceolate, laterally expanded to auriculate at
base, mm wide, acute, entire to slightly
erose, with sinus acute to narrowly concave. Inflo-
rescences erect to somewhat deflexed; peduncles
1.5—6.5 cm long; panicles pyramidal, 8-14 X 6-
12 cm excluding corollas, with secondary axes 10-
12 pairs, with flowers pedicellate in cymules of 3-
7; bracts narrowly triangular, acute, those subtend-
ing secondary axes 4— mm long, those
subtending pedicels 1.5-3 mm long; pedicels 1-6
mm long; peduncle, axes, bracts, and pedicels mod-
erately to densely hirsutulous, purple to magenta;
flowers with hypanthium 1-1.5 mm long, turbinate,
moderately to densely pilosulous; calyx limb pilo-
sulous, 1.5—1.8 mm long, divided nearly to com-
pletely to base, lobes lanceolate to elliptic or ovate,
usually slightly unequal on an individual flower,
obtuse to acute; corolla tubular, blue to purple, a
little swollen and gibbous at base, generally straight
there and in tube, externally glabrous or minutely
puberulous on lobes, internally glabrous except for
a pilose ring ca. 1 mm wide at ca. % of length of
tube above base, tube 13-14 mm long, ca. 3.54
mm diam. near middle, lobes deltoid, ca. 2 mm
long, acute, adaxially and abaxially a little thick-
ened at apex; anthers in short-styled form ca. 4 mm
long, partially exserted; stigmas in short-styled form
ca. 3 mm long, positioned just above middle of co-
rolla tube; disk ca. 1 mm high. Infructescences sim-
ilar to inflorescences; fruit subglobose to ellipsoid,
ca. 5 X 4.5 mm, hirtellous; pyrenes with low round-
ed longitudinal ridges. Cloud forest at 2800 m, Ec-
uador and probably western Colombia. [Subg. Mon-
tanae, Sect. Montanae, Ser. 4, Subser. e.]
This species is distinguished by its well-devel-
oped stipules with a truncate continuous sheath an
lanceolate, laterally expanded lobes, chartaceous to
subcoriaceous leaves with the margins typically a
little revolute, pyramidal purple inflorescences, са-
lyx limb 1.5-1.8 mm long and divided nearly or
completely to the base, tubular purple corollas, and
subglobose to ellipsoid fruits. It is similar to Pali-
courea toroi of Colombia, which has calyx limbs
.54 mm long; and to P. stipularis, which has nar-
rowly triangular stipule lobes and calyx limbs 0.9-
.2 mm long. This handsome species is named in
honor of botanist Wilson Devia of Tuluá, Colombia,
who showed it to me in the Cordillera Central in
Valle, Colombia, although only the one collection
from Ecuador has been seen. In Colombia P. deviae
flowers in the rainy season, when relatively fent
collections have been made, which may explain in
part why it is so poorly represented in herbaria.
Palicourea gentryi C. M. Taylor, sp. nov. TYPE:
Ecuador. Napo: 17 km W of Lumbaque [sic;
Lumbaqui] (70 km W of Lago Agrio), 1130 m,
Volume 84, Number 2
1997
Taylor
Conspectus of Palicourea
4 Nov. 1974, A. Gentry 12437 (holotype,
MO-2791014; isotypes, MO-2779141, MO-
3131952). Figure 4F
с species a Palicourea subalata inflorescentia mi-
nore "comgestioregoe bracteas longiores gerente, hypanthio
cupuliformi majore, limbi calycini lobulis subaequalibus
atque corolla extus villosa distinguitur.
Flowering at 2 m tall, to 4 m tall; stems glabrous.
Leaves paired; blades elliptic, 10-19 х 3.5-7.4 cm,
at apex acute to usually acuminate with tips 8-18
mm long, at base acute to somewhat tapered, pa-
pyraceous, adaxially glabrous, abaxially sparsely to
moderately hirtellous, more densely so on costa and
secondary veins; secondary veins 11-18 pairs, gen-
erally extending weakly to unite with margins, with-
out or with 1-2 weak intersecondary veins some-
times present between pairs of secondary veins,
adaxially costa thinly prominulous and remaining
venation plane to a little thickened, abaxially costa
prominent, secondary veins prominulous, and retic-
ulated minor venation plane; margins ciliolate; pet-
ioles glabrous, 5-30 mm long; stipules glabrous,
united around the stem in a continuous truncate
sheath 1.5—3 mm long, lobes narrowly triangular,
5-7.5 mm long, acute. Inflorescences perhaps erect,
sometimes tripartite and appearing sessile; pedun-
cles 2-5.5 cm long; panicles rounded-corymbiform,
1-3 X 2-7 cm excluding corollas, with secondary
axes 1-3 pairs, with flowers pedicellate in cymules
of 4—9; bracts subtending secondary axes elliptic
to lanceolate, 6-10 mm long, acute, ciliolate, those
subtending pedicels narrowly triangular to lanceo-
late, acute to acuminate, 1—4 mm long; pedicels 3—
6 mm long; peduncle, axes, bracts, and pedicels
glabrous, perhaps yellow; flowers with hypanthium
cupuliform, glabrous, 1.5-2 mm long; calyx limb
glabrous, 1-1.2 mm long, divided for %—%, lobes
deltoid, entire; corolla tubular to somewhat funnel-
form, white to yellow, a little swollen at base, gen-
erally straight there and in tube, externally mod-
erately to densely villous with linear multicellular
trichomes ca. 1 mm long, internally glabrous except
for a densely villous ring ca. 1 mm wide at ca. 4
mm above base, tube ca. 13 mm long, ca. 2 mm
diam. near base, lobes triangular, ca. 3 mm long,
acute, slightly thickened at apex; anthers in long-
styled form ca. 2.5 mm long, positioned in upper
% of tube; stigmas not seen; disk ca. 0.8 mm hi
Inflorescences and fruit not seen. Wet forest at 800-
130 m, east-central Ecuador. [Subg. Montanae,
Sect. Montanae, Ser. 3, Subser. c
This species is similar to Palicourea subalata,
from which it can be distinguished by its smaller,
more congested inflorescences with longer bracts,
longer cupuliform hypanthium, calyx limb with
lobes subequal on an individual flower, and corollas
villous externally. This species is named in honor
of American botanist Alwyn H. Gentry (1945—
1993), whose work on the floristics and diversity of
the Andean flora has contributed to our knowledge
of many Palicourea species.
Paratype. ECUADOR. Napo: de la carretera entre
Reventador y Lumbaqui, 10 km al sur, Río Tigre, 00°05'S,
77°24'W, Neill 7532 (AAU).
Palicourea harlingii C. M. Taylor, sp. nov. TYPE:
Ecuador. Pastaza: cantón Pastaza, along road
between Mera and Río Anzu, which is 11.7
km N of main plaza in Mera, on Puyo-Baños
road, 01720'5, 760-06 W, 5 Apr. 1992, T. В.
Croat 73584 (holotype, MO-4990802). Figure
8A-C.
Haec species a Palicourea condorica stipulis laminari-
bus (non vaginantibus), inflorescentia rotundato-corymbi-
formi, bracteis floralibus longioribus atque limbo calycino
5-2 mm longo distinguitur.
Flowering at 2.2 m tall, to 5 m tall; stems gla-
brous. Leaves paired; blades elliptic to broadly so,
19-27.5 X 11-19 ст, at apex shortly acuminate
with deltoid tips 5-10 mm long, at base cuneate to
obtuse and sometimes attenuate, papyraceous,
adaxially glabrous, abaxially puberulous especially
on costa and secondary veins; secondary veins 16—
18 pairs, generally extending to unite with margins,
without or with 1-2 weak intersecondary veins
present between pairs of secondary veins, adaxially
costa prominulous and canaliculate and secondary
veins and reticulated minor venation plane to
slightly thickened, abaxially costa prominent, sec-
ondary veins thinly prominulous, and minor vena-
tion plane to thickened; margins thinly cartilagi-
nous; petioles glabrous to puberulous, 5.5-11 cm
long; stipules moderately to densely puberulous,
laminar, ovate in outline, with undivided interpe-
tiolar portion 3-5 mm long, lobes 20-25 mm long,
lanceolate to ovate, rounded to subauriculate lat-
erally, acute to acuminate, entire, with sinus con-
cave to truncate. Inflorescences erect, subsessile;
panicles rounded-corymbiform, rather congested,
3-5 4-6 ст excluding corollas, with seconda:
axes 2-3 pairs, with flowers pedicellate and sessile
together in cymules of 3-5; bracts ciliolate, those
subtending secondary axes 5-12 mm long, lanceo-
late to narrowly elliptic, acute, those subtending
pedicels 3-5 mm long, lanceolate to oblanceolate,
acute to obtuse; pedicels 0-5 mm long; axes,
bracts, and pedicels glabrous, yellow; flowers with
hypanthium 1.2-1.5 mm long, turbinate to cylin-
drical, glabrous; calyx limb glabrous, 1.5-2 mm
252
Annals of the
Missouri Botanical Garden
long, divided nearly or completely to base, lobes
triangular to ovate, acute to acuminate, ciliolate;
corolla tubular, yellow, somewhat swollen at base,
generally straight there and in tube, externally gla-
brous, internally glabrous except for a pilose ring
5-2 mm long at са. Y of length of tube above
base, tube ca. 16 mm long, ca. 3 mm diam. near
middle, lobes 1.5-2 mm long, triangular, adaxially
thickened and hooked, abaxially with appendages
0.5-1 mm long; anthers in long-styled form ca. 5
mm long, positioned ca. % of length of tube above
base; stigmas in long-styled form exserted, ca. 1
mm long; disk ca. 1 mm long. Infructescences and
fruit not seen. Wet forest at 1200-1380 m, north-
eastern Ecuador. [Subg. Montanae, Sect. Cephae-
loides, Ser. 11.]
This species is distinguished by its relatively
large leaves, relatively long, bilobed, laminar stip-
ules, subsessile corymbiform-rounded inflores-
cences, relatively well-developed floral bracts and
calyx limb, and tubular yellow corollas with the
lobes abaxially appendaged, especially in bud. It is
similar to Palicourea condorica, which can be dis-
tinguished by its stipules with continuous truncate
sheaths, pyramidal inflorescences, and shorter flo-
ral bracts, calyx limb, and abaxial thickenings on
the corolla lobes. This handsome species is named
in honor of Swedish botanist Gunnar Harling,
whose extensive work has contributed significantly
to our scientific knowledge of the Ecuadorian flora.
Paratype. ECUADOR. Pastaza: Colonia Játiva, ca. 7
km N of Mera, Harling & Andersson 16917 (СВ, MO).
Palicourea jaramilloi C. M. Taylor, sp. nov.
E: Ecuador. Loja: Loja-Saraguro Km 18,
03°54'08"5, 79*14'54"W, 21 Арг. 1994, P. M.
Jorgensen, C. Ulloa, H. Vargas & G. Abendaño
459 (holotype, MO-5006718; isotypes, LOJA
not seen, QCA not seen, QCNE not seen). Fig-
ure 5F, G.
Haec species a Palicourea azurea corollae luteae tubo
ca. 7 mm longo distinguitur.
Flowering at 1 m or perhaps less, to 2 m tall;
stems weak to scrambling, densely hirtellous.
Leaves paired; blades elliptic to rather narrowly so,
5-14.5 X 2-5.5 cm, at apex acute to usually acu-
minate with tips 3-8 mm long, at base cuneate to
obtuse, subcoriaceous, adaxially nitid and glabrous
or sparsely hirtellous along costa, abaxially mod-
erately to densely hirtellous; secondary veins 8-13
pairs, extending to unite with margins or often loop-
ing to interconnect with each other and margin in
a closely set reticulum, with 1–2(3) weak interse-
condary veins usually present between pairs of sec-
ondary veins, adaxially costa thinly prominulous
and secondary veins and reticulated minor venation
plane to thickened, abaxially costa prominent, sec-
ondary veins prominulous, and minor venation
thickened; margins cartilaginous, often rather
thickly so, entire or ciliolate; petioles hirtellous, 5-
15 mm long; stipules moderately to densely hirtel-
lous, often becoming glabrescent and indurate with
age, united around the stem in a continuous trun-
cate sheath 4—6.5 mm long, strongly quadrate, usu-
ally somewhat costate on angles, lobes narrowly tri-
angular, 5-8 mm long, acute, with sinus truncate.
Inflorescences erect, often tripartite and apparently
sessile; peduncles 2-4 cm long; panicles pyrami-
14 X 5-12 cm excluding corollas, sec-
ondary axes 5-12 pairs, with flowers subsessile to
shortly pedicellate together in cymules of 5-11;
bracts narrowly triangular to deltoid or ovate, acute,
those subtending secondary axes 3-8 mm long,
those subtending pedicels 1-3 mm long; pedicels
.5-4 mm long; peduncle, axes, bracts, and pedi-
cels moderately to densely hirtellous, yellow or
greenish yellow; flowers with hypanthium hirtellous,
turbinate to cylindrical, 1-1.5 mm long; calyx limb
hirtellous, 1-1.2 mm long, divided for ca. %, lobes
deltoid; corolla tubular to somewhat funnelform,
yellow or greenish yellow to red, a little swollen at
base, generally straight there, straight to curved in
tube, externally moderately to densely hirtellous
with trichomes to 0.5 mm long, in bud more densely
so on lower part of tube, internally glabrous except
for a villous ring ca. 1 mm wide at ca. % of len
of tube above base, tube ca. 7 mm long, 1.5-2 mm
diam. near middle, lobes deltoid, ca. 1.5 mm long,
at apex somewhat thickened abaxially; anthers in
short-styled form ca. 2 mm long, partially exserted;
stigmas in short-styled form ca. 2 mm long, pos!
tioned at ca. % of length of tube above base; disk
ca. 0.8 mm high. Infructescences similar to inflores-
cences; fruit ellipsoid, often somewhat flattened lat-
erally, ca. 5 X 4 mm; pyrenes with 3-5 low rounde
longitudinal ridges. Wet forest at 2600-3400 m,
southern Ecuador. [Subg. Montanae, Sect. Montan-
ae, Ser. 4, Subser. f.]
This species is distinguished by its dense hir-
tellous pubescence, stipules with truncate, contin
uous sheaths that are usually costate on the angles,
subcoriaceous leaves, yellow inflorescences and
flowers, rather long floral bracts, corollas with tubes
ca. 7 mm long and hirtellous externally, and ellip-
soid laterally flattened fruits. It is similar to Pali-
courea azurea, which has blue to purple corollas
with tubes 15-16 mm long. The well-developed
stipule sheaths with costate angles are similar 10
Volume 84, Number 2
1997
Taylor 253
Conspectus of Palicourea
those of P. flavescens. This species is named in hon-
or of Ecuadorian botanist Jaime Jaramillo, who has
collected extensively in Ecuador and contributed
significantly to our knowledge of the flora of this
country.
Paratypes. ECUADOR. Loja: carretera Loja-Zamora,
desde el Km 16 al A Jaramillo & Winnerskjold 5784
(AAU); Loja-Saraguro, Km 58, turnoff towards Fierro
Отсо, Km 1-2, 03°41' 198, 19?16'22"W, riis a et al.
483 (MO); road Pichig-Fierro Urcu, Km 11 (ca. 15 Km
03°41'5, 79?20'W, Madsen 85448
W of Loja, along road past University
towards La es van de We rff & Palacios 90 т (МО).
Zamora-Chine Loja-Zamora, Km 14, 0475,
79°90’ W, Deli е яучы а al. 3959 (AAU, 5).
Palicourea lugoana C. M. Taylor, sp. nov. TYPE:
Ecuador. Pastaza: Mera, 4 Mar. 1940, M. Lugo
14 (holotype, S). Figure 4E.
с species а congeneris folii costa prominente adax-
ee canaliculata, lobulis stipularibus e ex obtusis. rotun:
datis, limbo calycino 0.3-0.9 mm long
valde gibbosa curvataque distinguitur.
Flowering at 1.3 m tall, to 6 m tall; stems gla-
brous. Leaves paired; blades elliptic to elliptic-ob-
long, 7.5-23 X 4-12 cm, at apex acute to shortly
acuminate with deltoid tips 3-5 mm long, at base
cuneate to obtuse, papyraceous, glabrous on both
surfaces; secondary veins 9-17 pairs, spreading,
broadly curved, usually extending to unite with
margins at least in distal part of blade, with 1–2(3)
intersecondary veins usually present between pairs
of secondary veins, adaxially costa prominulous
and narrowly to broadly canaliculate, secondary
veins thinly prominulous, and minor venation retic-
ulated and plane, abaxially costa prominulous to
prominent, secondary veins plane to thickened, and
minor venation plane; margins thinly cartilaginous,
entire; petioles 1.5-3.5 cm long; stipules glabrous,
laminar, ovate to ligulate in outline, laterally round-
ed and overlapping, 5-10 mm long, bilobed for ca.
142, lobes ligulate to triangular, obtuse to round-
ed, with sinus concave, minutely ciliolate. Inflores-
cences erect, sometimes tripartite and appearin
sessile; peduncles 2.5-4.5 cm long; panicles
pyramidal, 10.5-21 X 11-25 cm excluding corol-
las, with secondary axes 8-12 pairs, the basal pair
typically 1.5-2 times as long as the next pair, with
flowers pedicellate in cymules of 3-7; bracts nar-
rowly triangular, acute, those subtending secondary
axes 2-6 mm long, those subtending pedicels 1-2
mm long; pedicels (2)4-8 mm long; peduncle, axes,
bracts, and pedicels moderately puberulous to
short-pilosulous, yellow; flowers distylous, with hy-
panthium turbinate, slender, puberulous to gla-
brous, са. 1 mm long; calyx limb glabrous to
puberulous, 0.3-0.9 mm long, divided nearly to
completely to base, lobes slightly to strongly un-
equal on an individual flower, lanceolate to trian-
lar, acute to obtuse, entire; corolla funnelform,
yellow, strongly gibbous and swollen at base,
strongly constricted and bent to 90° just above this,
generally straight in tube, externally glabrous, in-
ternally glabrous except for a narrow pilose ring at
the constriction, tube 8-10 mm long, ca. 2 mm
diam. near middle, lobes deltoid to triangular, 2—
2.5 mm long, acute; anthers in long-styled form ca.
1.3 mm long, included, positioned in upper % of
tube, in short-styled form partially exserted, ca. 1.3
mm long; stigmas in long-styled form ca. 0.8 mm
long, exserted, in short-styled form ca. 1.5 mm
long, positioned near middle of tube; disk ca. 0.8
mm long. Infructescences similar to inflorescences;
fruit obovoid, somewhat flattened laterally, ca. 5 Х
4.5 mm, glabrous; pyrenes with 3-5 low rounded
longitudinal ridges. Wet forest at 1160-1380 m in
central Ecuador, to 2000 m in northwestern Colom-
bia. [Subg. Montanae, Sect. Obovoideae, Ser. 6.]
This species is distinguished by its leaves adax-
ially with secondary veins prominulous and costa
prominulous to prominent and canaliculate, lami-
nar stipules that are divided for ca. 4—% of their
interpetiolar length with lobes obtuse to rounded,
pyramidal yellow inflorescences, calyx limb 0.3—
0.9 mm long and lobed nearly or completely to
base, corollas strongly gibbous at base and bent
and constricted just above this, and obovoid fruits.
It is similar to Palicourea gibbosa, which has stip-
ules that are divided for % or more of their inter-
petiolar length and with acute lobes, leaves with
generally longer and more slender tips and the cos-
ta not canaliculate adaxially, generally smaller in-
florescences with shorter peduncles, and longer ca-
lyx limb; and to P. asplundii, which has calyx limbs
2-2.8 mm long. The specific epithet honors Ecua-
dorian plant collectors Manuel and Holguer Lugo,
who have helped to document much of the Ecua-
dorian flora with excellent specimens.
Paratypes. COLOMBIA. Antioquia: municipio Fron-
tino, region of Murrí, ca. 13 road-km from Nutibara,
06°40'N, 76720'W, McPherson 13368 о). ECUADOR.
Pastaza: cantón Pastaza, along Puyo an
Baños, ca. 3 km W of Mera at second boe. Croat 49714
(MO); along road between Mera and Río Anzu, 11.7 km
N of main plaza in Mera, 01720' W, 78706" W, Croat 73584
= Mera, ^ у ass: Harling 3189 (5); Mera, Harling
95 (5).
Palicourea luteonivea С. М. Taylor, sp. nov.
PE: Ecuador. Napo: Puerto Misahuallí, 8
km río abajo, margen derecha del río Napo,
Reserva Florística Jatún Sacha, 01%04'S,
254
Annals of the
Missouri Botanical Garden
77°36'W, 2 Oct. 1986, J. Zaruma 776 (holo-
type, MO-3596709; isotypes, AAU, GB,
QAME). Figure 6D, E.
Haec species a Palicourea thyrsiflora inflorescentiae ra-
mis luteis atque corolla alba distinguitur.
Flowering at 3 m tall, to 6 m tall; stems glabrous.
Leaves paired; blades elliptic to elliptic-oblong, 10—
22 X 2.538 cm, at apex acuminate with slender
tips 1-1.5 cm long, at base cuneate to obtuse or
somewhat rounded, papyraceous, adaxially gla-
brous, abaxially puberulous especially on costa and
secondary veins; secondary veins 9-12 pairs, usu-
ally looping widely to interconnect near margins,
with 1-3 intersecondary veins usually present be-
tween pairs of secondary veins, adaxially costa
prominulous and remaining venation reticulated
and raised, abaxially costa prominulous to promi-
nent, secondary veins prominulous, and minor ve-
nation raised; margins thinly cartilaginous; petioles
1-2 cm long, glabrous; stipules glabrous, united
around the stem in a continuous truncate sheath
1.5-3 mm long, lobes deltoid to ligulate, 1-1.5 mm
long, acute to obtuse, entire. Inflorescences erect,
often tripartite and apparently sessile; peduncles
4.5-10 cm long; panicles pyramidal, 10.5-15 x
10-16 cm excluding corollas, with secondary axes
5-8 pairs, with flowers pedicellate in cymules of 3—
9; bracts triangular to narrowly so, acute, entire to
ciliolate, those subtending secondary axes 3-5 mm
long, those subtending or borne along pedicels 0.3—
1 mm long; pedicels 2-4 mm long; peduncle, axes,
bracts, and pedicels glabrous, yellow; flowers ap-
parently distylous, with hypanthium turbinate to el-
lipsoid, somewhat flattened, glabrous, ca. 1 mm
long; calyx limb glabrous, 0.5-1 mm long, divided
for %—% of its length, lobes deltoid, entire to cili-
olate; corolla tubular, white, at base swollen and
somewhat gibbous, somewhat bent there, generally
straight in tube, externally glabrous, internally gla-
brous except for a pilose ring ca. 1.5 mm wide just
above basal swelling, tube ca. 11 mm long, ca. 2.5
mm diam. near middle, lobes triangular, 2.5-3 mm
long, acute, sometimes somewhat thickened at
apex; anthers in long-styled form included, posi-
tioned at ca. % of length of tube above base, са.
2.5 mm long; stigmas in long-styled form exserted,
ca. 0.8-1 mm long; disk ca. 1 mm high. Infructesc-
ences apparently similar to inflorescences; fruit el-
lipsoid to somewhat obovoid, laterally flattened, ca.
4 X 4.5 mm; pyrenes with low angled longitudinal
ridges. Wet forest at 450-1440 m, southern Colom-
bia to Ecuador. [Subg. Montanae, Sect. Montanae,
Ser. 3, Subser. c.]
This species is distinguished by its stipules with
continuous truncate sheaths and relatively short
lobes, pyramidal yellow inflorescences, relatively
short, shallowly lobed calyx limb, and white corol-
las. It is similar to Palicourea thyrsiflora, which has
yellow to orange or red inflorescence branches and
corollas. One fruiting specimen, Jørgensen 56420,
is provisionally placed here; if this is correct, then
P. luteonivea also differs from P. thyrsiflora in its
ellipsoid to somewhat obovoid fruits with less
marked ridges. Palicourea luteonivea may represent
only a distinctive color form of P. thyrsiflora, but
the consistent color differences support its recog-
nition. The specific epithet refers to this distinctive
combination of yellow inflorescence branches and
white corollas. Palicourea luteonivea is also similar
to P. petiolaris and P. leuconeura, with similarly col-
ored flowers and inflorescences, but these latter
species can be separated by their funnelform co-
rollas that are bent strongly, to as much at 90", in
the lower part of the tube.
Paratypes. COLOMBIA. Caquetá: municipio de Flo-
rencia, carretera Florencia-Suaeza, Km 28, vereda "Las
Brisas,” 01%36'N, 75?37'W, Ramírez et al. 4984 (JAUM,
MO). ECUADOR. Morona-Santiago: eastern slopes of
the Cordillera, valley of Ríos Negro and Chupianza (on
trail from Sevilla de Oro to Méndez), Camp 829 (S); Сог-
ега Cutucú, ridge just S and W of Río ltzintza, ca.
02*40'S, 78°W, Camp 1292 (S); road Limón-La Unión,
Bomboiza, Misión Salesiana
79*08'W, Jorgensen 56420 (AAU); Nangaritza, Pachicut-
za, camino al Hito, Cordillera del Céndor, 04
78°37'W, Palacios et al. 8348 (MO, QCNE).
Palicourea prodiga Standley ex C. M. Taylor, sp.
nov. TYPE: Ecuador. Napo: cantón de Quijos,
margen derecha del Río Cosanga, 3 km arriba
del puente de Cosanga, 00°36’S, 77°52’W, 18
Oct. 1990, W. Palacios 6385 (holotype,
MO-5006739; isotype, QCNE not seen). Fig-
ure 6F, G.
Haec species a Palicourea flavescente foliis stipulisque
sat amplis, partibus omnibus pubescentia densa vestitis
atque inflorescentia floribusque luteis distinguitur.
Flowering at 2 m tall, to 4 m tall; stems densely
hirtellous to usually hirsute. Leaves paired; blades
elliptic to somewhat broadly so, 13.5-28 X 6.5-18
cm, at apex acuminate with deltoid to slender tips
8-25 mm long, at base cuneate to obtuse or round-
ed, papyraceous, hirtellous throughout, usually
Volume 84, Number 2
1997
Taylor 255
Conspectus of Palicourea
ary veins prominulous, and reticulated minor ve-
nation plane to usually thickened; margins thinly
cartilaginous, sparsely to moderately ciliate; petioles
1–3.8 cm long, hirtellous to hirsute; stipules mod-
erately to densely hirtellous to usually hirsute, unit-
ed around the stem in a continuous truncate sheath
8-22 mm long, this generally quadrate, costate on
the angles, weaker in interpetiolar portions and fre-
quently splitting there, the costate angles terminat-
ing in narrowly triangular to linear lobes 6-10 mm
long, acute. Inflorescences erect; peduncles 3.5-7
cm long; panicles pyramidal, 10-19 X 7.5-19 cm
excluding corollas, with secondary axes 8—10 pairs,
with flowers pedicellate in cymules of 5—11; bracts
triangular to narrowly so or lanceolate, obtuse to
acute, entire to ciliolate, those subtending second-
ary axes 2-16 mm long, those subtending pedicels
1-2 mm long; pedicels 1-3 mm long; peduncle,
axes, bracts, and pedicels densely pilosulous to hir-
tellous, pale green to yellow; flowers distylous, uh
hypanthium turbinate, sparsely hirtellous, ca. 1 m
ea calyx limb sparsely to moderately наос
0.8-1.2 mm long, divided for %—%, lobes triangular
to deltoid, acute, ciliolate; corolla funnelform, yel-
low, a little swollen at base, generally straight there,
straight to slightly curved in tube, externally
sparsely to moderately hirtellous, internally gla-
brous except for a pilose ring ca. 0.5 mm wide at
ca. % of length of tube above base, tube 5-6 mm
long, ca. 1.5 mm diam. near middle, lobes trian-
gular to ligulate, 2.5-3 mm long, acute; anthers in
long-styled form ca. 1.2 mm long, just included, in
short-styled form ca. 1.5 mm long, exserted; stig-
mas in long-styled form ca. 1 mm long, exserted,
in short-styled form ca. 2.5 mm long, positioned ca.
2 of length of tube above base; disk ca. 1 mm high
Infructescences similar to inflorescences except col-
or unknown; young fruit ellipsoid, perhaps some-
what flattened laterally, ca. 4.5 X 4.5 mm; pyrenes
with 3-5 low rounded longitudinal ridges. Wet for-
ests at 2100-2900 m, central Ecuador. [Subg. Mon-
tanae, Sect. Montanae, Ser. 3, Subser. d.]
This species is distinguished by its relatively
large leaves, relatively large stipules with costate
quadrate sheaths and narrow lobes, dense pubes-
cence on all parts, yellow inflorescences, and yel-
low relatively short corollas. The corollas appear to
be unusually short for Palicourea, but while those
of one specimen (Ownbey 2663) are 5—6 mm long,
the collection notes describe the tubes in the field
as “12 mm long,” indicating that the corollas of this
species shrink significantly when dried. Palicourea
prodiga is similar to P. flavescens, which has small-
er leaves and stipules, pilosulous rather than hir-
tellous or hirsute pubescence, flowers white to yel-
low becoming blue when old, and a generally
higher elevational range, 2785-3200 m. This spe-
cific epithet was proposed by Standley in sched.
but never published; it apparently refers to the un-
usually large leaves and stipules. Standley also
wrote this name on several specimens that are here
excluded from P. prodiga, in particular specimens
of a purple-flowered plant from Peru with somewhat
spathaceous stipules.
Paratypes. ECUADOR. Napo: slopes of Спарта Urcu,
on the loma above upper Río Borja, 00°28'S, 77°44’W,
Holm-Nielsen et al. 27026 (AAU); upper slopes of Guagra
rcu, 28'S, 77°44'W, Holm-Nielsen et al. 27163
(AAU), 27250 (AAU); Guagra Urcu, the pass between Río
Borja and Río Suno, 00°28'5, 77°43'W, Holm-Nielsen et
al. 27295 (AAU); upper Rfo Suno, near Guagra Urcu,
28'S, 77°42'W, Holm-Nielsen et al. 27525 (AAU),
27550 (AAU); Quijos River region below Baeza, region of
Río San Juan, 15 km NW of Chaco, Ownbey 2663 (MO).
Palicourea subalatoides C. M. Taylor, sp. nov.
TYPE: Ecuador. Pastaza: Misahuallí to Tena to
mel 16026 (holotype, MO-4990805). Figure
B.
,
Haec species a congeneris stipularum vaginis truncatis
ac lobulis bene ренин pra — inflorescentia
rotundato-corymbifo a, pedicello et limbo
calycino bene вала atque inr carnosa tubulari dis-
i bau
Flowering at 1 m tall, to 8 m tall; stems with
angles sharp to somewhat costate or shortly winged,
glabrous to sparsely hirtellous. Leaves paired;
blades elliptic, 5-22 X 2.5-8.2 cm, at apex acute
to usually acuminate with slender tips 5-15 mm
long, at base cuneate to acute and sometimes at-
tenuate, papyraceous, adaxially glabrous, abaxially
glabrous to sparsely hirtellous; secondary veins 7—
21 pairs, usually extending at least weakly to mar-
gins, with 1(3) rather weak intersecondary veins
usually present between pairs of secondary veins,
adaxially costa thinly prominulous and remaining
venation plane to a little thickened, abaxially costa
prominent, secondary veins prominulous, and less-
er venation plane to thickened; margins thinly car-
tilaginous; petioles glabrous to sparsely hirtellous,
4-10 mm long; stipules glabrous to sparsely
hirtellous, united around the stem in a continuous
truncate sheath 1.5—4 mm long, lobes triangular to
slightly lanceolate, 4-16 mm long, acute, entire.
Inflorescences erect, sometimes tripartite and ap-
parently sessile; peduncles 1-8 cm long; panicles
rounded-corymbiform, 1.5-6.5 Х 2.5-9.5 cm ex-
cluding corollas, with secondary axes 2—4 pairs, as-
256
Annals of the
Missouri Botanical Garden
cending and usually equal to or exceeding the axis,
with flowers pedicellate in cymules of 2-7; bracts
entire, those subtending primary branches 5-10
mm long, triangular to lanceolate, acute, often with
two short acute lobes at base, those subtending
pedicels 1–3.5 mm long, narrowly triangular to lan-
ceolate or oblanceolate, acute to obtuse; pedicels
3-10 mm long; peduncle, axes, bracts, and pedicels
glabrous, red to orange or yellow; flowers distylous,
with hypanthium cylindrical to cupuliform, gla-
brous, ca. 1.5 mm long; calyx limb glabrous, 2-3
mm long, divided for ca. %—% its length, lobes
ovate to elliptic or suborbicular, slightly imbricated,
entire, obtuse to rounded; corolla tubular, yellow,
slightly swollen and asymmetric at base, generally
straight there and in tube, carnose, externally gla-
brous, internally glabrous except for a pilose ring
ca. 2 mm wide at ca. % of length of tube above
base, tube ca. 15 mm long, ca. 5 mm diam. near
middle, lobes triangular to deltoid, ca. 1.5 mm long,
acute, at apex a little thickened; anthers in long-
styled form ca. 3.5 mm long, positioned ca. % of
length of tube above base, in short-styled form ca.
3.5 mm long, partially exserted; stigmas not seen;
disk ca. 1 mm high. Young infructescences similar
to inflorescences; young fruit subglobose to obo-
void, ca. 5 X 5 mm, glabrous; pyrenes with very
low, perhaps rather sharp longitudinal ridges. Wet
forest at 800-1160 m, eastern slopes of Andes in
central to southern Ecuador. [Subg. Montanae,
Sect. Montanae, Ser. 3, Subser. c.]
This species is distinguished by its stipules with
relatively well-developed, truncate sheaths and
well-developed triangular to lanceolate lobes,
rounded-corymbiform red to yellow inflorescences,
well-developed pedicels and calyx limb, and car-
nose tubular corollas. It is similar to Palicourea
subalata, which has narrowly triangular stipule
lobes 2-6.5 mm long, calyx limb 0.5-1.5 mm long
with narrowly triangular lobes that are unequal on
an individual flower, and less carnose corollas with
lobes that are thickened abaxially at apex. The spe-
cific epithet refers to this similarity. The corollas
appear to elongate in the tube markedly shortly be-
fore anthesis.
Paratypes. ECUADOR. Morona-Santiago: 2-4 km N
of Arapicos, Lugo 5955 (GB, MO); 2-6 km S of Arapicos,
Lugo 5989 (GB, MO). Napo: cantón Gonzalo Pizarro, río
Tigre, afluente del río Dashiño, de la carretera Lum
& Palacios 7666 (MO). Pastaza: Mera, Asplund 18928
(S); along road between Puyo and Diez de Agosto and
N : IE
of Diez de Agosto, 01°27'S, 77°51'W, Croat 59024 (MO);
cantón Pastaza, between Shell and Mera, 5.3 km NW of
center of Shell, along gravel road 1.1 km N of highway,
01°27'S, 78°04’ W, Croat 73527 (MO); Shell-Mera, Fager-
lind & Wibom s.n. (S); 3.5 km N of Puyo, Fagerlind &
Wibom 1193 (S); 2 km from Puyo, road to Tena, Harling
3235 (S); Mera, Allpayacu, Harling 3342 (S); Mera, Har-
ling 3728 (S); Puyo-Puerto Napo road, San José ca. 17
km NE of Puyo, Harling & Andersson 17107 (GB); ca. 18
km E of town of Rio Negro towards Puyo, Humbles 6103
MO); Puyo-Macas road, 5 km after Veracruz, 01%33'S,
TT54'W, Jorgensen € Legaard 56476 (AAU); Canelos,
Lugo 4493 (GB, MO); 10-20 km N of Canelos, Lugo 4579
(GB, MO); Puyopungo-Pomona, ca. 3 km E of Puyopungo,
Lugo 5124 (GB, MO); Hacienda San Antonio de Barón
von Humboldt, 2 km al МЕ de Mera, 01°27'S, 78°06'W,
Neill et al. 6117 (МО); Madre Tierra, between Hacienda
Zulay and Río Pastaza, 01%34'S, 78°02'W, @llgaard
99587 (AAU); 5 km al NE de Mera, carretera al río Anzu,
01°26'S, 78°06'W, Palacios et al. 171 (MO); Km 10 on
Puyo-Puerto Napo road, 01%25'S, 78°00'W, Stein 2994
(MO).
~
Palicourea subtomentosa (Ruiz & Pav.) C. M.
Taylor, comb. nov. Basionym: Psychotria sub-
tomentosa Ruiz & Pav., Fl. peruv. 2: 61, t. 210,
fig. a. 1799. Cephaélis subtomentosa (Ruiz &
9. 1825
bus nemorosis ad Chinchao, Mesapata et May-
chainio runctationes, fl. Jun et Jul,” Ruiz &
Pavón s.n. [type, MA not seen photo
(Rockefeller/F neg. 29657) МОЈ.
Palicourea subtomentosa is distinguished by its
usually relatively small vegetative parts with pilo-
sulous to hirtellous pubescence throughout, broadly
pyramidal to rounded inflorescences, and pilosu-
lous funnelform corollas with tubes 6–10 mm long.
It is similar to P. buchtienii, and these two species
were apparently confused by Standley in his orig-
inal description of that species. Palicourea buch-
tienii is distinguished from Р. subtomentosa by its
glabrescent leaves, white to yellow corollas that are
moderately pilosulous to glabrous externally with
tubes 11-14 mm long and lobes with cylindrical
abaxial appendages 0.3-0.8 mm long, and yellow
inflorescence branches; in the areas where P. buch-
tienii and P. subtomentosa both grow, P. subtomen-
tosa has blue flowers and inflorescences. Standley
(1936) treated Psychotria subtomentosa as a poorly
known species of Psychotria that was narrowly en-
demic in Peru, apparently overlooking the pilose
ring inside the poorly preserved corollas on the
specimens that he examined. Based on this feature
and supported by the colored inflorescences and
corollas, this species is here transferred to Palicou-
rea. Two subspecies of Palicourea tomentosa ate
here distinguished:
1. Inflorescences blue to purple or lavender with the
Volume 84, Number 2
1997
Taylor
y 257
Conspectus of Palicourea
internodes of the primary axis usually at least
shortly developed; corollas blue to purple or Јау-
ite with blue or purple trichomes;
eru to central Bolivia, in cloud forest at
—
—
=
zh
=
@
2
о
c
5
e
O
Ф
~
o
E
о
>
5
—
т
=
=
[87]
=
1
л
©
Andersson 21367 (holotype, МО-4278926; iso-
type, GB). Figure 2B.
Haec subspecies a subspecie typica inflorescentiae ra-
mis luteis atque corolla ex lutea alba differt; in Aequitoris
australi tantum crescit.
Flowering at 0.5 m tall, to 5 m tall; stems some-
times weak to clambering, often suffrutescent, mod-
erately to usually densely pilosulous to hirtellous.
Leaves paired; blades elliptic, 4-15 X 1.2–6 cm, at
apex acute to infrequently acuminate with tips ca.
5 mm long, at base acute to infrequently cuneate,
papyraceous, moderately to densely pilosulous
throughout, usually more densely so on costa an
secondary veins; secondary veins 5-15 pairs, usu-
ally uniting with margins or infrequently looping to
interconnect, without or sometimes with 1-2 weak
intersecondary veins present between pairs of sec-
ondary veins, adaxially venation plane or costa
slightly raised, abaxially costa and secondary veins
prominulous and minor venation plane or thick-
ened; margins thinly cartilaginous, entire; petioles
moderately to densely hirtellous or pilosulous, 3—
12 mm long; stipules moderately to densely hirtel-
lous to pilosulous, united around the stem in a con-
tinuous sheath 1.5-3 mm long, lobes narrowly tri-
angular, 1.5-3 mm long, acute, entire to slightly
erose, with sinus concave to subtruncate. Inflores-
cences erect or perhaps somewhat deflexed; pedun-
cles 2.5—4.5 cm long; panicles broadly pyramidal
to usually broadly rounded, 1.5-3 X 3-5 cm, with
secondary axes 1—3 pairs, with lower internodes of
primary axis shortly or hardly expanded and sec-
ondary axes appearing subverticillate, with flowers
sessile to shortly pedicellate together in cymules of
5—7; bracts narrowly triangular to linear, acute,
those subtending secondary axes 3-6 mm long,
those subtending flowers 0.8-2 mm long; pedicels
.5 mm long; peduncle, axes, bracts, and pedi-
cels moderately to densely pilosulous to hirtellous,
yellow; flowers distylous, with hypanthium turbinate
to cylindrical, moderately to usually densely pilo-
sulous, 0.6-1 mm long; calyx limb moderately to
densely pilosulous, 0.6-1.2 mm long, lobed for ca.
4, lobes deltoid to narrowly triangular or ovate,
acute, sometimes unequal on an individual flower;
corollas funnelform, white to yellow, slightly swol-
len at base, straight there and in tube, externally
densely pilosulous with trichomes often blue or
purple, internally glabrous except for a rather dif-
fuse villous ring ca. 2 mm wide, tube 6-10 mm
long, 1.1-1.2 mm diam. near middle, lobes trian-
gular, 1.2-2 mm long, at apex smooth or sometimes
with thickened projections to ca. 0.3 mm long abax-
ially; anthers in short-styled form ca. 3 mm long,
partially exserted; stigmas in short-styled form ca.
1.5 mm long, positioned near middle of tube, in
long-styled form ca. 0.3 mm long, exserted; disk
0.8-1 mm long. Infructescences similar to inflores-
cences or often peo internodes of primary axis
expanding; fruit ellipsoid, 5-6 X 4.5-5 mm,
laterally flattened, Sr blue; pyrenes with 3—
5 distinct longitudinal angles. Wet forest at 2210—
2700 m, southern Ecuador. [Subg. Montanae, Sect.
Montanae, Ser. 3, Subser. c.]
Plants from southern Ecuador differ from plants
of Peru and Bolivia only in this separate geographic
ange and their inflorescence development and in-
florescence and corolla colors, which are yellow
rather than blue to purple. No other features sep-
arate these two populations, nor is any gradual vari-
ation evident between them. Therefore, the plants
m Ecuador are here recognized as a separate
subspecies. The subspecific epithet commemorates
the city and province of Loja, in the region from
which this subspecies is known.
~
Paratypes. DOR. Loja: Nudo de Sabanilla,
northern me Harling et al. 20570 (GB, MO); Nudo de
ue а ern slope on road sve. o Pi i god Har-
ng & D H ba (GB); N
e Yangana on road to Valladolid,
m SE of Yangana, Harling & Andersson
23825 (GB, MO); Cerro Bangala, ca. 10 km E of Yangana,
Harling 25328 (GB). Zamora-Chinchipe: сапбп Zamo-
a, Podocarpus National Park near El Tambo, 40 km NW
of Zamora on road to Loja, 03%58'S, 79*07'W, Gentry
79947 (MO)
Palicourea ulloana C. M. Taylor, sp. nov. TYPE:
Ecuador. Morona-Santiago: 17-18 km N of
Gualaquiza on road to Indanza, 17 Apr. 1985,
G. Harling & L. Andersson 24226 (holotype,
GB; isotype, МО-4278740). Figure БА, B, C.
ies a congeneris stipularum vaginis mem-
Haec s
branaceis truncatis ac lobis cartilagineis coarctatis, inflo-
rescentia ex anguste pyramidali cylindrica, limbo calycino
Annals of the
Missouri Botanical Garden
0.8-2.2 mm longo, corollae roseae rubraeve tubo 8-13
mm longo atque fructu pyrenas quattuor continente distin-
itur.
Flowering at 0.5 m tall, to 2 m tall; stems often
weak or clambering, moderately to densely hirsu-
tulous sometimes becoming glabrescent with age.
Leaves paired, blades elliptic, 4.5-14 X 1.4—5 ст,
at apex acuminate with slender tips 0.8–1.8 cm
long, at base acute to cuneate, papyraceous, adax-
ially glabrous or sparsely hirtellous along midrib,
abaxially glabrous or moderately to densely hirtel-
lous along midrib and sometimes on secondary
veins; secondary veins 10-12 pairs, extending to
unite with margins, with 1—2(3) intersecondary
veins present between pairs of secondary veins,
adaxially costa thinly prominulous, secondary veins
thickened to prominulous, and reticulated minor
venation plane to thickened, abaxially costa prom-
inent, secondary veins prominulous, and minor ve-
nation plane to thickened; margins thinly to strong-
ly cartilaginous; petioles glabrous to hirtellous, 3—
10 mm long; stipules glabrous to hirtellous, united
around the stem in a membranaceous, truncate
sheath 1—2.5 mm long, sometimes splitting intra-
petiolarly, interpetiolarly with a cartilaginous tri-
angular portion 2-3 mm long, this costate on mar-
gins and terminating in 2 closely set lobes 2-3.5
mm long, narrowly triangular, acute, entire to usu-
ally ciliolate. Inflorescences deflexed to pendulous;
peduncles 3-9 cm long; panicles narrowly pyra-
midal to cylindrical, 3.5-10.5 х 2-3 cm excluding
corollas, with secondary axes 4—8 pairs, not much
developed, usually terminating in 1 cymule or sol-
itary flowers, with flowers pedicellate in lax cym-
ules of 2-3; bracts triangular to narrowly so, entire
to ciliolate, acute, those subtending secondary axes
2-5 mm long, those subtending pedicels 1-3 mm
long; pedicels 3-7 mm long; peduncle, axes, bracts,
and pedicels glabrous to densely hirtellous, purple
to red; flowers distylous, with hypanthium glabrous,
са. 1 mm long, turbinate to cupuliform; calyx limb
glabrous, 0.8-2.2 mm long, divided nearly to com-
pletely to base, lobes narrowly triangular to narrow-
ly ligulate, subequal to usually strongly unequal on
an individual flower, acute, entire or ciliolate; co-
rolla tubular to somewhat funnelform, red to pink
or purple, slightly swollen at base, somewhat to
strongly bent there, straight in tube, externally gla-
brous, internally glabrous except for a pilose ring
ca. 1 mm wide at ca. % of length of tube above
base, tube 8-13 mm long, 2.5-4 mm diam. near
middle, lobes 2-3 mm long, acute, triangular, a lit-
tle thickened at apex; anthers in long-styled form
ca. 3 mm long, positioned ca. % of length of tube
above base, in short-styled form ca. 3 mm long,
partially exserted; stigmas 4, in long-styled form ca.
1 mm long and just exserted, in short-styled form
not seen; disk ca. 0.8 mm high. Infructescences sim-
ilar to inflorescences; fruit oblate, ca. 4 X 3 mm,
glabrous, blue; pyrenes 4, triangular with outer sur-
face rounded, smooth. Wet and cloud forest at
1 m, north-central Ecuador to northern
Peru. [Subg. Montanae, Sect. Montanae, Ser. 3,
Subser. c.
This species is distinguished by its leaves with
the secondary veins usually extending to the mar-
gins, which are rather thickened, its unusual stip-
ules with a triangular cartilaginous portion termi-
nating in closely set narrow lobes, narrowly
yramidal to cylindrical inflorescences with the
branches simple, calyx limb 0.8-2.2 mm long with
the lobes often strongly unequal on an individual
flower, pink to red tubular to somewhat funnelform
corollas, and oblate fruit with four pyrenes that are
smooth on the outer surface. The four pyrenes and
stigma lobes are unique in subgenus Montanae; the
unusual stipule morphology resembles that of Pal-
icourea vulcanalis. Palicourea ulloana is similar in
aspect to P. myrtifolia, which has laminar stipules
with broader lobes and corollas with tubes that are
5-9 mm long and gibbous and strongly bent at the
base; and to P. calantha, which has calyx limbs 0.8
mm long and corollas yellow. This distinctive, at-
tractive species is named in honor of Ecuadorian
botanist Carmen Ulloa Ulloa, whose work has con-
tributed significantly to our knowledge of the mon-
tane regions where P. ulloana lives.
Paratypes. ECUADOR. Morona-Santiago: road from
Limón (General Plaza) 2. Cuenca, са. 4 Кт above Plan
del Milagro, 03%00'S, 78°30-40'W, Stein 2833 (МО).
Napo: cantón Quijos, Pl Azul (Agrícola Industrial Río
Aragón), a de Tundal, 00%40'S, 77°54’ W, Alvarez el
al. 309 (Mo ‚ ОСМЕ), 339 (MO, QCNE); с
parroquía de Baeza, comunidad de Santa Lucía de Ber
mejo, i Pte S, 1155 W, Mei et at big (MO, QCNE);
slopes of Guagra Urcu, on loma above upper Río Borja,
00°28'S, 77°44’ W, Holm. Nielsen et pi 26995 (AAU); car-
retera Baeza-Tena, a 17 km de Baeza en la localida d de
gma Playa, Jaramillo et ra 12216 (GB); € river
region below Baeza, region of Río San Juan 15 k NW
of Chaco, Ownbey 2662 (MO); Archidona-Tena region, 5
W of Tena, Own-
bey 2740 (MO); сатбп Quijos, Cosanga, Hacienda Gua-
camayos, Río Cosanga, microcuenca
00*36'S, 77%51'S, Palacios & Freire 5034 (MO). Т
ahua: El Mirador, Sierra de León, valley of Río Pastaza
below Baños, Steere & Camp 8274 (F). Zamora-
chipe: Nudo de Sabanilla-Valladolid, horse-trail to o Cas
erío Quebrada Honda, Harling 27144 (СВ), ae T (9:
new road Loja-Zamora, trail to Podocarpus patch E
brada del Diablo, P. der Werff & Palacios 9241 (MO)
9240 (MO). PERU. Amazonas: Bagua province, са
km (by trail) E of ha Peca, еы 2818 (МО); Bagua
Volume 84, Number 2
1997
Taylor
Conspectus of Palicourea
259
province, Cordillera Colán SE of La Реса, Barbour 4161
(MO).
Palicourea vulcanalis Standl. ex C. M. Taylor, sp.
nov. TYPE: Ecuador. Napo: Cordillera de Gua-
camayos above Urcusiqui on trail between
Baeza and Tena, 6000 ft., 11 Mar. 1944, M.
Ownbey 2699 (holotype, F-1150284; isotype,
MO-1600100). Figure 6B, C
Haec species a congeneris stipularum vaginis bene ev-
olutis ac lobulis coarctatis anguste lanceolatis 12-15 mm
une unoquoque glandulam ad basim gerente distingui-
Shrubs, height not recorded; stems glabrous or
puberulous becoming glabrescent with age. Leaves
paired; blades elliptic, 11.5-20 X 3-7.8 cm, at
apex acuminate with slender tips 1.5-2 cm long, at
base cuneate to acute, papyraceous, adaxially gla-
brous, abaxially puberulous throughout but more
densely so on costa and secondary veins; secondary
veins 10—14 pairs, generally looping to intercon-
nect, with 1—2 weak intersecondary veins usually
present between pairs of secondary veins, adaxially
costa thickened to prominulous and lesser reticu-
lated venation thickened, abaxially costa promi-
nent, secondary veins prominulous, and minor ve-
nation thickened; margins thinly cartilaginous,
entire; petioles glabrous to puberulous, 7-15 mm
long; stipules glabrous to puberulous, united around
the stem in a truncate continuous sheath 7-9 mm
long, lobes narrowly to very narrowly lanceolate,
12-15 mm long, 1-2 mm wide near base, finely
nerved, acute to acuminate, entire, closely set, each
subtended at the leaf-side base by a gland 0.5-1.5
mm long, with sinus acute. Inflorescences apparently
ascending or perhaps deflexed or with peduncle
bent at base; peduncles 2-8 cm long; panicles py-
ramidal, 5-10.5 X 14-18 cm excluding corollas,
with primary axis apparently sometimes flexuous,
secondary axes 10-15 pairs, the basalmost fre-
quently reflexed and ca. twice as long as next pair,
with flowers pedicellate in cymules of 5-9; bracts
narrowly triangular to linear, acute, those subtend-
ing secondary axes 6-18 mm long, those subtend-
ing flowers 3-6 mm long; pedicels 3-6 mm long;
peduncle, axes, bracts, and pedicels puberulous,
red; flowers with hypanthium glabrous, cylindrical,
1.2-1.5 mm long; calyx limb glabrous, 0.8-1.2 mm
long, divided for % to most of its length, lobes lan-
ceolate to ovate, obtuse to acute, often somewhat
unequal on an individual flower; corolla funnel-
form, white, a little swollen at base, generally
straight there and in tube, externally glabrous, in-
ternally glabrous except for a villous ring ca. 1 mm
wide at ca. % of length of tube above base, tube
ca. 8 mm long, ca. 1.2 mm diam. near middle, lobes
triangular to deltoid, 1-1.2 mm long, acute; anthers
in short-styled form ca. 1.2 mm long, en in
throat, included or partially exserted; stigmas
short-styled form ca. 1.5 mm long, ы i ca.
24 of length of tube above base; disk ca. 0.5 mm
high. Infructescences similar to inflorescences ex-
cept violet; fruit ovoid, laterally flattened, ca. 5 X
4.5 mm, glabrous; pyrenes with 3-5 smooth angles.
Wet forest at 1800-2000 m, east-central Ecuador.
[Subg. Montanae, Sect. Montanae, Ser. 3, Subser.
This species is distinguished by its unusual stip-
ule morphology, with the truncate sheath well de-
veloped and the lobes closely set, relatively long
and narrow, finely nerved, and each subtended on
the leaf side by a gland to as much as 1.5 mm long.
This stipule morphology is similar in general aspect
to that of Palicourea toroi and P. deviae, but in both
of these latter species the stipules are laminar and
the lobes lack well-developed glands at the base.
Palicourea vulcanalis is also distinguished by its
relatively long inflorescences with the lowermost
branches usually reflexed and about twice as long
as the next pair, relatively long narrow bracts, white
corollas with tubes ca. 8 mm long, ovoid laterally
flattened fruits, and pyrenes with smooth angles
rather than longitudinal ridges. This epithet was
applied by Standley to the specimen designated
here as the type, but never published; it apparently
refers to the habitat of this species, in highlands of
volcanic origin.
Paratype. ECUADOR. Napo: cantón El здр mar-
gen derecha del rio Quijos, Finca ve ” de Se-
gundo Pacheco, 00°12'5, 77°39’ W, Palacios 5395 (MO).
INDEX TO SPECIES CLASSIFIED
ras
лаа (*) indicate names published in this work,
above.
Palicourea abbreviata Rusby II, H, 8
. M. Taylor II, 1, T
m II, E, 3
Palicourea affinis Standl. 1, C, 2
Palicourea albert-smithii Standl. Il, E, 4, e
Palicourea albocaerulea C. M. Taylor Il, E, 4, f
Palicourea alpina (Sw.) "€ т Е, 3, с
Palicourea amapaensis Ste LB
Palicourea amethystina (Ruiz & Pav.) DC. П, E, 4, e
Palicourea amplissima (Standl. ex Steyerm.) C. M. Taylor
ЕТ
Palicourea — Standl. П, E, 7
Palicourea anderssoniana C. M. к” П, E, 3, c
Palicourea andrei Standl. Il, Н. 9
260
Annals of the
Missouri Botanical Garden
Palicourea angustifolia Kunth H, i
Palicourea anianguana C. M. Tay
*LA
[orc anisoloba (Müll. "eed nif & M. T. Campos
Med antioquiana Standl. II, E. 4, f
Palicourea apicata Kunth IL, F, 7
Palicourea aschersonianoides (Wernham) Steyerm. II, H, 8
Palicourea asplundii C. M. Taylor* ЊЕ 6
Palicourea attenuata Rusby I, А
alicourea azurea C. M. Taylor* II, E, 4, f
Palicourea bella (Standl.) Dwyer II, I, 10
Palicourea bellula C. M. Taylor П, I, 10
Palicourea bo
Palicourea chimboracensis Standl. IL, E, 4, e
Palicourea chiriquina Standl. II, F, 5
Palicourea condorica С. M. Taylor* И, I, 11
Palicourea conferta (Benth.) Rana ECEB
Pa
licourea coriacea Mart. I, a
Palicourea ge (Standl. ex Steyerm.) C. M. Tay-
lor П, I, 11
Palicourea guianensis Aubl. I, A
Palicourea herzogii Standl. I, C, 1, a
Palicourea heterantha Standl. Ц, E, 3, c
Palicourea heterochroma K. Schum. & K. Krause II, E, 4, f
Palicourea holmgrenii Standl. П, E, 3, c
Palicourea hospitalis Standl. II, F, 6
Palicourea huberi Steyerm. Í, C, 1, a
Palicourea ionantha Standl. II, E, 3, с
i C. M. ul Il, E, 4, f
Palicourea killipii Standl. IL, 1, 10
Palicourea lachnant tandl.
„А
Palicourea lancifera Standl. & E O. Williams II, E, 4, f
Sm. Il, Е, E с
Palicourea lobbii Standl. II, E, 3, c
Palicourea longepedunculata Cardner LG,2
Palicourea longiflora (Aubl.) A. Rich. I, C, 1
Palicourea longistipulata (Müll. Arg.) Standl. pa С, 18
) aylor* II, F, 6
s. Il,
Palicourea macrobotrys (Ruiz & Pav) DC. I, C, 2
Palicourea macrocalyx Standl. II, E, 4,
Palicourea mansoana (Müll. Arg.) Standl. I, A
Palicourea mello-barretoi Standl. I, C, 1, a
Palicourea mexiae Standl. II, F, 7
Palicourea montivaga Standl. II, F, 5
Palicourea myrtifo lia K. тюл " K. Krause II, F, 6
Palicourea nigricans
Palicourea nitidella (Мий. ren ul LG. и
7
Palicourea pachycalyx Standl. 1, A
Palicourea padifolia (Willd. ex Roem. & Schult.) C. M.
pe
Palicourea pennellii Standl. П, E, 3, c
i rangularis Wernham Il, E, 4, €
Palicourea ponasae K. Krause II, E, 3, c
Palicourea prodiga Standl. ex C. M. Taylor* II, E. 3. d
Palicourea psittacorum Stand). ЇЇ, E, 3, c
a
Volume 84, Number 2
1997
Taylor 261
Conspectus of Palicourea
Palicourea pulchra Griseb. II, H, 8
Palicourea punicea (Ruiz & Pav.) робе ДЬ
e
quadri. ‘sae is C. M. Taylor II, E, 4, g
Palicourea rigida К nth I, A
s rigidifolia ( (Dwyer & M. V. Hayden) Dwyer II,
Palita salicifolia Standl. II, F, 7
Р
alicourea semirasa Standl. 1,
Palicourea stenosepala Stand | 11, i
ee ا
Palicourea i яғуни $0 M. aylor* Il, E, A
ndens Standl. ex "pers .H, G
ri,
heis subtomentosa (Ruiz & Pav) C. M. Taylor* II,
риба sulphurea (Ruiz & Pav.) DC. IL F, 7
alicourea tamaensis (Standl. € Steyerm.) Steyerm. II, I, 11
Palicourea tectoneura K. Schum P K. Krause Il, E, З, d
Palicourea tepuicola Steyerm
Palicourea tetraphylla Cham. & Schltdl C2
Palicourea thermydri J. H. Kirkbr. II, E, 4, g
Palicourea thyrsiflora (Ruiz & Pav.) DC. I, E, 3, c
Palicourea tilaranensis C. M. Taylor H, H, 8
Palicourea virens (Müll. Arg.) Standl. I, D
Palicourea vulcanalis Standl. ex C. M. Taylor* II, E, 3, c
Palicourea weberbaueri K. Krause II, E,
Palicourea wilesii C. D. Adams II, H, 8
Palicourea zarucchii C. M. Taylor II, H, 8
Literature Cited
Bacigalupo, N. M. 1952. Las especies argentinas de los
géneros Psychotria, [eden y Rudgea (Rubiaceae).
& C. M. Taylor. 1993. Flora costaricensis:
Family 4202. с bong Fieldiana, Bot. п. 8. 33: 1–
343,
E J. D. 1980. Flora of Panama Part IX. Family 179.
Rubiaceae. Ann. Missouri Bot. Gard. 67: 1—522.
пое 6. 1952. Pollen жан Sd and Plant Taxon-
y. Angiosperms. Chronica Bo
Gentry, A. H. 1982. Neotropical мы ке ge
togeographical connections between Cen
America, Pleistocene climatic fluctuations, or an acci-
dent of the Andean orogeny? Ann. Missouri :
69: 557-593.
1988. Changes i in plant conum. imd иза
аан io
nviranment
о
gradients. hn Missouri Bot. Gard. ts 1-34.
— —. 1995. Patterns of diversity and floristic compo-
sition in neotropical montane forests. Pp. 103-126 in
S. P. Churchill, H. Balslev, E. Forero & J. L. Luteyn
(editors), Biodiversity and Conservation of Neotropical
Montane Forests. The New York Botanical Garden,
Bronx, New York.
Grisebach, A. H. R. 1861. Flora of ei British West In-
ae Islands. Lovell Reeve, London.
ammen, T. van der. 1974. The Pleistocene changes of
vegetation and climate in tropical South America. J.
rest Š
35-50 in 5. P. Churchill, H. Balslev, E. Forero & + L
Luteyn (editors), Biodiversity and Conservation of N
tropical Montane Forests. The New York Botanical ed
den, Bronx, New ^
Kappelle, M M. & N. Zamora. 1995. Changes in woody s
cies richness along an altitudinal gradient in Tisi:
can montane Quercus forests, Costa Rica. Pp. 135-148
in S. P. Churchill, H. Balslev, E. Forero & J. L. Luteyn
(editors), Biodiversity and Conservation of Neotropical
Montane Forests. The New York Botanical Garden,
Bronx, New York.
Kiehn, M. 1986. Karyosystematische Untersuchungen
und DNA-Messungen an Rubiaceae und ihre Bedeu-
tung für die — a Familie. Ph.D. Disser-
tation, University of Vie
Liogier, H. A. & L. F. Legi odi 1982. Flora of Puerto
Rico and Adjacent Islands: A Systematic Synopsis. Ed-
re de la Universidad de Puerto Rico, San Juan.
r, J. J. 1881. Rubiaceae. In: C. Martius, Flora
gree аб 6(5): 1485. F. Fleischer.
Schumann, K. 1891. Rubiacese. In: A. Engler & K.
Prantl (editors), Die natiirlichen Pflanzenfamilien 4(4):
Enge ea ipzig.
P. A. & J. E. Ramos-Pérez. 1995.
Floristic a and phytogeography of the Cerro
del Torrá, Chocó, Colombia. Pp. 169-186 in S. P. Chur-
chill, H. Balslev, E. Forero & J. L. Luteyn (editors),
Biodiversity and Conservation of Neotropical Montane
Forests. The New York Botanical Garden, Bronx, New
York
ork.
Sobrevila, C., N. Ramfrez & N. X. de Enrech. 1983. Re-
productive. biology of работна nei and P. petio-
laris (Rubiaceae), heterostylou bs of a tropical
ela a. ا 15: 161-169,
Standley, P. C. 1936.
itor), Ene а Peru. Field Mus. Nat. Hist., Bot. Ser
13(6
Vea eng Flora of Costa Rica: Rubiaceae. Publ.
Field Mus. Nat. Hist., Bot. Ser. 18: 1264-1380.
. 1972. Psychotria. In: B. M. Maguire &
Collaborators, Flora of the Guayana Highlands. Mem.
New York Bot. Gard 406—717.
. 1974. Rubiaceae. Jn: Т. Lasser (editor), Flora de
Venezuela Б 1-2070. Instituto Botánico, Dirección de
ec aturales qus iari Ministerio de Agricul-
aracas, Vene
SR bine s of the World. Univ.
rke
, Reconsideration of the generic place-
ment of Ae урне domingensis (Rubiaceae: Psycho-
trieae). Ann. Missouri Bot. Gard. 74: 447—448.
——. ev of Palicourea hei in
Mexico and Central America. Syst. Bot. Monogr. 2
102.
262
Annals of the
Missouri Botanical Garden
——. 1990. A new species of Palicourea Me )
from ees Rica. Ann. Missouri Bot. Gard. 77: 215-216.
3. Revision of Palicourea ње Psy-
e Ар in the West Indies. Moscosoa TE 201-241.
Overview of the P:
in the Ar и Opera Bot. Belg. 7: 261-270.
кока . W. 1995. Cretaceous іо Tertiary geologic and
osperm paleobiogeographic و of the Andes.
Pp. 3-10 in S. P. Churchill, H. Balslev, E. Forero & J.
L. Luteyn (editors), Biodiversity and Conservation of
Neotropical Montane Forests. The New York Botanical
Garden, Bronx, New York
\
Webster, С. L. 1995. The panorama of neotropical cloud
forests. Pp. 53-78 in S. P. Churchill, H. Balslev, E.
Forero & J. L. Luteyn (editors), Biodiversity and Con-
servation of Neotropical Montane Forests. The New
York jean Garden, Bronx, New York
Young, K. R. 1995. Biogeographical paradigms useful for
the nagd a tropical montane forests and their biota.
n S. P. Churchill, H. Balslev, E. Forero &
LL Me (editors), Biodiversity and Conservation of
Neotropical Montane Forests. The New York Botanical
Garden, Bronx, New York
PALY NOLOGY,
PHYLOGENETIC
RECONSTRUCTION, AND
CLASSIFICATION OF THE
AFRO-MADAGASCAN GENUS
ARISTEA (IRIDACEAE)!
Peter Goldblatt? and Annick Le Thomas?
ABSTRACT
Examination of the pollen grains of Aristea using light and scanning electron microscopy shows that this Afro-
Madagascan genus of ca
sculpturing ranges ien ит cutis to rugulate to areolate with lumina ranging from
variable, with grains ranging from medium-sized (45-60
. 90 species is one of the palynologically most diverse genera of the family Iridaceae. Exine
large to geve Overall size is also
.5
pm equatorial diameter) to large (7 pm poete
diameter). Monosulcate apertures are plesiomorphic, and би sulculate (apertural membranes obscured
of exine), zonasulculate, 2-zonasulculate, 3-sulcate, and spiral apertures are present in various species. Pollen d
acters were combined with morphological and anatomical features in a matrix and subjected [о cladistic analysis. Trees
obtained using successive weighting procedures were used as the basis for a revised i f Ariste
in which three subgenera are recognized. Two subgenera are restricted to the winter-rainfall zone (the Саре Нопвис
Cape Region. Apomorphic character states are discussed in
nd Madagascar but is barely
relation to the biology of the genus, =
we suggest that some derived aperture types may be adapted to the desiccating climate of the Cape Region
Information now available on various aspects of
the pollen morphology of the Afro-Madagascan ge-
nus Aristea indicates that it is among the most vari-
able genera of Iridaceae (if not the most variable)
as regards pollen-grain apertures and exine sculp-
turing. The first re of pollen morphology in
Aristea (Radelescu, 1970; Schulze, 1971a), using
light microscopy, made it clear that the genus was
unusual palynologically. More detailed study using
scanning electron microscopy (SEM) of six species
occurring on Madagascar (Straka & Friedrich,
1984; Goldblatt & Le Thomas, 1992a) confirmed
the apertural diversity, even in this limited geo-
graphical area and with the low species diversity
for the genus. With some 50 species (Weimarck,
1940; Vincent, 1985; Goldblatt, 1995a, 1995b;
Goldblatt & Manning, 1997), Aristea is the largest
of the six genera of Nivenioideae, one of four sub-
families currently recognized in Iridaceae (Gold-
blatt, 1990, 1993). Aristea is unusual in Iridaceae
in having a particularly wide geographic range. It
is one of only a handful of the 35 African genera
of the family that extend across all of sub-Saharan
Africa, and one of only three that are shared with
Madagascar, the others being Crocosmia (9 spp: 1
in Madagascar) and Gladiolus (ca. 255 spp: 8 in
Madagascar), both members of subfamily Ixioideae.
Aristea is also among the most variable genera of
Iridaceae as regards fruit and seed morphology. In
this paper we extend our earlier palynological sur-
vey of Aristea to the African species and combine
the variation in pollen-grain characters with fea-
tures of fruit, seeds, and other morphological struc-
tures in a cladistic analysis in order to increase our
understanding of the phylogeny of the genus. The
results provide valuable insights into the evolution
and geographic radiation of the genus and are used
to refine the infrageneric classification.
MATERIALS AND METHODS
PALYNOLOGY
Pollen samples (Table 1) of 32 of the estimated
50 species of Aristea were collected in the wild,
! This study was s
ta d ith the SEM, and N. d'Amico (E.P.
gn Loir yu eed Mg mes National d'Histoire de for the use of their facilities, and James A. Doyle
their rivis with the phylogenetic analysis
re de Phanéro,
and fue € C. Hoch for
1* B. A.
upported in part by E.P.H.E. We утах with gratitude D. Guillaumin (C.N.R.S.) for her
ог preparation of the material. We also thank the
Krukoff Curator of African Botany, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166,
S.A
U.S.A.
* E.P.H.E., Laboratoire de Phanérogamie, Muséum National d'Histoire Naturelle, 16 rue Buffon, Paris 75005, France.
ANN. MISSOURI Bor. GARD. 84: 263-284. 1997.
264 Annals of the
Missouri Botanical Garden
Table 1. Species of Aristea examined, with voucher data and pollen grain dimensions. Voucher specimens are
located at MO, with additional duplicates often at NBG, P, and PRE (herbarium acronyms after Holmgren et al. (1990));
collectors are abbreviated as follows G = Goldblatt, M = Manning. Species are arranged alphabetically within the
sections recognized by Weimarck (1940). Aristea fimbriata, unknown to Weimarck, is assigned to section Racemosae,
where he would presumably have placed it.
Taxon Grain size, jum Voucher information
Section Singulares Weim.
A. singularis Weim. 59.1 х 51.0 S. Africa, W. Cape, Taylor 11009 (MO)
Section Eucapsulares Goldblatt (— sect. Euaristea Weim.)
A. angolensis Baker 60.0 x 58.5 S. Africa, Natal, G & M 8360; 9818; 9870
А. grandis Weim. 54.3 X 51.6 S. Africa, Natal, G & M 9857
А. montana Baker 60.0 х 57.0 S. Africa, Natal, G & M 9858
A. nyikensis Baker 61.0 x 51.0 Malawi, Bidgood et al. 1310
A. polycephala Harms 51.0 X 45.0 Tanzania, la Croix 4:
A. ranomafana Goldblatt 57.7 X 40.0 Madagascar, Pcia 1336
A. €: laena Harv 66.7 X 60.0 S. Africa, E. Cape, G & M 9588
A. w 56.5 S. Africa, E. Sibi G & M 9815; 9831
ii N. E. Br. У
(А. acer Baker, A. humbertii H. Perr., A. kitchingii Baker, and A. madagascariensis Baker (Goldblatt & Le
Thomas, 1992a) all have grains smaller than 60 jum equatorial diameter.)
Section Trilobatae Weim.
A. platycephala Baker 60.6 X 59.6 5. Africa, E. Cape, van Wyk & Mathews 7727
Section Ancipites Weim.
A. abyssinca Pax 6143x555 S. Africa, Natal, G & M 9855
A. anceps Ecklon 54.0 X 46.9 S. Africa, E. Cape, G & M 9530
Section Cladocarpae Weim.
A. cladocarpa Baker 48.5 X 45.0 Madagascar, Bosser 16695 (see
(
also Goldblatt & Le Thomas, 1992a)
Section Pseudaristea Pax
A. biflora Weim 71.2 X 70.5 S. Africa, W. Cape, Goldblatt 8898
A. ыле Goldblatt & J. Manning 69.2 х 66.4 S. Africa, W. Cape, G & M 10284
A. ecklonii Baker 63.0 x 59.0 S. Africa, E. Cape, ex hort
A. lugens (L.f.) Weim. 92.2 X 81.0 S. Africa, W. Cape, Oliver 4739
A. pauciflora W. Dod. 13.5 X 63.5 S. Africa, W. Cape, G & M 10102
A. pusilla (Thunb.) Ker 59.6 x 58.5 S. Africa, W. Саре Bayliss 7635
А. simplex Weim 41,1 067.5 S. Africa, W. Cape, С & M 9754
A. spiralis (L.f.) Ker Gawl. 73.5 X 63.5 S. Africa, W. Cape, G s.n. (Cape Point)
A. teretifolia Goldblatt & J. Manning 103 ХЛ 5. Africa, У. Cape, Bean 2785; Nanni s.n.
A. sp. 2 aff. pauciflora 81.2 х 76.2 S. Africa, Cape, Drewe 466
Section Racemosae Weim.
A. confusa Goldblatt 54.0 X 51.1 S. Africa, W. Cape, G s.n. (Hout Bay)
A. fimbriata Goldblatt ined. 73.5 х 70.5 5. Africa, №. Саре, С & М 10167
А. juncifolia Baker 52.7 х 45.0 5. Africa, W. Cape, Williams 891; Orchard 354
A. major Andrews 45.0 X 42.0 S. Africa, Cape, G s.n. (Hout Bay
A. monticola Goldblatt 54.0 X 49.5 S. Africa, W. Cape, G & M 9476A
A. racemosa Baker 52.5 X 48.0 5. Africa, Cape, Oakes s.n.
Section Aristea
A. africana (L.) Hoffmsg. 95.2 X 92.2 S. Africa, W. Cape, С & M 9352;
82.5 X 81.0 G & M 9505;
8750x832 Bean 2789;
88.5 X 77.2 G & M 9750
A. dichotoma (Thunb.) Ker Gawl. 075 X 930 S. Africa, W. Cape, С & M 9503; С & M 10154
A. glauca Klatt 82.5 X 75.0 S. Africa, W. Cape, G & M 9595A
A. oligocephala Baker 97.5 X 85.5 S. Africa, W. Cape, Barker 412
iei CEN А а ВН
Volume 84, Number 2
1997
Goldblatt & Le Thomas
Palynology of Aristea
fixed in FAA, and subsequently stored in 70% eth-
anol, or were taken from herbarium specimens in
the collections at the Laboratoire de Phanérogamie,
Paris (P), or the Missouri Botanical Garden, St.
Louis (МО). All samples were examined under the
light microscope (LM), without acetolysis, and
mounted in glycerine jelly, but after rehydration of
anthers in a wetting agent and subsequent washing
for herbarium samples. For SEM study, non-aceto-
lyzed rehydrated herbarium samples and spirit ma-
terial were fixed in 2.5% glutaraldehyde solution,
washed in a 0.1 M cacodylate buffer, then dehy-
drated and critical-point dried. Pollen of Aristea is
difficult to study owing to the fragility of the exine,
of which the foot layer is extremely thin. It is im-
portant to examine non-acetolyzed grains under the
light microscope in the hydrated, or rehydrated,
state so that the apertures can be more clearly in-
terpreted by the presence of thickened intine. In
some cases, acetolyzed grains were examin
to amplify our observations. Grain size (Table 1) is
the average of measurements of 10 grains mounted
in glycerin jelly. The terminology used is, as far as
possible, in accordance with the Glossary of Pollen
and Spore Terminology (Punt et al., 1994) and the
special sulcus types described for monocots by
Halbritter and Hesse (1993).
PHYLOGENETIC ANALYSIS
Aristea is relatively uniform in floral morphology,
fairly diverse in the arrangement of the inflores-
cence units (binate rhipidia) on the flowering stem,
and remarkably variable in capsule and seed mor-
phology. The current classification of the genus into
eight sections is based largely on fruit and seed
characters (Weimarck, 1940). Species are, however,
recognized in many instances on inflorescence
characters and sometimes on the basis of vegetative
features including overall size and stem compres-
sion. Using Weimarck’s monograph of the genus as
the primary source of morphological data, supple-
mented where necessary from herbarium material
and new observations (Goldblatt, 1995a; Goldblatt
& Manning, 1997), we have identified 24 phylo-
genetically informative morphological features (Ap-
pendix 1). These have been polarized exclusively
using outgroup comparison. Some of the features
are quantitative multistate characters that arguably
should not be included in a phylogenetic analysis.
We believe they do have some phylogenetic infor-
mation and are readily polarized; hence we have
included them. Our outgroup is a combined one,
four of the remaining five genera of subfamily Niv-
enioideae (Goldblatt, 1990, 1993). These are the
Australasian Patersonia and three shrubby south-
ern African genera, Klattia, Nivenia, and Witsenia.
Where characters in the outgroup genera are not
congruent, we determined the ancestral state by
comparing the character to genera of Iridaceae out-
side the subfamily. A sixth genus of Nivenioideae,
Geosiris, was not taken into consideration for char-
acter polarization because it is a highly specialized,
leafless and achlorophyllous plant, most features of
which have no homologue in Aristea
We also included two nontraditional characters
in the analysis, one from leaf anatomy and one re-
lating to floral nectaries. The outgroup genera have
leaves with strands of subepidermal sclerenchyma
in the leaf margins (Rudall & Burns, 1989), but the
seven species of Aristea known anatomically lack
this tissue (Rudall, 1995; J. Manning, pers. comm.),
instead having columnar epidermal cells with
thickened anticlinal walls. Both anatomical condi-
tions are probably universal for the genus but are
scored only for the species in which they are
known. Septal nectaries are considered to be the
plesiomorphic condition in Nivenioideae (Gold-
blatt, sees They occur in Patersonia (P. Rudall,
pers. comm.) and in Nivenia, Klattia, and Witsenia
(Goldblatt, 1993). Flowers of nearly all species of
Aristea lack nectar (unpublished observations), but
A. spiralis does secrete nectar, which has been
found to be produced from perigonal nectaries lo-
cated near the tepal bases (J. Manning, pers.
comm.), an autapomorphy for that species. The
above three characters contain no information rel-
evant to the relationships of members of the ingroup
but seem unambiguously to establish the monophy-
ly of Aristea, which has few apomorphic morpho-
logical features. The only apparent specialized fea-
ture in Aristea is the vestigial perianth tube.
Papillate outer epidermal cells of the seed coat,
present in the few species examined for the char-
acter, may be another. No other genus of Iridaceae
is known to have such epidermal cells in the seed
coat bro data).
aracters of the species of Aristea examined
ut were combined with data for several more for
which there is palynological information (Goldblatt
& Le Thomas, 1992a), assembled in a matrix (Table
2), and analyzed using the Hennig86 package of
programs for phylogenetic analysis (Farris, 1988).
The data set, including 33 characters and up to 36
taxa, took considerable time to run using the ie
option (generating trees by implicit enumeration).
We thus decided to use the mh* bb* options for
the analysis, which applies branch swapping to
each of the initial trees and saves all the shortest
trees generated. Tree lengths, and consistency and
266 Annals of the
Missouri Botanical Garden
Table 2. Data matrix and character list for Aristea. Multistate characters are 1, 4, 9, 11, 12, 13, 14, 15, 18, 19,
20, 24, and 33; of these 4, 9, 12, 14, and 18 are additive or arbitrarily divided grades in a more or less continuous
range of variation and are treated as ordered states. Characters are scored ? if unknown or intermediate between two
states [Note: some scorings change for the matrix that includes Ixioideae, and an additional character, 34, is added for
this analysis.]
1. Pollen grain apertures restricted to one surface, usually HU TEES rarely trichot | (0) i leulat
(1); 2-zonasulculate (2); grains 3-sulcate or spiraperturat
2. Aperture membrane smooth to granular, without exine карыны (0)—арепиге membrane with exine fragments,
sometimes forming an operculum-like median band (1)
3. Aperture border not diffuse (0)—aperture border diffuse (1)
4. Exine lumina large n (1); small (2)
5. Exine surface (muri or areolae) smooth (0)—ех пе surface — (1)
6. Exine surface without perforation: dp ы, with perforations (1)
7. Pollen grains smaller (diameter less than 65 рт) (0)—larger = more than 70 jum) (1)
8. Pollen always shed in monads (0)—pollen often shed in tetrads (1)
9. Exine reticulate (0)—Trugulate (1); rugulo-areolate (2)
10. Rhipidial spathes prominent, enclosing the entire inflorescence (0)—spathes not prominent and buds and floral
bracts evident (
11. Capsules round to lightly 3-lobed in section (0)—capsules 3-winged (1); capsules 3-lobed (2); deeply divided into
3 lobes, each widest in the middle (3)
12. Capsules in profile more or less ovoid (0)—capsules oblong (1); capsules elongate (2)
13. Capsule walls brittle and dehiscing normally (0)—walls thin, dehiscing on the septa (1); walls thick and woody,
tardily dehiscent (2)
14. Seeds numerous to several per locule (0)—seeds usually 4 per locule (1); seeds 2 per locule (2)
15. Seeds rounded to prismatic (0)—seeds radially compressed (1); seeds nearly cylindric and obliquely truncate at
both ends (2); seeds globose with an aril (3)
16. Anthers ovoid-oblong (0)—Aanthers narrrow and linear (1)
17. Anthers a dehiscent (0)—anthers apically dehiscent (рогове) (1)
18. Lateral rhipidia all terminal on branches, thus stalked (0)—some lateral rhipidia sessile (1); all lateral rhipidia
normally sessile (2)
19. Compound in , Spicate, or subpaniculate (0)—subdichot ly branched (1); 1-2 subterminal
flower clusters (2)
20. Stem lightly compressed to terete in upper half (0)—stem strongly compressed and broadly winged throughout (1);
narrow and ancipitous (2
21. Outer tepal whorl similar to inner (0)—outer and inner whorls differentiated in size and color (1)
22. Flowers more or less upright (0)—flowers secund (1)
23. Style lobed apically and the lobes fringed (0)—style minutely notched apically (1)
24. Rhipidial spathes and floral bracts green to brownish and chartaceous (0)—spathes and bracts silvery and scarious
with entire margins (1); with regularly fringed, rust-colored margins (2)
25. Septal nectaries present (0)—septal nectaries absent (1 )
26. Flowering stem bearing a few leaves decreasing in size above (0)—basal internode very long, leafless except for a
subterminal bract-like leaf (1)
27. Flowering stems exceeding the leaves and plants not cushion-like tufts (0)—flowering stems short and plants low
cushion-like tufts
28. Inflorescences with several to few flower clusters (0)—compound inflorescence massively enriched (1)
29. Seed surface with primary sculpturing, usually rugose (0)—surface without primary sculpturing (1)
30. margins with subepidermal sclerenchyma (0)—without subepidermal sclerenchyma (1)
31. Perianth tube well developed (0)—vestigial (1)
32. Outer tepals symmetric about the midvein (0)—asymmetric (2)
33. Tepals uniformly deep blue (0)—tepals pale ed to whitish (1); tepals shades of mauve to turquoise (or cream) (2)
Additional character when Ixioideae is added to m
34. Flowers lasting at least one day and not déliqueicg on fading (0)—flowers fugaceous, lasting less than a day nd
deliquescing on fading (1) [scored 0 for Nivenia and Ixioideae, 1 for all species of Aristea]
[Autapomorphies not included: exine with supratectal verrucae (A. spiralis); perigonal nectaries (A. spiralis); flower
алате pendent (A. singularis); spathes and bracts regularly fringed (A. africana); spathes and bracts lacerate
woodi
lar qp. NE
Моште 84, Митрег 2
1997
Goldblatt & Le Thomas 267
Palynology of Aristea
Table 2. Continued.
Character Number
1 TELLE 113 22228 22223 383
Тахоп 12345 67890 12345 67890 12345 67890 123
Nivpat 00010 00000 00020 00020 00000 00000 000
africana 30001 11001 10121 00010 00021 01001 100
anceps 11121 10010 01000 00221 00001 1000? 100
teretifolia 20000 11000 32202 10200 10001 0000? 112
biflora 20000 10000 32202 10200 10001 01001 112
cantharophila 20101 10000 32202 10200 00001 0000? 102
monticola 00020 00011 10011 00100 00101 00101 100
juncifolia 00010 00001 11011 00200 00101 0001? 100
fimbriata 00010 ?1001 #1771 00100 00121 01022 100
confusa 00020 00001 11011 00100 00101 0011? 100
dichotoma 30001 11001 10121 00012 00011 0100? 100
ecklonii T1413 10111 21000 00101 00001 00001 100
auca 30000 11001 10121 00012 00011 01002 100
lugens 20000 11000 32202 10200 10001 0000? 101
major 00020 00001 10021 00100 01101 0011? 100
spiralis 21120 01000 32202 10201 01001 00001 101
simplex 21341 11000 32202 10200 01001 0000? 101
schizolaena 00120 00001 00000 00200 00001 0000? 100
oetzel 11120 10021 00000 00201 00001 0000? 100
humbertii 11120 109871 00000 01110 00001 0010? 100
cladocarpa 11321 10011 21000 00101 00001 00002 100
madagascariensis 11120 10021 00000 01200 00001 0000? 100
kitchingii 11120 10021 00000 01100 00001 0010? 100
angustifolia 11120 10011 00000 00200 00001 0000? 100
nyikensis 11120 10021 00000 00100 00011 0000? 100
ranomafana 11120 10021 00000 01010 00001 0010? 100
polycephala 11120 10021 00000 00010 00011 00002 100
racemosa 00010 00001 11011 00200 00101 001?? 100
pusilla 11121 10110 21000 00101 00001 0000? 100
angolensis LILLE 10011 00000 00100 00011 00001 100
oligocephala 30001 11001 10121 00010 00011 00002 100
platycephala 1112? 10011 20020 00201 00001 0000? 100
abyssinica 11111 10011 00000 00121 00001 1000? 100
singularis 00020 00011 10123 00010 00001 01002 100
pauciflora 20001 11000 32202 00200 00001 00001 100
woodii 11111 10011 00000 00200 00001 0000? 100
montana 11111 10001 00000 00100 00001 0000? 100
grandis 11120 20011 00000 00200 00001 00002 100
retention indices (Figs. 47, 49), are automatically
calculated by Hennig86 (Farris, 1988). Because the
initial trees were poorly resolved, the successive
weighting option was also invoked. Successive
weighting, recommended by Farris (1969) for situ-
ations where unreliable (homoplasious) characters
outnumber reliable ones, is one way to improve tree
resolution. The method selectively weights those
characters that are more consistent at the expense
of those that are homoplasious. The method may be
extremely helpful especially when more conven-
tional analyses yield large numbers of trees and
consensus trees are poorly resolved. Cladograms
were then analyzed using CLADOS (Nixon, 1992)
and the trees presented here were generated using
this program. We were unable to determine a mea-
sure of confidence in the trees obtained by succes-
sive Mp by bootstrap analysis (Swofford,
1991) because the number of trees generated usin,
the method was so large in some replicates that it
became impractical to to perform.
А second series of cladistic analyses was per-
formed after the addition of one more taxon, sub-
family Ixioideae, to the matrix. This was because
in recent molecular systematic studies using se-
quences from two chloroplast genes, rbcL and rps4
268
Annals of the
Missouri Botanical Garden
(M. Chase et al., 1995, and pers. comm.; Souza-
Chies et al., 1997), the subfamily has been found
to be nested in Nivenioideae, as a sister clade ei-
ther to Nivenia plus Witsenia or Aristea. Conse-
quently, we were interested to see how this might
affect the topology of the trees.
RESULTS
PALYNOLOGY
Shape and Size. In hydrated or rehydrated pol-
len, grains are ellipsoid to nearly spherical. Pollen
is typically shed in monads, but in Aristea ecklonii
and A. pusilla the tetrad stage is evidently pro-
longed, and most pollen is shed as tetrads, a spe-
cialized condition (Figs. 1, 2).
Species of Aristea fall fairly well into two size
classes for pollen-grain size (Table 1). Nineteen of
the species examined have medium-sized to mod-
erately large grains (equatorial diameter 45-64 jum)
and 13 species have large grains (equatorial di-
ameter more than 69—97.5 jum). Aristea schizolae-
na, with grains 66.7 X 60.0 jum, falls between the
two groups. Pollen-grain size among the species ex-
amined ranges from a minimum of 45.5 X 42.0 um
i major, to a maximum of 97.5 X 85.5 jum in
A. oligocephala, and 97.5 X 93.0 jum in A. dicho-
toma. Species of the outgroup have grains that fall
in the medium to moderately large size category
(Manning & Goldblatt, 1989), hence this is pre-
sumed to be the plesiomorphic condition. Size is
correlated with some aspects of Weimarck's (1940)
infrageneric classification (Table 1). The four spe-
cies of section Aristea examined all have grains ex-
ceeding 80 X 77 jum. The remaining species of that
section, А. recisa, has grains of comparable size ac-
cording to measurements given by Schulze (1971a).
Only A. lugens and A. teretifolia (sect. Pseudaristea)
have comparably large grains. Most other species
of section Pseudaristea (as redefined by Goldblatt,
1995a) have somewhat smaller grains, more than
71 X 70 рт, and are thus still classified as large
according to our definition. Grains of species of
section Racemosae, excepting A. fimbriata, are me-
dium-sized, as are those of sections Eucapsulares,
Trilobatae, Cladocarpa, and Ancipites (Table 1), in-
cluding those of the Madagascan species (Goldblatt
& Le Thomas, 1992a). Except for A. schizolaena,
grains of members of these sections have an equa-
torial diameter less than 65 jum. Pollen grains of
the outgroup are less than 65 jum in diameter,
hence medium-sized grains are scored 0 and large
grains as 1. Aristea schizolaena, which has grains
intermediate in size between the two size classes,
is scored as 7.
Apertures. Pollen-grain apertures are remark-
ably variable for a genus the size of Aristea. In
many cases, correct terminology (sensu Punt et al.,
1994) cannot be established because we lack de-
velopmental stages necessary for its definition. In
our analysis, we recognize four main apertur
types. These are treated as four unordered states of
one character. The aperture types are as follows.
1. The monosulcate type, which is the plesio-
morphic condition for the family based on outgroup
comparison (e.g., Goldblatt, 1990). Apertures have
been observed in some species at the tetrad stage
and determined to be distal, and can reasonably be
assumed to be so whenever apertures are unambig-
uously sulcate (Figs. 3-6, 10, 11). We include in
this category the trichotomosulcate aperture of A.
fimbriata (Fig. 37) because, although the aperture
in the species is recognizably different from truly
monosulcate apertures, it seems to us to belong in
the same general class, being restricted to one face
of the pollen grain.
Monosulcate grains are present in five of six spe-
cies of section Racemosae examined (Aristea con-
fusa, A. juncifolia, A. major, A. monticola, A. race-
mosa) and in A. schizolaena and A. singularis.
Among the species examined by Schulze (1971a),
A. macrocarpa (sect. Racemosae) also has monosul-
cate pollen grains. Aristea fimbriata, also section
acemosae, alone has trichotomosulculate grains
(Fig. 38). As in grains with monosulcate apertures,
the aperture is generally wide, reaching the ends
of the grain and sometimes extending a short dis-
tance onto the non-apertural face. The aperture
membrane is typically smooth, that is without sub-
stantial amounts of exine on the surface, and the
margins are clearly defined. Occasionally tiny frag-
ments of exine are present, visible only using LM
(e.g., in A. fimbriata).
2. The sulculate type, which includes l-zona-
sulculate and disulculate grains, the distinction, m
some cases, being difficult to establish because
they may occur together in the same species, even
in the same sample, and there are often forms 1n
termediate between the two, for example, in species
like A. kitchingii (Goldblatt & Le Thomas, 1992a)
(Figs. 13-24).
Sulculate grains are the most common type 12
the genus and are found in 19 species of tropical
and eastern southern Africa as well as in 6 of the
7 species that occur in Madagascar. In these grains
the aperture is typically obscure and the apertural
membrane is covered by more or less disorganized
exine, sometimes consisting of an almost complete
band, free or not at the ends of the apertures (thus
an operculum or pontoperculum) especially in spe
Моште 84, Митбег 2
1997
Goldblatt & Le Thomas 269
Palynology of Aristea
Figures 1-12. —1. Aristea ecklonii; mature grains in tetrads. — 2. Aristea pusilla: tetrads of mature pollen. 3-12. Mono-
sulcate "DE grains of sections Касеток ae, Singularis, and E
majo, А. puer jlia.—5 onfusa.—6 & 9. .
po patus (sect. Singularis). PARR bar: whole grains 10
cies (Goldblatt & Le Thomas, 1992a) of Madagas-
car. The aperture margins are usually diffuse and
the ends of the apertures are often difficult to see,
apparently sometimes extending around the entire
А. dai
кт. with ехте detail for selected species.—3. A.
. А. mpa na (sect. Еш 'apsulares).—11, 12.
pm; se ا detail 1 p
q
~
grain (as а zonasulculus—Figs. 13, 15, 19) or in-
terrupted by exine (then evidently disulculate—
Fig. 24).
3. The 2-zonasulculate type (Le Thomas &
270 Annals of the
Missouri Botanical Garden
Figures 13-24. Sule V pollen grains of section Eucapsulares with exine sculpturing in selected species.— —18,
14. Aristea grandis.—15. А. ее 7. A. montana.—18. A. abyssinica.—19, 20. A. goetzei.—21. А. nyiken-
sis.—22, 23. A. ranomafana.—24. , A. polycephala Arrows indicale the apertural zone when not obvious. Scale bar:
whole grains 10 jum; sev ulpturing detail lu
Goldblatt, 1994; Le Thomas et al., 1996) in which Two-zonasulculate apertures occur in all seven
there are two clearly defined zonasulculi (Figs. 25- species of section Pseudaristea (Table 1). Here, the
34), an extremely rare type of pollen grain in flow- margins are usually clearly defined (Figs. 26, 29,
ering plants. 31, 34), but in A. spiralis (Fig. 35) and A. pauciflora
Моште 84, Митбег 2 Goldblatt & Le Thomas
1997
Palynology of Aristea
Figures 25-36. Two-zonasule ri pollen grains of section Pseudaristea as redefined here, with exine "m aperture
detail for selected specie с . Aristea еа. ау and polar view of grains.—28-30.
equatorial and polar view of grai ا 32. А. lugens.—33 & 36. РА 35
arrows s pointing to the two dials uli, and aperture ‘detail showing diffuse margins, apertural ex
supratectal exine gemmules. Scale bar: whole grains and apertural detail of A. spiralis 10 рт; sculpturing detail 1 pm.
A. biflora,
5. А. spiralis, cea grain with
ine fragments, and
(Le Thomas et al., 1996) they are somewhat diffuse of A. simplex, 2-zonasulculate grains occurred to-
and the apertural membrane is covered with small, gether with some 1-zonasulculate grains. The
scattered fragments of exine. This aperture type is 1-zonasulculate condition may represent the inter-
usually constant within a species, but in our sample mediate phase in the evolution of the 2-zonasul-
Annals of the
Missouri Botanical Garden
culate grain. Pollen of an unidentified plant, ap-
parently closely allied to A. pauciflora, but lacking
both well-preserved flowers and mature capsules
and seeds, has exclusively l-zonasulculate grains
with clearly defined margins and without diffuse
exine fragments on the apertural membrane (Fig.
45). These grains recall the 1-zonasulculate grains
found in our sample of A. simplex. The plant also
bears a fair resemblance to A. zeyheri, but speci-
mens identified with this species (Weimarck, 1940)
clearly belong to section Racemosae, close to A.
racemosa. Like it, they have monosulcate grains
(three collections checked under the light micro-
scope). Because of the uncertain identity of the spe-
cies with these 1-zonasulculate grains, and because
so many of its morphological characters are un-
known, we were unable to include it in the phylo-
genetic analysis. This plant, evidently representing
an undescribed species, should be investigated in
more detail when opportunity arises.
4. The 3-sulcate (or more or less spiraperturate)
type (Le Thomas & Goldblatt, 1994; Le Thomas et
al., 1996, Goldblatt et al., 1997), known elsewhere
in Iridaceae, with well-defined apertural zones sep-
arating equal or unequal plates of exine and the
sulci always confluent (Figs. 39-41).
Three-sulcate grains are restricted to three of the
four species of section Aristea examined, and a fifth,
А. recisa, examined by Schulze (1971a), probably
also has this aperture type although it was not spe-
cifically so described by him. In A. oligocephala of
the section, the apertures are difficult to define and
are more or less spiraperturate, consisting of sev-
eral confluent sulci separated by large plates of ex-
ine of varying size. The tendency for the spirali-
zation of the aperture apparently corresponds to
Schulze’s (1971а) observations for the species
based on examination of acetolyzed pollen.
In addition to the aperture itself, two more ap-
ertural characters are recognized, the nature of the
margin and the apertural membrane. Margins are
either sharply defined (e.g., Figs. 3-6), the plesiom-
orphic condition based on outgroup comparison
(Manning & Goldblatt, 1989), or diffuse (e.g., Fig.
35). The ancestral condition, again based on out-
group comparison (Goldblatt & Manning, 1989), for
the aperture membrane is without fragments of ex-
ine (i.e., smooth) (e.g., Figs. 3-5, 39, 40), and in
the derived state the membrane is more or less ob-
scured by irregularly shaped, fairly large fragments
of exine (e.g., Figs. 13, 15, 16, 21) that sometimes
form an operculum-like band (Goldblatt & Le Tho-
mas, 1992a).
Surface sculpturing. The exine is tectate-colu-
mellate and the exine patterning ranges from retic-
ulate (Fig. 7, 8, 42-44), to rugulate (Fig. 9, 12),
and rugulo-areolate (Fig. 20, 23), depending on the
size and shape of the lumina and the orientation of
the muri with respect to one another as well as their
overall shape. The character states form a contin-
uum and the distinction between them, although
clearly defined by terminology (Punt et al., 1994),
is arbitrary. The states thus constitute an ordered
series of one character. Lumen size is regarded as
a separate character from basic exine pattern, and
outgroup comparison suggests that lumina of mod-
erate size (1-2 ¡um in diameter), as for example, in
Aristea confusa and A. montana (Figs. 7, 17), are
the plesiomorphic condition for the genus. А retic-
ulum with large lumina (2-6 ¡um in diameter) char-
acterizes А. africana, A. glauca, and A. cantharo-
phila (e.g., Figs. 27, 42-44), and in these species
the bases of the lumina are often prominently ver-
rucate or baculate. Small lumina (less than 1 jum,
ie., microreticulate) characterize A. juncifolia, А.
angolensis, and A. spiralis, among others (e.g., Figs.
8, 15, 35). Lumina are normally fairly similar in
size, all or most falling in the same size class for a
particular sample. Aristea cantharophila is, how-
ever, exceptional in having mostly large lumina
mixed with small lumina on the distal exine plates
(Fig. 26). The three states of lumen size are arbi-
trary divisions of a continuum and intrinsically ad-
ditive, and the character is treated as ordered.
The exine muri are variable in shape and sculp-
turing, most often angular and smooth, especially
when the lumina of the reticulum are of moderate
or small size, but more or less wavy, sculpted, and
with perforations when the lumina are large. The
species of section Africana and some of section
Pseudaristea are notable for their ridged exine muri
(Figs. 27, 42, 44). A remarkable feature of the ex-
ine of A. spiralis is the presence of large, irregularly
distributed, supratectal gemmules on the reticulum
(Figs. 34, 35), a condition reported for this species
by Radelescu (1970). This recalls the exine of one
species of the outgroup, Witsenia maura (L)
Thunb., which is autapomorphic in having large
verrucae lying on a more or less microreticulate
tectum (Manning & Goldblatt, 1989). A typical ru-
gulate sculpturing with more or less elongate exine
elements, free or incompletely fused, occurs ш ™
monticola and A. singularis (Figs. 9, 12), but im
some species in this category the rugulae are con
torted, very irregular in shape, and more ог less
fused (e.g., А. grandis—Fig. 14). In this type, the
surface of the muri may be smooth or sculpted.
the Madagascan endemic species (Table 3), plus A.
goetzei and the tropical African endemic A. nyiken-
Volume 84, Number 2
Goldblatt & Le Thomas 273
Palynology of Aristea
Figures 37-46. 37, 38. Pollen grains of . Aristea — وپ (sect. Racemosae), section Aristea and A. sp. (aff. A.
A gr
pone iflora) with « exine ue for selected species.—
сас —3 1. africana.—A0 & 43. A. s i et
ale bar: о d grains 10 jum; sculpturing detail 1 jum.
TY showing er and non- а surfaces
А. oligocephala.—45, 46. A. sp. (aff. pauciflora).
274 Annals of the
Missouri Botanical Garden
Table 3. Species of Aristea with their geographic ranges arranged according to the revised classification proposed
here. There are 50 species currently recognized in the genus, one or possibly two undescribed species in section
Racemosae, and the plant referred to in the text as G. sp. aff. pauciflora may be an undescribed species of section
Pseudaristea. Species marked with * are unknown palynologically; those with @ are at least known for the aperture
but were not included in the phylogenetic analysis because other pollen characters are unknown. Figures in parentheses
are total species in the taxon. The type species of the subgenera and sections are indicated in bold type.
Taxon Geographical range
Subgenus Eucapsulares Goldblatt (24)
The A. angolensis group (= sect. Aristea sensu Weimarck) (12)
A. angolensis Baker (incl. A. flexicaulis Baker) Cameroon and Ethiopia to eastern S. Africa
A. angustifolia Baker dagascar
A. compressa Buch. ex Baker* eastern S. Africa
А. galpinii N. E. Br. ex Weim.* eastern S. Africa
A. gerrardii Weim.* (closely related to
A. compressa and possibly not distinct) eastern 5. Africa
A. grandis Weim. eastern S. Africa
er eastern S. Africa
A, goetzei Baker (= A. nitida Weim.) Tanzania and Madagascar
A. nyikensis Baker (incl. southern tropical Africa
A. hockii de Wild)
A. polycephala Harms southern tropical Africa
A. schizolaena Harv. ex Baker east 1
A. woodii N. E. Br. southern tropical and eastern S. Africa
они group (4)
A. humbertii Н. Perr. Madagascar
A. kitchingii Baker Madagascar
А. madagascariensis Baker Madagascar
A. ranomafana Goldblatt Madagascar
The A. ecklonii group (incl. section Trilobatae Weim. and section o Weim.) (5)
A. cladocarpa Baker Madag
А. ecklonii Baker с. and Uganda to eastern 5. Africa
A. ensifolia Muir* southern Cape Region
А. platycephala Baker eastern S. Africa
А. pusilla (Thunb.) Ker Сам!. eastern S. Africa and Cape Region
The A. anceps group (= section Ancipites Weim.) (3)
A. abyssinica Pax (incl. Ethiopia to eastern S. Africa
A. cognata N. E. Br.)
A. alata Baker* eastern tropical Africa
A. anceps Ecklon eastern S. Africa
Subgenus Pseudaristea Pax (7)
A. biflora Weim. Cape Region
A. cantharophila Goldblatt
& J. Manning Cape Region
A. lugens (L.f.) Weim. Cape Region
А. pauciflora W. Dod. Cape Region
A. spiralis (L.f.) Ker Gawl. Cape Region
А. simplex Weim Cape Region
А. teretifolia Goldblatt & J. Manning Cape Region
Subgenus Aristea (21)
Section Racemosae Weim. (13)
A. bakeri pem Peng distinct from A. confusa) Cape Region
A. confusa Go Cape Region
A. cuspidata E Cape Region
A. fimbriata Goldblatt & J. Manning Cape Region
A. inaequalis Goldblatt & J. Manning@ Cape Region
А. juncifolia Baker Cape Region
А. macrocarpa G. Lewis@ Cape Region
Volume 84, Number 2
1997
Goldblatt & Le Thomas 275
Palynology of Aristea
Table 3. Continued.
Taxon Geographical range
A. major Andrews Cape Region
A. monticola Goldblatt Cape Region
А. racemosa Baker Cape Region
A. rigidifolia G. Lewis* Cape Region
A. rupicola Goldblatt & J. Manning? Cape Region
A. zeyheri Baker? Cape Region
Section Singulares Weim. (1)
Weim Cape Region
Section Aristea (6)
A. africana (L.) Hoffmsg. Cape Region
A. dichotoma (Thunb.) Ker Gawl. Cape Region
A. glauca Klatt Cape Region
A. oligocephala Baker Cape Region
А. palustris Schltr.* Cape Region
A. recisa Klatt@ (closely Mies to and perhaps Cape Region
not distinct from A. africana)
Uncertain position (Lewis, 1952)
A. latifolia G. Lewis* Cape Region
sis (Figs. 20, 21), have a rugulo-areolate sculpturing
consisting of very closely aligned rugulae, which
sometimes conform to the strict definition of areo-
ae.
In addition to the basic exine patterning we rec-
ognize two additional exine characters: surface
smooth or sculpted, and with or without perfora-
tions. Outgroup comparison indicates that both the
smooth condition and the absence of perforations
are ancestral.
PHYLOGENY
Preliminary cladistic analyses quickly showed
that the data are homoplasious for many characters
and that numerous (usually over 1000) trees were
obtained whatever option was chosen from the Hen-
ning86 package for the analysis. The resulting con-
sensus trees, however poorly resolved, always em-
bodied a number of distinct clades (Fig. 47A). Most
notable among them are the clades at nodes 1 and
2 of Figure 47A. Node 1 includes those species of
Weimarck's section Pseudaristea that have 2-zona-
sulculate pollen grains combined with elongate
capsules, locules deeply lobed longitudinally, and
truncated seeds. Node 2 includes Weimarck's sec-
tions Aristea and Racemosae (capsules with promi-
nent narrow wings, radially flattened seeds, and a
reduced seed number per locule), plus A. singularis
(sect. Singularis), x has rounded seeds with an
aril but winged caps
Successive = eR саны a dramatic reduc-
tion in the number of trees and a rise in the con-
sistency index. In the final analysis we obtained
just 24 trees that differed minimally, and the strict
consensus tree (Fig. 47B) is relatively well re-
solved. Most significantly, what we call the core
species of section Pseudaristea (clade 1), also a
clade in the unweighted consensus tree, formed a
sister clade to the remaining species of the genus
(clade 2). The first bifurcation of clade 2 divides
its constituent species fairly evenly. Clade 3 of Fig-
ure 47B corresponds exactly to clade 2 of the un-
weighted consensus tree, that is, sections Aristea,
Racemosae, and Si is. The three sections each
constitute a single clade (Fig. 47B, clades 5, 6, and
7). The division corresponds closely to Weimarck’s
(1940) sectional composition. Clade 3 is supported
by fruit and seed characters (winged capsules, ra-
dially flattened seeds, few seeds per locule) (Fig.
48
).
The poorly resolved sister clade at node 4 in-
cludes species of five of Weimarck's sections: all
the members of his sections Eucapsulares (which
he called Euaristea) and Ancipites; the monotypic
sections Trilobatae (A. platycephala) and C.
pae (A. cladocarpa); and two species of his section
Pseudaristea, A. ecklonii and A. pusilla. There is
little support for any large species clusters in clade
4. Notable smaller clades include those at nodes 8,
9, and 10. Clade 8 includes two species of Wei-
marck's section Pseudaristea, plus the single spe-
cies each of his sections Cladocarpae and Triloba-
276 Annals of the
Missouri Botanical Garden
Subgenus Subgenus Subgenus
Pseudaristea Aristea ucapsulares
-—— Саре ---- -——— Саре === . ---- Е. sthn/tropical Africa, Madagascar ----
| | -|
5 | 3 ls E 3 E E
sÀ ы а» Е es E apes
Seon Bs, sgag rE ‚Шы FECE ES
SSSI ESELSSSSIE LSESESSESSSESES SLs 3 8
ASSISTS o9 SESHSOSONeE FOS eas rs Ces
SSSR г зеш LE 0с FESS So So S 5 Ss $F 5565
~ g E. ~ ~ = = 23923 ERES © i=} Я =
CFEREETEEESEEEEHEREEHTEHETSEEE БЕЗ
сл
о | О
~
> .. |
| со
anceps
gl abyssinica
goetzei
nyikensis
bh
O
ОЈ
°
Nivnat
DA pur
schizolaena
oligocephala
grandis
biflora
goetzei
nyikensis
A
Figure 47. Cladograms showing species relationships in Aristea using the matrix in Table 2 with the outgroup
Nivpat.—A. Strict consensus tree of 1012 equally parsimonious trees obtained using the mh* and bb* options of
Hennig86, length (L) 107, ~ index (CI) 45, retention index (RI) 80. Arrows Kit clades present in all pm
Clade 1 "epe ne to the core species of Weimarck's section Pseudaristea and 2, to sections Aristea, Racemosae.
Singularis. —B. Strict consensus tree of 24 equally parsimonious trees (CI 76, RI 93) obtained using the mh* and bb*
options of Hennig86 followed by successive weighting (xs w). The three subgenera of the revised classification pro
as a result of the analysis are shown above the main clades. Geographical distribution of the subgenera is also indicated.
tae. Clade 9 includes the two species of section is discussed below. The topology of the weighted
Ancipites included in our analysis, and clade 10 has e tree produced using the matrix that in-
the four Madagascan species with porose anther de- cluded the extra taxon, Ixioideae, differs only
hiscence. In the tree selected for plotting characters slightly. In this tree (Fig. 49) Ixioideae falls outside
Fig. 48), clades 8 and 9 are united (compare with Aristea and the three main clades (nodes 1, 2, an
the strict consensus tree, Fig. 47B). The isolated 4) are the same as in the analyses without Ixioideae
position of Aristea schizolaena in clade 4 (Fig. 478) (compare with Fig. 47B). The position of clades 1
is difficult to explain in terms of morphology and and 2 with respect to clade 3 is, however, reversed
~
277
Goldblatt & Le Thomas
Palynology of Aristea
'peurejqo
Zuleq 1[nsa1 aures эш Чим ѕәшц ст paguenieal Ајшоривл әләм exe 'sedueuo лојовлецо јојјелед = papddns (иоптриоо pozi[eroads e jo sso “3°2) as orqdaourorso[d әц 0} јевлолол
= 1#ә[ә ‘984S o1ydiowode əy} ој одивцо = ооо рцов "Surpeus seq ‘g әре], ur влодшти лодовлецо Əy} 0} puodsauoo aan ayi uo sq ayi ansoddo siaquiny (ар "dr ur имоце зод
snsuosuoo) поппашатр лојовлецо Zutmoys *gg3ruusy jo suondo «qq pue „уш əy} pue Випудгом aatssaoons рив z o[qe] ur хеш ay) Sursn роштејдо seen pz Jo ә) ‘gp amig
¿LUMEN
Volume 84, Number 2
1997
E IE
ms
© > |
14 de
= +E GB
F> =
з= E I PI
T о pe
– Ё н = К 15
„ср = hu p „з“
SR 2р =
oe ne TI о ps
А en х * 9 ds б
ps m >
pe 6 PT E 5
LS ES he p
no e oc d
A ~
ca - gno
ЕЧ аў ds P
ch
5 де У
^ eu. = E
С БЕЗЕ ЗОВЕ ОВ И БЕН АНТЕ ЕН ЋЕ:
e oe - с <<] е 0 Š е i ~ . 4
SRRRSPSRRIEBBBRBSÓISBRRBPIARSÓOBSPRPPRSRSASBBSSEZRE:
таб раса назва A Я 655 КЕ Ss £
33% - 8. SEE WI BER © See” RU E
8 ~ ~ |, = E = 5 ~ S 5 3 = 5 S а
8 >
Annals of the
Missouri Botanical Garden
Subgenus Subgenus
istea Pseudaristea
ris
------------ Саре ------------- , ------ Cape ------
|
| 5
3 os
a = 3 3 3.3 si a S
Оза = "= з 2s 5 2 =
: =
аза Зы], СЕРЕН
2 tom 55 E © Y SER S
= .
EEES БЕБЕ
= БОЉЕМ 55555
E
=
T 3
з 3 " 3 = ma
oS арапа SSB
з зб = = З WES. E бсш
~ = Vim из & > > зоћао ва
RRE ITE
3 = S S $9 Re
RS 2's $351 3353 25 Ss
SESE АИ
Ё
Figure 49. Strict consensus tree of 1012 equally parsimonious trees obtained using the matrix that included the
extra taxon, Ixioideae, and using the mh* and bb* options of Hennig86, followed by successive weighting (CI
and 48 are present, but the positions of subgenera Africana
RI = 80). The
same three major clades as in Figure 47B
= 45,
(clade 1) and Pseudaristea (clade 2) are shifted in relation to subgenus Eucapsularis (clade 3).
compared with their positions in Figures 47B and
48. Within clade 4 of Figure 49 A. schizolaena
again assumes its isolated position, A. ecklonii, A.
pusilla, А. cladocarpa, and A. platycaulis are a
clade, but now link basally to A. goetzei and the
two species of section Ancipites. The four Madagas-
can species with porose anthers also remain a sin-
gle clade.
DISCUSSION
INFRAGENERIC CLASSIFICATION
The results of the phylogenetic analysis confirm
the monophyly of Aristea, assumed in Material and
Methods, above. The results also indicate the need
for substantial revision of Weimarck’s (1940) infra-
generic classification. We propose subgeneric status
for the three main clades present in the weighted
trees we have generated (Fig. 47B, 49). Preserving,
where possible, Weimarck’s nomenclature, we call
these subgenera Aristea, Eucapsulares, and Pseu-
daristea. The first of these, clade 3 (Fig. 47B), in-
cludes three of Weimarck’s sections, Aristea (called
by him Cyaneae) at node 6, Racemosae at node 7,
and Singulares at node 5. The first two are evi-
dently monophyletic and their constitution is sup-
ported by the analysis. The third, section Singu-
lares, which is monotypic, is nested between these
two sections. It does not, as Weimarck assumed,
appear to be a taxonomically isolated and primitive
member of the genus. As explained elsewhere
(Goldblatt, 1995a), the only species of the section,
А. singularis, was incompletely known to Weimarc
and in some respects misunderstood. We hesitate
to include A. singularis in section Aristea (it falls
at the base of that clade) because of its odd array
of features. It accords with both sections Aristea
and Racemosae in its narrowly winged capsules.
The small stature and divaricate inflorescence
match section Aristea, but the plesiomorphic small
pollen grains with monosulcate apertures and apo-
morphic rugulose exine correspond with section
Racemosae and contrast starkly with the trisulcate
or even more complex grains with reticulate exine
of section Aristea. On balance, it seems best to
maintain section Singulares. The unique rounded,
and dorsiventrally compressed seeds with a col-
lapsed chalaza, funicular elaiosome, and pendent
inflorescence units certainly represent an assem-
blage of unique features for the genus and support
the recognition of a section despite its being mono"
typic. It is possible that the monosulcate pollen
grains represent a reversal to the ancestral state
and that A. singularis is actually nested in section
Volume 84, Number 2
1997
Goldblatt & Le Thomas 279
Palynology of Aristea
Aristea. Elsewhere (Goldblatt, 1995a) the species
has been compared to A. dichotoma. Future study
may throw light on this question.
The second subgenus we recognize (clade 1 of
Fig. 47B), subgenus Pseudaristea, contains the type
and closely related species of Weimarck’s section
of that name. Two more species of Weimarck’s sec-
tion, A. ecklonii and A. pusilla, fall in the third
subgenus, and A. zeyheri, not included in the anal-
ysis, is probably a specialized species of section
Racemosae (Goldblatt & Manning, 1997). These
three species were included by Weimarck in sec-
tion Pseudaristea largely on the basis of their elon-
gated capsules. All three have different pollen mor-
phology from the other members of section
Pseudaristea and differ morphologically in details
of capsule morphology. The elongate shape of the
capsules, the character used by Weimarck to define
his section, must be assumed to be convergent. In-
cluding the species described since the publication
of Weimarck’s monograph, subgenus Pseudaristea
has just seven species. Their synapomorphies in-
clude large pollen grains, extraordinary 2-zonasul-
culate apertures, exine with a reticulum of very
large lumina, on the one hand, and the unique
leathery-woody, elongate, tardily dehiscent cap-
sules and cylindrical, truncate seeds combined with
long inflorescence spathes on the other.
The third subgenus, Eucapsulares (clade 4 of Fig.
47B), incorporates the remaining four sections of
Weimarck’s classification plus the residue of sec-
tion Pseudaristea, in all some 24 species, 19 of
which are included in the cladistic analysis. The
species relationships are poorly resolved and our
results do not support the recognition of any formal
taxonomic groupings within the subgenus. Instead,
we suggest using an informal solution, referring
simply to “species groups” for well-defined clades.
One of these is particularly noteworthy. Aristea eck-
lonii and A. pusilla are linked to the Madagascan
species A. cladocarpa (sole species of sect. Clado-
carpa) and then to A. platycephala (sole species of
sect. Trilobatae). The latter, an eastern South Afri-
can endemic, was assigned its own tribe by Wei-
marck because of its distinctive short, warty cap-
sules. It is otherwise fairly similar to A. ecklonii
morphologically, and its position on the same clade
is not surprising. We suspect that one more species,
the southern Cape A. ensifolia, which is confined
to forested habitats, will fall in this clade too, but
pollen is so far unavailable and the species could
not be included in the phylogenetic analysis. Aris-
tea ensifolia also elongated capsules, but they
are unusual in being indehiscent. The relatively
soft capsule walls simply decay with age, gradually
releasing the seeds around the parent plants.
A second small clade, the Aristea anceps group,
includes the two species of Weimarck’s section An-
cipites, A. abyssinica and A. anceps (a third, A. ala-
ta, will probably also fall here). A common syna-
pomorphy with the A. ecklonii group, compressed
and broadly winged stems, is the reason the two
groups are linked in Figure 49 in a single clade.
Another clade that appears in the successively
weighted trees of both analyses, the A. madagas-
cariensis group, includes the four Madagascan spe-
cies that have apically dehiscent anthers, A. hum-
bertii, A. kitchingii, A. madagascariensis, and A.
ranomafana (Goldblatt, 1991, 1995b). The remain-
ing species of Weimarck’s section Eucapsulares ex-
hibit little evidence of relationship. Species not in-
cluded in our analysis, A. compressa and A.
gerrardii, appear to us to be fairly closely allied to
A. angolensis, and we are confident that they would
fall close to this species in phylogenetic analyses.
The isolated position of Aristea schizolaena in
subgenus Eucapsulares is difficult to explain in
terms of morphology. The species is sister to the
remainder of the Eucapsulares clade, and its basal
position is the result of its having plesiomorphic
pollen grains the sulcate apertures of which are
smooth. The remaining species have sulculate pol-
len grains with more complex apertures (disulculate
or zonasulculate) always covered with irregularly
shaped exine masses. The apparently plesiomorph-
ic pollen grains may represent a reversal, but an
alternative position for A. schizolaena would result
in longer (less parsimonious) trees. The remaining
species of the subgenus have sulculate pollen
grains, with more complex apertures covered with
irregularly shaped exine masses. Morphologically,
A. schizolaena is unexceptional. It appears to be
closely related to the eastern southern African A.
grandis, A. montana, and A. woodii and is readily
confused with A. woodii. We are reluctant to ascribe
any taxonomic significance to the basal position of
the species.
The remaining species of subgenus Eucapsulares
form a residual group that includes the type species
of section Eucapsulares, A. angolensis. We suggest
referring to these species as the A. angolensis
group.
PHYTOGEOGRAPHY AND PHYLOGENY
There is an extremely close correlation between
the phylogeny (and the revised classification based
on the phylogeny) and geographical distribution
(Fig. 50). All three sections of subgenus Aristea oc-
Annals of th
eem Botanical Garden
Africa and Madagascar pae the distri-
Figure 50
bution of infrageneric taxa of Aristea.— tion Aristea
of subgenus Aristea.—2. Section plone of subgenus
Aristea.—3. Subgenus Pseudaristea. 4. Subgenus Eucap-
sulares. The Cape Floristic Region nearly corresponds to
the range of section Racemosae (2).
cur in the winter rainfall climate zone of southern
Africa, the Cape Floristic Region (Bond & Gold-
of the Greater Cape Flora Region.
With the exception of one species (or two, depend-
ing on taxonomy), members of the subgenus are
actually restricted to the southwest of the region.
y A. confusa and A. bakeri (if that species is
separate from A. confusa) extend eastward to abut
the main range of subgenus Eucapsulares.
All the species of section Pseudaristea also occur
in the southern African winter-rainfall zone, thus
within the Cape Floristic Region proper. Again,
species concentration is highest in the west, re-
peating the pattern in sections Aristea and Race-
mosae of subgenus Aristea.
The geographical distribution of subgenus Eu-
capsulares is far wider, extending across all of sub-
Saharan Africa and Madagascar. Species diversity
exhibits a pattern fairly typical for predominantly
temperate-adapted taxa of fewer species as one
zone (including Lesotho, southern Mozambique,
eastern South Africa, and Swaziland), of which sev-
en are endemic there. Seven species occur in the
Flora Zambesiaca Region (including Malawi, north-
ern Mozambique, Zambia, and Zimbabwe) plus
southeastern Zaire. Species endemic to this region
plus adjacent southwestern Tanzania include A.
nytkensis and A. polycephala. There are eight spe-
cies in East Africa-Ethiopia, of which A. alata is
endemic to the wetter highlands of Kenya, northern
Tanzania, and southern Ethiopia. A second species,
A. goetzei, until recently thought to be endemic to
Tanzania, is now considered conspecific with the
Madagascan A. nitida, a later name (Goldblatt,
1996a). Just three species occur in West Africa,
ognized). Distribution patterns of more widespread
mainland tropical African species of Aristea are
also informative. The non-endemic species of each
of the broad tropical African regions circumscribed
above are shared with southern Africa or, in the
case of A. goetzei with Madagascar, and never with
the adjacent region of tropical Africa alone. Thus,
A. abyssinica and A. ecklonii both extend from the
contact zone between the South African summer-
and winter-rainfall zones northward to Ethiopia and
with isolated populations in the highlands of Cam-
eroon. Aristea angolensis has a somewhat wider
range, extending from eastern southern Africa to
Ethiopia in the north and Senegal in West Africa.
Madagascar has seven species of Aristea, six of
which are endemic. Only A. goetzei is shared wi
southwestern Tanzania. These endemics include
species of at least two clades and possibly three
(the phylogenetic position of A. angustifolia is un-
certain). Thus it appears that Aristea has colonized
Madagascar at least three times from ances
stock on the African mainland, a notable departure
from the usual pattern of African Iridaceae, which
are otherwise poorly represented on that island
(Goldblatt, 1991, 1994)
BIOLOGY AND ADAPTIVE RADIATION
About the remarkable diversity of pollen types
in Aristea very little can be said, and nothing con-
clusive. Pollen variability does not seem to be cor-
related with any aspect of floral ecology or habitat.
Floral presentation in Aristea is remarkably uni-
form. Except for microscopic details of the style
branches, flowers of most species seem to be for all
intents and purposes identical. The similarity ex-
tends even to phenology. Flowers typically open
shortly after sunrise and collapse and deliquesce in
the early afternoon. Pollination of species wi
these flowers is, as far as known (Goldblatt & Man-
ning, 1997), by pollen-collecting female bees of the
families Anthophoridae, Halictidae, and as is often
the case in Africa, Apis mellifera (Apidae).
The only exceptions to this pattern are t
(possibly five) species of section Pseudaristea that
appear to be adapted for pollination by beetles
СОУ ЕВИ МА ИВЕТА ви SS TOt
Volume 84, Number 2
1997
Goldblatt & Le Thomas 281
Palynology of Aristea
(Scarabaeidae: Rutilinae: Hopliinae), a strategy ev-
idently unique to the winter-rainfall region of south-
ern Africa (Goldblatt & Manning, 1996 a
M ги These species have flowers that open
n mid morning, last until late afternoon, and are
vale blue, mauve, or cream with dark contrasting
markings. One additional species of the section, A.
spiralis, is pollinated by long-tongued flies (Phil-
oliche: Tabanidae) (Johnson, 1992; Goldblatt &
Manning, 1997), but at least A. pauciflora of the
section has the plesiomorphic pollination syndrome
of the genus (Goldblatt & Manning, 1997). Yet all
species of the section have virtually the same type
of specialized pollen.
We surmise that climate may be a factor with
regard to pollen aperture in the section. The winter-
rainfall and summer-dry climate is one that is par-
ticularly stressful to plants, many of which flower
after the wet season and when atmospheric aridity
is high and desiccating winds frequent. Drying of
delicate floral parts including stigmatic surfaces
and pollen grains may put reproductive success at
risk. Pollen grains with germination grooves or
pores not restricted to one surface of the grain may
be an adaptive advantage under such circumstanc-
es, once they are transported to a receptive stigma.
Two-zonasulculate, trisulcate, or spiraperturate
grains like those of sections Africana and Pseudar-
istea are more likely to have an aperture in direct
contact with the moist stigmatic surface than mon-
osulcate grains with the aperture restricted to one
surface. Perhaps it is no accident that the most pe-
culiar pollen grains in Aristea, if not in the entire
Iridaceae, occur in species restricted to this partic-
ular climate zone. In this respect we note that the
subfamily of Iridaceae that has radiated most prom-
inently in this region also has unusual pollen for
the family, the monosulcate aperture almost invari-
ably having an operculum (Goldblatt et al., 1991).
It must be noted, however, that not all the spe-
cies of the southern African winter-rainfall region
have specialized pollen grains. Apparently ple-
siomorphic grains characterize all the species of
sections Racemosae and Singulares, and this pollen
type is barely represented in the genus outside the
winter-rainfall region. In regard to the peculiar sul-
culate grains with the apertures largely obscured
by exine fragments that characterize species of sec-
tion Eucapsulares, we can offer no suggestion of its
adaptive value.
Unusual floral features such as tepal markings
and elongate anthers, restricted to some species of
section Pseudaristea (e.g., A. biflora, A. cantharo-
phila, A. lugens, A. teretifolia), are evidently di-
rectly related to their pollination system. Hopliine
5.
beetles are attracted to dark markings on а раје
background, and although they are successful pol-
linators, they also forage on pollen. The larger an-
thers of species of the section evidently compensate
for some loss in pollen by the production of more
grains. In A. spiralis secretion of nectar and its par-
ticularly long stamens are similarly a direct adap-
tation to a pollination strategy, in this case to long-
tongued fly pollination. The flies are nectar feeders,
and the long filaments are necessary to permit
placement of the pollen-bearing anthers sufficiently
distant from the nectar source that they brush
against a foraging insect’s body as it feeds.
Variation in capsule and seed morphology, so
conspicuous in Aristea, seems more amenable to
evolutionary speculation. High wind velocities are
common in the southern Africa winter-rainfall zone
at the time when capsules ripen, exposing the
seeds. Flattened seeds are more likely than small
rounded or angular ones to offer sufficient wind-
resistance to be blown some distance away from the
parent plant. We suggest that the flattened seeds of
species of subgenus Aristea are simply adapted for
wind dispersal. In the case of A. singularis of the
subgenus, the rounded seeds with an elaiosome
must be dispersed by ants, a strategy well known
in the winter-rainfall zone of southern Africa (Bond
& Slingsby, 1983; Linder, 1985). We assume that
this is a novel adaptation in a subgenus in which
the flattened seeds are evidently adapted to wind
dispersal. Aristea singularis grows in rather shel-
tered sites such as in the lee of rocks and in shade
where wind dispersal is unlikely to operate effec-
tively.
The elongate, leathery-woody, tardily dehiscent
capsules of section Pseudaristea suggest another
strategy for survival in the southern African winter-
rainfall zone. Flowering occurs from September to
November in the various species, but the seeds are
not shed for at least six months after ripening and
thus are held in the hard-walled capsules until the
rainy season, which begins in April or May. The
capsules decay under wet conditions and then re-
lease their seeds at a time most favorable for ger-
mination. The tough capsules presumably serve as
protection from both desiccation and insect pre-
dation during the long period that the seeds are
retained.
We suggest that these alternative strategies for
survival, seed dispersal and/or pollination in dif-
ferent lineages explain to some extent the peculiar
adaptations of pollen apertures, flower coloring, an-
ther enlargement, and seed morphology that have
permitted species in different lineages of Aristea to
survive and radiate in the comparatively hostile en-
282
Annals of the
Missouri Botanical Garden
vironment in the southern African winter-rainfall
zone. In this tiny area, less than 1% of the entire
range of the genus, there are 28 species of Aristea
(25 of them endemic), belonging to two subgenera,
one with three sections. The only species of the
remaining subgenus, Eucapsulares, that occur in
the winter-rainfall zone are A. ensifolia and A. pus-
illa. Both occur in the more mesic southern part of
this zone, the former restricted to moist forest or
streamside habitats.
Radiation of subgenus Eucapsulares clearly fol-
lows highland areas in both southern and tropical
Africa as well as Madagascar, where it is restricted
to areas of relatively high rainfall. The species do
not exhibit any particularly striking adaptations in
their reproductive morphology. Elongated capsules
are restricted to some species of forest or forest
margin habitats. It is not clear why the production
of more seeds per capsule should occur in this hab-
itat. Eastern southern African and tropical African
species show very discrete habitat preferences.
Open grassland, short grassland, marshland, and
forest each harbor particular species. Except for
some species of forested habitats that have elongate
Eucapsulares thus appears largely habitat-driven.
To put the above observations in perspective, we
restate the extraordinary degree of variation in pol-
len characters in this genus of only moderate size.
This variation equals or even exceeds the range of
variation found across all the remaining ca. 1700
species of Iridaceae. Much larger genera, including
Gladiolus (ca. 255 species) and Moraea (ca. 130
species), for example, are constant for mc type
and show limited size and exine sculpt vari-
ation (Schulze, 1971a; Goldblatt, 1996b; С Coldblatt
et al., 1991; Goldblatt & Le Thomas, 1992b). Only
Iris (ca. 220 species) comes close to Aristea in hav-
ing a range of aperture types and moderate varia-
tion in exine patterning (Schulze, 1971b). Why pol-
len should be so variable in Aristea remains
ultimately puzzling. Part of the reason may lie in
the geological age of the genus. We suspect that it
is one of the oldest genera of the family, most likely
of pre-Tertiary age. Time alone may have allowed
variation in pollen apertures and exine patterning
to accumulate, and ultimately to be selected for
during periods of climate change such as the es-
tablishment during the Pliocene of semiarid and
ultimately Mediterranean climate at the southwest-
ern extremity of the African continent.
Literature Cited
Bond, P. & P. Goldblatt. 1984. Plants of the Cape Flor
A Descriptive Catalogue. J. S. African Bot. Suppl. 13:
1-455.
Bond, W. J. & P. Slingsby. 1983. Seed dispersal by ants
in shrublands of the Cape province and its evolutionary
"C Ies S. African J. Sci. 79: 231-233.
Chase, , M. R. Duvall, H. G. Hills, J. 6. Conran,
abii bs ef Lilisnes. Pp. 109-137 ¡ in P. J. Rudall, P.
Cribb, D. F. Cutler & C. J. Humphries (editors), Mono-
cotyledons: Systematics and Evolution. Royal Botanic
Gardens, ne
Farris, J. S. A successive weighting approach to
character weighing Syst. Zoo E 18: 374—385.
——. 1988. ig86 version 1.5. Program and soft-
ware а аа Published by the author, Port Jef-
ferson Station, New
Goldblatt, P. 1990. opos and classification of Iri-
daceae. = Missouri Bot. Gard. 77: 607-627.
1 Iridaceae-Famille 45 (2те édition). In
Flore de | de pe et des Comores. 1 Laboratoire
de Phanérogamie, Muséum National d'Histoire Naturel-
le, Paris.
1993. The Woody Iridaceae: Systematics, Him
ogy onal Evolution of Nivenia, Klattia and Witsen:
Timber uro Portland, Oregon.
4. Geography of African Iridaceae. Pp. 931–
940 i ii H. Seyani & A. C. Chikuni (editors), Pro-
Midas of the XIIIth plenary Meeting AETFAT, Ma-
lawi, Volume 2. National Herbarium and Botanic
Gardens of Malawi, Zomba.
1995a. Notes on Aristea Aiton (Iridaceae: b
enioideae): Taxonomy, chromosome cytology, and p
logeny. Ann. Missouri Bot. Gard. 82: 139-145
————. 19955. Aristea ranomafana сома а new
species from Madagascar. Bull. Mus. Hist. Nat., sér. 4,
sect. B, ae 17: 19
. Iridaceae. In R. Polhill (editor), Flora of
Topical суву Africa. a a, Rotterdam.
1996b. Gladiolus in Tropical Africa. Timber
fi uim Portland, Oregon.
& J. C. Manning. 1996. Aristeas and beetle pol-
lination. Veld & Flora 82: 17-19.
& New Species of Aristea (lri-
daceae) from South Africa and notes on the taxonomy
and pollination biology of section Pseudaristea. Novon
7: 137-144.
homas. 1992a. Pollen morphology 0 of
Madagascan Aristea and Geosiris (Iridaceae-Nivenioi-
deae) in rain to systematics and phylogeny. Bull.
Mus. Hist. Nat., 4 sér., sect. B, Adansonia 14: 223-
233
mem A . 1992b. Pollen apertures, exine scup-
turing and phylogeny i in Iridaceae subfamily Iridoideae.
Rev. Palaeobot. & Palynol. 75: 301-315
Bari 6 hikes 991. Sulcus vari-
einen in the pollen grains of Iridaceae subfamily Ixioi-
e. Ann. Missouri Bot. Gard. 78: 961.
Thomas & M. Suarez-Cervera. Phylogeny
and radiation of the genus Aristea (lri ата donées
palynologiques. Geol. Phy. Quartern. In p.
Halbritter, H. & M. Hesse. 1993. Sulcus онај in
some monocot families. Grana 32: 87-99.
Holmgren, P. K, N. H. Holmgren & L. C. Barn 1990.
Index Herbariorum edition 8. New York ie Gar-
den, а New Yor!
Johnson, S. 1992. Plant —animal relationships. ЭЕ РЕ
175-205 іп a Cowling tcd The Ecology of Fynbos
Oxford Univ. Press, Cape Tow:
Volume 84, Number 2
1997
Goldblatt & Le Thomas 283
Palynology of Aristea
Le Thomas, A. & P. Goldblatt. 1994. Deux ene na
р.
nouveaux chez les eben ас Palin
types polliniques originaux dans le Iri
daceae-Nivenioideae): Implications phylogéniques.
Grana 35: 87–96.
Lewis, G. ^ 1952. Aristea latifolia. Ann. S. African Mus.
60: 9-1
Linder, E P. 1985. Gene flow, speciation and species
diversity patterns in a species-rich area: The Cape Flo-
ra. Pp. 53—57 in E. S. Vrba (editor), Species ien Spe-
ciation. Transvaal Mus. Monogr. 4. Transvaal Museum,
retoria.
Manning, J. C. & P. Goldblatt. 1989. Pollen freni
of the shru bby Iridaceae, Nivenia, Klattia, and Witsenia
nn. Missouri Bot. Gard. 76: 1103-1108.
991. Seed coat structure in the
shrubby Cape Iridaceae, Nivenia, Klattia and Witsenia.
ot. J. Linn. Soc. 107: 387. т
Nixon, К. 1992. CLADOS 1.2 IBM base sean char-
acter analysis program. Published M the
Punt, W., S. Blackmore, S. Nilsso ју отав.
1994. Glossary " bw: and аро Terminology. LP
uri 1; . Laboratory of Рајаеођоћапу &
Palynology, шык:
Radelescu, D. 1970. “Re cherches morphopalynologiques
sur les mam d'Iridaceae. Lucr. Grad. Bot. Bucuresti
1968: 311-3
Rudall, P. Tog Anatomy of the Monocotyledons. Irida-
ceae. Clarendon Press, Oxford.
ms. 1989. Leaf gern Me of the woody
South African Iridaceae. Kew Bull. 44: 5
Schulze, W. 1971a. Beitráge zur edm der
Iridaceae und ihre Bedeutung für die Taxonomie. Fed-
des Repert. 82: 101-124.
— ———. 1971b. Beitráge zur Pollenmorphologie der Gat-
tungen um /ris L. Feddes Repert. 81: 507—517.
Souza-Chies, T., G. Bittar, S. Nadot, L. Carter, E. Besin &
‚ Lejeune. 1997. Phylogenetic analysis of Iridaceae
with parsimony and distance methods, using the plastid
gene rps4. Pl. Syst. Evol. 204: 109-123.
Straka, H. & B. Friedrich. 1984. Palynologica madagas-
sica et mascarenica. Fam. 17-49. Trop. € Subtrop.
: 401-470.
Swofford, D. L. 1991. PAUP: Phylogenetic Analysis Us-
ing Parsimony, Version 3.1. Illinois Natural History Sur-
vey, Champaign.
rtial revision of the genus
h of the genus Aristea.
Acta Unis: Lund (Lunds Univ. Arssk.) N. F. Avd. 2, 36
1): 1-140.
APPENDIX 1. Discussion of morphological characters
and their polarization and division into character states.
Numbers refer to sequence of the character in the matrix.
1-9. Discussed fully in main text.
1 ;
ally as long as or longer than the spathes in these species.
The exception: species of section Pseudaristea have large
rhipidial spathes that exceed the floral bracts and conceal
the buds.
11. Capsules of the outgroup are rounded in section or
barely ее Derived
d. Eucapsudares) are scored as 2. The states are re-
as uno
12. Capsule shape: in profile is treated as a separate
character oh sha
2. The character states are additive and
hence treated as ordered.
sules are cartilaginous to nearly woody and lo-
culicidally dehiscent for at pna one-third their length in
most species of section
s of s dim Mast are cartilaginous
with. enini "iin ‘ined walls of the wings and de-
hiscent ч near the — and often — «m the
u
septa as well. These capsules are scor
are sisi uni ia in A. singularis y seit соп-
orm to type 1 and аге so scored. Species of section Pseu-
daristea have leathery to woody, itr. dehiscent to in-
dehiscent capsules, scored 2. The character states are
treated as unordered.
14. Seed number is largely independent of capsule
length. Seed number per locu le is few (usually one) in the
seeds per l acemosae
ecd to four seeds per locule, scored 1, and some
ve two (or one) per ш as do all speci
Aristea, scored 2. The states are additive and treated as
unded to segmental (prismatic) seeds are the
most common type in Aristea and in the entire family.
Patersonia has seeds of the same type, scored as 0, but
Klattia, Nivenia, and Witsenia have tangentially flattened,
shield-sha seeds, unique in lridaceae (Manning &
Goldblatt, 1991). Radially compressed 8 oe
sections Aristea and Racemosae, scored as 1. The more o
ee cylindric seeds with truncated or oblique ends are
ored as 2, and the globose, vertically compressed, a
у seeds of A. singularis are — $ The р насывай
states of seed sha treated as un
16. Anthers are ——— short A им | in the out-
group genera and mos of Aristea as well. Species
of section Pse айал "ехгер ji pauciflora, have linear
anthers, regarded as derived.
17. Longitudinal iis dehiscence is plesiomorphic in
Iridaceae and is - usual condition in Aristea. Four Mad-
agascan species have apomorphic apically dehiscent an-
thers (Goldblatt, 1991. 1995b).
18. When flowering stems have more than just one ter-
lateral inflorescences are sessile, and
mal condition is for all lateral inflorescences to be sessile.
Annals of the
Missouri Botanical Garden
It seems useful to distinguish two states, some lateral in-
florescences consistently sessile (1), or all normally sessile
(2). The character is additive and hence is regarded as
order
19. The arrangement of individual inflorescence units
is highly variable in Nivenioideae. The ancestral condition
is most likely racemose (Goldblatt, 1993), with lateral in-
florescences bo short lateral branches (sessile lat-
units crowded at the stem apex, scored 2. The dicitor
states are unordered.
20. The outgroup genera have lightly compressed or
terete flowering stems, and this is the condition in most
species of Aristea. A few isolated species have strongly
compressed, more or less two-winged flowering stems (an-
cipitous sensu Weimarck, 1940), including A. spiralis
(sect. Pseudaristea) and A. abyssinica, A. alata, A. anceps,
A. cladocarpa, and A. platycephala (sects. Ancipites, Cla-
docarpae, and Trilobatae of Weimarck). In A. dichotoma
and A. glauca the upper part of the ae stem is also
compressed, but the stems are narrow and only obscurely
wing he two states are scored as 1 and 2, чане
апа аге » treated as unordere
Most species of днева as well as those of the out-
group genera have the inner tepal whorl similar to the
outer in length and color (flowers of Patersonia have very
short to vestigial inner tepals, clearly apomorphic for that
genus). Only A. lugens and A. teretifolia have the outer
tepals substantially smaller than the inner and differently
colore:
22. Flowers are more or less upright in the outgroup
and all but two species of Aristea. Only A. nes and
spiralis have secund flowers that face to the si si
Style es of N notch
apically, broadly lobed apically, or divided i into slender
recurved arms. The pattern in the
and we prefer to score the outgrou га as 7. In
species have lobed and fringed style branches, scored 0,
and a few species, all section decias have the styles
ча notched apically, se
e rhipidial spathes, de ually the floral bracts,
eous in the outgrou
, or opaque,
scored 2 (Table 2). The character e are regarded as
unordered.
25. Septal nect
aries are present in all pey of the
outgroup prague: 1998; Rudall, pers. comm.). Species
of Aristea ce nectar, thus yes par Seg ex-
do
cept for A. spiral which has perigonal nectaries (J. C.
прн pers. comm.). Absence of nectaries is regarded
as
26. "The ancestral condition in Iridaceae, including
nearly all species of Aristea, is a flowering stem bea aring a
few leaves decreasing in size above. A few species, in-
cluding А. abyssinica and А. anceps (Table 2), have a flow-
ering stem consisting of one long basal internode and
bearing a short subterminal bract-like lea
27. A low, cushion-like growth form is rare in Iridaceae
forming low, cushion-like tufts with the flowering stems
often not raised much above the leaves.
Few to several inflorescence units (or only one) per
e units are
feature of a few species (Table 2), notably A. major and
A. confusa (sect. Racemosae).
29. Seed surfaces in the outgroup genera typically have
primary sculpturing (Manning & Goldblatt, 1991), as do
most species of Aristea, the surface usually being more or
less rugose (unpublished data), treated as the plesiom-
orphic condition. Seeds of species of section Racemos
examined (Table 2) lack primary sculpturing and are treat-
ed as derived.
30. Species of the eic have leaves with a strand
d Burns,
€ lack subepidermal marginal sclerenchyma (Rudall,
5; J. Manning, pers. comm.), regarded as a derived
са а (0).
. А perianth tube is present in all genera of the out-
and is presumably plesiomorphic for Nivenioideae.
All species of Aristea have the tepals united basally in a
mee tube, 1-2 mm long, scored as
. Tepals of the outgroup and most ‘members of the
adl n tepals symmetric Mt M midvein. The
inner tepals are bacs usly asy ric about the mid-
vein in A. us and A. teretifolia. ga: apomorphic for
че sym
"The s Ed
cate flower qs. in Рта ЈН is most
color is treated as an un
(0), pale blue (1), mauve Бы crea
Additional character when besides i is included in the
analysis:
all Ixioideae. Flow a last les
(u sually fading px aer лал and iu on fad-
ing, the derived st
CHROMOSOME CYTOLOGY
OF IRIDACEAE—PATTERNS
OF VARIATION,
DETERMINATION OF
ANCESTRAL BASE
NUMBERS, AND MODES OF
KARYOTYPE CHANGE!
Peter Goldblatt? and Masahiro Takei?
ABSTRACT
Against a background of 100 original counts for 95 species in 34 genera of Iridaceae, we review chromosome
008 o
пон for the entire family. Counts are now available for во
and all but 5 of ca. 78 genera are known from at least one count. We suggest ancestral base
than x = 10. N
оте 1
unusually low frequency in Africa, the center of diversity for the family. Changes in basic number, frequent in а few
genera, are evidently the result of dysploid reduction. In all but a few possible examples, correlated AA
specialization suggests that dysploid reduction is involved i in stepwise change in base number. Major dysploid se
us, as well as Gladiolus and Lapeirousia (all ан.
are restricted to a few genera, includin
and Iris, Moraea, and Sisyri inchium (Iridoideae). Al other. genera have a single base number or d variation evident
or two species. Patterns of chrom
ш у т variation appear comple
osomal variation are
re counts are needed in the Australian Patersonia, the
and the н tribe гаа for all of which ancestral base number remains uncertain and patterns of
e particularly complex in /ris rocus and await
South данынан нна ли Orthro-
Chromosome cytology in the petaloid monocot
family Iridaceae is unusually varied in features of
the karyotype including basic number, ploidy level,
overall size, and chromosome arm ratios. It is thus
an important factor in considerations of the system-
atics and evolution of this family of some 1750 spe-
cies in ca. 78 genera (Goldblatt, 1990a, 1991). Ir-
idaceae are consequently comparatively well known
chromosomally. Including several new counts pub-
lished here for a range of genera and species, some
until now poorly or not at all known cytologically,
there are now counts available for at least half the
species in the family, including a good sampling of
three of the four subfamilies currently recognized:
Nivenioideae, Iridoideae, and Ixioideae. The fourth
subfamily, Isophysidoideae, which is monotypic, re-
mains uncounted, an unfortunate gap in our knowl-
edge because this taxon appears to occupy a basal
or near basal position in the family (Goldblatt,
1990a; Chase et al., 1995 and unpublished). An-
cestral chromosome numbers can be suggested for
almost all genera, and patterns of chromosomal
variation within genera, and sometimes among re-
lated genera, now seem to be reasonably well un-
dersto
Here, patterns of chromosomal variation are in-
terpreted against a background of the systematics
and morphology of the Iridaceae. We also draw at-
tention to those genera that are uncounted or poorly
known and require further investigation in order to
interpret the variation in their karyology. As part of
our review of chromosome number and morphology
of the family, we also review the available data on
genome size.
Methods used are described elsewhere (Goldblatt,
1990a; Goldblatt et al., 1993; Goldblatt & Takei,
1993) and invariably involved squashes of root tips
or shoot apices, pretreated in a variety of ways.
port for this study by grants BSR 85-00148 and 89-06300 from the U.S. National Science Foundation is
cue acknowledged. We also
Bous Dillon, W. Hahn, J. C. Manni
thank E those who have contributed seeds or plants for our ies including M.
g. M. Spurrier, R. Ornduff, J. Solomon, and G. Williamso
. Kru koff Сокмо of. African beds. Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166,
U.S.
2 Biological Institute, Oita University, Oita 870-11, Japan.
ANN. Missouni Bor. GARD. 84: 285-304. 1997.
286 Annals of the
Missouri Botanical Garden
RESULTS Iridoideae, nevertheless, differ considerably in
Original counts are presented in Table 1, accom-
panied by voucher information. Karyotypes are not
described individually; instead, results are illus-
trated and notable karyotypic details are described
under Discussion.
DISCUSSION
PATTERNS OF VARIATION AND DETERMINATION OF
ANCESTRAL BASE NUMBERS
1. NIVENIOIDEAE
This relatively primitive subfamily of six genera
and ca. 85 species (Table 2) consists of the large
African and Madagascan Aristea, the Madagascan
saprophyte, Geosiris, three shrubby southern Afri-
can genera, and the Australasian Patersonia. De-
rived features of the subfamily (Manning & Gold-
blatt, 1991) are a binate rhipidial inflorescence, a
blue perianth that is fugacious and deliquescent in
some genera, and а 2-3-seriate outer seed integu-
ment. Aristea and the shrubby southern African
Klattia, Nivenia, and Witsenia have a common base
number, x = 16 (Table 2), and small chromosomes
(Figs. 1-5) with a size range 0.8-2.5 jum. Pater-
sonia is inadequately sampled (Goldblatt, 1979a)
and also has small chromosomes (Fig. 6), compa-
rable in size to those of Aristea. Diploid numbers
of 2n = 62, 42, and 22, the latter the most frequent,
have been reported in the 6 species of Patersonia
counted out of a total of 20 in the genus. From these
counts we assume the most likely ancestral base
number for Patersonia may be x = 11. It seems
premature to speculate about how the base number
in Patersonia relates to x = 16 in the remaining
genera of Nivenioideae. Patersonia has been sug-
gested to be nested within Nivenioideae and to be
the sister genus to the lineage that includes Niv-
enia, Klattia, and Witsenia (Manning & Goldblatt,
1991). If x = 11 is correct, then ancestral base
number in Patersonia must be derived. Additional
counts in the genus are needed before the situation
here is clarified. Geosiris is unknown cytologically.
2. IRIDOIDEAE
Iridoideae share with Ixioideae specialized xy-
lem vessels in the roots with simple perforations,
likely a specialized condition (Cheadle, 1963;
Goldblatt et al., 1987), similar and putatively de-
rived anther endothecial thickenings (Manning &
Goldblatt, 1990), and a more specialized spectrum
of flavonoid compounds than in Nivenioideae
(Goldblatt, 1990a). Possibly a clade, Ixioideae and
many features, including pollen grain morphology,
leaf anatomy, and inflorescence structure (Gold-
blatt, 1990a, Goldblatt et al., 1991). Iridoideae are
defined largely by a fugacious perianth and hollow,
terminally stigmatic styles that divide below the an-
thers. These two morphological synapomorphies are
supported by the presence of an unusual chemical
character, free meta-carboxy amino acids (Gold- |
blatt, 1990a), not present elsewhere in the family.
In three of the four tribes of Iridoideae the style
branches are further specialized in being thickened
or compressed, and the apices are elaborated into
crests or similar structures (Goldblatt, 1990a; Ru-
dall, 1994).
Of the four tribes (Table 2), Sisyrinchieae are
basal, while the remaining Irideae, Mariceae, and
Tigridieae may be a monophyletic lineage defined
by several morphological and chemical synapomor-
phies (Goldblatt, 1990a). Chromosomes of many
Sisyrinchieae, including Libertia and Orthrosan-
thus, are small, generally 1-2 jum long (Figs. 7-9),
and comparable in size to those of Nivenioideae. In
Sisyrinchium, the largest genus of the tribe, the
chromosomes are usually somewhat larger (Kenton
& Heywood, 1984), sometimes substantially so in
Olsynium (Kenton et al., 1987, reported as Phaio-
phleps and Sisyrinchium sections Eriphilema and
Nuno). Bobartia, usually regarded as the only Af-
rican genus of Sisyrinchieae, has been transferred
to Irideae as a result of molecular data (Souza-
Chies et al., 1997). Its comparatively large chro-
mosomes and x = 10 match the plesiomorphic fea-
tures of the latter tribe. i
Basic chromosome number in Libertia is Orr
dently x = 19, established for four Australasian
species (Hair et al., 1967), but the only South
American species counted has 2n = ca. 72 (Kenton
& Heywood, 1984), making this possibly tetraploid
if it, too, is based on x = 19. Base number for
Libertia in South America still needs to be verified.
In the only other American and Australasian genus
of Iridaceae, Orthrosanthus, base number is uncer
tain. We have established 2n = 84 in the Austra-
lian O. polystachyus (Fig. 7), also reported as 2n =
40 (Kenton & Heywood, 1984). In the South Amer-
ican O. chimboracensis and O. acorifolius (Figs. 8-
9) we found 2n = 54. This accords with an earlier
count of n = 27 for O. chimboracensis, based on à
different collection (Goldblatt, 1982a). Kenton and
Heywood (1984), however, have reported 2n — 50
in both O. chimboracensis and O. exsertus (as var
exsertus). The difference in the counts seems to
concern the identity of four small chromatin bodies,
regarded as satellites by Kenton and Heywood, but
===
Volume 84, Number 2 Goldblatt & Takei 287
1997 Chromosome Cytology of Iridaceae
Table 1. New chromosome numbers in Iridaceae (Figs. 1-46) with collection data. Unless stated to the contrary,
vouchers are at the Missouri Botanical Garden (MO)—herbarium acronyms are abbreviated according to Holmgren et
al. (1981) and are indicated in parentheses after the collection number. Species are arranged alphabetically within the
subfamilies and tribes recognized by Goldblatt (1990a).
Diploid
Species number 2n Collection data
Subfamily Nivenioideae
Aristea abyssinica Pax 64. Zimbabwe, Nyanga, Goldblatt 9070
A. anceps Eckl. ex Klatt 32 S. Africa, E. Cape, near "аи : mouth, Goldblatt 9053
A. angolensis Baker 32 Malawi, Zomba Mt., Goldblatt 4528
А. juncifolia Baker 32 S. Africa, W. Cape, Cape Point eene Goldblatt 9056
A. angustifolia Baker 32 Madagascar, Andringitra, Goldblat
Patersonia sericea R. Br. ex 22 Australia, NSW, Clarence, Hind са ud
Ker Gawl.
Subfamily Iridoideae
Tribe Irideae
v: deii (N.E. 20 S. Africa, N. Cape, Richtersveld, Viviers s.n.
Br.
) Go
Dietes hisce: (F. 60 Lord Howe Island, Pickard 3377 (NSW)
Muell.) Klatt
Ferraria glutinosa (Baker) 40 Namibia, near Rehoboth, Goldblatt & Manning 8808
Rendle
Tris tridentata Pursh 40 U.S.A., Florida, Leon Co., Henderson 92-135
Homeria hantamensis Gold- 12 S. Africa, N. Cape, Hantamsberg, Goldblatt & Manning 10355
blatt & J. C. Manning
Moraea anomala G. J. Lewis 20 S. Africa, W. Cape, Elim, Goldblatt 361 (BOL)
. bella Harms 12 Tanzania, Ufipa, Goldblatt et al. 8289
M. bipartita L. Bolus 12 S. Africa, W. Cape, Goldblatt 4948
M. callista Goldblatt 12 Tanzania, Kitulo Plateau, Lovett & Congdon 2905; Lusitu Ridge,
ombe, Spurrier s.n. (no voucher)
M. carsonii Baker 12 на Copperbelt, Goldblatt 7544
M. deserticola cael 12 S. Africa, N. Cape, Knersvlakte, Goldblatt 9128
M. elsiae Goldblat 20 S. Africa, W. Cape, Kenilworth, Esterhuysen 32358 (BOL)
M. fergusoniae L. als 12 5. Africa, W. Cape, Caledon district, Snijman 820 (NGB); Mier-
kraal, Bredasdorp, Goldblatt 6186; E. of Heidelberg, Goldblatt
4956; Bontebok Park road, Goldblatt 4316A
M. fugax (D. Delaroche) Ker 10 S. Africa, N. Cape, Wallekraal, Goldblatt s.n. (no voucher)
Gawl.
M. incurva G. J. Lewis 12 S. Africa, W. Cape, near Tulbagh, Hansford 1 (NBG)
M. namibensis Goldblatt 20 Namibia, Süd Witputs, Lavranos 21257
M. neglecta G. J. Lewis 20 S. Africa, W. Cape, Klaasjagersberg, Goldblatt 5268
M. tulbaghensis L. Bolus 12 S. Africa, W. Cape, near Tulbagh, Burgers 2756 (STE)
M. schimperi (Hochst.) Pic. 12 Zaire, Shaba, Schaijes 5108
Serm
M. dole. Ker Gawl. 12 S. Africa, W. Cape, Goldblatt 5865
Tribe Mariceae
Neomarica caerulea (Ker 32 ex hort., origin unknown, Goldblatt 5930
Gawl.) Sprague
N. cf. northiana (Schneev.) 18 Paraguay, Amambay, Cerro Corá, Solomon et al. 6950
Sprague : i :
Trimezia martinicensis (Jacq. 80 Venezuela, Anzoátegui, Serranía de Turimiquire, Davidse & Gon-
Herb z 19456; Venezuela, Mérida, Páramo de Mucubaji, Berry
Tribe Sisyrinchieae
Orthrosanthus polystachyus 84 Australia, ex Hort. Bot. Melbourne, Goldblatt s.n.
Benth.
288
Annals of the
Missouri Botanical Garden
Table 1. Continued.
Diploid
Species number 2n Collection data
O. acorifolius (Kunth) Raven- 54 Venezuela, Mérida, Grifo & Hahn 361 (BH)
na
O. chimboracensis (Kunth) 54 Venezuela, Dorr et al. 5044 (NY)
Baker
Sisyrinchium micranthum 48 Nicaragua, Henrich s.n.
Cav.
Tribe Tigridieae
Alophia drummondii (Gra- 28 U.S.A., Texas, Bastrop Co., Lee sub Goldblatt s.n.
ham) R. C. Foster
Calydorea azurea Klatt 28 Uruguay, Treinta y Tres to Tacuarembó, Castillo 1146 sub Gold-
n.
C. amabilis (Ravenna) Gold- 14 Argentina, Entre Ríos, Concepción del Uruguay, Goldblatt s.n.
blatt & Henrich
C. pallens Griseb. 28 Argentina, Córdoba, Cerro Colorado, Goldblatt s.n.
C. xiphioides (Poepp.) Espino- 42 Chile, Coquimbo, Hoffmann s.n
sa
Cipura paludosa Aubl. 14 Brazil, Maranhao, near Imperatriz, Plowman et al. 9305
Cobana gua 1 28 Honduras, Nelson s.n. (no voucher)
(Standl.) Ravenna
Cypella fucata Ravenna 14 Uruguay, Maldonado, Punta del Este, Castillo s.n.
C. herbertii subsp. brevicrista- 14 Uruguay, Artigas, northern suburbs, Castillo s.n. (no voucher)
L nna
subsp. wolfheugelii (Hauman) 14 Argentina, Buenos Aires, Cerro Ventana, Lamberto € Méchel s.n.
Ravenna (BB 3881)
C. е subsp. opalina 14 Argentina, Misiones, Garruchos, Castillo s.n. (FAA)
гота ет (Сп- 14 Bolivia, Тагіја, Arce, Solomon 9972
seb.) Rav
E. foliosus (Kunth) Ravenna 14 Peru, Dillon 4514 (F)
Herbertia lahue Molina 42 Chile, Ornduff 9153 (UC); U.S.A., Louisiana, Shreveport, Hei-
amp, s.n.
H. aff. lahue 14 Brazil, Rio Grande do Sul, Rosario do Sul, Castillo s.n.
H. pulchella Sweet 42 Uruguay, Maldonado, Punta del Este, Castillo sub Goldblatt s.n.
H. quareimana Ravenna 28 Brazil, Rio Grande do Sul, Uruguaiana, Goldblatt s.n
Subfamily Ixioideae
Tribe Watsonieae
Micranthus junceus (Baker) 20 S. Africa, W. Cape, Cape Point Reserve, Goldblatt s.n. (no
N.E. Br. voucher)
Thereianthus нагар var. li- 20 S. Africa, W. Cape, Cape Point, Goldblatt 5400
nearifolius G. J. Le
Watsonia dubia Eckl. e ex a 18 S. Africa, W. Cape, Malmesbury, Goldblatt 8708
W. hysterantha Mathews & 1. 18 1B S. Africa, W. Cape, Langebaan, Snijman 71 (BG)
Bolus
W. minima Goldblatt 18 S. Africa, W. Cape, near Greyton, Goldblatt 8047
Tribe Ixieae
Crocus longiflorus Raf. 28 Italy, Sicily, Eloro, Goldblatt 5073
Dierama inyangense Hilliard 20 Zimbabwe, Nyanga, Clarke s.n. (no voucher)
Geissorhiza са ма Goldblatt 26 S. Africa, W. Cape, near Grayton, Goldblatt 8680
G. foliosa Baker 26 S. Africa, W. Cape, Strawberry Hill, Goldblatt 7948
G. roseoalba (G. J. Lewis) 26 S. Africa, E. Cape, Pootjeshoogte, Vlok 1663
ldblatt
Gladiolus aquamontanus 30 S. Africa, W. Cape, Rust-en-Vrede, Vlok s.n.
Goldblatt & Vlok
PSone e NN
Volume 84, Number 2
Goldblatt & Takei
Chromosome Cytology of Iridaceae
289
С. Manning
Table 1. Continued.
Diploid
Species number 2n Collection data
G. inandensis Baker 30 S. Africa, Kwazulu-Natal, Inchanga, Goldblatt & Manning 10057
G. longicollis Baker 30 S. Africa, E. Cape, Naudes Nek, Goldblatt & Manning 9542;
Mpumalanga, Long Tom Pass, Goldblatt & Manning 9823
G. ochroleucus Baker 30 S. Africa, E. Cape, East London, Batten 1123
G. oppositiflorus Hook.f. 30 S. Africa, E. Cape, near Kei Road, Batten s.n.
G. permeabilis subsp. edulis 28 S. Africa, E. Cape, Rhodes, Batten 1118
(Ker Gawl.) Oberm.
G. phoenix Goldblatt € J.C. 30 S. Africa, W. Cape, Bain’s Kloof, Goldblatt & Manning 9423
Manning
G. pole-evansii Verdoorn 30 S. Africa, Mpumalanga, Denilton, Goldblatt & Manning 9808
G. pretoriensis O. Kuntze 30 S. Africa, Gauteng, Pretoria, Goldblatt & Manning 9799
G. saundersii Hook. | 30 S. Africa, E. Cape, Naudes Nek, Batten 1114
G. stefaniae Obe 30 S. Africa, W. Cape, Montagu, Manning s.n. (NBG)
G. sulcatus Gol Tanzania, Njombe, Lusitu Ridge, Spurrier 804
G. wilsonii ae а 30 S. Africa, E. Cape, Nahoon, Batten s.n.
G. woodii rog 30 S. Africa, Mpumalanga, Belfast, Goldbaltt & Manning 9811
Hesperantha baurii Baker 24 Swaziland, near Mbabane, Goldblatt
H. luticola Goldblatt 26 + 1B S. Africa, N. Cape, Hantamsberg, Goldblatt s.n. d voucher)
H. radiata (Jacq.) Ker Gawl. 26 Lesotho, Blue Mountain Pass, Schmitz 8531 (ROM
Ixia brevituba G. J. Lewis 20 9. Ao. W. Cape, Roggeveld, Goldblatt & шей 8645
І. latifolia var. ramulosa G. J. 20 S. Africa, N. Cape, Namaqualand, Goldblatt 9241
wis
1. paniculata D. Delaroche 20 8. Africa, W. Cape, Piekenierskloof, Goldblatt 5117
Melasphaerula graminea (L.) 20 S. Africa, W. Cape, near Soutpan, Goldblatt s.n. (no voucher);
N.E. Br. near Clanwilliam, Goldblatt 3832
Radinosiphon leptostachya 30 Malawi, Golomoti Mts., Boussard s.n.; Mt. Malosa, Goldblatt
(Baker) N.E. Br. 9201
Romulea cameroonensis Baker 26 Malawi, Zomba Plateau, Goldblatt 9094; Snijman s.n.
(7 R. campanuloides Ba-
ker
R. doe Es ca.78 Rwanda, Virunga Mts., D'Arcy 7887
R. c.f. linaresii Parl. a.39 Greece, Athens, Goldblatt 5051
Rad puce n Gold- 20 S. Africa, W. Cape, Nardouw Pass, Goldblatt 6162
9; prior (G. J. Lewis) 40 5. Africa, W. Cape, without precise locality (no voucher)
Goldb
8. ае subsp. meteler- 20 S. Africa, W. Cape, Algeria, Goldblatt s.n.
kampiae (L. Bolus) Gold-
blatt
Syringodea bifucata de Vos 12 S. Africa, Gauteng, farm Sandfontein, Davidson 3108
Tritonia chrysantha Fourc. 22 S. Africa, E. Cape, Alicedale, Bayliss
T. florentiae (Phill.) Goldblatt 20 S. Africa, W. Cape, Ceres Karoo, Goldblatt 3912
T. laxifolia Benth. 20 Tanzania, Makambako, Spurrier 601
T. squalida (Aiton) Ker Gawl. 22 S. Africa, W. Cape, S. of Riversdale, Goldblatt 5090
Xenoscapa fistulosa (Klatt) 22 S. Africa, N. Cape, Richtersveld, Williamson & Hamer 4428
Goldblatt & J. C. Manning
X. uliginosa Goldblatt & J. 22 S. Africa, N. Cape, Namaqualand, Goldblatt 9244
loid. This does not explain the counts of both 2n
— 40 (by Kenton & Heywood) and 84 (by us) in
the same Australian species. This species also dif-
fers from the South American species of the genus
which we consider to be chromosomes. Ancestral
base number for the genus is probably x — 9, not
x = 10 as suggested by Kenton and Heywood, mak-
ing the species counted in South America hexap-
Annals of the
Missouri Botanical Garden
Ties 2. Chromosome numbers in genera of Nivenioideae and Iridoideae. Data are largely from Goldblatt (1971,
82a) who included reviews of previous reports for the subfamilies, and Kenton & Henmed (1984). Sources other
а Goldblatt (1971) are е below.
out of the total in the genus; parenthetical entries in the base-number odit
Numbers in parentheses after the genera
mn indicate secondary hase numbers.
Presence of triploids, unless known to be stabilized, and B chromosomes are not included. Genera are arranged
alphabetically in subfamilies and tribes according to Goldblatt (1990a) excepting that Bobartia has been moved from
Iridoideae: Sisyrinchieae to Irideae in light of molecular data (Souza-Chies et al., 1997; M. Chase, pers. comm.).
Basic and diploid numbers
Genus x 2n Reference
Nivenioideae
Aristea (12/52) 6 32, 64 Goldblatt, 1995
Geosiris (0/1) uncounted
attia (1/3) 32
Nivenia (4/10) 16 32
Patersonia (6/20) — ?11 22, 42, ca.62 Goldblatt, 1979a
Witsenia (1/1) 16 32
Iridoideae
Tribe Sisyrinchieae
Diplarrhena (2/2) 16
Libertia (5/8) 19
Olsynium (5/11) 10(9, 8, ?11)
incl. Phaio-
phleps)
Orthrosanthus (4/8) ?9(?12, 10)
Sisyrinchiu 9(8, 5, 17)
са.
Solenomelus (1/2) 28
Tapeinia (0/1) uncounted
Tribe Mariceae
Neomarica (4/12) ?9(8)
Pseudotrimezia uncounted
(
Trimezia (5/20) ?14(13, 10 or 20)
Tribe Tigridieae
Ainea (0/1) uncounted
Alophia (3/5)
Calydorea (4/10) 7
(incl. Catila)
Cardenanthus (0/8) uncounted
7
1 1
Cypella (5/20) 7(5)
Ennealophus (3/5) 7
(incl. Tucma
Fosteria (1/1) 14
Gelasine (2/5) 7(6)
Herbertia (4/8) 7
(=Trifurcia)
Kelissa (0/1) uncounted
Mastigostyla (3/16) 1
32
38, 76, 112
20, 22, 40, 60, 18, 96
54, 40, 84, 50
18, 36, 72, 16, 32, 48, 64, 96,
34
18, 16, 32
28, 26, 52, 40, 60, 80
28
14, 28, 42
14, 28'
28
14, 28, 10
12:
14
28
14, 12
14, 28, 42
28
PF —Á————— e
Kenton & Heywood, 1984
Hair et al., 1967; Kenton & Hey-
(as Phaiophleps and Sisyrinchium
sects. Nuno and Eriphilema, e.g.
T et al., 1986; Rodriguez,
u nton d Heywood, 1984
Rudall et al., 1986; Kenton et al.,
1987
Rudall et al., 1986
Goldblatt, 1982a; Kenton & Heywood
1984
Goldblatt, 1982a; Kenton & Heywood
984
Goldblatt, 1982a; Kenton & Heywood
1984
Goldblatt, 1982a; Kenton & Heywood
1984
Goldblatt & Henrich, 1987
Ravenna, 1974
Goldblatt & Snow, 1991
Kenton & Heywood, 1984; Martinez
Molseed, 1970
Kenton & Rudall, 1987
Winge, 1959; Kenton & Heywood,
1984
Huynh, 1965; Martinez & De Azkue.
1987
d === AN
Моште 84, Митбег 2 Goldblatt & Takei 291
1997 Chromosome Cytology of Iridaceae
Table 2. Continued.
Basic and diploid numbers
Genus х 2п Кеѓегепсе
Nemastylis (3/5) 7 14, 28, 56 Goldblatt, 1982a
Onira (0/1) uncounted
Sessilanthera (2/4) 14 28 Cruden, 1972
Tigridia (14/35) 7 14, 28 Molseed, 1970; Kenton & Heywood
1984
Tribe Irideae
Barnardiella (1/1) 10 20 Goldblatt, 1977
Belamcanda (1/1) 16 32, 64 Nakajima, 1936; Hsu, 1971
Bobartia (8/14) 10 20 Strid, 1972
] 10 20, 40, 60 Goldblatt, 1981а
Ferraria (10/10) 10 20, 40, 60 de Vos, 1979
Galaxia (14/15) 9(8, 7, 6) 18, 16, 14, 12 Goldblatt, 1979c, 1984a
Gynandriris (7/9) 6 12, 24 Goldblatt, 1980b
Hermodactylis (1/1) 10 20 Simonet, 1932
Hexaglottis (6/6) 6(7, 5) 12, 14, 10 Goldblatt, 1987
отепа (34/34) 6(5, 4) 12, 24, 10, 9, 8 Goldblatt, 1980a, 1981b
Tris (185/ca. 225) ? € diploid numbers occur in this large genus; both polyploidy and dysploidy are fre-
ent)
Moraea (95/130) 1009, 8,7,6,5) 20, 40, 18, 16, 12, 24, 48, 10 Goldblatt, 1976, 1986a, 19866
Pardanthopsis (1/1) 16 22 Simonet, 1932
Roggeveldia (2/2) 6 12, 24 Goldblatt, 1992
Eres & Snow (1991) have shown that the plants associated with the counts of 2n — 12(-14) for Cipura paludosa
— 14 for Eleutherine bulbosa (Sharma & Talukdar, 1959) were confused with one another. Counts of 2n — 14
are d C. paludosa, while those of 2n = 12(-14) are for E. bulbosa. The count of 2n = 14 for the latter reported by
Rao (1969) is probably also for C. paludosa.
Figures 1-9. Mitotic Basi in Iridaceae subfamily | — new and Patersonia) and олар чр
a (Orthrosanthus).—1. Aristea abyssinica (2n = 64).—2. A. anceps (2n = 32).—3. A. angolens
32).—4. A. june ifolia (2n = 32).—5. A . angustifolia e = 32)—6 ш ама sericea (2n = 22).—7. TD E
раба coal 84).—8. 0. acorifolius (2n = 54).—9. 0. chimboracensis (2n = 54). Vouchers as given in Table
Scale bar, 1
292
Annals of the
Missouri Botanical Garden
in having a bimodal karyotype with the larger chro-
mosomes 2-2.3 jum long, and the smaller 0.5—1.2
um long, a pattern also noted by Kenton and Hey-
wood. More counts are needed in Orthrosanthus, for
both Australian and South American taxa. Chro-
mosomal characters will likely be relevant to the
systematic and phylogenetic understanding of the
genus.
We include only one original count in Sisyrin-
chium (Table 1), 2n = 48, for the annual S. mi-
cranthum. This agrees with past reports for the spe-
cies. Ancestral base number for Sisyrinchium is
most likely x = 9. Subgenus Echthronema has spe-
cies based on either х = 9, 8, or 17, but most
species of subgenus Sisyrinchium have x = 8 (Ken-
ton & Heywood, 1984) and one annual species has
n = 5. The related Olsynium (including Sisyrin-
chium subg. Eriphilema) has base numbers of x =
10, 9, 8, and perhaps 11, but the ancestral base
number may well be x = 10. We conclude that the
ancestral base number for Sisyrinchieae may be x
= 10, based on the pattern so far reported in the
tribe (Table 2). The Australian Diplarrhena, cur-
rently included in Sisyrinchieae, has large chro-
mosomes and x = 16. On the basis of chromosome
size it accords better with lrideae. Number alone
provides no information about its relationships.
Large chromosomes and a base number of x =
10 have been postulated to be ancestral for Irideae
(Goldblatt, 1990a), and Bobartia (usually included
in Sisyrinchieae), Dietes, and Ferraria (Table 2)
conform to this pattern. The count here of 2n = 60
in D. robinsoniana (Table 1), the Lord Howe Island
endemic and only non-African species of Dietes,
conflicts with an earlier count of 2n = 20 in the
species (Goldblatt, 1979a). It seems unlikely that
D. robinsoniana is heteroploid. We question the
identity of the plant counted earlier under that
name, unfortunately without a voucher. The closely
related D. bicolor from South Africa is also poly-
ploid, 2n = 40 (Goldblatt, 1971). The four other
species of Dietes are diploid, 2n = 20 (Goldblatt,
198 1a).
Our count here of 2n = 40 for the tropical Af-
rican Ferraria glutinosa confirms a report that the
species is tetraploid in Namibia (Goldblatt, 1979).
Interestingly, a diploid population of the species
has been reported from Zaire (De Vos, 1979), thus
establishing that the species is heteroploid. Of the
remaining nine species of Ferraria, seven are dip-
loid, F. divaricata has diploid and tetraploid sub-
species, and two more have both tetraploid and
hexaploid populations. Ferraria is one of the few
ican genera of Iridaceae in which polyploidy ap-
pears to have played a significant role in its evo-
lution.
Ancestral base number in the large African ge-
nus, Moraea, is also x = 10 (Goldblatt, 1976,
1986a). Our counts for numerous species of the ge-
nus with this base number show some degree of
variation in their karyotypes (Figs. 11-13). Notably
in subgenus Visciramosa, M. elsiae has a distinctive
karyotype in which 5 of the 10 chromosome pairs
are telocentric (Fig. 13). The other four species of
subgenus Visciramosa also have this unusual and
presumably derived karyotype (Goldblatt, 1971 and
unpublished).
Dysploid reduction in at least four lineages of
Moraea has resulted in the independent evolution
of the secondary base of x = 6 (Goldblatt, 1986a).
Counts reported here from subgenus Vieusseuxia,
section Polyanthes, and subgenus Grandiflora pro-
vide examples of the three main groups with x =
6 (Figs. 15-18, 21-24). In a fourth lineage, M. fu-
gax (subg. Moraea) offers a remarkable example of
dysploid reduction from x = 10 (presumably the
ancestral base) to n = 8, 7, 6, and 5 within a single
polymorphic species (Goldblatt, 1986b). The count
here of 2n = 10 (Fig. 19) is representative of one
of several cytotypes in the species.
In subgenus Vieusseuxia, karyotypes typically
have marked size variation (Figs. 21, 22, 24), the
shortest chromosome pair, 7-8 jum long, being
about two-thirds to about half as long as the longest
pair, 12-14 um. One of the two longest pairs 1s
always more or less metacentric and the other is
acrocentric. A small satellite is consistently present
on one of these long pairs. Chromosomes of sub-
genus Grandiflora are consistently larger than else-
where in the genus and 10-14 ¡um long. In Moraea
schimperi, typical of the subgenus (Fig. 23), the
chromosomes are acrocentric to nearly telocentric
and a small satellite is located on one of the nearly
telocentric chromosome pairs. Based on measure-
ment of total chromosome length, we estimate that
genome size in M. schimperi is some 20-30% larget
than in subgenus Vieusseuxia or subgenus Moraea.
In section Polyanthes, e.g., M. bipartita, M. cal-
chromosome size variation is modest.
mosomes are 7-12 рт long and all are more or
less acrocentric. This pattern resembles closely that
of the karyotypes of the southern African genera,
Hexaglottis, Homeria, and Roggeveldia (Goldblatt,
1980a, 1987). The karyotypic similarity par
these three taxa suggests common ancestry wi
Moraea sect. Polyanthes. This possibility should be
considered in future studies of the phylogeny Г
Irideae. We have noted an unusual condition 1n ™
QA — ай
Volume 84, Number 2
Goldblatt & Takei 293
Chromosome Cytology of Iridaceae
Figures 10-20. "weg metaphase in
m oraea anomala — 20).—12. M.
rien carsonii (2n =
E
—1 —16. |
bec ie (2n = 12).—19. M. fugax (2n = "N —20. эз ы (2п =
10 pm
са за. One population examined has a karyotype
consisting of ma tching pairs (Fig. 46A), but another
is Se nei sal (Fig. 46B) and has
one long metac chromosome and one very
short metacentric. arena em in this tropical
M. elsiae (2n = 20).—14. Barnardiella spiralis
ee (2n = 12).—17. M. bipartita (2n = 12).—18. M.
12). Vouchers as given in Table 1. Scale bar,
Iridaceae — er tribe Irideae.—10. Iris tridentata (2n = 40).—
ini ibe А аи —13.
African species are needed to elucidate the signif-
icance of this situation.
Unusual in Iridaceae, the only three species ~
Homeria that exhibit хи рел Н. pallida, п =
and 4, and H. tenuis and Н. flavescens, 2п = ~
Annals of the
Missouri Botanical Garden
Figures 21-31. Mitotic metaphase in Iridaceae subfamily Iridoideae tribes Irideae чару x Maries (Neomarin
and Trimezia), y Tigridiese (€ ypella, Calyaoren, and Herbertia). M. tulbag
(2л = 12).—23. M. schimperi (2n = 12).—24. M. unguiculata Qn = 12).—25. Neomarica ier ts 32). —»
V. ef. northiana (2л = 18).—2
| Final ia c ensis (2n = 80).—28. Cypella herbe row и Фа“
репа her пи subs sp. Y
14). —29. Calydorea pallens а = 28).—30. Herbertia lahue (2n = 42).—31. H. pulchella (2n = 28). Vouchers 29
given in Table 1. Scale bar, 10 ат
Volume 84, Number 2
1997
Goldblatt & Takei
Chromosome Cytology of Iridaceae
9, and 8, are complex (
1980c). The complex heterozygosity in these spe-
cies is associated with dysploidy, itself evidently
unequal reciprocal ion. In
specialized Н. has popu-
display a dysploid with
a € OS nae
bt the bo he pre oer
(Goldblatt, 1979.
EN ilio we
genera of Iridaceae, is
base number x = 20 is frequent in the less spe-
cialized iris, but subgenus /ris al-
most certainly has х = 12 ( idy,
the most common numbers are n = 12, 11, 10, and
9). Based on an analysis of the many counts avail-
able in the (some-
times treated as a genus, Iridodictyum) has x = 10
(n = 10, 9, 8); Xiphium may have « =
18 (n = 18, 17, 16, 14); and subgenus Scorpiris
(syn. Juno) appears to have x = 12 (n = 12, 11,
10, 9, and even 13 and 15).
The single
American species was available
to us. The count of 2n = 40 and large chromosomes
3.3-6.3 pm long (Fig. 10) are consistent with sev-
eral species of section Limniris, common numbers
for which аге 2n * 44 and 40. However, our count
differs from the available reports for both /. setosa
Se ee ar Mt
species. specialized T.
kii has п = 26 and T. martii has n = 14, and the
vegetatively specialized T. fosteriana has n = 13,
More counts are needed in Trimena before it will
be possible to make sense of the pattern bere. Ten-
tatively, we suggest that x = 14 may be the ances-
296
Annals of the
Missouri Botanical Garden
Alophia drummondii (2n = 28).—
malensis
Figures 32—44. Mitotic e in Iridaceae subfamily iy tribe Tigridieae and mem ides 32.
28).
3. Cobana guatem
= 30 | 42.
Romulea cameroonensis (2n
laxifolia ac
tral base for the genus. Cycles of polyploidy and
dysploid reduction then would best explain the re-
maining base numbers.
The related Neomarica, included in Trimezia by
Ravenna (1976), is specialized in having a winged
aerial stem and branches crowded apically. Neo-
marica northiana has 2n = 18 (Fig. 26), a count
confirmed here, as does N. cf. vittata (Kenton $
Heywood, 1984). Reports for N. gracilis include 2n
— 18, 40, and 32. Some of these counts are almost
certainly from misidentified plants. The Central
American N. variegata (often confused with N.
gracilis in the literature), specialized in its red,
fleshy seeds, has 2n — 16 and N. caerulea has 2n
— 32 (Fig. 25). The ancestral base number for the
.— 936. Xenoscapa fistulosa is = 22)—37. X.
= 20).—39. Ixia latifolia (2n = 20 ).—40. S
= 26).—43. Gladiolus голи a (2n =
20). Vouchers as given in Table 1. Scale bar, 10 pm
uncifolius (2 20).—35.
20).—38. Dierama
—A1. Radinosiphon leptos-
30).—44. Tritonia
paraxis parvifolia (2n =
genus is most likely x = 9, and x = 8 is evidently
derived by dysploid reduction. The variation al-
ready evident in imezia and Neomarica
makes it essential that chromosome cytology be in-
cluded in future systematic and phylogenetic stud-
ies of these genera.
Unlike Mariceae, the pattern in the other exclu-
sively American tribe Tigridieae is reasonably cleat
(Figs. 28-31). The ancestral base number appears
to ђе х = 7, and this is the only base number in
most genera of subtribe Cipurinae counted, and the
only base in the other subtribe, Tigridiinae (Table
2). The base of x = 7 is strongly conserved despite
a fair degree of karyotypic diversity. Bimodality is
common and is also conserved despite variation in
,
Моште 84, Митбег 2
1997
Goldblatt & Takei 297
Chromosome Cytology of Iridaceae
ho ЧИ
E IS ИС]
e 45. Mitotic metaphase in Ennealophus foliosus (2n = 14) ee cae ra —Figure 46. Metaphase
cnomosome of Moraea callista (2n = 12) "iHe и. A: populat
on from Lusitu Ridge with normal chro-
e complement; d population from the Kitulo Plateau showing per heterozygosity. Vouchers as given in
Table 1 1. Scale bar, 5 р
chromosome and karyotype size (Kenton et al.,
1990). Among seven species of Cypella and Hes-
peroxiphion (a genus sometimes included in Cypel-
la), all with х = 7, chromosome size varies consid-
erably. A more than three-fold difference in total
DNA among the species has been demonstrated by
Kenton et al. (1990). Yet bimodality and relative
sizes of the chromosomes within the karyotypes of
each species are maintained.
A particularly distinctive karyotype is present in
Ennealophus (including Tucma) (Fig. 45). All three
species counted (of a total of five) have one long
metacentric pair, ca. 7 jum long, and six much
shorter pairs, ca. 1.5 jum long, one of which has a
satellite larger than the short chromosome to which
it is attached (see also Kenton & Heywood, 1984).
Cypella plumbea, sometimes segregated as Phal-
ocallis (Ravenna, 1977), has 2n — 10 and is ex-
ceptional here (counts for the few other species that
have been placed in Phalocallis would be interest-
ing). The two species of Eleutherine, an apparently
very derived genus, have x — 6 (Goldblatt & Snow,
1991) as does Gelasine intermedia (= G. azurea).
The latter is an autogamous, complex heterozygote
(Kenton & Rudall, 1987) with an apparently de-
rived floral morphology. The only other species of
Gelasine counted has 2n — 14 (Ravenna, 1984).
Polyploidy is common in Tigridineae, and a few
genera are exclusively polyploid. In Tigridia most
species so far counted (e.g., Figs. 32, 33) have x —
14 (Table 2), including all those from Central
America and Mexico (Molseed, 1970; Kenton &
Heywood, 1984), but at least two South American
species have n = 7. The only counts in Alophia
(the species of which are sometimes confused no-
menclaturally with those of Herbertia), Fosteria,
Cobana, and Sessilanthera are 2n — 28. At least
the three latter, all small genera closely related to
Central American Tigridia, and probably nested
within it, may be exclusively tetraploid. Despite
their polyploidy, karyotypes in Cobana, Alophia, Ti-
gridia, and their allies consistently have one long
metacentric chromosome pair and one submetacen-
tric pair. In Cobana (Fig. 33) it is possible to tell
by careful observation that the long and apparently
submetacentric chromosome pair is actually acro-
centric with a long satellite attached to the short
chromosome arm. The mismatch of the four long
chromosomes suggests early chromosomal rear-
rangement in a polyploid ancestor of the group or
a hybrid origin from parents with different karyo-
ypes.
e
3. IXIOIDEAE
Small chromosomes, usually 1-2.5 pm long
(Figs. 34—44), are plesiomorphic in this, the largest
subfamily, and are found in most genera (Goldblatt,
1971). Ixioideae are otherwise defined by several
morphological and palynological features (Gold-
blatt, 1990a; Goldblatt et al., 1991). By fusion one
or more larger chromosome pairs have evolved in
some genera, notably Lapeirousia (Goldblatt & Tak-
ei, 1993) and Crocus (Brighton, 1976; Brighton et
al, 1983). In Lapeirousia the two long chromo-
somes present in all species are 4.1—6.5 pm long,
and in most species the remaining 12-18 chromo-
somes are 1-2 рт long, thus typical of Ixioideae.
Basic numbers are generally high in the subfamily
(Table 3). Dysploidy and the development of sec-
ondary base numbers, known in a few genera, are
discussed below.
Freesia, including the species of Anomatheca
transferred here (Goldblatt & Manning, 1995),
and Xenoscapa (Figs. 36, 37) have x — 11 and
all species counted are diploid, 2n = 22 (Table
3). Similar karyotypes with x — 11 characterize
most species of Tritonia and Crocosmia, genera
that are closely allied to one another and prob-
ably closely related to Freesia (Goldblatt, 1971,
298 Annals of the
Missouri Botanical Garden
Table 3. Chromosome numbers in genera of Ixioideae. Data largely from Goldblatt (1971), who included references
to earlier studies. Other sources are referenced below.
. Numbers in parentheses after the genera indicate number of
species counted out of the total in the genus; parenthetical numbers in the base number column indicate secondary
se numbers. Presence of triploids, unless known to be stabilized, and В chromosomes are not included in the table.
Basic and diploid numbers
Genus x
2n
Reference
Tribe Pillansieae
Pillansia (1/1)
Tribe Watsonieae
10
Lapeirousia (32/40) 10(9, 8, 6, 5, 4, 3)
Micranthus (3/3) 10
Savannosiphon (1/1) 8
Thereianthus (3/7) 10
Watsonia (31/52) 9
Tribe Ixieae
Babiana (20/64) 7
(incl. Antholyza)
Chasmanthe (2/3) 10
Crocosmia (6/9) 11
Crocus (79/ca.80)
26 (uncertain: many base
40 Goldblatt & Takei, 1993
20, 18, 16, 12, 10, 8, 6 Goldblatt, 1990b; Goldblatt &
Takei, 1993
Goldblatt & Marais, 1979
Goldblatt, 1989a
numbers recorded including much polyploidy, sometimes within
a species, Mathew, 1982)
Devia (1/1) 10
Dierama (7/44) 10
Duthieastrum (1/1) 10
Freesia (13/15) (incl. 11
Anomatheca)
Geissorhiza (43/82)
Gladiolus (85/255) 15(14, 13, 12, 11)
(incl. Homoglos-
sum
Hesperantha (32/ca.
13(12)
65) (incl. Schizos-
tylis)
Ixia (14/45) 10
Melasphaerula (1/1) 10
Radinosiphon (1/1)
15
Romulea (ca.80/ca. 13 or 12(14, 11, 10, 9)
90
Sparaxis (10/13) 10
(inel. Synnotia
ngodea (7/8) 11)
Tritonia (15/28) 11(10)
Tritoniopsis (6/22) ?16(15)
(inel. Anapalina)
Xenoscapa (2/2 11
Zygotritonia (1/4) z
Goldblatt & Manning, 1990
20
20 de Vos, 1974b (as Duthiella)
22 Goldblatt, 1972 (as Anomathe-
ca), 1982
26, 39, 52 Goldblatt, 1985
30, 60, 90, 120, 28, 26, 24, Goldblatt et al., 1993
22
26, ca.50, са.72–76 Goldblatt 19845
20, 40
20
(not x = 11, Goldblatt, 1971)
30
26, 28, 24, 22, 20, 18, 30, 52, de Vos, 1972
78
20, 40
12, 22 de Vos, 1974a
22, 20, 44 de Vos, 1982
32, 30 Goldblatt, 1990c
22 Goldblatt, 1972 (as Anomatheca)
14 Goldblatt, 1989b
1991; Shneyer, 1990; Rudall & Goldblatt, 1991).
Counts here for Т. florentiae and Т laxifolia (Fig.
44), 2n = 20, are the first records for both spe-
cies and the only counts in the genus not based
on x = 11. We assume that these are examples
of dysploid reduction by fusion. Tritonia floren-
tiae is specialized in Tritonia in being acaules-
cent, or almost so, and it does not seem closely
related to T. laxifolia, indicating independent
dysploid reduction twice in the genus. Chasman-
the and Devia, evidently closely related to Tri-
tonia and Crocosmia (de Vos, 1984; Goldblatt &
Manning, 1990), have x = 10, additional exam-
ples of dysploid reduction in this alliance. Du-
Volume 84, Number 2
1997
Goldblatt 4 Takei
Chromosome Cytology of Iridaceae
299
thieastrum, which may be another specialized de-
rivative of this group, also has x =
Counts here for Dierama, Ixia, and Sparaxis, all
x = 10, are consistent with several other reports
for these genera (Goldblatt, 1971). There is, how-
ever, an important difference between their karyo-
types. In Dierama and Ixia the karyotypes are bi-
modal, with two relatively long pairs 2.3-2.6 pm
long, and eight shorter pairs 1-1.7 ¡um long (Figs.
38-39), but in Sparaxis (Fig. 44) the chromosomes
are all small, 0.9-1.6 jum long (Fig. 40). In two of
the species of Ixia in which we could distinguish
satellites, /. polystachya and I. latifolia, they were
attached to the long arms of long chromosomes (Fig.
39). How consistent this is in [xia is uncertain, but
in /. acaulis (Goldblatt & Manning, 1993) the sat-
ellites (not illustrated there) are in the normal po-
sition on the distal end of the short arms of two
long chromosomes. In Dierama and Sparaxis the
satellites are in the conventional position on the
short arm of a long chromosome pair. Earlier re-
ports of the satellites on short chromosome arms in
Ixia (Goldblatt, 1971) should be confirmed using
more refined techniques.
The difference between the karyotypes of Spar-
axis (now including Synnotia) and Dierama and
Ixia, the two genera with which it has often been
closely associated (Lewis, 1954, Goldblatt, 1969),
suggests that the affinities of Sparaxis may lie else-
where, or at least are not close to the aforemen-
tioned genera. Fundamental differences in their
leaf margin anatomy (Rudall & Goldblatt, 1991)
support this contention. The count here for S. par-
viflora (Goldblatt, 1991) is the first record of poly-
ploidy in the genus (but this should be confirmed
for additional populations).
Our counts for two populations of the monotypic
African genus, Radinosiphon, 2n = 30 (Fig. 41),
confirm past reports for this genus (Table 3), ap-
parently related to Gladiolus, which has the same
ancestral base number and similar karyotype
(Goldblatt et al., 1993). Total chromosome length
in Radinosiphon is some 10-15% less than in sev-
eral species of Gladiolus that we have examined,
but the significance of this in connection with their
possible close relationship cannot yet be assessed.
Counts here for Gladiolus, 2n = 30 for 12 species,
2n = 28 for G. permeabilis subsp. edulis, and 2n
= 90 for the tropical African G. sulcatus, include
the first reports for nine species. The karyotypes
are consistent with the majority of counts for the
genus. The count of 2n = 28 for G. permeabilis does
not accord with previous counts for the species.
Most likely the population we examined is dysploid
and not representative of the entire species or even
subspecies.
In the monotypic, western southern African Me-
lasphaerula we found 2n = 20 in two separate pop-
ulations (Table 1). We suspect that the only other
count for the genus, 2n = 22 (Goldblatt, 1971), is
incorrect. Melasphaerula is believed to be most
closely related to Geissorhiza and Hesperantha, with
which it shares woody corm tunics. The two latter
genera have an ancestral base number of x = 13
(Goldblatt, 1971, 1984b), a marked difference from
the base number in Melasphaerula. Additional
counts here (Table 3) for Geissorhiza and Hesper-
antha confirm x = 13 and the predominance of
diploidy, 2n = 26, in both genera. Our count for
the eastern southern African Н. baurii, 2n = 24,
establishes a new, and evidently, secondary base
number in Hesperantha. More counts for this wide-
spread and common southern African species and
its close relatives are needed before the signifi-
cance of our report of dysploidy can be gauged.
Our observations here for Romulea cameroonen-
sis (Fig. 42) and R. congoensis are consistent with
the small chromosome size (са. 0.9-2.2 шт) and
range of base numbers reported for the genus (de
Vos, 1972). However, our count of 2n = 26 in R.
cameroonensis (Fig. 42), based on plants from trop-
ical Africa, differs from de Vos’s report of 2n = 22
for the species (as R. campanuloides) in southern
Africa. The possibility that tropical and southern
African plants assigned here are different species
should be considered. Romulea congoensis, 2n =
ca. 78 and not before counted, appears to be hexa-
ploid on a base of x = 13. The population of R.
linaresii from Greece with 2n = ca. 39 studied here
may be triploid (on a base of х = 13) or alterna-
tively tetraploid on a base of x = 10. Particularly
small chromosomes in the species make accurate
counting difficult.
The ancestral base number for Romulea was sug-
gested by de Vos (1972) to be x = 12, but x = 13,
which seems to be an equally likely ancestral base,
is common. Lower base numbers in Romulea, x =
11, 10, and 9, seem almost certainly to be derived
and occur in specialized lineages. The question of
ancestral base number in the genus should be re-
examined in the light of phylogeny, at present poor-
ly understood. If Romulea is related to Geissorhiza
and Hesperantha as suggested by Goldblatt (1991)
then an ancestral base number of x = 13 seems
likely for the genus on the basis of outgroup com-
parison.
The single count reported here for Crocus (Table
1), 2n = 28 in С. longiflorus, confirms past records
for that species, as does the count for Syringodea
Annals of the
Missouri Botanical Garden
bifucata (= S. bicolor), 2n = 12. The latter count
is for plants from Gauteng Province, South Africa,
where the genus was not recorded until 1983 (de
Vos, 1983). Basic number in Syringodea is most
likely x = 6.
The ancestral base number for Ixioideae is most
likely x = 10, evidently the base number for tribes
Pillansieae and Watsonieae (Table 3). The ancestral
base number in Ixieae may also be x = 10, but
genera less specialized as regards leaf anatomy and
seed characters (Goldblatt & Manning, 1995) have
higher base numbers, and it is equally likely that
x = 10, present in a few genera of the tribe, is
secondary. At least Geissorhiza, Gladiolus, Hesper-
antha, Radinosiphon, Romulea, and Tritoniopsis, all
with ancestral base numbers between x = 16 and
13 (Table 3), are paleopolyploid. We suspect that
Syringodea (x = 6) is a dysploid derivative of an
ancestor shared with Romulea. Both have inflores-
cences reduced to solitary flowers and similar
etric corms with woody tunics. The single
species of Syringodea with x 11 (Goldblatt,
1971; de Vos, 1976) is almost certainly a secondary
hypotetraploid. Notably low base numbers charac-
terize Babiana and Zygotritonia (Table 3); although
both have x = 7, they are probably not closely
related (Goldblatt, 1989b).
Extensive dysploid series in Ixioideae are re-
stricted to just 4 genera, Lapeirousia, Gladiolus, Ro-
mulea, and Crocus, out of a total of 28. Limited
intrageneric dysploidy occurs in Syringodea, as
noted above, and is reported here for the first time
in two species of Tritonia and one of Hesperantha.
Thus, the pattern outlined by Goldblatt (1971) for
Ixioideae of genera each having a single base num-
ber must be modified. Although most genera do
have a single base number and exhibit little or no
polyploidy, significant dysploidy has been discov-
ered in Gladiolus since that review (Goldblatt et al.,
1993). Most species of the genus have x = 15, but
among the small-flowered tropical African species
numbers include n = 14, 13, 12, and 11. The pat-
tern of dysploidy in Lapeirousia has also been
found to be more extensive than previously thought
(Goldblatt, 1990b). As hypothesized by Goldblatt
and Takei (1993) there is a striking example of dys-
ploid reduction from х = 10 to 4 followed by poly-
Bd: inerease and further reduction Котл = 6 to
=-3 in Lapeirousia виђе; “Расай: - Gladiolus
d Fapeirousia mast; EEE: zhe, added: to the
in their respective genera, the first specialized in
its acaulescent habit and the second in having a
solitary and terete leaf. In Gladiolus all the dys-
ploid species appear more specialized than those
with the presumably ancestral х = 15 (Goldblatt et
al., 1993). Moreover, they appear to belong to at
least three, and probably four, separate lineages.
Likewise, in Romulea members of primitive sec-
tions have x = 12 or 13, and those of derived sec-
tions have х = 11, 10, or 9 (de Vos, 1972).
Cytological patterns in Crocus, last of the genera
of Ixioideae with extensive dysploidy, are complex
(Mathew, 1982) and remain to be satisfactorily ex-
plained. Both dysploidy and polyploidy have been
significant in the cytological evolution of the genus.
Provisionally we suggest an ancestral base of x =
6 for the genus, a hypothesis based on outgroup
comparison (the immediately related Syringodea
has x = 6) and on the pattern of counts in section
Crocus (summarized by Mathew, 1982), which in-
cludes the more primitive members of the genus.
From this base we assume dysploid reduction to x
= 4, perhaps in several lines, and repeated poly-
ploidization on bases of 6, 5, 4, and 3.
REVIEW OF GENOME SIZE
Genome sizes have been established for a num-
ber of Iridaceae and, as in many other families,
have been found to vary considerably, even within
genera. In Iridoideae, species of Sisyrinchium — =
(Kenton et al., 1987) have basic genome sizes (ad-
justed for polyploidy), i.e., 1C values, of 0.48-0.73
pg in subgenus Sisyrinchium (= sect. Bermudiana)
and 0.25-2.10 pg in subgenus Echthronema. The:
related Olsynium (= Phaiophleps plus Sisyrinchium
sections Filifolium and Nuno) typically has larger —
genomes among the temperate southern South--~
American species with 2.66-3.26 pg, but the oC-- >.
toploid North American member of the genus, 0.
douglasii, has a basic genome size of 0.49 pg.
Among Tigridieae, which typically have larger
chromosomes than Sisyrinchium, 1C genome sizes
range from 2.03-2.39 pg in Cypella and 1.24-1.34
pg in three species of Hesperoxiphion (Kenton et
al., 1990), but H. huilense has a genome size of
4.38 pg, despite also being diploid. These genome
sizes were determined cytophotometrically against
a standard, Hordeum vulgare, genome size of which
is 11.12 pg (2C) or 5.56 pg (1C), a value recently
confirmed by Arumuganathan and Earle (1991).
Comparable genome sizes of 1.47-2.48 have also
been reported by Martinez and De Azkue (1987)
for five species of Eleutherine, Ennealophus, an
i4 A | Whi
ПР РРА УК
-
hi3
TEF]
Volume 84, Number 2
1997
Goldblatt & Takei 301
Chromosome Cytology of Iridaceae
Mastigostyla, but Calydorea sp. has a smaller ge-
nome of 0.98 pg.
Among Old World Iridoideae, Irideae have com-
paratively larger genomes than Tigridieae, although
genome sizes vary considerably among diploid
members of the tribe (Goldblatt et al., 1984). Ge-
nome sizes reported by Goldblatt et al. (1984) using
Zea mays as a standard need correcting. Genome
size in maize has now been found to be C = 4.75—
5.63 pg (Arumuganathan & Earle, 1991; Bennett
& Smith, 1991), somewhat lower than the estimate
of 6.3 pg used by Goldblatt et al. The 4C values
published by Goldblatt et al. (1984) thus require
adjustment by a factor of 0.75-0.84. We make a
correction of 0.75 here in the figures given below
(the strain used in the Goldblatt et al. study had a
karyotype without knobs or structural additions).
Basic 1C genome sizes (corrected for polyploidy) in
Irideae are 1.20 pg in Galaxia to 12.24 pg in Iris
histrio.
Moraea, the only genus investigated extensively,
has larger basic genomes in subgenus Vieusseuxia,
4.93-6.81 pg (x = 6) than in subgenus Moraea,
3.74–4.32 (x = 10). In Homeria (including Sessi-
listigma) and Hexaglottis (both x = 6) 1C genomes
range from 3.87 to 5.94 pg in the five species ex-
amined. The larger genomes evident in Moraea
subg. Grandiflora (discussed above) have not yet
been measured for total DNA. One species of Iris,
I. histrio (2n = 20), has a 1C genome size of 12.24
pg. Iris itself, or at least subgenus Reticulata to
which I. histrio belongs, may be paleopolyploid.
The specialized, acaulescent Galaxia has the
smallest reported genome in Irideae, 1.2 pg, about
one-third the size of that in its closest allies (Mo-
raea subg. Moraea), a puzzling observation.
Genomes are smaller in Ixioideae (Olszewska &
Osiecka, 1982; Goldblatt et al., 1984; Goldblatt &
Takei, 1993). Using the above correction factor
when necessary, 1C genome sizes range from 1.01
pg (or 0.54 pg for the basic genome) in the paleo-
tetraploid Pillansia to 0.60 pg in Gladiolus (even
lower estimates for Hesperantha and Anomatheca
obtained by Goldblatt et al. (1984) are probably not
reliable). In Lapeirousa genomes ranging from 1.27
to 1.45 pg (corrected for intrageneric polyploidy)
were established by Masterson (in Goldblatt & Tak-
ei, 1993) using Nicotiana tabacum L. as a standard.
Lapeirousia is evidently unusual in Ixioideae in
having genomes as large as those in Galaxia (Iri-
deae) and some Tigridieae. Genome sizes reported
by Olszewska and Osiecka (1982) in general con-
firm the above pattern, but as chromosome numbers
were not provided for the plants that they studied,
corrections for polyploidy are not possible. Their
estimate of a 1C value of 1.1 pg for Crocosmia
Xcrocosmiiflora (as Tritonia) is, however, consistent
with the pattern for Ixioideae. Their reports for Cro-
cus vernus (Ixioideae) of 11.56 pg and 2.1 pg for
Iris sibirica (Iridoideae) may be reversed. Iris sibir-
ica, normally 2n = 28, has much larger chromo-
somes than Crocus vernus, in which diploid num-
bers of 2n = 8, 10, 12, 16, 18, and 20 are known
(Brighton, 1976).
Genome size sometimes reflects taxonomic rela-
tionship, but it is clearly influenced by environ-
mental factors as well, and gross differences be-
tween taxa may have little taxonomic significance
above species rank. On the other hand, there seems
to be a correlation between genome size and spe-
cies relationships in Moraea. In subgenus Vieus-
euxia (x = 6) species have consistently larger ge-
nomes than do species of subgenus Moraea or the
related genus Dietes (both x = 10). Still larger ge-
nomes characterize subgenus Moraea and subgenus
Grandiflora (x = 6
е
MODES OF NUMERICAL CHANGE
Most generic ancestral base numbers in Irida-
ceae are relatively high (Tables 2, 3), which sug-
gests an early burst of polyploidy in the evolution
of the family or a paleopolyploid origin for the fam-
ily. Low ancestral base numbers for Tigridieae (x =
7), and for Babiana and Zygotritonia (x = 7) and
Syringodea (x = 6) (all Ixioideae), appear to rep-
resent paleodiploidy, but we suspect that these base
numbers are most likely derived. The latter would
be more consistent with their levels of morpholog-
ical specialization. Neopolyploidy (defined here as
intrageneric polyploidy) is common in Northern
Hemisphere representatives of the family and is es-
timated here to be above 60%. For example, in
Gladiolus 5 polyploid species, 1 of them with dip-
loid races, are known in the 80 species counted
from sub-Saharan Africa and Madagascar, but all
the 6 to 10 species of the genus that occur in Eur-
asia and North Africa are tetraploid (4x) to dode-
caploid (12x). Neopolyploidy in African members
of the three subfamilies of Iridaceae occurring in
sub-Saharan Africa is remarkably low on a world
scale. Some 5% of the species of Ixioideae and
10% of Iridoideae are polyploid, and 10% more
species of the latter have both diploid and polyploid
populations. No species of African Nivenioideae
are exclusively polyploid, but 3 species of 24
counted (12%) have diploid and tetraploid races.
Polyploidy appears to be more important in the
evolution of South and Central American Iridaceae.
Nearly all Central and North American Tigridieae
302
Annals of the
Missouri Botanical Garden
are tetraploid or hexaploid, including the 10 spe-
cies of Tigridia counted, 2 of 3 species of Nemas-
tylis counted, and all 3 species of Alophia in the
region. One population of a Mexican species of Ti-
gridia has a diploid race, and at least two South
American species of this genus are diploid. The
genera Fosteria and Cobana, closely allied to Ti-
gridia, are apparently exclusively polyploid. In oth-
er genera that are moderately well known cytolog-
ically, like Herbertia and Cipura, more than half of
the species counted are polyploid. Polyploidy
seems equally important in other tribes of Iridoi-
deae in South America, but except for Sisyrin-
chium, the genera are too poorly known cytologi-
cally for useful comparisons to be made. For
Sisyrinchium, subgenus Sisyrinchium seems to have
over 90% of the species polyploid, while subgenus
Echthronema has about 75% of the species tetra-
ploid.
Dysploidy (stepwise change in base number) is
important in several genera of Iridaceae, and exten-
sive dysploid sequences are described in more detail
above for Crocus, Gladiolus, Lapeirousia, and Ro-
mulea (Ixioideae) and Galaxia subg. Eurystigma and
Moraea (Iridoideae). Outgroup comparison and/or
correlated morphological specializations in dysploid
species consititute compelling evidence that de-
creasing dysploidy has been the dominant or exclu-
sive factor in numerical change in these genera. Less
extensive dysploid changes in Gelasine, Hesperantha,
omeria, and Tritonia should also be regarded as
having proceeded from higher to lower base number
for the same reasons. In the one species of Hexa-
glottis mentioned above there are examples of both
descending and apparent ascending dysploidy. Thus
in Iridaceae, all but one of the numerous examples
where the direction of dysploid change can be de-
termined, the direction is downward. This is consis-
tent with Jones's (1977) hypothesis that most cases
of dysploid change in plant families are downward.
Ascending dysploidy is probably a rare phenomenon
in natural populations, and arguing from the infor-
mation about dysploidy in Iridaceae and a few other
families, we question whether it has played a sig-
nificant role in the evolution of any plant group.
Literature Cited
— K. & E. D. Earle. 1991. Nuclear DNA
ent of some important plant species. Pl. Molec
Biol. Reporter 9: 208-218.
Bennett, М. D. € Ј. B. Smith. 1991.
amounts in angiosperms. Philos. Trans.,
309-345.
Brighton, C. A. 1976. Cytological problems in the genus
Crocus repa The Crocus vernus aggregate. Kew
32-46.
pag ig
Ser. B.
Bull. 31:
, B. Mathew & P. Rudall. 1983. A detailed study
of Crocus speciosus and its ©; С. pulchellus (Iridaceae).
Pl. Syst. Evol. 142: 187-
Chase, M. W. ‚ Duvall, УН: С. Hills, J. 6. шене
systematics of Lilianae. Pp. 109-137 in P. J. Rudall, P.
ribb, D. F. Cutler & C. J. Humphries (editors), Mono-
ийын: жее and Evolution. Royal Botanic
Gardens,
Cheadle, V. I. “1963. Vessels in Iridaceae. Phytomorphol-
ogy 13: 245-248.
Cruden, R. 1972. ТОРВ chromosome number reports 35.
Taxon 21: 161-166.
De Vos, M. P. 1972. The genus Romulea in South Africa.
J. S. African Bot. Suppl. 9.
1974а. Die Suid-Afrikaanse genus Syringodea.
5. Micah Bot. 40: 201-254.
. 1974b. Duthiella, 'n nuwe genus van die Irida-
ceae. 1, 5. African Вог. 40: 301-309.
1976. Die Suid-Afrikaanse genus Syringodea. J.
S. African Bot. 40: 201—254.
1979. The African genus Ferraria. J. S. African
Bot. 45; 295-315.
1982. The African genus Tritonia Ker-Gawler
ilzidacene): Part 1. + 5: agnor Bot. ee kat
Syringodea. In O. A. ner (editor),
Flora: of ordei Alfica 7(2): hu ans Research
Institute.
. The African genus Crocosmia Planchon.
FS . African Bot. 50: 463-5
Goldblatt, Р. 1969. The genus Sparaxis. J. S. African Bot.
5: 219-252.
. 1971. Cytological and morphological studies in
the southern African Iridaceae. J. S. African Bot. 37:
o
972. A revision E the genera Lapeirousia Pour-
ret and Anomatheca Ker in the winter rainfall region of
South Ок Contr. Bolus Herb. 4: 1-111.
976. Evolution, cytology d subgeneric clas-
sification in Moraea (Iridaceae). Ann. Missouri Bot.
Gard. 63: 1-23.
Barnardiella: А new pere of the Irida-
ceae and its оа to and Moraea.
Ann. Misso t. Gard. 63: 300-313.
19 9792. a а. cytology of Australasian Ir-
idi eus. Ann. Missouri Bot. Gard. 66: 8 55.
1979b. MS ded chromosome counts in
angiosperms; II. Including new family and generic re-
cords. Ann. Missouri Bot. Gard. 66: 856—861.
9 hromosome cytology and karyotype
hana d in Colonia (Iridaceae). Pl. Ci Evol. 133: 6l-
69
Redefinition of Homeria pisaa in
——. 1980a.
the light of re data, with Rheome gen.
3: 85-95.
Bot. ا
b. Systematics of Gynandriris (idee а
Gatco Бак ern African disjunct. Bot. Not.
133: 239-260.
1 Uneven diploid numbers and complex
heterozygosity in Homeria (Iridaceae). Syst. Bot. 5:
337-340.
a. Systematics, phylogeny and evolution of
Dietes бен Ann. Missouri Bot. Gard. 68: 13
153.
981b. Systematics and biology of Homeria (Ir-
idaceae). Ann. Missouri Bot. Gard.
-
|
|
|
|
EK <
Айдаш. م ونس
>
-
Volume 84, Number 2
1997
Goldblatt & Takei 303
Chromosome Cytology of Iridaceae
. 1982a. Chromosome cytology in relation to su-
prageneric systematics of neotropical lridaceae. Syst.
Bot. 7: 186-198.
1982b. Systematics of Freesia Klatt (Iridaceae).
! African Bot. 48: 39-9].
New species of Galaxia (Iridaceae) and
notes on сера pe evolution in the genus. Ann. Mis
souri Bot. Gard. 71:
1984b. A revision ۳ Беан л in the winter
rainfall region of southern Africa. J. S. African Bot. 50:
15-141.
. 1985. Systematics of the southern African genus
Gélisorhisd | жайуу ушл Ann. Missouri Вог.
Gard. 72: 277-447,
. 1986a. The genus Moraea in southern Africa.
Ann. Kirstenbosch Bot. Gard. 13.
Cytology and systematics of the Moraea
fugax complex гый Ann. Missouri Bot. Gard.
73: 140-157.
. 1987. Systematics of the southern African genus
Hexaglottis (Iridaceae—Iridoideae). Ann. Missouri Bot.
569.
The southern н genus Watsonia.
Ann. Kirstenbosch Bot. Ga
1 Revision of the QN African genus
Zygoirinia (Iridaceae) Bull. Mus. Natl. Hist. Nat., B,
212.
Adansonia
а. Phylogeny and classification of Irida-
ceae. Ann. Missouri Bot. Gard. 77: 607-627
3 b. Cytological variability in the African ge-
nus Lapeirousia (lridaceae-Ixioideae). Ann. Missouri
Bot. Gard. 77: 375-382.
. Status of the southern African Anapalina
and Antholyza (Iridaceae), genera based solely on char-
са е Ыга polli arre and a new species of
is. S. ies J. Bot. 56: 577-
overview of the е эуен, phylogeny
and er of nua African Iridaceae. Contr. Bolus Herb.
due New species, chromosome cytology and
SA on the southern African Iridaceae—Irideae: Mo-
raea, Roggeveldia and Homeria. S. African J. Bot. 58:
209-214.
— ———. 1995. Notes on Aristea Aiton (Iridaceae: Niv-
enioideae): Taxonomy, chromosome cytology, and phy
logeny. Ann. Missouri Bot. Gard. 82: 139-145.
— —— & J. E. Henrich. 1987. Notes on Cipura (Irida-
ceae) in South and Central America, and a new species
from Venezuela. Ann. Missouri Bot. > . 74: 333-340.
G Manning. 1990. Devia morpha, a new
us and species of eine from the Cape
Province South Africa. Ann. Missouri Bot.
359-364.
————. 1993. Ixia acaulis, a new acaulescent
species of Iridaceae—Ixioideae from the Knersvlakte,
Namaqualand, South Africa. Novon 3: 148-151.
E . 1995. Phylogeny of the African gen-
era Anomatheca and Freesia ie and
. Marais. 1979. Savannosiphon g „а
segregate of — Пино Si) "dne
Missouri Bot. Gard. 66: 845—850.
& N. Snow, 1991. a and chromosome
cytology of Eleutherine Herbert (Iridaceae). Ann. Mis-
: 942-949,
& M. Takei. 1993. Chromosome cytology of the
African genus Lapeirousia (lridaceae-Ixioideae). Ann.
Missouri Bot. an 80: 961—973.
Bari & J. C. Manning. 1991. Sulcus vari-
rend in the pollen grains of е err Ixioi-
e. Ann. Missouri Bot. Gard. 7.
, M. Takei & 7. A. M 1993. а
cytology in tropical African Gladiolus (Iridaceae). Ann.
Missouri Bot. Gard. 80: 461—470.
—— ———, V. Walbot & E. A. Zimmer. 1984. Estimation of
genome size (C-value) in e by cytophotometry.
nn. Missouri Bot. Gard.
, P. Rudall, V. I. ms L. oh Dorr & C. A. Wil-
liams osiris, Iridaceae or Burmanniaceae.
Bull. Mus. Natl. Hist. Nat., B, Adansonia 9: 239-
Hair, J. B., E. J. Beuzenberg & B. Р 1967. Con
romosome atlas of ithe New Zealand
flora. New Zealand J. Вог. 5: 185-196.
Holmgren, P. K., W. Keuken 8 E. K. Schofield. 1981.
Index Herberiorum, Part 1, ed. 7. Regnum Veg. 106:
1-452.
Hsu, C.-C. 1971. Preliminary studies on the vascular
plants of меч (IV). Taiwania 16: 123-136.
Huynh, K. . Contribution à l'étude caryologique et
socal ye des phanérogames du Pérou. Denkschr.
Schweitz. Naturf. Ges. 85: 1-178.
Jones, K. 1974. Chromosome evolution by Robertsonian
translocation in Gibasis (Commelinaceae). Chromosoma
Bern 45 = 353-368.
——. 7. The role of Robertsonian change in karyo-
type на in higher plants. Chromosomes Today 6:
21-129.
к А. & С. A. Heywood. 1984. Cytological studies
n South American aie Pl. Syst. Evol. 146: 87-
— & P. Rudall. 1987. An unusual case of complex-
heterozygosity in n Gelasine azurea (lridaceae), and its
implications for reproductive biology. Evol. Trends PI.
1: 95-1 gd
. Rudall & A. R. Johnson. 1987. Genom
size unn in Sisyrinchium L. (Iridaceae) and its re-
lationship to phenotype and habitat. un Gaz. 147:
342-354
"+ B. Dickie, D. H. Langton & M. D. Bennett.
1990. Nuclear DNA amount and karyotype symmetry
n Cypella m pecto ко (Tigridieae; Iridaceae).
Evol. trends P
Lewis, G. J. Y^ ana and anatomical studies
in the South African Iridaceae. Ann. S. African Mus.
40: 1
Manning, J. C. & P. Goldblatt. 1990. Endothecium in
ridaceae and its systematic implications. Amer. J. Bot
77: 527-532.
991. Seed coat structure in the
shrubby Cape Say Nivenia, Klattia and Witsenia
Bot. J. Linn. Soc. 107: 387-404.
Martínez A. & D. De Azkue. 1987. Variacíon ys conten-
ido de ADN ue su implicancia evolutiva. Pp. 55 in
Anales del C Latinoamericano de n vol.
> Simposio Citología y Citotaxonomía. Bogotá, Colom-
ia [for Congreso Latinoamericano de Botánica, Medel-
Ме Colombia 1986].
Mathew, B. 1981. The /ris. Batsford Press, London.
. 1982. The Crocus. Batsford Press, London.
Molseed, E. 1970. The genus Tigridia (Iridaceae) of Mex-
ico and central America. Univ. Calif. Publ. Bot. 54. 1-
Chien, M. J. & R. Osiecka. 1982. The relationship
Annals of the
Missouri Botanical Garden
between 2C DNA content, life cycle type, systematic
of roots in some monocotyledonous species. Biochem
Physiol. Pflanzen 177: 319-336.
Nakajima, G. 1936. er numbers in some crops
p. J. Genet. 12: 211-218.
9. таласа interchange іп Eleutherine
plicata Herb. Genetica 40: 417—420.
Ravenna, P. 1974. Cobana, a new genus of central Amer-
ican Iridaceae. Bot. Not. 127: 104—108.
1976. Neotropical species threatened and en-
dangered by human activity in the Iridaceae, Amaryl-
lidaceae and allied bulbous families. Pp. 257-266 in
С. T. Prance 4 T. S. Elias (editors), Extinction is For-
ever. joe > Botanical Garden, New York.
. Notas sobre реке У. Not. Mens. Mus.
Nac. TA x 21(249): 7
——. 1984. The delimitation of Gelasine (Iridaceae),
and G. uruguaiensis sp. nov. from Uruguay. Nordic J.
Bot. 4: 347—350.
Rodríguez, R. 1986. Die Chilenischen Arten der Gattung
Sisyrinchium L. (Iridaceae). Mitt. Bot. Staatssamml.
München 22: 97-201.
co P. 1994. incertae "n systematics of Iridaceae.
t. J. Linn. Soc. 114:
& P. Goldblatt. 1991. Leaf wes in Iridaceae-
Ixioideae. Bot. J. Linn. Soc. 106: 329-345
, A. Y. Kenton & T. J. Lawrence.
tomical and chromosomal M rtr of Sisyrinchium
and allied genera. Bot. Gaz. 147: 466-477.
Sharma, А. K. & C
1959. Cyto-taxonomic
the Iridaceae with speci
reference to the structural кеја of Cipura pal-
udosa Aubl. Nucleus (Calcutta) 11: 63-84.
Shneyer, V. S. 1990. A serotaxonomic study of tribe
Ixieae (Iridaceae). Bot. Zhurn. (Moscow & Leningrad)
75: 1657-
Simonet, M.
tiques chez les /ris. Bull.
1932. Recherches cytologiques et géné-
Biol. France Belgique 66:
Souza-Chies, T., G. Bittar, S. Nadot, L. Carter, E. Besin
. Leje 1 ylogenetic analysis of Iri idac
a гану and distance aire s, using the laid
gene rps4. Pl. Syst. Evol. 204: 3.
Strid, A. 1972. A taxonomic revision of Вобата L. (lr-
idaceae). a Bot. 37: 1-45.
Winge, H. 195 Studies o on the cytotaxonomy and poly-
morphism T the genus Herbertia (Iridaceae). Revista
Brasil. Biol. 19: 195-201.
CHROMOSOMAL
OBSERVATIONS ON THE
ALZATEACEAE (MYRTALES)!
Frank Almeda?
ABSTRACT
Alzatea verticillata Ruiz & de subsp. amplifolia S. A. Graham of the monotypic neotropical family Alzateaceae has
a haploid chromosome number of n — 14.
the 14 core families of the Mitos. Given that x — 12 appears to
ave arisen as an ascending dysploid or as a tetraploid derived from an ancestral bas
rs is given for all families currently recognized in the vo id except the Cryp-
of ir basic chromosome numbers
teroniaceae, which remains Mom cytologically.
This first report for the family is unique as a putative
base number among
be the basic number for the I n = 14 pons
se number of x = 7. А summary
Alzatea is a monotypic genus of shrubs or small
trees restricted to low montane cloud forest habitats
from Costa Rica and Panama south to Andean
South America from Colombia to Bolivia. The
placement of this genus in eight families in five
orders since its discovery nearly 200 years ago
(Graham, 1984) has stimulated considerable re-
search in the last two decades aimed at assessing
its relationships and taxonomic status. Several re-
cent papers have brought to light data from macro-
morphology, anatomy, chemistry, embryology, paly-
nology, DNA sequencing, and cladistic analysis
that support the recognition of Alzatea as a distinct
family within the angiosperm order Myrtales (Dahl-
gren & Thorne, 1984; Graham, 1984; Graham &
Averett, 1984; Johnson & Briggs, 1984; Keating,
1984; Patel et al., 1984; Tobe & Raven, 1984; van
Vliet & Baas, 1984; Conti et al., 1996; Conti et al.,
in press). I here add chromosome information for
the Alzateaceae that fills a notable gap in the
knowledge of cytology in the order.
MATERIALS AND METHODS
Flower buds were collected from a natural pop-
ulation in the field, fixed in modified Carnoy’s so-
lution (4 chloroform, 3 ethanol, 1 glacial acetic
acid, v/v/v) for 24 hours, then transferred to 70%
ethanol for refrigerated storage. Anthers were
teased open and the contents squashed in 1% ferric
aceto-carmine. Counts were made from pollen
mother cells using a Zeiss light microscope with
phase contrast and a 100X oil immersion objective.
Drawings of meiotic configurations were made by
camera lucida at a magnification of 1500. The
voucher collection for the count reported here is as
follows:
Alzatea verticillata Ruiz & Pav. subsp. amplifolia
S. A. Graham. n = 14. Panama. Chiriquf: Fortuna
Dam area in the vicinity of Centro de Investiga-
ciones Jorge Arauz, 1200 m, 8 Feb. 1996, Almeda
et al. 7545 (CAS).
OBSERVATIONS AND DISCUSSION
The first chromosome count for this monotypic
family, n = 14, is based on diakinesis figures and
one telophase I figure (Fig. 1). Meiosis was regular
in all cells examined. Chromosome clumping at
metaphase I and telophase I presented some inter-
pretation difficulties, but examination of numerous
diakinesis figures provided clear and consistent
counts о
Chromosome numbers are now known for rep-
resentative species of all core families in the Myr-
tales except the Crypteroniaceae. Excluding the Al-
zateaceae, base chromosome numbers аге
established with some certainty for 9 of the other
13 families in the order (Table 1). Four or possibly
five of the families have x = 12, the number most
commonly postulated as the original base number
for the order (Raven, 1975; Johnson & Briggs,
1984). As noted by Graham et al. (1993), х = 12
is also the only base number occurring in each of
the major evolutionary lineages hypothesized for
the Myrtales by Johnson and Briggs (1984). The
lowest base numbers in the order are x = 8 for the
' Fieldwork for this study was made possible, in part, by the G. Lindsay Field Research Fund of the California
1
Academy of Sciences. I am grateful to pis Correa
suppo al Valdespino, T. F. Daniel, and J. L.
and the Smithsonian Tropica
Luteyn for — in E field; Fei-Mei Chuang for technical laborato
assistance: and Shirley Graham for helpful comments on the
? De
esearch Institute for v. eni
uscrip
epartment of Botany, California Academy of Sciences, Golden с Park, San Francisco, California 94118, U.S.A.
ANN. MISSOURI Bor. GARD. 84: 305-308. 1997.
306 Annals of the
Missouri Botanical Garden
Figures 14. Camera lucida drawings of meiotic
1-3. Diakinesis (n = 14).—4. Telophase I (iX M chromosome figures of Alzatea verticillata subsp. amplifolia.—
Table 1. Base chromosome numbers of core families in the Myrtales.
Family Base number (x) References
Alzateaceae 14? This paper
Combretaceae 12 Raven, 1975
Crypteroniaceae ? —
Heteropyxidaceae 12 Fernandes, 1971
Lythraceae 8 am, pers. comm. in Raven, 1975;
Tobe et al., 1986
Melastomataceae 9 or 12 Almeda, unpublished
Memecylaceae Favarger, 1952, 1962; Solt Я en 1980
Myrtaceae 11 Smith-White, 1959; Rye, 19
Oliniaceae 10? Goldblatt, 1976
Onagraceae 11 Raven, 1975
Penaeaceae 10 Dahlgren, 1968, 1971
Psiloxylaceae 12 Johnson & Brigg
Rhynchocalycaceae 10 Goldblatt, 1976; redd & Briggs, 1984
Trapaceae 12 Raven, 1975
Lythraceae and possibly x — 7 for the Memecyla-
ceae (Table 1), both of which were very likely de-
rived by descending dysploidy from the ancestral
base. The chromosome number reported here for
the Alzateaceae would not be discordant with what
is known about base numbers in the order. The
process by which it evolved, however, is open to
speculation.
Accepting the premise that chromosome num-
bers higher than n — 13 are considered to be of
polyploid derivation (Grant, 1963, 1981), then n —
14 for Alzatea could be interpreted as a tetraploid
based on x — 7. Recent collections from South
America have extended the geographic ranges for
each of the two subspecies of Alzatea, and, not sur-
prisingly, diminished the perceived morphological
distinctions between them (Graham, 1995). Chro-
mosome information for the nominate subspecies of
Alzatea is needed before x — 7 can be ruled out as
an extant base number for the family since intra-
specific polyploidy is known in species of other
myrtalean families such as the Lythraceae (Graham,
1989), Melastomataceae (Almeda, in press), Ona-
graceae (Berry, 1982; Raven & Tai, 1979), and
Myrtaceae (Rye, 1979).
Given the range of base numbers in the Myrtales,
another less parsimonious scenario that could ac-
count for the origin of n — 14 is descent from a
tetraploid (n — 16) based on x — 8. A parallel
example has been postulated by Graham (1992) to
account for x — 15 in Diplusodon of the Lythraceae.
n alternative scenario that could also account
for the origin of n — 14 would be dysploid increase
from the ancestral base of x — 12. Given that as-
cending dysploidy is believed to be four times less
common in the flowering plants than descending
dysploidy (Jones, 1970; Goldblatt & Poston, 1988),
one could argue that this scenario is less likely, but
it certainly is not implausible.
А base number of 7 or 14 would serve to un-
derscore the uniqueness of Alzateaceae but would
provide no additional insights about relationships
that have not already been demonstrated using oth-
er lines of evidence. Information amassed to date
shows that the Alzateaceae and the Rhynchocaly-
caceae share a common ancestor that, in turn, is
closest to the Penaeaceae, Oliniaceae, and Cryp-
teroniaceae (Graham, 1984; Johnson & Bri
1984). Chromosome data are lacking for the latter
family. The Penaeaceae and Rhychocalycaceae
both have x — 10 (Table 1). The Oliniaceae are in
need of additional study to confirm the few known
counts that are also suggestive of x = 10 (Goldblatt,
1976). The fact that Alzatea is sister to the three
families with x — 10 or 10? in the clade Alzatea-
ceae-Rhynchocalycaceae-Penaeaceae-Oliniaceae
(Conti et al, 1996) lends further support to de-
scending dysploidy as the likely mechanism for
chromosome number changes in Alzatea. Chromo-
some information can sometimes inform us in ways
that morphological data cannot. In this instance, if
x = 14 is ultimately established as the base num-
ber for Alzatea, it may prove to be the only family-
level lineage in the order that originated via poly-
ploidy.
Literature Cited
ress. Chromosome numbers and their evo-
lutionary significance in e neotropical and paleo-
tropical Melastomataceae. BioLlania 12.
The systematics and evolution of Fuch-
sia sect. Fuchsia (Onagraceae). Ann. Missouri Bot.
Gard. 69: 1-198.
308
Annals
ictum n Garden
Conti, Е., A. Litt & K. J. Sytsma. 1996. Circumscription
of Myrtales and their relationship to other rosids: Evi-
dence from rbcL sequence data. Amer. J. Bot. 83: 221-
————, S. A. Graham, A. Litt, P. С. Wilson, B. С. Brigg
L. А. 5. Johnson & K. J. Sytsma. In press. Interfamilial
relationships in Myrtales: Molecular phylogeny and pat-
terns of morphological evolution. Syst. Bot
Dahlgren, R. 1968. Studies on Punsencene. Part 11. The
genera Brachysiphon, Sonderothamnus, and Saltera.
Opera Bot. 18: 1-72
1971. Studíós on Penaeaceae. Part VI. The ge-
nus Pigos. Opera Bot. 29: 1–
& R. F. Thome. 1984. Myrtales: Circumscription,
variation, and relationships. Ann. Missouri Bot. Gard.
99.
1: 63
F mes С. 1952. Recherches sur quelques Mélasto-
es d'Afrique осш. Ber. Schweiz. Bot. Ges
62: 5-65.
1962. Nouvelles recherches cytologiques sur les
Mélastomatacées. Ber. Schweiz. Bot. Ges. 72: 290-305
Fernandes, А. 1971. Contribution а la connaissance du
genre Heteropyxis Harv. Mitt. Bot. Staatssamml. Miin-
chen 10: 207-234.
ew P. 1976. New or noteworthy chromosome re-
s in ^ad angiosperms. Ann. Missouri Bot. Gard. 63
+ M. E. Poston. 1988. Observations on the chro-
mosome си ову of Velloziaceae. Ann. Missouri Bot.
Gard. 75: iar 195.
Graham, S. A. 1984. Alzateaceae, a new family of Myr-
tales in yo a tropics. Ann. Missouri Bot. Gard.
71: m
1 Ghivbanpius numbers in Cuphea geri
cosy counts and a summary. Amer. J. Bot. 7
1530-1540.
1992. New chromosome counts in Lythraceae—
Systematic and evolutionary implications. Act. Bot.
Mex. 17: 45-51.
. 1995. Two new species in Cuphea Чүкө,
and а note on Alzateaceae. Novon 5: 272-27
& J. E. Averett. 1984. Flavonoids of ACA
(Myrtales). Ann. Missouri Bot. Gard. 71: 855-8
1993,
Chromosome numbers in Sonneratia and Duabanga
(Lythraceae s.l.) and their E PE es. Tax-
1:
, K. Oginuma, P. H. Raven d H. Tobe.
Grant, y. 1963. The Origin of Adaptations. Columbia
Univ. Press, New York.
1981. Plant Speciation, 2nd ed. Columbia Univ.
Ре New York.
Johnson, L. A. S. & B. G. Briggs. 1984. Myrtales and
Myrtaceae—A phylogenetic analysis. Ann. Missouri
Bot. Gard. 71: 700-756.
Jones, K. 1970. Chromosome changes in plant evolution.
Taxon 19: 172-179.
Keating, R. C. 1984. Leaf histology and its contribution
to relationships in the Myrtales. Ann. Missouri Bot.
Gard. 71: 801-823.
Patel, У. C., J. J. Skvarla & P. H. Raven. 1984. Pollen
characters in relation to the delimitation of Myrtales.
Ann. Missouri ug Gard. 71: 858—969.
. The bases of sapo phylogeny:
gy. Ànn. Miss Bot. Gard. 62: 724—764.
а W. L. Tai. 1979. сна of chromosomes
in Ludwigia as Ann. Missouri Bot. Gard. 66:
862
Rye, B. L. 1979. Chromosome number variation in the
Myrtaceae and its taxonomic implications. Austral. J.
Bot. 27: 547-573.
Smith-White, S. 1959. Cytological Evolution in the Aus-
tralian Flora. Cold Spring Harbor Symp. Quant. Biol.
24: 273-289.
Solt, M. L. & J. J. Wurdack. 1980. Chromosome numbers
in the Melastomataceae. Phytologia 47: 199-220
Tobe, Н. € P. H. Raven. 1984. The embryology and re-
lationships of Alzatea Ruiz & Pav. (Alzateaceae, Муг-
tales). Ann. Missouri Bot. Gard. 71: 844-852.
. H. Raven & S. A. Graham. 1986. Chromo-
some pa for some Lythraceae sens. str. (Myrtales),
and the base number of the family. Taxon 35: 13-20.
Vliet, G. J. C. M. van & P. Baas. 1984. Wood anatomy
and classification of the Myrtales. Ann. Missouri Bot.
Gard. 71: 783-800.
Zu TM ч =. Ай: Ашу 37 3. 24 9 E eee
NOTICE
PLANT EVOLUTION AND DOMESTICATION
26-27 SEPTEMBER 1997
Indiana University will hold a weekend symposium
in honor of Dr. Charles Heiser’s prominent contribu-
tions to Botany during his 50 years at IU. The sym-
posium is entitled “Plant Evolution and Domestica-
tion,” and will take place Friday evening, September
26 and all day Saturday, September 27. Speakers in-
clude Greg Anderson, John Doebley, Jeff Doyle, Don
Levin, Barbara Pickersgill, Charles Rick, Loren Rie-
seberg, Doug Soltis, and Herb Wagner. Registration
fees are $75.00 for regular participants and $25.00
for students. For further information contact Angi Bai-
ley or Jennifer Jones, Department of Biology, Indiana
University, Bloomington, IN 47405-6801. Phone:
812-855-6283, FAX: 812-855-6705, email: abail-
ey@bio.indiana.edu.
Editor’s Note
With this issue of the Annals, we witness a change in editorship. Henk van der Werff, who has
successfully led the journal through the past five years, passes the editorial pen to Michael H. Grayum.
Beginning in 1992 with volume 79 issue 1 and continuing with contributions to the current issue, Henk
has provided leadership, sound scientific judgment, and a keen editorial eye, for which the readership of
the Annals and those of us responsible for its production are grateful. It is with heartfelt thanks that we
acknowledge Henk’s significant contributions to the journal, and his continued involvement as a member
of the Editorial Committee. We also welcome with great enthusiasm the new editor, and wish him well in
the many challenges that lie ahead.
Editorial Staff of the Annals of the Missouri Botanical Garden
Editors of the Annals of the Missouri Botanical Garden
This information was taken from the covers of the Annals or from separately published volume front
matter. No individual editor(s) can be identified for volumes 1-13. The title for what I call the editor
varies slightly through the years, e.g., Editor of Publications, Editor, Editor-in-Chief. Through most of its
history, a single individual produced the Annals: reviewed manuscripts, edited them, saw them through
production, and got the issues distributed, although editorial asssistants were listed irregularly through
the years. Beginning in the mid-1980s an assistant did much of the copy-editing. The journal is now
edited by a scientific editor, responsible for technical review of manuscripts, and a managing editor, who
along with with an editorial assistant, handles the manuscripts from the time they are accepted through
their production and distribution in issues of the Annals. The Annals has usually been a quarterly; volumes
53-61 were complete in three numbers, and volumes 50 and 51 contained a single issue, numbered 14
1914-1915: George T. Moore, Benjamin Duggar, Jacob R. Schramm, Editorial Committee, vols. 1, 2.
1916-1926: George T. Moore, Benjamin Duggar, Editorial Committee, vols. 3-13.
1919-1957: Nell C. Horner, vols. 6-44.
1958-1962: Robert L. Dressler, vols. 45-49.
1963: No editor listed, vol. 50.
1964—1968: Walter Н. Lewis, vols. 51-55.
1969: Derek Burch, vol. 56(1, 2).
1969-1974: Marshall R. Crosby, vols. 56(3)-61.
1975-1981: Gerrit Davidse, vols. 62-68(1-3).
1981-1986: Nancy Morin, vols. 68(4)-73.
1987: George K. Rogers & Nancy Morin, vol. 74(1).
1987-1990: George K. Rogers, vols. 74(2)-77(2).
1990-1991: Marshall R. Crosby, vols. 77(3)-78.
1990- : Amy Scheuler, Managing Editor, vols. 77(2)- . [Amy Scheuler McPherson, 1994—.]
1992-1997: Henk van der Werff, vols. 79-84(2).
1997- : Michael Н. Grayum, vols. 84(2)-.
Marshall R. Crosby
April 30, 1997
Volume 84, Number 2, pp. 167-310 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN
was published on June 17, 1997.
A NI A Num rrt wt ue T
—
- ——— — пи =
a E a
Experimental and Molecular Approaches to Plant Biosystematics _
The proceedings of the Fifth International Symposium of the International Organization of Plant
Bio aiii (IOPB)
Edited by Peter C. Hoch and A. G. Stephenson
Twenty-three original contributions that span the breadth of biosystematics, a dynamic field of dide
that bridges the realms of systematics and population biology. The papers are arranged in four groups,
reflecting the original four symposia of the 1992 meeting. DNA and Plant Biosystematics presents
innovative work that uses the rapidly developing nucleic acid methods adapted from molecular biology.
Plant Growth Patterns and Biosystematics includes comparative and developmental analyses of plant
Annals of the Missouri Botanical Ceci, Volume 82, Number 2: Alternative Genes
for Phylogenetic Reconstruction in Plants |
A symposium cosponsored by the American Society of Plant Taxonomists and the Botanical Society of .
America, organized by Pamela S. Soltis and Douglas E. Soltis, and presented at the 1993 AIBS meetings.
таку the chloroplast репе rbcL has been successfully че? to reconstruct plant phylogeny, many
important questions of plant phylogeny and evolution cannot be addressed using it. The contributors to
this issue of the Annals explore the potential of eight alternative genes or DNA regions for phylogenetic
reconstruction at a variety of hierarchical levels. Both nuclear and chloroplast genes are evaluated. Three
regions of the nuclear ribosomal RNA cistron are explored: the 18S gene, the internal transcribed spacers
signal: the phytochrome gene family and the small heat shock gene family. Three genes
roplast genome are also considered: atpB, ndhF, Са, ide Each paper describes the location, size,
structure, and rate of evolution of the chosen gene and discusses its potential for чари главна study. This
issue also contains: “The Comparative Pollination and Floral Biology of Bao (A аса-
ceae)" by David A. Baum and “In Memoriam: Peter G. Martin.” Annals 82(2) 1995. 174 рај pages. $27.50
U.S. $28.00 Non-U.S.
To order, please indicate method of payment below. Checks or money orders should be in U.S. funds, DE.
payable through a U.S. bank, to Missouri Botanical Garden. Orders must be prepaid. $3. 00 handling
fee on all orders. Additional $2.00 handling fee will be added to orders
are made until RAYS is received. Phone: Dr 971-9534, fax: (314) 577-9594, e-mail:
dept11@mobot.org
Send order Ne d Please send — copy (ies) of {Monograph ре A
Гереро Eleven, Sinai Botanical бан Please send ___ copy(ies) of Annals 82(2)
Ss pons MO 63166-0299, U 5. А.
O Check/money order л n euer I Send books to:
O Send invoice ($2.00 fee will be added to total) E Wr Md Ad
L] Charge card number (MasterCard/Visa) |
Мате |
Expiration date _ © ы ы 2 Addr "d
Taya as it appears on IN
Telephone number (daytime) -
Postal Code Country
Prices А ARE наг то CHANGE Wrmout Norice UPS 842)
СОМТЕМТ5
The Genus Lycianthes (Solanaceae) in Venezuela
Carmen Benitez de Rojas & William С. П'Атсу
Dialypetalanthus fuscescens Kuhlm. (Dialypetalanthaceae): The Problematic Taxonom- _
ic Position of an Amazonian Endemic
Conspectus of the Genus Palicourea (Rubiaceae: Psychotrieae) with the Description
of Some New Species from Ecuador and Colombia —. Charlotte М. Taylor
Palynology, Phylogenetic Reconstruction, and Classification of the Afro-Madagascan
Genus Aristea (Iridaceae) Peter Goldblatt & Annick Le Thomas
Chromosome Cytology of Iridaceae— Patterns of Variation, Determination of Ancestral
Base Numbers, and Modes of Karyotype Change
Peter Goldblatt & Masahiro Takei
Chromosomal Observations on the Alzateaceae (Myrtales) _______ Frank Almeda
Notice
Editors Note and Editors of the Annals of the Missouri Botanical Garden ______
Cover illustration. Eccremocarpus viridis Ruiz & Pavón, by Phyllis Bick.
Frederic Piesschaert, Elmar Robbrecht & Erik Smets |
Ref.
Qh)
Annals |
of the
Missouri
Botanical |
Garden —
seu Y
Se "m
~ =
Volume 84, Number 3
Summer 1997
Annals of the
Missouri Botanical Garden
The Annals, published quarterly, contains papers, primarily in systematic botany, -
contributed from the Missouri Botanical Garden, St. Louis. Papers originating out- —
side the Garden will also be accepted. All manuscripts are reviewed by qualified,
independent reviewers. Authors should write the Managing Editor for information
concerning arrangements for publishing in the ANNALS. Instructions to Authors are
| printed in the back of the last issue of each volume.
Editorial Committee
Michael H. Grayum
Editor,
Missouri Botanical Garden
Amy Scheuler McPherson
Managing Editor,
Missouri Botanical Garden
Diana Gunter |
Editorial Assistant,
Missouri Botanical Garden
Vicki Couture
Secretary
Teresa Johnson
Publications Order Processor
Ihsan A. Al-Shehbaz
Missouri Botanical Garden
Gerrit Davidse
Missouri Botanical Garden
Roy E. Gereau
Missouri Botanical Garden
Peter Goldblatt
Missouri Botanical Garden
Gordon McPherson
Missouri Botanical Garden
P. Mick Richardson
Missouri Botanical Garden
Henk van der Werff
Missouri Botanical Garden
For subscription information contact Depart-
ment Eleven, P.O. Box 299, St. Louis, MO
63166-0299. Subscription price is $110 per
volume U.S., $115 Canada and Mexico, $135
all other countries. Four issues per volume. The
journal Novon is included in the subscription
price of the ANNALS.
amcpherG'admin. mobot.org (editorial duis
dept] еня = олари
http:/A
O Missouri Botanical Garden 1997
The ANNALS or THE MISSOURI BOTANICAL —
` GARDEN (ISSN 0026-6493) is published quar — —
terly by the Missouri Botanical Garden, 2345 —
Tower Grove Avenue, St. Louis, MO 63110. Pe- — ;
riodicals postage paid at St. Louis, MO and ad- |
ditional mailing offices. POSTMASTER: Send ad- |
dress changes to ANNALS OF THE MISSOURI
BOTANICAL GARDEN, Department Eleven, P.O. —
. Box 299, St. Louis, MO 63166-0299.
The mission of the Missouri Botanical Garden is to discover and share knowledge about pan pu Дд |
their environment, in order to preserve and enrich life. чо:
Ө Thi: xL да m ^ P
У УУ |
=
: 239.48 1972 Pomanence ol Papen:
Volume 84 Annals
Number 3 of the
1997 Missouri
Botanical
Garden
A REVISION OF Thomas B. Croat?
PHILODENDRON SUBGENUS MISSOURI BOTANICAL
PHILODENDRON (ARACEAE)
FOR MEXICO AND CENTRAL OCT 3 1 1997
AMERICA!
GARDEN LIBRARY
ABSTRACT
This is the first revision of Philodendron subg. Philodendron since that of K. Krause in Das Pflanzenreich i in 1913.
Philodendron subg. Philodendron, the largest of the three subgenera, includes 103 taxa (95 species and 8 subspecies
4 ие roat, P. :
copense Croat, P. correae Croat, P. cotobrusense Croat & Grayum, P. cotonense Croat & perm P. s Croat
pésudauriculatum Croat, P. purulhense Croat, P. roseospathum Croat, P. scalarinerve Croat & Grayum, P. sousae Croat,
P. squamicaule Croat & Grayum, P. squamipetiolatum Croat, P. straminicaule Croat, P. sulcicaule Croat € к Р,
thalassicum Croat € Grayum, P. tysonii Croat, P. ubigantupense Croat, P. utleyanum Croat, P. serapasense Croat, P.
wilburii Croat & Grayum, P. zhuanum; 6 infraspecific taxa: P. davidsonii Croat subsp. bocator t, P. hederaceum
(Jacq.) Schott var. kirkbridei Croat, P. ligulatum Schott var. heraclioanum Croat, P. ligulatum Schott var. ovatum Croat,
P. roseospathum Croat var. angustilaminatum Croat, P. wilburii var. К ыл Сгоа! & dnm and 2 new
combinations: P. hederaceum (Jacq.) Schott var. oxycardium (Schott) Croat, P. radiatum Schott v r. pseudoradiatum
i Philodend
With the к, of Belize. which has 1 endemic, no other country in Middle America has any endemic species.
Only 26 species (28 taxa), a total of 26% of all Central American species, range into South America, 8 species (7%
of ili total) only to Colombia.
! This study was completed with support from National Science Foundation grant BSR-8614777.
? P. A, Schulze Curator of Botany, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S
ANN. MISSOURI Bor. GARD. 84: 311—704. 1997.
312 Annals of the
Missouri Botanical Garden
INTRODUCTION considered endemic. Mexico and Costa Rica each
Philodendron is, after Anthurium, the largest ge-
nus in the Araceae, with 700 or more species (Croat,
1979, 1983a, 1988, 1990). This is the first major
revision of P. subg. Philodendron for Central Amer-
ica since Kurt Krause’s (1913) generic treatment in
Das Pflanzenreich. Philodendron is one of the most
important genera in the Neotropics, inhabiting a
wide range of mesic habitats from sea level to over
2000 m and in life zones (Holdridge, 1967) ranging
from Tropical moist forest to Premontane rain forest.
While most species occur in virgin humid forests,
the genus is known from freshwater swamps, stream
banks, regrowth forest, rock outcrops, and road-
banks. It is not only one of the largest genera in
the Neotropics but also often constitutes the most
conspicuous element of the vegetation because of
its abundance, primarily climbing habit, and fre-
quently large, showy leaves. The genus provides a
wide variety of choice ornamental plants for horti-
culture, including most of the species treated here.
Unfortunately, it is also still poorly known taxonom-
ically, especially in the South American Andes.
Philodendron has 119 Central American species
comprising 128 taxa, distributed in two subgenera
of Philodendron. This Central America revision en-
compasses only members of P. subg. Philodendron,
with 103 taxa, including 95 species and 8 varieties
or subspecies (Appendix 1, Geographic Distribu-
tion of Central American Philodendron subg. Philo-
dendron). A total of 68 taxa are new to science: 62
species, 6 subspecies or varieties, and 2 combi-
nations. Alternatively, P. subg. Pteromischum, re-
vised separately by Grayum (1996), contains 21
species (26 taxa) for Central America. That revision
encompasses all species from Pacific and Carib-
bean tropical America, ignoring only species from
the region of the Guianas and from the Amazon
drainage of South America.
Philodendron is a distinct genus, not easily con-
fused with any other, though closest to Homalo-
mena, which differs in having a consistently terres-
trial habit, frequently spiny petioles, a sap smelling
of anise, and staminodia among the pistillate flow-
ers.
Species diversity of Р. subg. Philodendron in
Central America shows a general diminution from
Mexico to Middle America with the lowest totals
just north of the San Juan depression, followed by
a marked increase approaching the South American
continent. Mexico has 21 taxa, Guatemala 15, Be-
lize 9, El Salvador 5, Honduras 13, Nicaragua 18,
Costa Rica 48, and Panama 82. Endemism is high,
especially for Panama where 39 taxa are currently
have 7 endemic species. With the exception of Be-
lize, which has 1 endemic, no other country in Mid-
dle America has any endemic species.
Most of the Central American species of P. subg.
Philodendron (Appendix 3, Sectional Composition
of Philodendron subg. Philodendron in Central
America) are in P. sect. Calostigma (Schott) Engl.
with 52 taxa (48 species) and P. sect. Philodendron
with 62 taxa (58 species). Other sections repre-
sented in Central America are: P. sect. Tritomo-
phyllum (Schott) Engl. with 6 species; P. sect. Po-
lytomium (Schott) Engl. with 3 species (4 taxa); and
P. sect. Macrogynium Engl. with 1 species. Sec-
tions not represented in Central America are: Schi-
zophyllum (Schott) Engl.; Camptogynium K. Krau-
se; and Philopsammos G. S. Bunting. A key for the
sections of P. subg. Philodendron is included under
"Taxonomy."
MATERIALS AND METHODS
This revision is based on more than 25 years of
field studies in Central and South America, be-
tween 1967 and 1993. All but 15 of the 95 species
were studied in the field or under cultivation at the
Missouri Botanical Garden. Those only known from
herbarium material are: P. breedlovei, P. brewster-
ense, P. chirripoense Croat & Grayum, P. cotobru-
sense, P. dwyeri, P. folsomii, P. hammelii, P. jefense,
> madronense, P. roseospathum var. angustilami-
natum, P. sousae, P. ubigantupense, P. utleyanum,
and P. verapazense Croat. Except for these, all de-
scriptions have been prepared from both living and
dried specimens. The use of (“dried”) preceding all
or any part of the description is an indication that
all that follows is based on herbarium material only.
Morphological characters were coded directly into
a computerized database to ensure parallel an
sortable descriptions. The aroid descriptions data-
base, completely rewritten since the publication of
my revision of Anthurium sect. Pachyneurium
(Croat, 1991), contains 892 character states used
to describe the morphological diversity expressed
in Philodendron. A total of 108 of these are use
exclusively for description of the bisexual inflores-
cence (and thus were not used in the descriptions
of Philodendron), while 220 describe unisexual in-
florescences. The database also allows for sorting
of characters for use in writing keys or in providing
useful lists of characters for preparing а cladistic
survey. In addition, the database can be put to fu-
ture use for the preparation of floristic surveys or
for adding additional newly discovered species. The
description database is tied directly to the nomen-
ан |
Volume 84, Number 3
1997
Croat 313
Philodendron Subgenus Philodendron
clatural database in TROPICOS (Crosby, 1986;
Crosby & Magill, 1986). Finally, discussions and
references to illustrations as well as exsiccatae are
stored separately but tied to a particular species
description and to the nomenclatural information
by a unique taxon number. Specimens can be add-
ed to the exsiccatae almost until the time of pub-
lication because they are automatically presorted to
localities before being printed. Species descriptions
are decoded into narrative text automatically before
final editing for style.
Terminology and usage in the descriptions in this
revision are largely defined by Croat and Bunting
(1979). Further definitions of petiole cross-sectional
shapes are defined and illustrated in Croat (1983a).
The colors reported in the description frequently
are taken from the color chart by Berlin and Kay
(1969) and are referenced in the text as B & K.
This color chart, available from the University of
California Press, is a reproduction of the Munsell
Color Array of 40 hues, at maximum saturation,
with nine degrees of brightness. This represents 40
hues in the vertical columns and 9 degrees of
brightness in the horizontal columns. Colors are ar-
ranged in 10 basic clusters with 4 different hues
per cluster, ranging from red through yellow, green,
blue, purple, and finally red-purple. The four col-
umns for each color cluster are numbered 2.5, 5,
7.5, and 10. These numbers are repeated for each
basic color type. The colors from the B & K color
chart are read by first reporting the color, then the
row followed by the column. For example, the third
color in the fifth row is Red 5/7.5. The second color
in the eighth row is Red 8/5.
Ecological zones, though sometimes estimated
from my own experience with Central American
vegetation, are largely taken from Holdridge life-
zone maps (Holdridge, 1967; Holdridge et al.,
1971), where they exist for Central American coun-
tries. Vegetation types for Mexico are taken from
the “Mapa de tipos de vegetación de la República
de Mexico” (Flores et al., 1971).
Each life zone is represented by a full textual
statement and abbreviation which appear on life
zone maps. The Holdridge Life Zones of Central
America and areas where Philodendron occur are
listed here, arranged in a generally drier to wetter
order: Tropical thorn woodland (T-tw); Tropical dry
forest (T-df); Tropical moist forest (T-mf); Tropical
wet forest (T-wf); Tropical rain forest (T-rf); Premon-
tane thorn nd (P-tw); Premontane moist for-
est (P-mf); Premontane wet forest (P-wf); Premon-
tane rain forest (P-rf); Tropical Lower Montane wet
forest; (TLM-wf); Tropical Lower Montane rain forest
(TLM-rf); Tropical wet forest transition to Premon-
tane wet forest (T-wf/P-wf); Premontane wet forest
transition to moist forest (P-wf/mf); and Premontane
wet forest transition to rain forest (P-wf/rf).
Herbarium material has been widely distributed,
and original field vouchers are cited for all herbaria
whose material was seen. Herbarium material may
consist of one of three kinds: (1) complete original
sets (wild collected); (2) sterile original material
with an inflorescence added from a cultivated plant
of the same number; and (3) material collected en-
tirely from cultivated plants. Specimens based en-
tirely or in part on cultivated material are clearly
indicated on the herbarium label.
Herbarium specimens were borrowed from most
major herbaria including: AAU, B, BBS, BISH,
BM, BR, CAS, CAY, CM, COL, CR, DAV, DUKE,
DS, EAP, ECON, ENCB, F, FLAS, FSU, FTG, G,
GH, HBG, ISC, K, L, LA, LL, LE, M, MEXU,
MICH, NY, PMA, RSA, S, SCZ, SEL, TEX, U, UC,
UMO, US, VEN, and WIS.
Descriptions are mostly parallel and as complete
as possible. Descriptions of the pistils, vitally im-
portant in the infrageneric classification of Philo-
dendron, are particularly detailed. In order to avoid
repetition, description references are made to Style
Types discussed by Mayo (1986). These style types
are discussed and illustrated (Fig. 469) in the in-
troduction under the section on *Morphology of Re-
productive Structures—Gynoecium."
Infraspecific categories in this work adhere to the
following definitions. Subspecies are those infra-
specific taxa that are mostly or entirely allopatric
either geographically, elevationally, or ecologically.
Varieties, though morphologically distinct in one or
several characters, are apparently geographically,
elevationally, or ecologically sympatric. All infra-
specific taxa within Central American Philodendron
subg. Philodendron are ecologically sympatric.
ACKNOWLEDGMENTS
This revision could not have been completed
without the help of many people, most important
among them my dedicated and able co-worker, Pe-
tra Malesevich, who participated in nearly every
phase of the work. I could count on her to continue
the research unabated during my long field trips.
Along with Bob Magill, she revised, updated, and
improved the more powerful description program
that we used for the Philodendron revision. Jo Ann
Beiser played an important role for nearly two
years, providing assistance in the preparation of de-
scriptions. Kathy Pickett Upton, former Research
Greenhouse manager, played a major role in main-
taining the live collections, making pollinations,
314
Annals of the
Missouri Botanical Garden
and in dealing with label problems. Very special
thanks also go to my colleague and fellow aroid
specialist Mike Grayum, whose knowledge of the
Costa Rican Araceae is without parallel. He has
not only made many of the best collections of Philo-
dendron, but his breadth of experience in that part
of Central America and his vivid insights into the
interrelationships of species were very rewarding.
In addition, his skills as an editor and his knowl-
edge of Latin and rules of nomenclature contributed
much to this work. Simon Mayo, like Mike Grayum,
conducted extensive work on Philodendron. Their
work was concurrent or preceded my own, and their
ideas and interpretations of phenomena in the ge-
nus were ever insightful. My own work has bene-
fited greatly from their work. Dan Nicolson was al-
ways generous with his time in helping to solve
nomenclatural problems as well as problems with
translations of difficult Latin phraseology. Gordon
McPherson provided pickled material of many spe-
cies of Philodendron during his tenure as our res-
ident botanist in Panama. Eleanor Sauer has proof-
read or written Latin diagnoses for nearly all of the
new species. Joseph Tosi, of the Tropical Science
Center in San José, Costa Rica, assisted us in stan-
dardizing our use of the Holdridge Lifezone System,
which we have added to our computerized data-
base. Leland Russell, summer volunteer and later
summer employee while vacationing from his un-
dergraduate studies at Carleton College in Minne-
sota, became very skillful at dissections of Philo-
dendron inflorescences. His critical comparative
study of the Central American species of Philoden-
dron went a long way in increasing our understand-
ing of the differences within the genus. Finally, I
would like to thank my wife, Patricia, who has with-
out complaint allowed me to spend several months
each year wandering the tropics of Central and
South America for almost three decades. She was
always available to solve every computer glitch and
has also helped with the computerized sorts of in-
formation which led to the construction of the keys.
HISTORY OF THE GENUS PHILODENDRON
Although Philodendron apparently figured in
pre-Columbian folklore, art, and medicine during
the 16th century, and herbarium material was col-
lected by Georg Marcgraf as early as 1644 (Mayo,
1990), it was Charles Plumier who made the first
effective introduction of the genus to European sci-
entists (Mayo, 1990). He collected five or six spe-
cies from Martinique, St. Thomas, and Hispaniola,
giving phrase names beginning with “Arum” or
“Dracunculus.” See Mayo (1990) for a detailed ac-
count of Philodendron collections made by other
late 17th-century and 18th-century explorers, in-
cluding Hans Sloane in Jamaica and N. J. Jacquin
in the West Indies, Colombia, and Venezuela.
Plumier's expeditions resulted in the first species
of Philodendron to be published as new to science,
albeit as an Arum, A. lingulatum L., a member of
Philodendron subg. Pteromischum. His trips also
resulted in Philodendron hederaceum, the first
member of P. subg. Philodendron to be published
new to science, as Arum hederaceum Jacq.
HEINRICH WILHELM SCHOTT
One hundred thirty-six years passed from the
time Plumier first introduced plants of what came
to be known as Philodendron to European scientists
in 1693 and the first circumscription of the genus
by Schott in 1829. During this interim a number of
workers, including Carl Linnaeus, worked with ge-
neric concepts and decided that not all aroids be-
longed in the same genus. Arum came to be used
only for the European plants it now represents and
Arisaema, Dracunculus, and Colocasia were also
separated by the early 18th century (Hermann,
1698; Tournefort, 1700). By the fifth edition of Gen-
era Plantarum Linnaeus (1754) had recognized
also Calla, Dracontium, and Pothos as well as
Orontium, Pistia, and Acorus, although not recog-
nizing the last three as related to Arum.
By the middle of the 18th century the exploration
of the Neotropics was well under way. The intro-
duction of so many new plants from the New World
tropics, including many Philodendron species,
made it clear that further separation was needed.
Because of the uncritical acceptance of many ab-
errant elements into Arum during the course of the
18th century a number of taxa now recognized as
Caladium, Philodendron, and Syngonium were 1n-
corporated into Arum. Ventenant (1800) solved part
of the problem by recognizing Caladium, but Willd-
enow (1805) erred in transferring into Caladium
four of Jacquin’s West Indian Philodendron species
(all considered Arum at the time).
Even by the time of the 16th edition of Systema
Vegetabilum Sprengel (1826) did not distinguish
Philodendron but had increased the number of gen-
era to 12, adding Ambrosina, Arisarum, Caladium,
Gymnostachys, and Zantedeschia. +
Resolution of many of the remaining generic
problems with the Araceae awaited Austrian bota-
nist Heinrich Wilhelm Schott, who was the first to
focus on the taxonomy of the Araceae (Nicolson,
1960). Schott was uniquely qualified in this regard»
having spent four years in Brazil collecting plants
е өөө өөө toU LAM A дин и aA conem _ ден мо“ ll, NWN т т ы
Volume 84, Number 3
1997
Croat 315
Philodendron Subgenus Philodendron
and with access to the large collection of living
plants at the Imperial Gardens [of the Hapsburg’s]
at Schónbrunn Palace in Vienna where he worked
as the director. These included his own Brazilian
collections as well as those of N. J. Jacquin from
the Caribbean. In 1829 Schott described the genus
Philodendron (published as Philodendrum) Schott
in one of his first publications after returning from
Brazil in 1821. The first species placed in the ge-
nus were: P. grandifolium (Jacq.) Schott, P. laci-
niosum Schott, P. tripartitum (Jacq.) Schott, P. lac-
erum (Jacq.) Schott, P. pinnatifidum (Jacq.) Schott,
and Р. hederaceum (Jacq.) Schott. In 1832 Schott
published a preliminary classification of the Ara-
ceae in a work entitled Meletemata Botanica, which
included many plant families and was done in con-
junction with his colleague S. Endlicher. In this
work he recognized 39 genera of Araceae including
Philodendron and some other by now well known
species-rich genera such as Anthurium, Syngoni-
um, Dieffenbachia, Aglaonema, and Spathiphyllum.
In this work he also published the first infrageneric
system of classification for Philodendron, although
it contained only three groups still recognized to-
day. Though Schott subsequently continued to work
on his circumscription of the genera and species of
the Araceae for the remainder of his career, he did
not publish any majer revisions of Araceae for 24
years.
Endlicher (1837), publishing alone but with the
obvious assistance of Schott (Mayo, 1990), pro-
duced a revision of Philodendron that differed from
the Meletemata Botanica account by Schott only in
having a more complete generic description, in-
cluding vegetative details which were presented for
the first time.
Shortly after Endlichers revision, K. S. Kunth
(1841) published in his “Enumeratio Plantar-
um...” the first species-level revision of Philoden-
dron. This revision included new species and new
combinations and for the first time included species
of the genus that would -— = recognized as mem-
bers of P. subg. Pterom
It was not until Schott’ (1856) publication of a
work entitled Synopsis Aroidearum that a fully de-
veloped infrageneric system of classification was re-
alized. This was Schott’s first species-level revision
of the genus, and the Synopsis divided the genus
into 22 “greges” grouped in 7 unnamed categories
arranged in increasing order of complexity of leaf
blade shape (Mayo, 1990). A total of 99 species
(which was recognized here for the first time as grex
Pteromischum) and 3 species of P. subg. Meconos-
tigma were included (two of them reported as mem-
bers of grex Sphincterostigma).
Schott’s final classification of Philodendron was
published four years later in the Prodromus Syste-
matis Aroidearum (1860), a more rigorous work that
came to be his last comprehensive self-published
work, since he died at the age of 71 in 1865. This
revision included 110 genera; almost all are still
recognized as genera or subgenera. The fact that
his work has stood the test of time is a testimony
to the serious nature of the research he had done
in Vienna during his long period of seeming inac-
tivity.
Schott’ treatment of Philodendron in the Prod-
romus differed from his revision in the Synopsis in
having more complete descriptions, using more in-
florescence characters; in having six rather than
seven unnamed categories of leaf shape to group
the “greges”; and in beginning to make use of the
cataphyll (prophyll in the strictest modern sense)
as a character. Schott used the term “subopposite
stipule” for the feature that came to be known as
the cataphyll in Engler’s usage. This has proven to
be one of the best and most reliable characters for
the genus. For a detailed listing of the key char-
acters for Philodendron used in Schott’s Prodromus
revision, refer to Mayo (1990: 50).
The Prodromus treatment included 135 species
of Philodendron included in the same “greges.”
Twenty species of the total were members of P.
subg. Pteromischum Schott and 6 species are now
placed in P. subg. Meconostigma (Schott) Engl. (3
in grex Meconosti and 3 in grex Sphincterostig-
ma). Thus Schott included 110 names of Р. sect.
Philodendron. Taking synonymy into account, only
76 species of P. sect. Philodendron were included
in this 1860 revision. Only 18 names pertained to
Central America. Of these, only 8 are accepted
members of P. sect. Pteromischu.
The species included by Schott (1860) in each
grex of the Prodromus for Central America are list-
ed below:
=
Grex Baursia Rchb. ex Schott: P. wendlandii Schott
Grex Pteromischum Schott: P. aurantiifolium
Schott, P. ligulatum (L.) K. Koch, P. seguine
Schott, P. inaequilaterum Liebm.
Grex Canniphyllum Schott: (no species represent-
e
Grex Glossophyllum Schott: (no species represent-
ed)
Grex Solenosterigma Klotzsch: P. oxycardium
ott, P. micans K. Koch, P. scandens K.
Koch & Sello — (P. hederaceum)
Grex Psoropodium Schott: P. tenue K. Koch & Au-
gustin, P. gracile Schott
Annals of the
Missouri Botanical Garden
Grex Achyropodium Schott: P. verrucosum L. Ма-
thieu ex Schott
Grex Platypodium Schott: P. pterotum K. Koch &
Augustin, P. fragrantissimum (Hook.) Kunth
Grex Cardiophylacium Schott: P. brevispathum
Schott, P. hederaceum Schott (= P. jacquinii
Schott)
Grex Belocardium Schott: P. hoffmannii Schott, P.
advena Schott, P. acrocardium Schott
Grex Cardiobellium Schott: (no species represent-
d
e
Grex Meconostigma Schott: (no species represent-
ed
Grex Eubelium Schott: (no species represented)
Grex Macrolonchium Schott: (no species represent-
ed)
Grex Macrobelium Schott: P. daemonum Liebm. =
P. sagittifolium Liebm., P. sagittifolium Schott,
P. tanyphyllum Schott — P. sagittifolium
Liebm.
Grex Imbéa: (no species represented)
Grex Oligophlebium Poepp.: (no species represent-
e
Grex Doratophyllum Schott: (no species represent-
ed)
Grex Schizophyllum Schott: (no species represent-
d)
e
Grex Tritomophyllum Schott: P. tripartitum (Jacq.)
Schott, P dagilla Schott = Р. tripartitum
(Jacq.) Schott, P. anisotomum Schott
Grex Polytomium Schott: P. subincisum Schott, P.
impolitum Schott — P. radiatum Schott, P. po-
lytomum Schott — P. radiatum Schott, P. war-
szewiczii K. Koch & Bouché
Grex Sphincterostigma Schott: (no species repre-
sented)
In all, only 18 of the 135 species of Philoden-
dron included in this 1860 revision were from Cen-
tral America, and 4 of the total were members of
P. grex Pteromischum. One of the four species, P.
ligulatum Schott, is a member of P. sect. Philoden-
dron, which Schott had inadvertently placed in his
grex Pteromischum. Five of Schott's names became
P. hederaceum (P. acrocardium, P. hoffmannii, P.
oxycardium, P. scandens, and P. micans), but the
count was reduced by only four since the name P.
hederaceum was involved in two currently accepted
species (namely, P. hederaceum and P. jacquinii).
Other reductions are P. gracile, which is a synonym
of P. tenue; P. daemonum and P. tanyphyllum,
which are synonyms of P. sagittifolium; P. dagilla,
a synonym of P. tripartitum; and P. impolitum and
P. polytomum, synonyms of Р. radiatum. Thus
Schott was dealing with only 16 (16.5%) of the 96
currently known Central American species.
ADOLF ENGLER
Litle was done with Philodendron following
Schott's death in 1865 until Adolf Engler, working
at the Universities of Kiel and Breslau (and finally
at the Berlin Botanical Garden), began his revi-
sionary work on the Araceae. Schott had laid the
groundwork, describing most of the genera that still
exist today, but he was dealing with only a small
portion of the species. Taking up his first positions
at Kiel and Breslau in 1871, at age 27, Engler
worked with various tropical families on the Flora
Brasiliensis project, publishing and working on a
general review of the vegetative and floral mor-
phology for the entire family (Engler, 1877). Later,
in his powerful position as Director of the Berlin
Botanical Garden, Engler commanded attention
and a wealth of herbarium specimens and living
material from all over the world during Germany's
preeminent period of domination in the botanical
world preceding World War II. Since Engler was
only 21 at the time of Schott's death, it is not likely
that the two ever met, but Engler would have had
access to some of the same material, including liv-
ing material from the Schónbrunn greenhouses, as
well as Schott's notes and illustrations made over a
40-year period (Engler, 1876). Unlike modern
workers, he had access to Schott's herbarium in Vi-
enna before it was destroyed during World War II.
This is important for a continuum of species con-
cepts in groups often described from inadequate
material of unknown origin and worse yet some-
times destroyed by war.
Engler's (1878) treatment of the Araceae in Mar-
tiuss Flora Brasiliensis included 116 species of
Philodendron, 95 of which were members of P. sect.
Philodendron (13 were in P. sect. Pteromischum
and 8 in P sect. Meconostigma (Schott) Engl.). Only
47 species of Philodendron were then known from
Brazil. The work also included sectional and spr
cies descriptions as well as a key to all existing
species of Philodendron. In this work Engler mod-
ified Schott’s system of classification for Philoden-
dron by recasting Schott’s “greges” as sections and
reducing the number from 22 to 10. He also syn-
onymized a number of species, reducing the total
from 132 to 116 species. Only a single Central
American species, P. oxycardium (= P. hederaceum
var. oxycardium) was reported for Brazil, a fairly
accurate picture as we know today. Only a couple
of other species, namely P. fragrantissimum and Р.
verrucosum, have been found to range into the Am-
Volume 84, Number 3
1997
Croat 317
Philodendron Subgenus Philodendron
azon drainage of South America. Two additional
species, P. glanduliferum Matuda and P. brevispa-
thum, have a subspecies or variety that occurs in
the Amazon basin of South America, but the same
do not occur in Central America.
In the following year Engler's treatment of Philo-
dendron for A. and C. DeCandolle's Monographie
Phanerogamarum (Engler, 1879) was essentially
unchanged, adding only 3 species to bring the total
to 120. Of these, 93 species were members of P.
subg. Philodendron and 20 of the epithets repre-
sented species currently known from Central Amer-
ica (now reduced to 15 through synonymy).
The final revisionary effort by Engler on Philo-
dendron published 20 years later (Engler, 1899)
was changed only slightly at the subgeneric level
from the 1878 work. One section was raised to sub-
generic status, and the remaining nine sections
were included in P. subg. Euphilodendron Engl.
(now P. subg. Philodendron). Despite minor
changes made by Krause (1913), it is essentially
Engler's classification that persists almost a century
later. Engler’s (1899) revision was substantially
larger than Schott's last revision. Engler's revision
contained 167 species, 134 of them in P. subg.
Philodendron, with 23 reported for Central America
(reduced to 15 species through synonymy in this
revision). One species, P. purpureoviride Engl., re-
ported for Ecuador, is now known for Central Amer-
ica.
The species of P. subg. “Euphilodendron” in-
cluded in Engler’s (1899) revision broken down by
section for Central America are listed below:
P. sect. Pteromischum Schott: P. aurantiifolium
Schott (as synonym of P. guttiferum Kunth), Р.
guatemalense Engl., P. inaequilaterum, P. se-
guine, P. talamancae Eng
P. sect. Baursia (Rchb. ex Schott) Engl: P. wen-
dlandii Schott
P. sect. Polyspermium Engl.
“Gruppe” Platypodium Schott: P. pterotum
“Gruppe” Solenosterigma Klotzsch: P. oxycardium
(= B hederaceum var. oxycardium), P. pur-
pureoviride (as P. purpureoviridis from South
America), P. micans K. Koch (= P. hedera-
ceum var. hederaceum forma micans), P. scan-
dens (= P. hederaceum var. hederaceum)
niet Cardiobelium Schott: P. brevispathum, P.
acile (= P. tenue), P. schottianum, P. tenue
“Gruppe” Achyropodium Schott: P. verrucosum
P. sect. Oligospermium Engl.
“Gruppe” Macrobelium Schott: P. sagittifolium,
> daemonum, P. sanguineum Regel (=
sagittifolium), P. mexicanum Engl.
“Gruppe” Belocardium Schott: P. ligulatum
Schott, P. advena, P. subovatum Schott (=
P. advena), P. smithii Engl.
“Gruppe” Oligocarpidium Engl.: P. рипеп Engl.
(= P. hederaceum)
P. sect. Tritomophyllum (Schott) Engl.: P. anisoto-
mum, P. tripartitum, P. fenzlii Engl. (= P. tri-
)
P. sect. Schizophyllum (Schott) Engl.: no species
represented
P. sect. Polytomium (Schott) Engl.: P. augustinum
K. Koch (7 P. radiatum), P. radiatum Schott,
P. warszewiczii K. Koch & Bouché
P. sect. Macrolonchium (Schott) Engl.: P. fragran-
tissimum
P. sect. Macrogynium Engl.: P. hoffmannii Schott
sensu Engl. (= Р. jacquinii Schott)
The turn of the century saw major activity with
Philodendron, no doubt due to Englers just pub-
lished revision. Engler made no changes in his re-
vision but went on to publish 26 additional species
(Engler, 1905b). In addition, seven species were
described by Alfred Barton Rendle, Ignaz Urban,
Ambroise Gentil, and N. E. Brown between 1901
and 1908
KURT KRAUSE
Kurt Krause, who began working with Engler at
the Berlin Botanical Garden on 1 January 1905,
described two additional species before preparing
his revision of Philodendron for Das Pflanzenreich
(Krause, 1913). The latter remains the most recent
revision of the whole genus. Krause's revision is a
slightly reworked version of Engler’s (1899) revi-
sion but did include the description of a new sec-
tion, P. sect. Camptogynium, with a single species
in P. subg. Philodendron (“Euphilodendron”) and
included 55 more species. There were 32 other new
species published in P. subg. Philodendron. Six of
these were in P. sect. Pteromischum (P. subg. Pter-
omischum), while one was in P. subg. Meconostig-
ma. The remaining 25 were in P. subg. Philoden-
dron. Most were members of P. sect. Baursia and
P. sect. Polyspermium (Philodendron) with a single
species each in the following sections: Oligosper-
mium (Calostigma), Schizoplacium, Macrolon-
chium; and three species in P. sect. Polytomium.
Only two species, P. grandipes K. Krause and P.
panamense K. Krause (both in current Р sect.
Philodendron), were from Central America. Krau-
se's treatment of P. subg. Philodendron included the
following sections and species for Central America
(or at least now known from Central America):
Names used by Krause (1913) and their current
status. Species numbers refer to those in Krause's
2. P. sect. Baursia (Rchb. ex Schott) Engl.
wendlandii Schott
95. P. scandens K. Koch & Sello = P hedera-
ceum
96. P. oxycardium Schott = Р. hederaceum var.
99. P. micans K. Koch = Р. hederaceum var.
hederaceum
K. Krause
117. P. gracile Schott = P. tenue K. Koch &
Augustin
116. P. tenue K. Koch & Augustin
Us P de = P. hederaceum (Jacq.)
5. ia eae M ds
cies represented i
7. P sect. Polytomium (Schott) Engl.:
195. P. radiatum
198. Р. augustinum К. Koch = P radiatum
Schott
199. P. polytomum Schott P. radiatum Schott
* Philodendron pedatum (Hook.) Kunth reported
Ја Hell off Corin Rica Suk de Diu a АТ y
available for confirmation.
davisianum G. S. Bunting, P. lancigerum Standl. &
tuda = Р. advena, P. P
Standl. & L. O. Williams (= P gran
dipes K. Krause), P. - p
radiatum var. (Matuda) Croat), and
P. trisectum Standl. (7 P. anisotomum).
Three additional species were described in
Philodendron but actually pertain to other genera
ond иде дин Mensa"
Many of the other taxa in Krause's revision were
synonymized or reduced in rank, and only 11 taxa
Philodendron remained.
in P. subg. were: P.
brenesii, P. radiatum var. auri-
culatum, P. microstictum, P. basii, P. glanduliferum,
P. dressleri, P. P. platypetiolatum, P.
davidsonii, and P. strictum.
Most of the Central American floristic work out-
side of Mexico was carried out by Paul C. Standley,
often working with his associate Louis O. Williams.
Standley worked initially at the Smithsonian and
later at the Field Museum in Chicago, then at the
herbarium of the Escuela Agrícola Panamericana,
where he died at Zamorano in Honduras. Standley
described P. brenesii and P. trisectum (= P. aniso-
tomum Schott) alone, and with L. O. Williams he
also provided the following epithets: P. armigerum
(= armigerum (Standl. € L. O. Wil-
liams) Croat), popa auriculatum, P. brevinodum (=
Monstera tuberculata Lundell var. brevinoda
(Standl. & L. 0. Williams) Madison), P. hastiferum
workers did not succeed in describing many of
them since of the nine described, three proved to
belong to other genera, and three others proved to
be synonyms of existing Philodendron names. This
is particularly surprising since both Standley and
L. O. Williams were astute observers who were very
familiar with the Central American flora in general.
Their mistakes point out the complexity of the tax-
onomy of Araceae and the bewildering array of ma-
terial available to them at that time. Even when 1
began my own work with the Araceae in the late
1960s there were few specimens (aside from types)
that proved to have the correct names. Specimens
nor
иі» Ul MCSE ELIE,
especie ef tho pl sora ex dick pius
understanding
America чеге ойей inade-
quately done, even if that
ава ‘The
treatment of the Flora of Panama (Standley, 1944),
for example, is woefully inadequate considering the
small percentage of the total aroid flora that is cov-
ered compared to what is now known to exist.
In his treatment of some genera Standley (1944)
seemed too willing to accept epithets of species de-
scribed in Colombia, regardless of how well they
“fit” Panamanian species. As a result many species
names in genera such as Anthurium, for example,
Philodendron subg.
he included in the Flora of Panama were: P. bre-
nesii, P. brevispathum, P. grandipes, P. panamense,
P. radiatum, P. tripartitum, and P. ~ Не
did not до so well with members of P. subg. Pter-
omischum, where P. karstenianum Schott was a
mixture of two species and P. guttiferum was а mix-
ture of three species. Perhaps the most curious
thing about Standley’s Flora of Panama treatment
is that by 1944, after Robert Woodson and his col-
laborators had already made several expeditions to
Panama, so few of the new species included in the
present revision had been collected. Standley had
collected some of the new species but failed to rec-
ognize them as new. These included: Р. crassispa-
thum (Standley & Valerio 51910), P. findens (Stan-
dley & Torres 52355), P. purulhense (Standley
89902), P. strictum (Standley 51371), P. verapazense
(Standley 91978), and Р. wilburii var. wilburii
(Standley 38300). See discussion of those species
for additional details. See also section on “Collect-
ing History of P. subg. Philodendron.”
The Flora of Guatemala (Standley & Steyermark,
1958b) was much more accurate and complete in
the percentage of the total taxa of P. subg. Philo-
Annals of the
Missouri Botanical Garden
dendron included. However, this probably has less
to do with the fact that it was published 14 years
later than it does with the fact that there are fewer,
generally more widespread species occurring there
than in Panama. The Flora of Guatemala treated
11 species of Philodendron, 8 of them members of
P. subg. Philodendron. These were: P. anisotomum,
P. hederaceum, P. hoffmannii (= P. jacquinii), Р.
radiatum, P. sagittifolium, P. smithii, P. tripartitum,
and P. warszewiczii. With 50% of the present total
taxa included and with all but one of the taxa still
properly bearing the name, the Guatemala treat-
ment remains generally more useful than that of the
Flora of Panama. Added to the flora since the 1958
revision are: P. advena, P. fragrantissimum, P. glan-
duliferum, P. jodavisianum, P. mexicanum, P. pu-
rulhense, and P. verapazense.
The Flora of Guatemala (Standley & Steyermark,
1958b) treated P. hederaceum correctly but treated
P. jacquinii under its synonym, P. hoffmannii. Cu-
riously, however, the illustration used represents
both species. The leaf and stem seem clearly to be
P. hederaceum, but the inflorescence clearly shows
the long-protruded styles of P. jacquinii.
Standley's (1937) treatment of the Araceae of
Costa Rica was reasonably good, partly because
many species had been described by Schott from
collections made by H. Wendland in Costa Rica.
Other widespread species whose taxonomy had
been well established were also a part of the flora.
Properly named Costa Rican species recognized by
Standley were: P. brenesii, P. ligulatum, P. pterotum,
P. radiatum, P. schottianum, P. tripartitum, P. ver-
rucosum, and P. wendlandii. Species now synony-
mized are: P. gracile (= P. tenue), P. hoffmannii (=
P. jacquinii), P. pittieri (= P. hederaceum), and P.
trisectum Standl. (= P. anisotomum). It has never
been determined which species he included under
the name P. panamense, but that species is not
known for Costa Rica. Thus, with 8 out of the 13
names correct and 3 additional species that at least
represent synonyms of currently recognized spe-
cies, Standley did pretty well. However, with only
13 of the current 49 species treated (2696), the
treatment was no more complete than that of the
Flora of Panama, which was written a few years
later.
Though no other floristic taxonomist had such a
prominent role with Central American Philoden-
dron as Standley, there were others who described
Philodendron during the course of their floristic
work.
Ivan M. Johnston of the Arnold Arboretum de-
scribed Philodendron erlansonii (= P. jacquinii)
and P. harlowii (= P. hederaceum) while working
on the flora of San José Island (Johnston, 1949) of
Panama.
In Mexico, Eizi Matuda, the local aroid special-
ist, described seven species (Matuda, 1954): P.
apocarpum (= P. jacquinii), P. basii, P. glandulifer-
um, P. latisagittium (7 P. mexicanum), P. miduhoi
(= P. hederaceum), P. monticola (= P. advena), and
P. pseudoradiatum (— P. radiatum var. pseudora-
diatum); and George Bunting described four spe-
cies during his investigations of Mexican Araceae
(Bunting, 1965): P. dressleri, P. jamapanum (= P.
advena), P. jodavisianum, and P. tuxtlanum G. S.
Bunting (= Р. sagittifolium).
Matuda's treatment of the Philodendron in Mex-
ico (Matuda, 1954) dealt with 16 species, 13 of
them in P. subg. Philodendron. While a number of
the species had the correct names, e.g., P. advena,
P. mexicanum, P. pseudoradiatum, P. radiatum, and
P. tripartitum, others had synonymous names, e.g.,
P. sanguineum and P. daemonum (both currently
recognized as Р. sagittifolium). Matuda also recog-
nized P. mexicanum under three different names,
namely P mexicanum, P. latisagittatum, and P.
sagittifolium (a distinct species that he treated as
both P. daemonum and P. sanguineum (a currently
recognized name improperly used; see above). Ma-
tuda redescribed two species using the names Р
apocarpum and Р. miduhoi (currently Р. jacquinú
and P. hederaceum, respectively). Finally, he de-
scribed P. monticola (now considered synonymous
with P. advena).
Other Central American species of P. subg.
Philodendron published since the time of Krause's
revision and prior to the beginning of this project
in 1986 are P. davidsonii and P. platypetiolatum,
the latter described from Ecuador during Mike
Madison's extensive work with the flora of Ecuador
during his tenure at the Selby Botanical Garden.
Thus, up to the initiation of the current study, only
26 species in P. subg. Philodendron were described
for Central America. With 96 species of P subg.
Philodendron now known for Central America, this
registers an increase of 70 species or a 27096 in-
crease.
Of the species of P subg. Philodendron de-
scribed since Krause's revision (not including the
present effort) only seven were described by non-
specialists (I. M. Johnston, P. C. Standley, and Stan-
dley and L. O. Williams). Standley, together with
L. O. Williams, described most of the new Centra
American species and wrote most of the floristic
treatments of the Araceae for Central America. De-
spite his extensive experience, 1 do not believe that
Standley fully understood the diversity of the Ara-
ceae. This is because he failed to recognize a rel-
Моште 84, Митбег 3
1997
Croat 321
Philodendron Subgenus Philodendron
atively large number of species that were new to
science. In Standley’s defense it must be stated that
the Araceae are a particularly complex family with
so much interspecific variation that proper deci-
sions often cannot be easily made without direct
comparisons of living material. In addition, collec-
tions in the past often had few or no field notes to
use for study.
COLLECTING HISTORY
In addition to the history of revisionary efforts it
is instructive to consider the collecting efforts in
Central America that have laid the groundwork for
the scientific work done with Philodendron subg.
Philodendron. Perhaps owing to their sometimes in-
timidating size, the difficulty of retrieval, and cer-
tainly due to the difficulty of preparing and drying
specimens, members of P. subg. Philodendron have
never been popular with botanical collectors. The
early neotropical collectors, i.e., Ruiz and Pavon,
Sessé and Мосто, Triana and Planchon, and others
collected few Araceae (or at least few survived to
modern times). Eduard Poeppig, working in Peru,
and Richard Spruce, working in the Amazon region
of Brazil, did somewhat better, collecting a number
of new species. In Central America few collectors
made many collections until modern times. Even
Standley and Julian A. Steyermark, two of the most
prodigious collectors in the region, did not make
many collections of Araceae. Both gathered well
over 100,000 collections in their careers. Yet in all,
Standley made only 146 collections of P subg.
Philodendron, comprising 21 species in all of Cen-
tral America. Of this total only 6 of the collections
(these previously mentioned) proved to be unde-
scribed species, none of which Standley recognized
as new. Standley was not avoiding collecting Philo-
dendron because they constituted a lot of work to
press. This is demonstrated by the fact that he col-
lected P. radiatum 13 times and P. шагзгеилсхи 12
times. These are among the most difficult plants to
prepare owing to their huge size and fleshy parts.
That so few new species of Philodendron were
collected was apparently due to the fact that in ear-
lier times relatively few roads led into areas of wet
forest.
Matuda, working exclusively in Mexico and con-
centrating on Araceae, accrued 27 collections of P.
subg. Philodendron and only 1 of these, P. glan-
duliferum, proved ultimately to be new to science.
George Bunting, also working only in Mexico (un-
der the numbers of Harold Moore of Cornell Uni-
versity), made 36 collections of P. subg. Philoden-
dron, including 2 new species (P. dressleri and Р
jodavisianum).
teyermark, though collecting many more Philo-
dendron in Venezuela later in his career, collected
only 26 Central American P. subg. Philodendron,
comprising 9 species, none of which were new. L.
O. Williams, who also worked on the Flora of Gua-
temala and collected for many years in Guatemala,
Honduras, and Costa Rica, made only 14 collec-
tions of P. subg. Philodendron. Two of these were
species that remained undescribed until this work,
namely Williams 628 (P. sulcicaule) and Williams
28387 (P. wilburii var. longipedunculatum Croat &
Grayum). Percy Gentle, collecting in Belize, made
20 collections. The Philodendron subg. Philoden-
dron collections of Pittier in Costa Rica totaled
nine, none representing new species. Adolf Tonduz,
also working in Costa Rica, made only five collec-
tions of this group. Alberto Brenes made four col-
lections in Costa Rica between 1926 and 1932. Two
of them, P. brenesii and P. bakeri, proved to be new.
Paul Allen made five collections of Philodendron
in Costa Rica, four of which were described as new
(though two were subsequently synonymized).
Aside from these few collections of new species
mentioned above, no other new species were col-
lected until the early 1960s. Roy Lent, living in
Costa Rica and collecting between 1964 and 1971,
made 24 collections of P. subg. Philodendron, in-
cluding 5 new taxa. These are: P. lentii Croat 4
Grayum, P. hederaceum var. kirkbridei, P. strictum,
P. thalassicum Croat & Grayum, and P. wilburii var.
wilburii. W. C. Burger, collecting in Costa Rica be-
tween 1968 and 1986, in part with one-time aroid
specialist Richard Baker, made 56 collections of P.
subg. Philodendron, including 5 new species, P.
bakeri, P. chirripoense, P. crassispathum, P. thalas-
sicum, and P. wilburii.
Though Panama is even richer in Araceae than
Costa Rica, the collecting activity there was not
particularly rewarding. The Philodendron collec-
tions of H. von Wedel, who worked in Bocas del
Toro Province in Panama, totaled only seven. Rob-
ert Woodson and his collaborators, Paul Allen, and
Carrol Dodge collectively made only 11 collections
before the Flora of Panama project was begun.
This occurred in a country that proved to have 96
species, 65 new to science.
Even James Duke, who regularly got into areas
of wet forest in many parts of Panama, made only
four collections of P. subg. Philodendron, none of
them new.
Collecting activities begun by Walter H. Lewis
and staff from the Missouri Botanical Garden in the
early 1960s were more aggressive by using heli-
322 Annals of the
copters, and made it into areas not previously ac- serving as the Missouri 's resident
cessible. Despite this greater penetration, even botanist in Panama, during 1970-1971, and while
total, 153 were of P. subg. for Central
His collections are particularly useful,
since they are accompanied by excellent field
notes. He was ible for collecting 13 unde-
given day proved to be new to science. Later, when
a aor a Ya думи ере
made in Central America, and 1594 of those wens
members of P subg Central
Amencan collections ба andes
FOSSIL. HISTORY
The fossil history of the Araceae was reviewed
1911) shared the subfamily with seven other tribes,
А р
onemateae, Dieffenbachieae, ha dé
phonodoreae, and Peltandreae. ee E
Philodendron v Homalomena and
Bogner and Nicolson (1991) left Engler's subfamily
Philodendroideae intact, but Grayum (1984) made
of
in its own tribe Phil-
dron close to H
Araceae (Grayum, 1990). Grayum believed that the
Phi are a sister group to the Pothoi-
deae (including Englers Monsteroideae), which
have in common the exclusive characteristics of ge-
cepts are still evolving. A comparison of the major
systems of classification at the suprageneric level
was made by Croat (1990[1992]). It included the
systems of Hotta (1970), Grayum (1990), and Bog-
ner and Nicolson (1991).
which to distinguish it completely from other gen-
era. He reported the genus to be only distantly re-
lated to other genera in the subfamily but that its
closest relatives were the African genera Culcasia
and Cercestis. In his survey of sclerotic hypodermis
in the roots of Araceae, French (1987a) provided
evidence to link Philodendron to the West African
genera Anubias, Culcasia, and Cercestis and the
sr es Montrichardia.
in the systems of
and Hay and Mabberley (1991), Mayo et al. (1995)
conducted another sweeping survey produced
a cladistic analysis. While maintaining essentially
the same alliances by Grayum (1990),
suggested
— PERRA
unisexual flowers i
restriction
Ariflorae by French et al. (1995) places Philoden-
dron as a sister group to Homalomena, suggesting,
according to Grayum (1996), that Homalomeninae
letic
«d Mayo et al. (1995) has taken into account all
the evidence to date including the extensive mo-
lecular studies by French
sometimes requiring
be thet determine te gms Siete p
—— Dieffenbachia are distant from one
ноо ње
spathe, while those of Philodendron аге closely com-
pacted, devoid of staminodia, and borne on a spadix
б only for dem he apart, in
visible. К шск кийме
noting terrestrial habit (consistently true of Dieffen-
much more problematic
are rarely ovate and never truly cordate, whereas this
blade shape is common in Philodendron. Philoden-
dron may, however, have blade shapes that closely
those of
subg. Philodendron; the latter generally has a very
short sheath, while it is rare that the sheath of Dief-
ia does not extend to the middle or above the
middle of the petiole.
Live material of neotropical Homalomena is not
easily confused with Philodendron because the for-
324
Annals of the
Missouri Botanical Garden
mer usually has anise-scented sap, while Philoden-
dron usually has a distinct turpentine-like aroma,
sometimes also like fresh carrots, but never anise-
scented. Philodendron and Homalomena may have
similar leaves, but the latter often has pubescence
on the blades and pubescence and/or scales and
spines on the petioles. These two genera have sim-
ilar inflorescences with unisexual flowers, sterile
and fertile sections of the staminate portion of the
spadix, and a close arrangement of pistillate flow-
ers, not to mention the similar constricted spathe
that persists after anthesis. However, Homalomena
can usually be determined by the presence of min-
ute, club-shaped staminodia sparsely scattered
among the pistils.
Sterile specimens of neotropical Schismatoglottis
may be confused with Philodendron by the novice
because the two genera share similar venation.
However, Schismatoglottis always occurs terrestri-
ally, often in somewhat marshy situations where
ilo ron rarely occurs. In fertile condition
they are easily separated by the spathe promptly
dehiscing above the tube in Schismatoglottis, with
the staminate portion of the spadix falling free. By
contrast, in Philodendron the spathe is thick and
persistent, usually reclosing over the staminate por-
tion of the spadix, which in turn rots away inside,
never really falling free until the spathe opens in
ruit.
Sterile material of Spathiphyllum has been con-
fused with Philodendron, but that genus differs by
its consistently terrestrial habit, long-sheathed pet-
ioles (exhibited ы Philodendron only in Р. subg.
Pteromischum), the presence of trichoscherieds,
and by its silo closely spaced primary lat-
eral veins.
INFRAGENERIC RELATIONSHIPS
Philodendron is currently divided into three sub-
genera. А subgeneric system of classification for
Philodendron was proposed as early as 1832 by
Schott, who recognized four unranked groups: Eu-
philodendron, Calostigma, Meconostigma, and
Sphincterostigma. The latter two were combined by
Engler (1899) as P. subg. Meconostigma. Schott's
Calostigma was later called P. sect. Oligospermium
Engl. (Engler, 1878) and is once more called P.
sect. Calostigma [(Schott) Pfeiffer] (Mayo, 1990).
uphilodendron became P. sect. Polyspermium in
Englers Flora Brasiliense treatment in 1878 and
must now be treated according to the Code (Greuter
et al., 1994) as P. sect. Philodendron (Mayo, 1990).
It was not until Kunth’s (1841) treatment that
members of what are now called P. subg. Pterom-
ischum were removed from Monstera and placed in
Philodendron. Schott recognized Pteromischum as
a grex in his 1860 Prodromus, and Engler first rec-
ognized the species occurring in this group as P.
sect. Pteromischum in his Flora Brasiliensis treat-
ment (Engler, 1878).
Phylogenetic and phenetic analyses by Mayo
(1986, 1988) have shown Philodendron to have
three subgenera distinct in vegetative and floral
morphology, floral anatomy, and to some extent by
distribution. Philodendron subg. Meconostigma,
with a predominantly southeastern South American
distribution, is highly apomorphic but cladistically
primitive in the genus (Mayo, 1990). Based on a
study of gynoecial morphology Mayo considers P.
subg. Meconostigma to have evolved in eastern Bra-
zil as a group adapted to open habitats and later
spread into the more humid Amazon basin. By the
same standard he assumed that P. subg. Philoden-
dron and P. subg. Pteromischum also arose later
and became predominant as hemiepiphytes in hu-
mid forests. He considered P. subg. Pteromischum
to be a sister group to P. subg. Philodendron and
that P. subg. Philodendron is the most advanced of
the three subgenera. The geological history of the
continent would probably support this since eroded
mountain plateaus of eastern Brazil are much older
than the current land surfaces of the Amazonian
basin. Most of the species of the genus, now so rich
on the Andean slopes of northern and western
South America, surely must have evolved since the
Andes arose during the late Cenozoic.
Mayo elevated P. sect. Pteromischum to the sta-
tus of subgenus (Mayo, 1989) and Grayum (1996)
subdivided the subgenus into two sections, P. sect.
Pteromischum (Schott) Engl. with sylleptic* sym-
A
* Sylleptic shoots are shoots that develop from lateral
meristems without any cessation of activity after initiation,
, growth is continuous with the lateral shoo 1
taking over from the main axis, which — in e
ering in most Araceae genera. Sylleptic s sympodial
characteristic of P subg. Philodendron, results qe
branching occurs from a non-resting lateral bud such that
the existing stem with its terminal inflorescence is im-
eii overtopped by continued growth of the axillary
branch in a manner that makes it appear that the gro
of m se is indeterminate and that the inflorescences
produced appear to be axillary to Ше) й produced by the
ncw stem segment. This type of gro though in reality
a series of branches, each with a Le pues (prophyll
of Ray), a single stem segment, and a single leaf (8 (sym-
podial foliage leaf of Ray; metaphyll of com ае ance
minated by an inflorescence, appears to be an u nc
stem producing a continuous series of cata maig ed
continuous series of what appears to be laterally sach
leaves -— with an axillary inflorescence (see hg fig.
Ray, 198
ај CITAS
1
|
4
1
——— "————————————————————————
MM OO a M ELEM TT а САО a Са و ы
Volume 84, Number 3
1997
Croat 325
Philodendron Subgenus Philodendron
podial growth and P sect. Fruticosa Grayum with
proleptic? sympodial growth (Ray, 1987b), a growth
form that is rare in the family, known only in Alo-
casia and а few species of Monstera (Grayum,
1996).
Philodendron subg. Philodendron is difficult to
define and is primarily distinguished by its nega-
tives, i.e., it lacks the specific characteristics of P.
subgenera Pteromischum and Meconostigma (see
key to subgenera below). There are relatively few
members of Р subg. Philodendron with а pachy-
caulous habit common to so many members of P.
subg. Meconostigma, i.e., with very stout, generally
erect stems and possessing conspicuous leaf scars.
Philodendron subg. Philodendron also lacks the
conspicuous, more or less triangular scales borne
in the leaf axils of P. subg. Meconostigma. Though
sometimes obvious (Fig. 11) in P. subg. Philoden-
dron, they are usually small and inconspicuous and
fall early. The species most similar to P. subg. Me-
conostigma is P. warszewiczii, but another species,
P. basii, is similar in being large with a thick, erect
stem.
Species of P. subg. Pteromischum have subtle
characteristics that to the expert permit immediate
recognition. These characters include the slender,
somewhat woody, brittle stem, a conspicuous peti-
ole sheath, thinner blades with rather pronounced
primary lateral veins, the presence of interprimary
veins, and the frequent presence of raphide cells
or stitch-like markings. Another feature that is of-
ten useful in separating P. subg. Pteromischum from
P. subg. Philodendron is the much higher incidence
of asymmetrical leaf blades in the former. Species
with asymmetrical — 2 at the base) blades
are not common in P. subg. Philodendron. Some
species of P. subg. Philodendron may have oblong
to elliptic, ans cie blades like those of P. subg.
Pteromischum, but they never have the fully
sheathed petioles of the latter (except in juvenile
condition, which may confuse the non-expert).
e three subgenera of Philodendron in general
can be most easily separated by the characters pre-
sented in the following key (modified after Mayo,
1991).
5 Proleptic sympodial growth, characteristic of members
of Philodendron sect. Fruiticosa (Grayum, 1996), results
relative
nflorescence
not axillary as in sylleptic sympodial pase The first few
internodes in proleptic growth are very short, lack buds,
and a tive series of Ils
leaves (see fig. 2 in Ray, 1987b).
KEY TO THE SUBGENERA OF PHILODENDRON
la. Stem of mature flowering plants with a succes-
subg. Pteromischum
lb. Stem of mature flowering plants w ith a succes-
th bearing а
the leaf axils; petioles of adult plants with short,
usually inconspicuous petiole sheath and borne
on the side of the stem, not encircling it at the
2a. Stems often arborescent; staminodial zone
tween
tere disini or КЕ than fertile zone;
stamens usually at least 3 times gr than
d P. subg. Meconostigma
Stems rarely “arborescent, her. scandent;
staminodial zone between staminate and pis-
tillate zones of spadix much shorter than the
fertile staminate zone; stamens less than 3
times longer than broad ..
t
4
P subg. Philodendron
There are also a number of anatomical charac-
teristics separating the subgenera. Vegetative buds
of Philodendron subg. Philodendron are always lo-
cated below the point of overlap in the sheath mar-
gins of the cataphyll, whereas they are lacking in
P. subg. Pteromischum (Ray, 1987b). Philodendron
subg. Pteromischum is distinct in having a style
with a shallow compitum* with a subepidermal con-
centration of raphide crystals (Mayo, 1986, 1989)
and a total lack of tannin cells in the stamens
(Mayo, 1986). In addition, while hypophyllous’
stem segments are typical for P. subg. Philodendron
they are ambiphyllous,* hyperphyllous,’ or pera-
phyllous'? in P. subg. Pteromischum. Philodendron
subg. Philodendron is characterized by having con-
tinuous parenchyma from the cortex to the center
of the stem. In contrast, P. subg. Pteromischum has
a central cylinder with a solid ring of fibers around
LN
-
While not definitive, there are a number of other
features that normally are useful to separate P.
* The common depression that leads to individual stylar
ls.
' Hypophyllous stem segments are those those in which
the petiole scar borders the lower == of the segment (see
figs. 3, 7 in Ray, 1987b).
е сетан к мет аон ит so short that the pet-
iole scars 4 s of the stem segment (see
-— 9-11 in Ray, "19870.
yllous stem ents have the petiole scar at
the xia edge of the stem segment (see (see figs. 2, 8 in Ray.
е Peraphyllous stem scars are those in which the inter-
node subtending the petiole “is elongated and supercedes
the point of attachment" (Ray, 1987b).
326
Annals of the
Missouri Botanical Garden
subg. Pteromischum. Its blades are typically more
or less oblong, moderately thin, and typically more
inequilateral than those of P. subg. Philodendron.
The stems of P. subg. Pteromischum are commonly
less than 1 cm in diameter and frequently with
rather long internodes. While there are many mem-
bers of P. subg. Philodendron with more or less
oblong leaf blades, blades of most species are cor-
date or subcordate at the base. Many members of
P. subg. Pteromischum have stems that branch and
spread away from their support before flowering;
this behavior is rare in P. subg. Philodendron.
Because of the usually conspicuously sheathed
petioles of P. subg. Pteromischum, the subgenus is
more likely to be confused with sterile specimens
of Rhodospatha than with the oblong-bladed spe-
cies of Р subg. Philodendron (and in such cases
the presence of trichosclereids beneath the epi-
dermis of Rhodospatha easily distinguishes it from
Philodendron).
RELATIONSHIPS WITHIN P. SUBG. PHILODENDRON
Discussion of subgeneric classification. Any at-
tempt to revise the subgeneric classification of
Philodendron subg. Philodendron is frustrated by
the lack of morphological characteristics that
correlate with one another throughout the sub-
genus. Engler (1899) separated the species of P.
subg. Philodendron primarily on the basis of leaf
shape, leaf blade venation, and the nature of the
pistil (i.e., number of locules per ovary, type of
placentation, and number of ovules per locule).
Most sections, all moderately small ones, were
separated on the basis of leaf morphology. These
are: P. sect. Tritomophyllum, P. sect. Schizophyl-
lum, and P. sect. Polytomium. Two sections, P.
sect. Macrogynium and P. sect. Camptogynium,
are based on the nature of the pistil. Following a
cladistic analysis of a relatively small number of
species, including some in P. subg. Philoden-
dron, Mayo (1986) concluded that Philodendron
should be divided into two to three sections in-
stead of the existing nine sections in the genus.
While I agree that P. sect. Macrolonchium should
be reduced, I think that a cladistic analysis mak-
ing use of the leaves as well would justify the
existence of the remaining sections recognized by
Krause with the possible exception of P. sect.
Camptogynium, which was not studied by Mayo.
The most time-honored way to separate species
in P. subg. Philodendron is based on number of
ovules per locule, a system first devised by Engler
stemming from his first revision of Philodendron
(Engler, 1878). Engler used the number of ovules
per locule to separate two large groups, which he
called P. sect. Polyspermium and P. sect. Oligosper-
mium (now P. sect. Philodendron and P. sect. Ca-
lostigma, respectively). These two sections together
comprise the largest percentage of species in the
subgenus. As they are constituted they are very di-
verse morphologically, and it is possible that the
number of ovules per locule will not prove to be
reliable for separation at the sectional level. It is
possible that species with relatively few ovules or
solitary ovules may have evolved independently
several times from ancestors with numerous ovules
having axile placentation. Since it has not yet been
determined if this is the case, the classification sys-
tem used here will in general be conservative.
Krause (1913) closely followed Engler' sectional
revision. His P. sect. Philodendron (as P. sect. Poly-
spermium) consisted of species with axile placen-
tation and “many” ovules per locule, while P. sect.
Calostigma (as P. sect. Oligospermium) consisted of
those species with sub-basal (or less frequently ba-
sal) placentation with “1 or few" ovules per locule.
Since the time of the last revision of Philodendron
many species have been added, and the distribu-
tion of ovules per locule for all species now forms
a more complete continuum. There is still a signif-
icant correlation between axile placentation and
moderately large numbers of ovules per locule and
the converse, basal and sub-basal placentation and
relatively low numbers of ovules. Both P. sect.
Philodendron and P. sect. Calostigma will be dis-
cussed below.
While these two groups, P. sect. Philodendron
and P. sect. Calostigma, constitute the largest per-
centage of Central American species, several other
sections are separated on the basis of leaf shape.
venation, and style type (see above key). Each of
these will be discussed in turn, despite the fact that
some do not occur in Central America. Krause
(1913) treated ten sections in his revision of Philo-
dendron. As previously discussed, P. sect. Pter-
omischum has been elevated to the status of sub-
genus, and P. subg. Macrolonchium Engl. has been
reduced to a subsection of P. subg. Philodendron.
1. Philodendron sect. Baursia (Rchb. ex Schott)
Engl, in Mart, Fl. bras. 3(2): 134. 1878.
Philodendron grex Baursia Rchb. ex Schott,
Syn. Aroid. 73. 1856. TYPE: Philodendron
crassinervium Lindl. (lectotype, designated by
Mayo, 1990
As defined by Engler and Krause, P sect. Baur-
sia consists of species with generally incon
primary lateral veins, but the group as constitute
Моште 84, Митбег 3
1997
327
Philodendron Subgenus Philodendron
by Krause remains highly variable in terms of its
ovules, habit, and leaf shape. The group comprises
species with moderately many ovules and axile pla-
centation, moderately many ovules with basal pla-
centation, a few ovules with basal placentation, and
solitary ovules with basal placentation. All of these
species have leaves purportedly devoid of primary
lateral veins. In reality this is not true of all species
included in the group. Most species have elongate,
simple blades, but three have 3-lobed or tripartite
blades.
With 33 species included by Krause the section
was third in size of the three major sections (P. sect.
Philodendron with 64 and P. sect. Calostigma with
53 species). One species, P. acreanum K. Krause,
is actually a member of P. sect. Pteromischum. Of
the remaining species in the section, those that best
fit the description of the group occur principally in
eastern South America and in the upper Amazon
basin and have more or less oblong leaf blades.
Philodendron crassinervium is the type of the sec-
tion. Except for P. crassinervium, P. longilaminatum
Schott (with axile placentation), P. bahiense Engl.,
and P. paxianum K. Krause (each with solitary
ovules per locule), species of P. sect. Baursia have
a few basal ovules per locule and oblong to oblong-
elliptic blades. Many but not all have indistinct pri-
mary lateral veins.
Some members of Krause's P. sect. Baursia, es-
pecially the species that are vines with a solitary
ovule per locule and occur in the Andes west of
the Continental Divide, such as P. lehmannii Engl.,
P. ellipticum Engl, P. pachycaule K. Krause, P.
chimboanum Engl., P. longipes Engl., and P. grav-
eolens Engl., do not seem to belong with the re-
mainder and should perhaps be put into another
section. The same is true of the three-lob
tripartite species, P. deltoideum Poepp. & Endl., Р.
panduriforme (Kunth) Kunth (Krause also included
here P. reichenbachianum Schott, now a variety of
P. panduriforme), and P. micranthum Poepp. ex
Schott. With the exception of P. micranthum, which
has primary lateral veins lacking or weak, these
species have primary lateral veins at least some of
ЊУ time (though they are indistinct in P. delto-
deum). None of the three species appear to have
any other features in common with the more typical
members of P. sect. Baursia, e.g., P. crassinervium
Lindl., P. linnaei Kunth, and P. callosum K. Krause,
among others.
Mayo (1986) believed that P. sect. Baursia
contained two groups of species, and he would
also separate P. deltoideum and related species
from the remainder, suggesting that Schott's grex
Oligophlebium be recognized to accommodate
these species.
Some species that were placed in P. sect. Baursia
will have to be reinvestigated to determine if they
belong instead in P. sect. Philopsammos G. S. Bun-
ting (1986). That group is often similar in having
elongated leaf blades, but it differs in having bi-
locular ovaries whereas those of P. sect. Baursia
are plurilocular.
By no means all of the species with more or less
oblong blades in P. sect. Baursia have primary lat-
eral veins weak or lacking. At least one species, P.
wendlandii, the only Central American species
placed in P. sect. Baursia by Engler, should be
placed in P. sect. Calostigma. It has distinct pri-
mary lateral veins and a spongiose petiole with a
distinct dark green annular ring like the other
members of P. subsect. Glossophyllum in Central
America. This leaves Central America without
members of P. sect. Baursia.
2. Philodendron sect. Philopsammos G. S.
Bunting, Phytologia 60: 306. 1986. TYPE:
Philodendron ptarianum Steyerm., Fieldiana,
Bot. 28: 99. 1956. [Philodendron callosum K.
Krause subsp. ptarianum (Steyerm.) G. S.
Bunting, Phytologia 64: 467. 1988.]
Philodendron sect. Philopsammos is restricted to
South America, known largely from the region of
the Guiana highlands with extensions into the Am-
azon basin, occurring principally on white sand sa-
vannas, sandstone outcrops, and on tepuis, rarely
in alluvium in lowland forests. It is characterized
by having usually terrestrial or epipetric, thick,
creeping stems with mostly short internodes, mostly
long persistent, mostly intact cataphylls, moderate-
ly long petioles, mostly erect, geniculate petioles,
more or less oblong, elliptic to narrowly ovate, co-
riaceous blades, usually lacking any prominent
posterior lobes and with usually distinct, sometimes
moderately obscure primary lateral veins. Inflores-
cences are moderately large with pistils bilocular,
rarely 3-locular, and ovaries moderately numerous
with axile placentation.
Included in the section are the following species:
P. canaimae G. S. Bunting, P. craspedodromum R.
E. Schult., Р. dunstervilleorum С. S. Bunting, P.
dyscarpium R. E. Schult., P. peraiense G. S. Bun-
ting, P. phlebodes G. S. Bunting, P. pimichinense G.
S. Bunting, P. callosum K. Krause, P. pulchrum G.
M. Barroso, P. remifolium R. E. Schult., P. sabu-
losum G. S. Bunting, P. steyermarkii G. S. Bunting,
and P. tatei K. Krause. Perhaps also to be included
328
Annals of the
Missouri Botanical Garden
in this group is P. englerianum Steyerm. No species
in the group occur in Central America.
In describing this section Bunting (1986) made
no mention of how the section is distinguished
from P. sect. Baursia (or any other section). This
is a critical point since both sections have spe-
cies with elongated blades and at least sometimes
have primary lateral veins not markedly more
prominent than the interprimary veins (secondary
lateral veins). The section is presumably distin-
guished from P. sect. Baursia on the basis of hav-
ing mostly two locules per ovary. As an indication
of its affiliation with P. sect. Baursia, Bunting, at
the time he described P. sect. Philopsammos,
specifically mentioned P. callosum (a species in-
cluded by Krause in P. sect. Baursia and initially
considered by Bunting to be distinct from P. ptar-
ianum) as being a possible member. Philoden-
dron ptarianum has proven to be closely related
to P. callosum, but that species was described by
Krause as being “plurilocula,” i.e., with many
locules per ovary and “pauciovulata,” i.e., with
few ovules per locule. If this is true, the single
character separating P. sect. Philopsammos from
P. sect. Baursia, namely the small number of loc-
ules per ovary, would break down even in two
subspecies (as now recognized by Bunting,
1995), one of which is the type of P. sect. Phil-
opsammos. One collection of P. callosum, Davidse
& Miller 27269, had 2-locular ovaries with 8—10
ovules per locule with unusual black, shiny
seeds. It would clearly appear to be a member of
P. sect. Philopsammos.
3. Philodendron sect. Philodendron. TYPE: P
grandifolium (Jacq.) Schott
Philodendron sect. Philodendron in Central
America is both large and diverse, and like P sect.
Calostigma (a discussion of which follows) it is fur-
ther subdivided here into subsections.
With 38 species (40 taxa) P. sect. Philodendron
is the second largest section in P. subg. Philoden-
dron in Central America. Philodendron sect. Philo-
dendron is characterized by having axile placenta-
tion and typically many ovules per locule but,
owing to its size and diversity, there are no other
characters that completely characterize the group.
Philodendron sect. Polyspermium (according to the
Code (Art. 21) it must now be P. sect. Philoden-
dron) was subdivided by Krause into six groups,
"Gruppen" (termed subsections by Mayo, 1990).
These subsections will be discussed here, especial-
ly in relation to the Central American species.
SUBSECTIONS OF P. SECT. PHILODENDRON
1. Philodendron subsect. Macrolonchium
(Schott) Engl., in Mart., Fl. bras. 3: 139. 1878.
Philodendron grex Macrolonchium Schott,
Prodr. syst. Aroid. 269. 1860. TYPE: Philo-
dendron simsii (Hook.) G. Don (lectotype, des-
ignated by Mayo, 1990: 64).
This is a small group of species characterized by
D-shaped or broadly and sharply sulcate petioles
and the presence of short stems with short inter-
nodes. The leaf blades typically are ovate-triangu-
lar. The cataphylls typically persist as fibers. The
ovaries are 5—10-locular with numerous ovules per
locule. This group was treated by Engler (1899) and
Krause (1913) as a section based on stems with
internodes shorter than broad versus scandent
stems, but the group is in no way warranted at the
sectional level based on this or any other character.
Though neither short stems nor D-shaped peti-
oles are unique to this group, it appears to be nat-
ural at the subsectional level. All of the species
have similar ovate-triangular blades and coarse
reddish brown persistent cataphyll fibers. The two
pinnately lobed species in the group, P. pinnatifi-
dum (Jacq.) Schott and P. robustum Schott, seem
unlikely members of this subsection. Those species
are very similar to P. fendleri K. Krause, which En-
gler placed in his P. sect. Polytomium. Philoden-
dron pinnatifidum and P. robustum should probably
be placed there as well. They seem to have little
in common with the other simple-leaved species.
Philodendron melinonii Brongn., Р. fragrantissi-
mum, P. simsii, and P. roraimae K. Krause all ap-
pear to be related. The only Central American spe-
cies in this subsection is P. fragrantissimum.
2. Philodendron subsect. Canniphyllum
(Schott) Mayo, J. Linn. Soc., Bot. 100. 168.
1989. Philodendron grex Canniphyllum
Schott, Syn. Aroid. 76. 1856. TYPE: Philo-
dendron fibrillosum Poepp.
This is a small group that included only five spe-
cies in Krause's revision and one of these, P. —
rulescens, proved to be a synonym of P. inaequila-
. Pteromischum.
no doubt distantly related P. blanchetianum Schott
is a species from eastern Brazil in Bahia. Philo-
dendron subsect. Canniphyllum has only a few spe-
cies in Central America. Both Philodendron creto-
sum and P. roseospathum have persistent fibrous
cataphylls and resemble the type species. P. fibril-
Volume 84, Number 3
1997
Croat 329
Philodendron Subgenus Philodendron
losum (selected by Mayo, 1990). The only other
Central American species that has characteristics
to fit in the subsection is P. chirripoense, which is
tentatively placed here, though it differs in having
longer, more slender internodes and lacks persis-
tent cataphylls. The presence or absence of persis-
tent cataphylls is highly correlated with the length
of internodes (persistent when internodes are short,
deciduous when internodes are long) and thus it is
not surprising that P. chirripoense would lack cat-
aphylls. Still, the latter is a most peculiar species
so its placement remains in doubt.
3. Philodendron ѕиһѕесі. Platypodium
(Schott) Engl., in Mart., Fl. bras. 3: 137. 1878.
Philodendron grex Platypodium Schott, Syn.
Aroid. 85. 1856. TYPE: Philodendron ptero-
tum K. Koch & Augustin (lectotype, designat-
ed by Mayo, 1990: 61).
This group was characterized by Schott (1856)
as having D-shaped petioles. While he included
both P. pterotum and P. fragrantissimum, Engler
later transferred the latter to his P. sect. Macrolon-
chium (Schott) Engl. Mayo (1990) designated the
only remaining species in Schott's section, namely
P. pterotum, as the type species.
In addition to P. pterotum, Krause (1913) placed
in P. sect. Platypodium four more species, P. cle-
mentis Wright, now considered a synonym of P. fra-
grantissimum (a member of P. subsect. Macrolon-
chium); P. splitgerberi Schott, a possible synonym
of P. fragrantissimum; P. lechlerianum Schott, a
close relative of P. purpureoviride (a member of P.
subsect. Solenosterigma); and P. ernestii Engl.
Thus, of the species listed by Krause, probably only
P. pterotum and P. ernestii belong here. It is un-
likely that P. ernestii is very closely related to P.
pterotum, but it does appear to be closely related
to P. brunneicaule, another species placed in this
section.
In addition to P. pterotum, other species of P.
subsect. Platypodium that occur in Central Amer-
ica are: P. brunneicaule, P. copense, P. findens, and
P. fortunense Croat.
4. Philodendron subsect. Psoropodium
(Schott) Engl., in Mart., Fl. bras. 3: 138. 1878.
Philodendron grex Psoropodium Schott, Syn.
Aroid. 84. 1856. TYPE: Philodendron orna-
tum Schott (lectotype, designated by Mayo,
1990: 61).
As defined by Schott (1856), this is a group that
has petioles verrucose at the apex. Mayo (1991)
appropriately selected P. ornatum as the lectotype.
Krause's description, *petioles semirounded above,
species in the genus Philodendron, but Schott's in-
tention for the subsection was clear. The only other
species included by Schott was P. rubens Schott,
now considered by some authors to be a synonym
of P. ornatum. Krause also expanded the group sub-
stantially, adding a number of plants that do not
have glandular petioles and others that have proven
to be synonyms of P. ornatum. The latter are: P.
muschlerianum K. Krause, P. dolosum Schott, P. as-
peratum K. Koch, and P. tobagoense Engl. The ab-
errant elements which, I believe, are unrelated to
P. ornatum include P. brevilaminatum Schott and
P. traunii Engl. (now both synonyms of P. fragran-
tissimum in P. subsect. Macrolonchium), P. gran-
dipes, and P. maximum K. Krause, a gigantic spe-
cies from the southwest Amazon basin that is
unlikely to be related in any way to P. ornatum. 1
have moved P. grandipes from P. subsect. Psoro-
podium to P. subsect. Philodendron, where it more
appropriately belongs.
Krause also included in his Psoropodium group
three other poorly known species, P. thaliifolium
Schott, P. brandtianum K. Krause, and P. bertae K.
Krause. These do not appear to be in any way re-
lated to P. ornatum. It is clear from these as well
as the other species included by Krause that the
section as defined by Engler and by Krause no lon-
ger resembles Schott's original description. Proba-
bly a number of the species will need to be moved
into other groups, or the section will need to be
more well defined. As currently defined no species
in P. subsect. Psoropodium occur in Central Amer-
ica.
5. Philodendron subsect. Solenosterigma
(Klotzsch ex Schott) Engl., in Mart., Fl. bras.
3: 139. 1878. Philodendron grex Solenoster-
igma Klotzsch, Syn. Aroid. 81. 1856. TYPE:
Philodendron scandens K. Koch & Sello [(—
Philodendron hederaceum (Jacq.) Schott (lec-
totype, designated by Mayo, 1990: 61)].
As treated by Krause (1913), this appears to be
a natural group, consisting of P. hederaceum, the
type, as well as P. consanguineum Schott and a
number of relatives. Philodendron fuertesii K. Krau-
se, P. krebsii Schott, and P. urbanianum K. Krause
all closely resemble P. consanguinem, and P. mar-
ginatum Urb., Р. prieurianum Schott, P. oxycar-
dium, P. micans, and P. melanochrysum Linden &
André are all synonyms or subspecies of P. heder-
aceum. These species share long, slender inter-
Annals of the
Missouri Botanical Garden
nodes, deciduous, mostly unribbed cataphylls,
more or less terete petioles, ovate-cordate leaf
blades, and solitary inflorescences.
Other species included by Krause in P. subsect.
Solenosterigma are P. purpureoviride (aside from P.
hederaceum and P. brevispathum, the only other
species of the group that occurs in Central Amer-
ica), P. jenmanii K. Krause, and P. scabrum K.
Krause. The latter two species are now considered
synonyms of P. muricatum Willd. ex Schott. Though
surely belonging in P subsect. Solenosterigma
(along with the two synonyms already placed here)
Krause placed P. muricatum in P. sect. Calostigma.
Philodendron brevispathum is transferred here
from P. subsect. Cardiobelium (Schott) Engl., since
it appears to be closely related to P. muricatum.
6. Philodendron subsect. Philodendron
Philodendron grex Cardiobelium Schott, Syn. Aroid. 88.
1856. Philodendron subsect. Cardiobelium (Schott)
Engl., in Mart., Fl. bras. 3: 139. 1878. Philodendron
“Gruppe” Cardiobelium (Schott) Engl., Bot. Jahrb.
Syst. 26: 522, 529. 1899. TYPE: Philodendron gi-
ganteum Schott (lectotype, designated by Mayo,
1990: 60).
Philodendron grex Eubelium Schott, Syn. Aroid. 92.
1856. Philodendron subsect. Eubelium (Schott)
Engl., in Mart., Fl. bras. 3: 140. 1878. TYPE: Philo-
dendron grandifolium (Jacq.) Schott.
As defined by Schott (1860), grex Cardiobelium
consisted of a single species, P. giganteum. The
group was greatly expanded by Engler (1899) and
by Krause (1913) as a subgroup within section Po-
lyspermium. Now with 20 species, it constitutes the
largest subsection in P. sect. Philodendron. The
characterization, as expanded by Krause, “petioles
smooth or lightly striate, asperate; blade cordate to
sagittate, with the primary lateral veins much more
conspicuous than the secondary veins,” is so broad
that many unrelated species might easily be con-
tained within it. Certainly to be excluded is P rub-
ens Schott (now a synonym of P. ornatum and the
core species in P. subsect. Psoropodium). Among
the well known and seemingly distinct taxa includ-
ed by Krause are Р. grandifolium, P. acutatum
Schott, and P. fraternum Schott, all with deciduous
cataphylls, and Р. tenue, P. schottianum, and P.
panamense, with persistent cataphylls. The type, P.
giganteum, also has conspicuous persistent cata-
phylls. A particularly unusual species included by
Krause is P. quitense Engl., with deeply three-lobed
leaves. It is probably a synonym of P. acuminatis-
simum Engl. in P. subsect. Doratophyllum.
The only Central American species included in
the section by Krause were: P. brevispathum, P.
panamense, P. schottianum, and P. tenue. Philoden-
dron brevispathum, with its scaly stems, is best ac-
commodated with P. muricatum in P. subsect. So-
lenosterigma.
Philodendron subsect. Philodendron has 20 spe-
cies in Central America. The size and diversity of
P. subsect. Philodendron warrant the recognition of
the five series presented below.
SERIES OF P. SUBSECTION PHILODENDRON
1. Philodendron ser. Philodendron. TYPE:
P. grandifolium (Jacq.) Schott
The series is not known from Central America,
but has several species in South America. This
group is characterized by thick stems, deciduous
cataphylls, subterete petioles, large ovate-triangular
blades conspicuously lobed at the base, several in-
floresences per axil, and 5—7-locular ovaries with
many ovules per locule. In addition to the type, P.
acutatum Schott, P. billietiae Croat, and P. megal-
ophyllum Schott appear to be typical members of
this group, having many ovules per locule, decid-
uous cataphylls, more or less terete petioles, and
ovate-triangular leaf blades.
2. Philodendron ser. Impolita Croat, ser. nov.
TYPE: P. strictum G. S. Bunting
Hala teres
ernodia brevia: catan
Ж.
lla istenti
vel D-formatus; lamina palida adaxialiter, glaucescens.
Etymology. From impolitus meaning unpol-
ished, i.e., matte, in reference to the matte and pale
lower blade surfaces.
Three Central American taxa, P. hebetatum, P.
strictum, and P. thalassicum, and at least one ad-
ditional undescribed species from South America
belong in this group. It is characterized by having
leaf blades dark to medium green above but very
pale, almost white beneath and covered with a min-
ute waxy covering making the surface matte. The
species have relatively short internodes, persistent
cataphylls that often dry yellowish or have patches
of yellowish epidermis persistent (not always true
for P. thalassicum). Petioles may be obtusely some-
what flattened to D-shaped and dry with a yellowish
epidermis (not always true for Р thalassicum).
Blades are ovate-cordate to ovate-triangular.
3. Philodendron ser. Velvetina Croat, ser
nov. TYPE: P. gigas Croat
Internodia brevia, maxime crassa; cataphylla D-formata,
persistentia in fibris tenuibus; petiolus subteres; me
ovata-cordata, bicolorata, velutina adaxiliter, impo ita
abaxialiter, 81-125 ст longa, 37-90 ст lata.
Volume 84, Number 3
1997
Croat 331
Philodendron Subgenus Philodendron
Etymology. From velutinus meaning velvet, re-
ferring to the velvety upper surfaces of the blades.
The subsection consists of a single species, P.
gigas, in Central America but would also include
P. andreanum Devansaye from northern Colombia.
The group is characterized by its stout stems, short
internodes, cataphylls that persist as thin fibers,
subterete petioles, and especially by the very large,
narrowly ovate-cordate blades, which are velvety
and bicolored on the upper surface with the midrib
and primary lateral veins paler, and paler and matte
on the lower surface. Inflorescences range up to 7
per axil.
4. Philodendron ser. Fibrosa — ser. nov.
TYPE: P. jodavisianum G. S. Bunt
rnodia brevia, cataphylla persistentia ut fibrae; pe-
E plerumque teres aut subteres; lamina ovato-cordata;
pistilla cum placentatione axiali; loculi pluriovulati.
Etymology. Fibrosa = composed of separable
fibers, in reference to the cataphylls persisting as
ers.
This represents the largest series in P. subsect.
Philodendron. It is characterized by thick stems,
short internodes, cataphylls that usually persist as
a mass of fibers on the stem, usually subterete pet-
ioles, and more or less ovate-cordate blades. Pistils
have axile placentation and many ovules per locule.
The following species of P. ser. Fibrosa occur in
Central America: Р alticola, Р antonioanum, P.
breedlovei, P. chiriquense, P. dodsonii, P. grandipes,
P. jodavisianum, P. lazorii, P. llanense, P. pana-
mense, P. pirrense, P. purulhense, P. scalarinerve, P.
schottianum, and P. tenue. Of these a few are still
doubtful. Philodendron grandipes, with a D-shaped
petiole, and P. jodavisianum, with a U-shaped pet-
iole, perhaps belong in P. subsect. Platypodium but
also do not seem to be related to the species in that
group. Philodendron breedlovei, which appears to
lack persistent cataphylls (specimen very incom-
plete), is at odds with the other species but fits
nowhere else. A few species are in related clusters
within the series; for example, P. lazorii and P. pan-
amense seem to be closely related as do P. gran-
dipes and P. jodavisianum.
Grayum (pers. comm.) believes that P. dodsonii
perhaps belongs with P. pterotum in P. subsect. Pla-
typodium.
5. Philodendron ser. Albisuecosa Croat, ser.
nov. TYPE: P. albisuccus Croat
Internodia brevia; succus albus, calcareus; cataphylla
persistentia in fibris tenuibus pallidis; petiolus subteres;
lamina ovato-cordata; inflorescentia solitaria; pistilla 5—6-
locularia; loculi cum 18-20 seminibus.
Etymology. Albus = white; succus = juice,
sap, in reference to the white sap of cut parts.
The series consists of a single species, P. albi-
succus, characterized by having copious white sap,
which turns chalky on drying. Only one other spe-
cies in Central America, P. cretosum, shares this
unusual feature. It may belong in the same series
despite its linear to oblanceolate leaf blades. It cur-
rently is placed in P. subsect. Canniphyllum.
7. Philodendron subsect. Achyropodium
(Schott) Engl., in Mart., Fl. bras. 3: 139. 1878
P. grex Achyropodium Schott, Syn. Aroid. 85.
1856. TYPE: P verrucosum L. Mathieu ex
Schott
This is a seemingly quite natural group charac-
terized by scaly petioles. As defined by Schott
(1856) it was represented only by the type, P. ver-
rucosum. Krause (1913) included six species (one
of which, Р. arcuatum К. Krause, is a synonym of
P. brevispathum in P. subsect. Solenosterigma). The
others in the subsection are: P. nanegalense Engl.,
P. pilatonense Engl., and P. gualeanum Engl. (all
of which, I believe, represent a single species), as
well as P. serpens Hook. f. All but P. verrucosum
were known only from South America. There are
several other undescribed species in P. subsect.
Achyropodium now known from South America.
Philodendron subsect. Achyropodium is largely
restricted to the northern Andes and lower Central
America (Costa Rica and Panama) but has one spe-
cies, P. verrucosum, that ranges as far south as Peru.
The subsection is represented in Central America
by P. glanduliferum subsp. glanduliferum, P. ham-
melii, P. malesevichiae, P. squamicaule, P. squami-
petiolatum, and P. verrucosum.
4. Philodendron sect. Calostigma (Schott) Pfeif-
fer, Nomencl. Bot. 2: 674. 1874. P. [rankless]
b. Calostigma Schott, in Schott & Endl., Me-
let. Bot. 19. 1832. TYPE: P. imbe Schott
Philodendron sect. Calostigma is the largest sec-
tion in Central America with 48 species comprising
52 taxa. It is characterized by having basal or sub-
basal placentation and typically solitary or few
ovules per locule. Owing to its size and diversity,
there are no other characters that completely char-
acterize the group.
There are some differences between Р. sect.
Philodendron and P. sect. Calostigma in Central
America that are statistically significant, even if not
without exception. For example, 8196 of those spe-
332 Annals of the
Missouri Botanical Garden
Table 1. Leaf blade breakdown by section. # = number of species.
Blades entire
With basal lobes Lacking basal lobes Blades not entire
Cordate Subcordate Ovate Oblong Incised-lobate
% of % of % of % of 96 of Section
Section # sect. # sect. # sect. # sect. = sect. total
Calostigma 28 56% 1 14% 1 2% 14 28% 0 0% 50
Macrogynium 1 100% 0 0% 0 0% 0 0% 0 0% 1
Philodendron 34 81% 3 7% 1 2% E 7% 1 2% 42
Polytomium 0 0% 0 0% 0 0% 0 0% 3 100% 3
Tritomophyllum 3 38% 0 0% 0 0% 0 0% 5 63% 8
cies in P. sect. Philodendron have cordate blades,
while only 7% have subcordate blades, and 12%
lack posterior lobes. In contrast, P. sect. Calostigma
has only 56% of its species with cordate blades,
14% have subcordate blades, and 30% lack pos-
terior lobes. See Table 1 for a complete breakdown
by section.
See other comparisons between P. sect. Calostig-
ma and P. sect. Philodendron under sections enti-
Чед “Ovary Locule Number" and *Ovules Per Loc-
ule” as well as in Appendix 2, Technical Data on
Pistils.
Philodendron sect. Calostigma was subdivided
by Krause into five groups (termed subsections by
Mayo, 1990). These subsections will be discussed
here, especially in relation to the Central American
species.
1. Philodendron subsect. Macrobelium
(Schott) Engl., in Mart., Fl. bras. 3: 143. 1878.
P. grex Macrobelium Schott, Зуп. Aroid. 96.
1856. TYPE: P. sagittifolium Liebm.
Philodendron subsect. Belocardium (Schott) Engl., in
Mart., Fl. bras. 3: 141. 1878. P. grex Belocardium
Schott, Prodr. Syst. Aroid. 255. 1860. TYPE: P. ad-
vena Schott (lectotype, designated by Mayo, 1990).
As reported by Krause, Philodendron subsect.
Macrobelium was the largest subsection in P. sect.
Calostigma, with 22 species. It was poorly defined
by Krause as consisting of species with somewhat
spongy petioles, 6—12-locular ovaries with relative-
ly few ovules per locule, and basal to sub-basal
placentation. In addition (although not stated by
Krause), the blades are all cordate, sagittate, or
hastate. Although Krause described the subsection
as having 2—5 ovules per locule, many more species
have been added to the group since the last revision
and several species that logically belong in P. sub-
sect. Macrobelium have more than 10 ovules per
locule, despite having sub-basal placentation.
Philodendron subsect. Belocardium must be syn-
onymized with P. subsect. Macrobelium since P. ad-
vena, clearly a close relative of P. sagittifolium, was
designated the lectotype of the former subsection,
and Schott's definition of grex Macrobelium differs
little from grex Belocardium, primarily since the
blades are described as elongate-sagittate with 6–
7 primary lateral veins instead of ovate-cordate
blades with 3—4 primary lateral veins for grex Be-
locardium. As defined originally by Schott (1860),
P. grex Belocardium comprised vining plants with
ovate-cordate blades with the posterior rib mostly
lacking and not at all naked along the sinus and
bearing solitary inflorescences with 1-2 ovules per
locule. The group consisted of P. subovatum Schott
(= P. advena), P. hoffmannii (= P. hederaceum), Р.
deviatum Schott (= P. jacquinii), P. advena, P. po-
pulneum K. Koch, P. erubescens Linden, P. jacqui-
nii, Р. lindenii Schott, P. acrocardium Schott, and
P. consanguineum Schott. Several of these species
are now known to be synonyms of P. sect. Oligo-
carpidium or P. sect. Macrogynium. Philodendron
lindenii would have better served as a lectotype for
the group and, along with P. erubescens, P. devia-
tum, and P. populneum, might have to be formed
into a group distinct from P. sect. Belocardium.
Engler (1913) synonymized Schott's grex Glos-
sophyllum with P. “Gruppe” Belocardium, but the
former group, as defined by Schott, appears to be
distinct and will be resurrected in this work. It
comprises more or less scandent plants with tumid
petioles and more or less oblong blades that are
frequently subcordate or cordulate at the base (see
that section for more details).
Because of its size (28 species) and the diverse
composition of the species comprising P subsect.
Macrobelium, it was deemed necessary to divide the
subsection into four series. A discussion of the new
series of P. subsect. Macrobelium follows.
Моште 84, Митрег 3
1997
Croat 333
Philodendron Subgenus Philodendron
1. Philodendron ser. Macrobelium (Schott)
Croat, ser. nov. TYPE: P. sagittifolium Liebm.
Philodendron ser. Macrobelium is the largest
group of species in P. subsect. Macrobelium and
includes the type. Philodendron ser. Macrobelium
is distinguished by having moderately coriaceous,
cordate to sagittate blades with the basal posterior
rib usually not naked or only weakly naked near
its base. In addition, cataphylls are usually decid-
uous or only briefly persistent. Most of the species
in the series have a strong resemblance to the wide-
spread P. sagittifolium. All members have type D
styles (see section on style type), relatively few
ovules per locule (usually 1-5, rarely to 6, 7, or 8).
In most cases the ovules are contained within a
translucent or transparent ovule sac.
Central American representatives of P. ser. Ma-
crobelium are: P. advena, P. annulatum, P. aroma-
ticum, P. coloradense, P. dwyeri, P. edenudatum, P.
ferrugineum, P. grayumii, P. knappiae, P. mexican-
um, P. PTE P. sagittifolium, P. subincis-
> verapazense, and P. zhuanum.
E platypetiolatum is unusual in having
a much-flattened petiole. Perhaps it warrants rec-
ognition as a separate series.
Philodendron mexicanum, long considered a
member of the group, is unusual in not greatly re-
sembling P. sagittifolium v in having blades
sometimes more or less hastat
It is possible that P пл је belongs in Р.
subsect. Glossophyllum because it sometimes has a
urple annular ring at the apex of the petiole (one
of the features characterizing this subsection) and
usually has only 1-2 ovules per locule (though
sometimes 4—5 ovules per locule
2. Philodendron ser. Ecordata Croat, ser.
nov. TYPE: P. brenesii Standl.
Internodia elongata vel pene ew Noha deci dua; у
tiola subteres; lamina acuta bcordata ad basim
via minores saepe “etched” in йе supra: айя ^il
14 locularia; loculi plerumque 4—14 ovulat
Philodendron ser. Ecordata represents a group of
species with elongate stems, internodes often longer
than broad, deciduous cataphylls, elongate petioles,
and blades that are ovate to ovate-elliptic, acute,
or at most subcordate at the base (hence the name
“ecordata,” meaning without a cordate blade), often
with the minor veins on the upper surface weakly
etched. The basal veins are either free to the base
or if they are united into a posterior rib, the latter
is not naked or is naked for only a short distance.
Pistils are 5—14-locular and locules are 4—14-ovu-
late
The Central American species of P. ser. Ecordata
are: P. brenesii, P. crassispathum, P. davidsonii, P.
Іепій, and P. niqueanum. A relationship between P.
brenesii and P. ser. Impolita is possible, based on
the glaucous lower blade surface in P. brenesii, but
no other member of the group has glaucous leaves.
Moreover, all the members of the above group usu-
ally have internodes longer than wide and have de-
ciduous, rather than persistent cataphylls.
3. Philodendron ser. Reticulata Croat, ser.
nov. TYPE: P. tysonii Croat
Internodia brevia; cataphylla persistentia in fibris ten-
uibus; petiolus lamina subequans aut longior, teres aut
subteres; lamina ovato-cordata; smi cum loculis 5-9-
ovulatis; ovulae plerumque 5-7 per ovaria.
Etymology. Reticulus, meaning netted, refer-
ring to the dried network of cataphyll fibers char-
acterizing this series.
hilodendron ser. Reticulata is the only group of
species in P subsect. Macrobelium with a few
ovules per locule, stems with short internodes, and
persistent cataphylls. Most species with short in-
ternodes and persistent cataphyll fibers are mem-
bers of P. sect. Philodendron. Species in this group
have terete or subterete petioles about as long as
the ovate-cordate pedes and pistils with 5—9 loc-
ules, each with 5—7 ovules.
Philodendron ser. но is represented in
Central America by only two species, P. jefense and
P. tysonii.
4. Philodendron ser. Pachycaulia Croat, ser.
nov. TYPE: P. basii Matuda
audex succulentus; internodia brevia, crassa; cata-
4 cm; pi ER 4—6-locularia; loculi 4—6 ovulat
Etymology. Pachy = thick, caule = stem, re-
ferring to the thick, succulent stems.
Philodendron ser. Pachycaulia is represented by
a single species and is characterized by its very
stout succulent stems with intact persistent cata-
phylls, terete petioles, ovate-cordate blades with
the posterior rib naked along the sinus to 4 cm.
The pistil has a type D style and is 4—6-locular
with 4—6 ovules per locule. It is believed that the
succulent stems evolved to store water, allowing the
plant to survive the long dry season in western
Mexico. The series is represented only by P. basii
from western Mexico.
Annals of the
Missouri Botanical Garden
2. Philodendron subsect. Glossophyllum
(Schott) Croat, comb. nov. Basionym: Philo-
dendron grex Glossophyllum Schott, Syn. Ar-
oid. 80. 1856. TYPE: P elaphoglossoides
Schott (lectotype, designated by Mayo, 1990).
1. Philodendron ser. Glossophyllum Croat,
ser. nov. TYPE: P. elaphoglossoides Schott
Philodendron subsect. Glossophyllum consists of
two new series, Glossophyllum and Ovata.
Philodendron ser. Glossophyllum, as defined
here, has the appearance of being a natural group
of species with more or less oblong leaves acute or
frequently cordulate or subcordate at the base. The
primary lateral veins are usually distinct. Stems are
typically somewhat scandent, though some mem-
bers of the group, such as P. auriculatum, P. bakeri,
P. dolichophyllum, P. ligulatum, P. morii, P. pseu-
dauriculatum, P. utleyanum, and P. wendlandii,
sometimes have the internodes scarcely longer than
broad. The petioles are usually spongy or subspon-
gy. usually subterete, and often bear a purplish or
greenish annulus around the circumference where
the petiole joins the blade; the cataphylls may be
unribbed or sharply 1-2-ribbed and are typically
deciduous (though persisting for a time in some
species with short internodes, e.g., P. auriculatum
and P. wendlandii). The style type is variable in the
group, with most having B or D type styles but with
one species, P. granulare, having an unusual type
E style. Many species in the series have orange
berries. Philodendron ser. Glossophyllum ranges
from Nicaragua to Colombia and Ecuador on the
Pacific slope and to the Guianas and the Amazon
basin.
Krause included this group of plants with his
section Belocardium, comprising both Schott's grex
Belocardium and grex Glossophyllum. The former
group consisted of plants with elongate internodes
and ovate-cordate to sagittate blades, and Krause
included P. subovatum Schott (— P. advena), P. lin-
denii Schott, P. weberbaueri Engl., P. smithii, P. sub-
hastatum Engl. & K. Krause, P. myrmecophyllum
Engl., P. pachyphyllum K. Krause, P. advena, and
P. viride Engl. The latter group included species
with mostly oblong blades. He characterized his
section Belocardium as having tumid petioles and
unilocular ovaries. Most of the species in the cor-
date-bladed group are not believed to be closely
related to P. ser. Glossophyllum and have been re-
ferred here to P. subsect. Macrobelium (see discus-
sion of P. subsect. Macrobelium). What remains is
a group that usually has more or less oblong blades
with tumid petioles that are purple-ringed at the
apex and ovaries usually unilocular.
Philodendron smithii the only other Central
American species among those mentioned above,
does indeed have tumid petioles and only a single
ovule per locule like most members of P. subsect.
Glossophyllum, but the great difference in leaf
shape in this species and others placed here war-
rant their separation into another series within P.
subsect. Belocardium (see below).
hough Krause's revision characterized P. sub-
sect. Belocardium as having a single ovule per loc-
ule, that is not in itself the defining feature of the
subsection. For example, several species typical of
the group have oblong leaves, purple-ringed peti-
oles, and the same general appearance but have
more than one ovule per locule. These are: P. au-
riculatum, with (3)4 ovules per locule; P. ligulatum
raclioanum, and P. ligulatum var. ovatum,
both with 2; P. pseudauriculatum, with 1–2(4); and
P. wendlandii, with 2. Philodendron bakeri some-
times has 2 ovules per locule. Other Central Amer-
ican species, each with 1 ovule per locule are Р
brewsterense, P. correae, P. dolichophyllum, P. fol-
somii, P. granulare, P. immixtum, P. ligulatum, P.
morii, P. ubigantupense, and P. utleyanum.
Typical South American species in P. subsect.
Glossophyllum presented by Krause (1913) are: P
cyrtocleum Diels (— P. ruizii Schott), P. longipetiol-
atum Engl., P. heterophyllum Poepp., P. uleanum
Engl., P. adhatodifolium Schott, P. elaphoglossoides
Schott, P. ruizii Schott (erroneously placed in sect.
Baursia by Schott), P. wittianum Engl., and P. an-
gustialatum Engler. Philodendron tenuipes Engl.,
placed in P. subsect. Belocardium by Engler, ap-
pears to be closely related to P. fibrillosum Poepp.
and probably belongs in P. subsect. Canniphyllum.
Other species of P. subsect. Glossophyllum de-
scribed since the last revision by Krause are E
acutifolium K. Krause, P. buntingianum Croat, P.
liesneri Croat, and P. wurdackii G. S. Bunting.
Though, as defined here, P. subsect. Glossophyl-
lum ser. Glossophyllum consists only of species with
more or less oblong blades, some of the species are
somewhat anomalous. Philodendron granulare 1$
much like other species in P. subsect. Glossophyl-
lum, but it has a style that is unique in the group
(Style Type E; see discussion of this under that spe-
cies). Philodendron brewsterense and P. ubigantu-
pense are still poorly known but appear to belong
here. Perhaps the most doubtful is P. dolichophyl-
lum, with 3-7 ovules, which is certainly high for
this group; however, it is otherwise similar in most
aspects to other members of the subsection. Philo-
dendron wendlandii, placed by Krause in P. sect.
Volume 84, Number 3
1997
Croat 335
Philodendron Subgenus Philodendron
Baursia, seems to fit best in P. subsect. Glosso-
phyllum. It differs from other members of subsec-
tion Glossophyllum in having a petiole that is usu-
ally broader than thick and sharply flattened
adaxially and lacks an annular ring.
2. Philodendron ser. Ovata Croat, ser. nov.
TYPE: P. smithii Engl.
Caudex longus vel brevis; internodia plerumque longiora
quam lata; cataphylla plerumque decidua; petioli sub-
aequantes laminam, teres Ка subteres, cum аппшо дећ-
слепи apice; lamina vel к, SM
vel sagittata ad basi; pistilla cum stylo ple
B," rare C; ovaria 4—8-locularia; loculi ован i raro
–5.
Philodendron ser. Ovata is а somewhat hetero-
geneous group characterized by blades ovate to
ovate-triangular and cordate to subcordate at the
base, with terete or subterete petioles lacking an
annular ring at the apex. Locule number varies
from 4 to 8, and each usually has a single ovule.
Berry color is greenish white to white, yellowish to
lavender for most species known, but P. cotonense
and P. wilburii have orange fruits. Though only sev-
en Central American species are known, the series
undoubtedly has South American representatives.
The relationship with other members of P. sub-
what spongy petiole, there is little resemblance be-
tween this group and typical members of P. subsect.
Glossophyllum, which have elongated, rather than
more or less ovate, blades. Philodendron domini-
calense is the most doubtful member of the group;
with the strongest possible similarity to P. dodsonii
except for having a solitary ovule per locule (rather
than about 20 ovules per locule as in P. dodsonii),
it would appear that it might belong elsewhere.
Philodendron cotonense, despite having 4—5 ovules
per locule, appears also to belong to this section,
considering its marked resemblance to P. wilburii
and even P. smithii.
Central American species of P. ser. Ovata are: P.
cotonense, P. dominicalense, P. microstictum, P. smi-
thii, P. straminicaule, P. sulcicaule, and P. wilburii.
With the exception of P. smithii, which ranges from
Mexico to Nicaragua, P ser. Ovata in Central
America ranges from Nicaragua to Panama.
3. Philodendron subsect. Oligocarpidium
(Engl) Mayo, J. Linn. Soc., Bot. 100: 168.
1989. TYPE: P. multispadiceum Engl. (lecto-
type, designated by Mayo, 1990).
As defined by Krause, this was a group of four
species that differed greatly from one another.
Philodendron deviatum Schott has proven to be a
synonym of P jacquinii (which was placed by
Krause in its own P. sect. Macrogynium). Another
species that he included, P. pittieri, is just poorly
preserved material of P. hederaceum, which Krause
placed in his P. *Gruppe" Solenosterigma in P. sect.
Philodendron. This leaves only P. multispadiceum
Engl. and P. muricatum which, in my opinion, are
quite unrelated. Philodendron muricatum is the
oldest name for two other synonyms, P. jenmanii K.
Krause and P scabrum, which were placed by
Krause in P. subsect. Solenosterigma. Though I am
doubtful of the affinity between P. hederaceum and
P. muricatum, it seems best to include P. murica-
tum in subsect. Solenosterigma because P. muri-
catum is clearly a member of P. sect. Philodendron
and not P. sect. Calostigma.
This leaves only P. multispadiceum, which Mayo
(1990) wisely chose as the type of P. subsect. Oli-
gocarpidium. There are two Central American rep-
resentatives of the subsection and a number of oth-
er undescribed South American species as well.
The Central American species are P. clewellii and
P. heleniae.
The subsection is distinguished by having a rel-
atively large number of small (usually 4—10 cm
long) inflorescences per axil. The plants are vines
with long internodes and long-petiolate, narrowly
ovate to ovate-cordate blades that have the basal
veins either lacking or all free to the base and with-
out the development of a posterior rib. Both Central
American species represented have a type B style
and an ovule sac surrounding the ovules.
. Philodendron subsect. Bulaoana Mayo [as
laca t J. Linn. Soc., Bot. 100: 168.
1989. TYPE: P. bulaoanum Engl. (lectotype,
designated by Mayo, 1989).
This subsection noxa two soi treated by
Krause, P. bulaoanum and Р. acuminatissimum.
They probably saine the same species, a plant
with deeply 3-lobed leaves and persistent, reddish
brown cataphyll fibers. Currently the subsection is
known only from South America. Krause treated
this subsection as P. *Gruppe" Doratophyllum, but
that name has no priority at the subsectional level
(Mayo, 1990).
This subsection can be easily confused with and
is perhaps inseparable from P. sect. Tritomophyl-
lum. А moderately large number of South American
species, including Р. barrosoanum G. S. Bunting,
P. cataniapoense G. S. Bunting (Bunting placed it
in Р. “Gruppe” Doratophyllum), P. effusilobum
336
Annals of the
Missouri Botanical Garden
Croat, P. holtonianum Schott, P. hylaeae G. S. Bun-
ting (P. sect. Calostigma), P. levelii G. S. Bunting
(similar to P. barrosoanum), P. panduriforme Schott,
and P. victoriae G. S. Bunting might all belong here.
Philodendron subsect. Bulaoana and P. sect. Tri-
tomophyllum are separated on weak features.
Krause described his *Gruppe" Doratophyllum as
having somewhat succulent petioles longer than the
blade, a hastate blade, and ovaries that are several-
ovulate near the base of the locule, whereas he de-
scribed P. sect. Tritomophyllum as having tripartite
blades with the lateral lobes erect or spreading, and
the primary lateral veins more prominent than the
minor veins. The ovary was described as 5-11-loc-
ular with the locules 1- or more-ovulate. The Cen-
tral American species that appear to be closely re-
lated have one or two ovules per locule, whereas
those of P. subsect. Bulaoana are described as hav-
ing several ovules per locule. These distinctions are
pretty weak, and further study of the members of
both groups is necessary to decide whether the two
groups should be merged.
5. Philodendron subsect. Eucardium (Engl.)
Mayo, J. Linn. Soc., Bot. 100: 168. 1989.
Philodendron “Gruppe” Eucardium Engl., Bot.
Jahrb. Syst. 26: 535, 542. 1899. TYPE: Р. wal-
lisii Regel ex Engl.
This is a subsection of dubious status, based only
on P. wallisii. The original description characterizes
the subsection as having scarcely succulent peti-
oles that are flattened to sulcate abaxially, cordi-
form blades, and 5—6-locular ovaries with a few
sub-basal ovules per locule. Unfortunately, P.
wallisii is a poorly known taxon for which, so far
as is known, no material is extant.
Following his treatment of Eucardium, Krause
discussed several poorly known species, all of
which lacked inflorescences and for which no prop-
er sectional placement was possible. Among these
were P. andreanum, which is almost certainly re-
lated to P. gigas, a member of P. ser. Velutina. List-
ed among these dubious species are P. latilobum
Schott and P. obtusilobum Miq. The former is a syn-
onym of P. panduriforme, possibly a member of P.
subsect. Bulaoana, which it most resembles. Philo-
dendron obtusilobum is a poorly known species of
unknown origin, known only from a single leaf. Its
affinities remain unclear, but it appears similar to
P. lindenii Schott or P. rubens Schott.
Also described among this group of unassigned
species were P. gloriosum André and P. mamei An-
dré. These two species, along with P. sodiroi Hort.,
appear to be closely related and probably constitute
a new section. Philodendron pastazanum K. Krause
has similar features and probably belongs here as
well. Another species known from the Amazonian
lowlands of Peru and believed to be new is also in
this group. All the species in this putative new sec-
tion are terrestrial plants with a unique growth form
for Philodendron. All have a short, repent, creeping
stem with erect leaves clustered near the apex. The
internodes are usually much broader than long and
have cataphylls persistent, sometimes persisting
somewhat intact. The petioles are frequently winged
or undulate-winged along adaxial margins (but not
P. gloriosum) and blades are typically quite attrac-
tive, sometimes mottled with paler green, sometimes
(as in P. gloriosum) somewhat velvety. The group is
restricted to South America chiefly in the upper Am-
azon region. Studies of the ovules are necessary to
confirm these speculations, and investigations will
be carried out as material becomes available.
5. Philodendron sect. Tritomophyllum (Schott)
Engl., in Mart., Fl. bras. 3: 144. 1878. Philo-
dendron grex Tritomophyllum Schott, Syn. Ar-
oid. 107. 1856. Baursia sect. Tritomophyllum
(Schott) T. Post & Kuntze, Lex. gen. phan. 63.
1903. TYPE: P. tripartitum (Jacq.) Schott (lec-
totype, designated by Mayo, 1990: 63).
The section is distinguished by having a scan-
dent habit, three-lobed leaf blades and 1-2 ovules
per locule. The section ranges from Mexico to South
America, ranging as far as Ecuador on the Pacific
slope and to Venezuela, the Guianas, and the Am-
azon basin. It is represented in Central America by
six species: P. angustilobum, P. anisotomum, P. co-
tobrusense, P. madronense, P. rothschuhianum
(Engl. & K. Krause) Croat & Grayum, and P. tri-
partitum. In South America there are a number of
species with 3-lobed leaves, including: P barro-
soanum, P. cataniapoense, P. effusilobum, P. holton-
ianum, P. hylaeae, P. levelii, P. panduriforme, and
P. victoriae, all now tentatively placed in P. subsect.
Bulaoana, which might belong in P. sect. Tritomo-
phyllum. Certainly P. hylaeae, with 1-2 ovules per
locule and a strong similarity with P. tripartitum,
would appear to be closely related to P. sect. Tri-
tomophyllum.
Bunting (1986) placed Р. сагатароепзе їп P.
sect. Oligospermium “Gruppe” Doratophyllum (now
P. subsect. Bulaoana) but this species has a single
ovule per locule, elongate internodes, and decidu-
ous cataphylls. It may properly belong with P. sect.
Tritomophyllum. |
Some of the above-mentioned South American
species are doubtfully included here, since they
Моште 84, Митбег 3
1997
Croat
Philodendron Subgenus Philodendron
have several (more than 1 or 2) ovules per locule.
Philodendron levelii has 2—4 sub-basal ovules, and
P. barrosoanum has 4–6 ovules attached above the
base. Philodendron victoriae is apparently closely
related to P. barrosoanum so it probably has similar
ovules. It is possible that none of these species be-
long to P. sect. Tritomophyllum, but on the other
hand they do not closely match P. bulaoanum, the
type of P. subsect. Bulaoana, either.
6. Philodendron sect. Schizophyllum (Schott)
Engl., in Mart., Fl. bras. 3: 144. 1878. Philo-
dendron grex Schizophyllum Schott, Syn. Aroid.
104. 1856. TYPE: P. pedatum (Hook.) Kunth
(lectotype, designated by Mayo, 1990: 63).
The section is a small but natural group of 6–7
scandent species with irregularly pedatisect leaves,
3—4 ovules per locule, and sometimes scaly petioles,
primarily restricted to eastern South America and
the Amazon basin, but with one species, Р pedatum,
more widespread and ranging to northwestern Co-
lombia. The species may occur in Central America,
according to horticulturist John Hall of Costa Rica,
who illustrated such a plant supposedly from the Osa
Peninsula. Characteristically, no material was col-
lected and this claim cannot be refuted.
7. Philodendron sect. Polytomium (Schott)
Engl., in Mart., Fl. bras. 3: 145. 1878. Philo-
cle ge grex Polytomium Schott, Syn. Aroid.
108. 1856. TYPE: P. radiatum Schott (lecto-
type, designated by Mayo, 1990: 63
The section is a small group of 7-8 species with
pinnately or bipinnately lobed leaves from the West
Indies, Central America, and northern South Amer-
ica. Plants are vines or appressed climbers with
more or less terete petioles, generally deeply lobed,
large, moderately coriaceous blades, and flowers
with several axillary or sub-basal ovules per locule.
Only one species, P. distantilobum K. Krause, was
reported for the Amazon basin, but another species,
P. pinnatifidum, placed erroneously, I believe, in P.
sect. Macrogynium, really belongs here as well. It
is also a species occurring in the upper Amazon
basin. Two additional species, Р. angustisectum
Engl. and P. elegans K. Krause, occur in Colombia.
Philodendron fendleri occurs in Trinidad and north-
ern Venezuela, and P. lacerum (Jacq.) Schott occurs
in the Greater Antilles (Cuba, Jamaica, and His-
paniola). One poorly known species, P. houlettian-
um Engl, has been reported from French Guiana
but no material exists to confirm what it is. The
most widespread species in the section, P. radia-
tum, occurs in Central America, ranging from Mex-
ico (San Luis Potosí) to Colombia (Antioquia).
Philodendron radiatum var. pseudoradiatum is en-
demic to the State of Chiapas in Mexico. Central
American species in P. sect. Polytomium are P.
warszewiczul, ranging from Mexico to Nicaragua,
and P. dressleri, which is endemic to Mexico.
Mayo (1986), following a cladistic survey of inflo-
rescence types, concluded that P. fendleri belonged in
a group with P. melinonii and P. pedatum. Based on
overall morphology I would conclude that the three
species are not closely related. The latter is, in my
opinion, a member of a distinctive section, Р sect.
Schizophyllum. Philodendron melinonii Brongn., a dis-
tinctive species with cordate blades and placed by
Krause in P. sect. Macrolonchium, is in my opinion,
not related to either of the sections with lobed leaves.
8. Philodendron sect. Macrogynium Engl., Bot.
Jahrb. Syst. 26: 553. 1899. TYPE: Р hoff-
mannii Schott.
Philodendron sect. Macrogynium is one of two
small sections treated by Krause (1913) (the other
being P. sect. Camptogynium). Philodendron sect.
M ium consists of a single species, P. jac-
quinii (treated by Krause as P. hoffmannii). Both
sections are represented by scandent species, dis-
tinguished by having the style prolonged and much
narrower than the ovary and a single ovule per loc-
ule. Despite these similarities the two sections are
very different from one another. Philodendron jac-
quinii (P. sect. Macrogynium) has setose stems and
thin, veiny, ovate-cordate blades which may be de-
ciduous during the dry season. Its spathe is bulbous
and roomy inside, quite unlike most species. The
style, though narrowed to the apex, has a typical,
hemispheroid stigma. For differences with P. sect.
Camptogynium, see below.
Philodendron brevispathum, with its similarity to
P. jacquinii, might be considered a relative, but the
former has branched scales, not simple trichomes
on stems and petioles; a normal, sessile style; and
6–14 ovules per locule with axile placentation,
rather than 2 ovules per locule and sub-basal pla-
centation for P. jacquinii.
This section appears, at least on the surface, to
be natural. No other species known is alike either
morphologically or ecologically. Its thin, veiny
leaves are deciduous in the dry season and the
large colorful infructescences are prominently dis-
played in a mostly deciduous environment.
9. Philodendron sect. Camptogynium K. Krau
in Engl., Das Pflanzenreich IV. 23Db (Heft 60):
3, 127. 1913. TYPE: P. longistilum K. Krause
338
Annals of the
Missouri Botanical Garden
In contrast to P. jacquinii, which has setose stems
and thin cordate blades with prominent veins, P.
longistilum (P. sect. Camptogynium) has glabrous
stems and oblanceolate, subcoriaceous blades with
obscure primary lateral veins. The chief reason for
its status as the only member of a section is the
peculiar prolonged style, which is deflected to one
side and has a cupular apex. It is perhaps not as
unique as Krause assumed. Other species are now
known to have prolonged styles deflected to one
side, among them two undescribed species with
cordate blades from western Ecuador, one based on
Camp 3701 and the other on Jaramillo et al.
25449. However, neither has a cupular style apex.
These perhaps represent another section, or more
likely Krause's P. sect. Camptogynium may have to
be incorporated in P. sect. Calostigma, where it
might be easily accommodated.
KEY TO THE SECTIONS AND SUBSECTIONS OF P. sUBG. PHILODENDRON
Pistils usually 2-locular, rarely 3
T Pistils 4-10-locular; throughout the range of the g
2a. dic with primary pes 1 le
ich is transferred to P. sect. Calos
af veins. s moderately obscure; all South American (excludes P enia
ma)
-locular; eastern €— America and Amazon basin ____ P. sect. Philopsammos
of
sect. Baursia
2b. Plants with primary lateral leaf veins же much more prominent than the minor ve
РМ ы wit pay style much narrowed and much prolonged beyond the body of the ovary pai much narrower
than
4a. Style 1 turned toward apex, perpendicular to the body of the ovary; South ee зма Веј n
only
t Cano
4b. Style straight, directed in the same axis as the ovary; Central — Saec Ameri
e Macrogyiun (P. jacquinii)
3b. za with style about as broad as the ovary, scarcely prolonged a body of ovary.
. Blades 3-lobed or deeply јего lobate
6a. Bade es 3- lobed: ovules
with 3-lobed blades “о on
1-3 per осше, е) ог бал [Note: Р. subsect. Dorie a
here. ussion after
oma hl iP opere Sun (perhap
P. subsect. Mocrelslium (Schott) Engl.), P. anisotomum, P. c
that subsection.] ----------
this belongs in
rusense,
murna аи P que
6b. ке incised-lobate; ovules 2—8 per locule, Meng or a
. Blades divided along the anterior lobe i
nto two or
terminating re ix smaller lobes; ied gri шике with t
5 only
outh Americ
8, еасћ division in turn
richome-like scales
sect. t Schizophyllum
они
Tb. Hades endi: or bipinnately lobed, the iaces uniform along the anterior lobe; petio
er scaly; Central and South
American
Ps
ect. рои [P. dressleri, Р. radiatum, Р. warszewiczii]
5b. — entire, sometimes with the anterior "ose markedly concave, but the blade not markedly
lo
d or incised-lobate
n Pistils with axile placentation: ovules usually 15 or more per locule, dice i as few
10 per locule, rarely
Phila
9a. Blades Sol А pario or sagittate at base; petioles va
10a. Рен di
oles + D-s
with the lateral margins г
Па. Ste
la. Stems with ашны Дев much Spr nce than lo
Ps х заң
llb. Stems with internodes
P. subsect.
than broa
ord: cher pug [P. brunneicaule, P. copense.
10b. Petioles terete or subterete.
12a. Petioles warty at apex;
neously placed here by Krause); P. ornatum Schott and relativ
aped in cross section, sharply Anova] prac sometimes also
long or longer
ics > finden: ,
P. fortunense, P. pterotum]
South American species (excluding P. ee er-
subsect. Psorop odium ин Engl.
12b. Petioles smooth or conspicuously scaly bit: ubl merely warty al
13a. Plants scandent; internodes much longer than bens ca sal usually
deciduous; blades cordate
14b. Plants with d
Cat
15a.
Aer
P. hederaceum]
0 E Р ый. А
Р landulerum, P. hammelii, P. m
petiolatum, Р. squamicaule, P. verrucos
P
ки ot scaly ____ > subsect. Philodendron
aphylls deciduous Aus or eg o (idis ng ome
k soon deciduous ______ i r. Philodendron
Volume 84, Number 3
1997
Croat 339
Philodendron Subgenus Philodendron
9b. Blades acute to obtuse at base; petioles subterete an
P. subs
l6a. Blades matte on lower surface
17a. Blades matte, not velvety on upper surface, whit-
ish and glaucous on lower surface _______
P. ser. Impolita Croat [P. hebetatum, P. strictum,
P. thalassicum
17b. Pan velvety on piper voit not whitish and
glaucous on lowe ra
P. ser. t rta Croat [P. gigas]
16b. Blades air sentiam never dd or glaucous,
te lower su
. Sap = to pie never He МЕРА САН E
Ble TS IA r. Fibro бош ae айко,
Р. antonioa E bree 1 ê Р,
Pris E rani, HH P jim, R as
P. Џапе ee e, P. ритепзе, P. p
ens регин B pito а n оц
18Ь. Sap white it} chalky
ДО к ser. Albisuccosa Croat |Р. albisuccus]
ect. дшше a (Schott) Mayo [P. Sea P. cretosum, P. roseospathum]
8b. Pistils with basal or sub-basal placentation; ovules few per locule, usually fewer than 5, seldom
up to 8, rarely as many as 12 (but with some locules in the same inflorescence with as few as
6 ovules per locule) P. sect. Calostigma
19a, Plants usually ap iret hemiepiphytic climbers; internodes hia ea as long as wide
or longer than wide: se idespread in both Central and South A
20a. Blades ss ply 3- lobed: cataphylls persisting in a деме, reddish brown = =
тем species only; P. acuminatissimum Engl. P.
Ps ubsect. Ro] Mayo
20b. Blades not at all 3-lobed, cordate, sagittate, or + oblong; es den deciduous or
persistent, fibrous or intact; widespread in both Central and merica.
21а. n nces both small and numerous, ed more bp: "убн ахі], less than
5 em long ——— P. subsect. Oligocarpidium (Engl.) Mayo v — Р —
21b. Inforscences of normal size, жыгар л — and more tha m lon
22a [s with nip ovules per locule; petioles me Г Gers a
x; bla on, or cordate at base ........ 2 subsect. Маи
234 Cataphylls deciduo
Leaf blades with minor veins not at all etched into upper
surfaeé meri r. Macrobelium (Schott) Croat
[Р. advena, P. annulatum, р агота tiii P inc «р
P. dwyeri, P. edenudatum, P. ferrugineum, P. gi LU
m" P. mexicanum, LP plat ypetiolatum. P ieri ti-foli.
m, P. sousae, P. subincisum, P. verapazense, P. негован
24b. Leaf њи with minor veins uci into upper surfac
r. Ecordata mel IP
brenesii, di laii ade Р кым, P. lentii, P. niqueanum]
23b. Cataphylls persi
25a. жй эи oia "d gene as über, ia
ticulata Croat (P. јејепзе, P. tysonii]
25b. Cataphylls persisting ена эга су-и а succulent ___
P. se redeam Croat Аа Сей
. Pistils with ovules solitary (or sometimes 2, as in P. wilburii) in
осше; petioles frequently with purple ring at apex (Р. subsect.
ria blades elongate (P. ser. Glossophyllum) or + deae (P. > чп
~
Y)
c
. Ovata
26a. Blades mostly + oblon ong and acute to oye — at base;
petioles frequently with a purple rin
DE ect. Glosso-
phyilunGGchot) Е mag 4 т, ser. —— Goo Сто И, аи-
тепзе, P. co ichophyll.
um,
P pog P gral 7А. immixtum, "Р he т, P. mo
P. pseudauriculatum, P. ubigantupense, P. ене E бета
26b. Blades ovate to хз Мазы гуни acd to subcordate at base;
petioles usually lacking a purple ring at apex ___________
. P. subsect. sg
phyllum; P. ser. Ovata Croat E cotonense, P. dominicalense
ictum, P. smithii, P. straminicaule, P. ida P бил]
19b. Plant habit unknown; da much зеб than garn Colombia (exact locality
unknown) Ps
ubsect. Eucardium (Engl.) Mayo
Note: For an outline of the sectional classification of Philodendron subg. Philodendron in Central America see
Appendix
340
Annals of the
Missouri Botanical Garden
MORPHOLOGY OF VEGETATIVE STRUCTURES
ANATOMY
Vegetative anatomy. (Section on Vegetative Anat-
omy contributed by R. C. Keating; vouchers will be
cited by Keating (in press)).
General comments. Philodendron has secretory
ducts present in all morphological parts. These oc-
cur mostly as 24 “files” in vascular bundles. La-
ticifers are of the non-anastomosing type, simple
and articulated (French, 1988). The genus also has
secretory ducts present in roots, stems, leaves, and
inflorescences. The roots have a sclerotic hypoder-
mis.
Leaf surfaces. Cuticle smooth or occasionally
rough on both surfaces. Epidermis: adaxial cells po-
lygonal (1–2:1), straight-sided. Abaxial cells simi-
lar to or larger than adaxial cells or more elongate
(2-4:1 Vw). Stomata: abaxial, randomly oriented,
brachyparacytic to brachyparahexacytic. Venation:
secondary veins parallel with reticulate higher or-
der veins.
Leaf Cross Section. Cuticle of medium thick-
ness, may intrude deeply along anticlinal walls of
epidermal cells. Abaxial cuticle may be thinner
and smooth or striate. Epidermal cells: large or
small, square to columnar or tile-like adaxial cells,
larger abaxially, or same size; outer and inner walls
flat. Stomata level with surface with conspicuous
double cuticular flanges often present. Hypodermis:
absent or as 1-3 layers present on adaxial side of
midrib, or as isodiametric cells beneath both epi-
dermal layers (in P. crassispathum, hypoderm of 1—
4 layers adaxially and 2-3 layers abaxially), cells
may contain dark material. Mesophyll: palisade 1—
4 layers of elongated or rounded cells, usually
packed with plastids; occupying 10-30% of lamina.
Spongy layers up to 15 cells deep. Air spaces: large,
substomatal cylinders perpendicular to surface;
30–90% space in spongy layer; or air cavities ir-
regular in some species. Paradermally, spongy tis-
sue with one layer of unarmed cells over abaxial
epidermis, then small cells or short-armed cells
surrounding large air cavities extending from sto-
mata to palisade cells; 3-5 spongy cells per uni-
seriate partition. Large cavities in midrib separated
by uniseriate partitions of ground tissue, partitions
3—4 cells long between junctions. Collenchyma: 4—
9 discrete subepidermal layers of small cells on
abaxial side of midrib, often extending to vascular
bundles.
Vascular bundles. One to several large bundles
in the midrib, often at corners of a net-like aeren-
chyma; well delimited, with 1—5 large metaxylem
cells, next to a small, round or elliptic phloem
strand. Sclerenchyma: fibers, usually as a phloem
cap 1-3 layers deep, and less commonly as a xylem
cap. Some laminar bundles ensheathed with fibers.
Fibers having up to 5 discrete wall layers. Xylem
cells often ringed with a single layer of small pa-
renchyma cells. Secretory ducts: occasional in mid-
lamina or midrib with 1-2 layers of epithelium. La-
ticifers: lateral to vascular bundles, 2-3 per bundle,
outside fibers, often with short processes protruding
between adjacent mesophyll cells; containing dark
amorphous contents. Starch: abundant in midrib
ground tissue, not seen in lamina. Crystals: raphi-
des common or uncommon in single packets in
short or elongate cells, ends protruding across раг-
titions into both adjacent air spaces, or in large
rounded cells of upper or lower mesophyll. Raphide
packets occasionally surrounded by dark sheaths
within the cell (in P. crassispathum crystal cells
elongated parallel with vascular bundles). Druses
rare or common, often coarse and sharp-pointed, in
rounded cells in palisade and spongy tissue parti-
tions. Crystal sand absent or common in most me-
sophyll cells.
Petiole Cross Section. Cuticle smooth and thin.
Epidermis of small cells, level or rounded on sur-
face and inner walls rounded to angular. Hypoder-
mis of 2-4 layers. Collenchyma beneath hypoder-
mis as a continuous band of 4—8 layers of angularly
thickened, longitudinally elongated cells (8-10:1).
Chlorenchyma: up to 5 rows of cells inside collen-
chyma, and in addition, in P jodavisianum, as per
pendicular partitions dividing the collenchyma
and, and reaching subepidermal layers. Ground
tissue of various sizes of parenchyma cells among
numerous small to medium-sized air cavities sep-
arated by uniseriate partitions of up to 3 cells long.
Cells often longitudinally elongated (3-6:1 И).
Vascular bundles numerous and scattered, mostly
oriented with phloem facing perimeter. Bundles are
parallel in longitudinal section with some higher
order angular connections. Xylem of 1-3 ргогоху-
lem cells per bundle, with spiral to annular thick-
enings, and occasionally one metaxylem element.
Phloem of sieve cells and companion cells forming
organized grid. In longitudinal section, sieve ele-
ments with horizontal to 45° oblique end walls.
Sclerenchyma: fibers forming phloem cap, 2-4 cells
deep, usually not fully surrounding phloem later-
y. In some species, fiber caps surrounding xylem
and phloem or ensheathing entire bundle.
Secretory ducts surrounded by epithelium and
1-2 rows of additional small cells, very common 1n
и ин reee
Моште 84, Митбег 3
1997
Стоа! 341
Philodendron Subgenus Philodendron
collenchyma and common in ground tissue. Latic-
ifers: 1-3, found laterally on one or both flanks of
vascular bundles; non-articulated. Starch common
throughout ground tissue, rare in collenchyma.
Crystals: narrow raphide cells occasional, oriented
across aerenchyma partitions with ends pointing
into air cavities, or cells randomly oriented. Druses
present subepidermally and in cells bordering air
cavities. Small groups of small prismatics common
in ground tissue.
Stem Cross Section. 1-3 cm diam. Cuticle of
medium thickness, smooth to rough-surfaced. Epi-
dermis of very small cells, outer walls rounded.
Cork: cells may be present subepidermally. Hypo-
dermis: a subepidermal layer of large cells, over 1—
2 layers of thick-walled fibers, followed by thin-
walled periderm cells. Collenchyma: angularly-
thickened cells, 3-10 layers deep, beneath peri-
derm. Cortical ground tissue of loosely packed pa-
renchyma. Air cavities: cell-sized or smaller in all
ground tissue. Central cylinder: weakly delimited
by a circle of vascular bundles, some of which tan-
gentially fused in pairs or threes by phloem fibers
and confluent phloem strands. Cortical bundles
randomly scattered. Vascular bundles collateral and
highly variable in form and orientation. Compound
bundles numerous in central cylinder with irregular
fiber strands in center of groups of 2-6 bundles.
mall phloem strands outside fibers facing groups
of 1-2 metaxylem elements on outer side. Simple
collateral bundles may have a large wide strand of
fibers surrounding phloem, a very small irregularly
shaped strand of fibers, or no fibers. Fibers having
up to 7 discrete wall layers visible.
Secretory ducts small and numerous in collen-
chyma and outer cortex, less common elsewhere.
Duct cavities surrounded by two modified layers of
small cells; contents of cavities testing positively
for lipids. Laticifers: absent or present as 1-3 cells
lateral to many vascular bundles, occasional in
phloem. Crystals: raphides as single packets in
elongate cells, most common in central cylinder.
Druses of all sizes common in small cells in cortex.
Vascular anatomy. The configuration of shoot vas-
culature of Philodendron has been extensively
studied by French and Tomlinson (1980, 1981,
1984). They reported the genus to be one of the
most diverse in the family from an anatomical point
of view. The axis of the vascular system of Philo-
dendron is continuous throughout the stem because
the renewal shoots develop precociously and be-
cause the morphologically terminal parts of the
stem soon become branches (French & Tomlinson,
1984). All species of Philodendron examined by
French and Tomlinson have an “independent cor-
tical vascular system of traces that pass from the
leaf into the cortex, but without entering the central
cylinder.” Generally the cortex is wide with 3-5 or
more indistinct series of bundles. Cortical bundles
are collateral, typically with a fibrous sheath next
to the phloem. Major leaf traces enter the cortex at
an acute angle and promptly enter the central cyl-
inder, while smaller traces enter the cortex at a less
acute angle and may enter the central cylinder well
below where they entered the cortex. No endoder-
mis was observed by French and Tomlinson for any
of the species.
Philodendron subg. Philodendron has secretory
resin canals occurring in the cortex. These consist
of a schizogenous space lined with 2-3 layers of
epithelial cells and contain a Sudan IV-staining
resin (French & Tomlinson, 1984). The central cyl-
inder is separated from the cortex by an arrange-
ment of vascular bundles into which the leaf traces
merge. Bud traces are equal on both sides and form
an arc near the periphery of the central cylinder
before joining with the axial bundles.
In their survey of 3-dimensional arrangement of
vascular bundles, leaf traces and axial bundles
were distinct, with the leaf traces consistently col-
lateral with protoxylem and usually with a promi-
nent sheath of sclerenchyma next to the phloem.
The axial bundles were divided roughly into five
groups, four of which pertain to P. subg. Philoden-
dron.
In the P. hederaceum pattern (French € Tomlin-
son pattern 1) there are simple collateral bundles
with some bipolar bundles in the central cylinder
of the internodes with thin-walled and non-lignified
ground tissue.
Another relatively rare pattern involving P. sect.
Baursia (but not a Central American representative
of the section) and P. jacquinii (pattern 3 of French
& Tomlinson) has compound bundles throughout
the central cylinder with individual compound bun-
dles consisting of strands of xylem and phloem sep-
arated from each other by sclerenchyma “in the
form of a partial or complete sheath” (French &
Tomlinson, 1984). In P. jacquinii leaf traces enter
the central cylinder but make a variety of config-
urations, sometimes including pairing before join-
ing the compound bundles. Compound bundles do
not make a particularly straight course, sometimes
bundling and pairing within and between com-
pound bundles.
A pattern of vascular bundle traces exhibited by
P. fragrantissimum and P. roseospathum (French &
Tomlinson pattern 4) is similar to pattern 3 except
that "the pattern of the vascular components with
342
Annals of the
Missouri Botanical Garden
compound bundles is less clear because compo-
nents are not separated by sclerenchyma” (French
& Tomlinson, 1984). The sclerenchyma occupies
the central core but does not isolate individual bun-
dle components.
The pattern for P. mexicanum and P. sagittifol-
ium (French & Tomlinson pattern 5), described as
the most common in the genus, has axial bundles
strictly amphivasal with a central core of phloem
and a peripheral region of xylem. The tracheary
elements may form a more or less continuous су!-
inder or may be arranged in clusters around the
phloem. Leaf traces penetrate the central bundle
and fuse with the central cylinder, while the scle-
renchyma sheath of the trace migrates to the center
of the axial bundle.
Нађи and growth patterns. In terms of growth
habit Philodendron is clearly one of the most vari-
able genera in the Araceae (Blanc, 1977a, 1977b,
1978, 1980; French € Tomlinson, 1981). The habit
ranges from terrestrial to epiphytic or hemiepiphy-
tic (primary or secondary). Secondary hemiepiphy-
tes may be vines or appressed-climbers.
Relatively few Philodendron species are terres-
trial, although a few are consistently terrestrial.
These include P. glanduliferum, P. grandipes, P.
malesevichiae, and perhaps Р. hammelii (known
only from a single collection). The stem of P. glan-
duliferum (Fig. 198) is usually repent. Other spe-
cies are mostly terrestrial, but sometimes hemiepi-
phytic: e.g, P. basii, P. roseospathum var.
roseospathum, and Р. warszewiczii. Philodendron
knappiae is about equally terrestrial or hemiepi-
phytic. Label data regarding habit on herbarium
collections are often suspect, with many herbarium
labels using the term “epiphyte” when in fact the
collections were probably rooted in the soil and
were therefore technically hemiepiphytic. In addi-
tion, although many species have collections re-
porting them to be terrestrial, the majority were
probably found on the ground as a result of acci-
dentally falling from trees. Many collections are
naturally made in virgin forest areas where man-
made disturbances such as road building give ac-
cess to the forest. In such areas, and especially in
the regrowth along road cuts, aroids are very com-
mon. Many Philodendron species, e.g., Р. lenti
(Fig. 261) and P. squamicaule (Fig. 382), persist in
excellent condition on steep roadbanks because the
steep clay slopes allow adequate drainage for the
plants to survive. Still, these are somewhat unusual
situations. On the other hand, stream banks often
provide similar habitats.
Many members of Р. subg. Philodendron are
hemiepiphytic, meaning that they are growing on
trees as appressed climbers or as vines, while being
rooted in the soil. There are two types of hemiepi-
phytes (Putz & Holbrook, 1986). Primary hemiepi-
phytes begin their lives when seeds germinate on
the host tree, in the same way as true epiphytes,
then go on to develop slowly until they reach suf-
ficient age and size to begin developing long aerial
roots that may eventually reach the ground. This
growth strategy enables the species to quickly at-
tain a height where light is adequate but nutrients
are more difficult to obtain. Not surprisingly, one of
the most successful species with this life form is P.
megalophyllum Schott, which often lives on ant
nests from which it obtains extra nutrients. No Cen-
tral American species are known to live exclusively
with associated ants, although occasional ant nests
occur among the cluster of roots of many species.
The roots hold the otherwise fragile ant nest to pre-
vent it from being washed away in the rain. Ex-
amples of this type of hemiepiphyte are P. goeldii
С. Barroso, P. solimoesense А. C. Sm., P. megalo-
phyllum (all from South America), and P. radiatum
(Fig. 1), the only species known with this growth
form in Central America.
Secondary hemiepiphytes (Putz & Holbrook,
1986) start their lives on the ground or on tree
trunks near the ground (where they may send roots
to the ground) and climb trees where they become
adults and may lose their connection with the
ground. Most Philodendron sect. Philodendron are
members of this group.
As is the case with most Araceae, adult plants
of P. subg. Philodendron have homeophyllous mon-
opodial growth!! (monophyllous sympodial of Rit-
terbusch, 1971) with each article bearing a cata-
phyll and a leaf. When the plant is mature, each
article is terminated by an inflorescence (Ray,
1986, 1987b) or its aborted remains (Engler, 1877;
Ritterbusch, 1971). The branching pattern for Р
subg. Philodendron has the prophyll following each
internode suppressed, with a single foliage leaf fol-
lowed by the inflorescences and with the internode
subtending the prophyll of the elongation shoot be-
ing suppressed. Growth patterns for Р. subg. Philo-
dendron were diagrammed schematically by Engler
n “In monopodial growth, flowering occurs on axillary
short shoots and does not interrupt the formation of the
main shoot by a single apex. In monophyllous sympodial
growth, after the shoot terminates in an inflorescence
terminating in another inflorescence, and being тер
by another continuation shoot with one foliage leaf” (Ray,
987b).
ЦД ACCC A LL! NL LLL LLL она Qn
E O
Volume 84, Number 3
1997
Croat 343
Philodendron Subgenus Philodendron
(1877) and Ray (1988) and discussed in detail by
French and Tomlinson (1981). In contrast to adult
plants of P. subg. Philodendron, the juvenile growth
phase (with a few exceptions) has monopodial (ac-
tually anisophyllous sympodial) growth with an ar-
ticle bearing an indeterminate number of foliage
leaves before terminating in the first (likely abort-
ed) inflorescence, marking the transition into ho-
meophyllous sympodial growth where each article
has a fixed number of leaves and is terminated by
an inflorescence or its aborted remains. This tran-
sition does not mean that stem has reached matu-
rity. It has only made a transition from monopodial
(actually anisophyllous sympodial) growth to ho-
meophyllous sympodial growth. At this point, the
plant is probably still not mature, and while every
article will end in an infloresence, these will all be
aborted. Later, after further thickening of the stem,
the shoot will become mature, and this will be in-
dicated by the fact that the inflorescences do not
always abort (T. Ray, pers. comm.).
In contrast to P. subg. Philodendron, P. subg.
Pteromischum has monopodial growth only up until
the time of flowering when it is followed by aniso-
phyllous sympodial growth, wherein the stem arti-
cles bear a variable number of leaves before pro-
ducing another inflorescence. Each new branch
forms from a bud in the “axil of the penultimate
leaf of the previous unit” (French & Tomlinson,
1981). Both P. subg. Pteromischum and P. subg.
Philodendron have development of the stem beyond
each terminal inflorescence. Thus although P. subg.
Philodendron appears to have an unbranched stem
with an inflorescence in each leaf axil it is really
producing a new branch after producing each leaf.
Close examination will show that a bud for the re-
newal shoot occurs on each article just below the
point of overlap of the sheath edges of the cataphyll
(prophyll of Ray, 1987a). It is from this point that
the new branch will form. There is also a second,
supernumerary bud which lies below (proximal to)
the bud for the renewal shoot. This acts as a reserve
meristem, lying dormant unless the stem is severed
just distally to it (French & Tomlinson, 1984).
Unlike P. subg. Philodendron, P. subg. Pterom-
ischum typically does not branch after the produc-
tion of each new leaf and therefore lacks compa-
rable branch buds. The inflorescences are never
produced terminally on determinate lateral branch-
es (Grayum, 1996).
Internode length and width are altered markedly
as the plant climbs, with the internodes getting ever
shorter and thicker. Ray (1986) has shown a direct
correlation between the length and width of an ar-
ticle (one segment of the stem) demonstrating that
the length-width ratio is fixed and varies according
to a set pattern. Ray (1987b) categorized four dif-
ferent types of stem segments based on where the
leaf was attached to the stem. In Р subg. Philo-
dendron all species had the petiole attached to the
lower end of the stem segments, and these are re-
ferred to as “hypophyllous segments.” Most species
of P. subg. Pteromischum have “hyperphyllous seg-
ments” where the petiole is attached to the upper
end of the stem segment. “Ambiphyllous seg-
ments,” where the stem segment is so short that the
petiole is attached across the entire segment, is also
known in P. subg. Pteromischum and is also the
the stem segments elongated and extended below
the point of attachment of the cataphyll.
Most members of P. subg. Philodendron have
leaves that turn from juvenile to adult gradually as
the plant climbs so that there is no abrupt transition
to adult foliage.
Although not as pronounced as in Monstera or
Syngonium, leaf dimorphy is present in some spe-
cies of Philodendron. For example, in P. hedera-
ceum var. hederaceum juvenile leaves have short
petioles tightly appressed to the tree, more or less
like the “shingle” leaves of Monstera. They are also
dark blackish green and velvety above and often
somewhat purplish violet beneath. The adult plants
have spreading leaves with longer petioles and
smooth, semiglossy blades. The velvety blades
seem to be associated only with the earliest growth.
Once the plant grows up high enough on the tree
the leaves become smooth and semiglossy. Further
growth, even when it represents a reversion to
smaller-leaved forms, results in the same smooth,
semiglossy texture as that of the adult.
Philodendron hederaceum and other scandent
species tend to climb high up the trunk of the tree
then spread into the canopy and finally often hang
down from branches before they flower. Philoden-
dron jacquinii often has a similar habit. Some scan-
dent species, such as P. brevispathum and P. sul-
cicaule, typically sprawl over lower vegetation
rather than high in the canopy.
The amount of internode elongation varies im-
mensely in some vining species, such as P. immix-
tum, P. hederaceum, and P. sulcicaule, with inter-
nodes 10-20 cm long (even longer on plants that
are juvenile or have reverted to a “searching mode”
as the result of dislodgement from the tree). Usually
internode length varies considerably depending on
the light and nutrients available for growth. Even
Annals of the
Missouri Botanical Garden
fully adult plants with very short internodes can be
induced to produce longer internodes in cases
where the plant is accidentally displaced from its
growing situation or if it becomes heavily shaded.
Some species have evolved the ability to produce
flagellate branches in an attempt to reposition
themselves. These have long slender internodes
and reduced leaves. At first the flagelliform growth
spreads laterally, but if no other growth support is
in the immediate vicinity the branch inclines to-
ward the ground where it may creep across the sur-
face and ascend another tree. Blanc (1980) referred
to these as “flagelles” or “stolons.” Ray (1987b)
indicated that these flagellate branches are usually
developed when a plant overgrows its support or is
accidentally dislodged, but in some species, e.g.,
P. fragrantissimum, the flagellate branches may de-
more slender flagellate stems, which ascend the
tree trunk. After a period of rapid growth resulting
in long, narrow internodes with the leaves reduced
to small scales, the plant produces a series of short
thick internodes, each of which is associated with
an increasingly larger leaf, and leaves arranged in
a tight rosette (Ray, 1987b). Ray (pers. comm.) re-
ports that “with only a few exceptions, flagellar
shoots do not occur in species with homeophyllous
growth. Flagellar shoots are almost universal among
climbing species with anisophyllous or intermittent
homeophyllous growth, but almost totally lacking
among climbing species with homeophyllous
growth.”
Stems. Stems vary considerably in length depend-
ing on the nature of growth. Vines, such as P. hed-
eraceum, may produce stems well over 30 m long,
whereas the more slow-growing appressed climbing
hemiepiphytes rarely have stems more than about
2 m long. The youngest part of the stem bears most
of the leaves and has the newest root growth in-
cluding the most active feeder roots. Older portions
of the stem have thick, strong roots usually tightly
fastened to the tree, anchoring it in place. Some
roots all along the stem extend downward along the
stem and may lead all the way to the ground. The
older portion of the stem is often devoid of any
leaves and usually lacks even cataphyll fibers. The
bare stem clearly shows the petiolar and cataphyll
scars (Figs. 9-12). See discussion below. Inspection
usually shows that the older portions of the stem
have been at some time attacked by root borers so
there may be surface damage or more likely active
root borers in the center of the stem. Eventually the
lower part of the stem rots away as the stem climbs
higher on the side of the tree.
Stems of Philodendron are typically unbranched,
but may be branched naturally, as in many species
of P. subg. Pteromischum, or through injury. Com-
monly internodes become increasingly shorter and
thicker toward the apex of the stem on appressed-
climbing plants, but some species, e.g., P. fragran-
tissimum, may from time to time begin to produce
long slender internodes to enable the plant to enter
a colonizing mode (see above section on “Нађи and
Growth Patterns").
Mayo (1991) described differences in the
branching pattern of P. subg. Philodendron and P.
subg. Meconostigma. While most P. subg. Mecon-
ostigma have very short internodes, some species,
e.g., P. leal-costae Mayo & Barroso and P. corco-
vadense Kunth, have elongated internodes. In these
species the elongated internode of each article is
the one between the cataphyll (prophyll of Ray,
1987a) and the succeeding foliage leaf, while the
internode preceding the cataphyll is abbreviated.
In P. subg. Philodendron the elongated internode is
the hypopodial internode (which precedes the cat-
aphyll), while the internode between the cataphyll
and the foliage leaf is the abbreviated one.
Stems of Philodendron are typically rich in tax-
onomic characters, and together with their associ-
ated cataphylls, yield some of the best key char-
acters for identification. Normally these stem
characteristics are useful only at the specific level,
but Grayum (1996) has found the dried stem color
to be useful in separating the two sections of P
subg. Pteromischum. Unlike most Anthurium spe-
cies, which have internodes so short that the epr-
dermal features are for the most part obscure or
very uniform, Philodendron has features important
both before and after drying. The internodes of most
appressed-climbing hemiepiphytes are actually
broader than long at maturity (Figs. 10, 394. 443),
but some species, such as P. advena (Fig. 38) and
P. straminicaule (Fig. 392), have internodes typ
cally longer than broad (though short enough that
e plants are not considered scandent). Only nine
species in Central America have internodes longer
than 25 cm on adult or preadult plants. Alterna”
tively, many species have short internodes, with
Central American species having internodes to only
5 cm long or less.
Internode width is somewhat less variable, rang-
ing from less than 3 mm diam. in P. chirripoense to
10 cm diam. in P. gigas. Only nine additional spe
cies (P. chiriquense, P. copense, P. dressleri, P. fer
Volume 84, Number 3 Croat 345
1997 Philodendron Subgenus Philodendron
rugineum, P. fortunense, P. grayumii, P. pterotum, Stems commonly dry with irregular (or some-
P. schottianum, and P. warszewiczii) have stems that
attain diameters greater than 7 cm. Most species
have relatively narrow internodes, with 44 species
having internodes of no more than 3 cm diam., and
16 species with internodes of less than 1.5 cm
iam.
Fresh stems often have characteristic surface
features. Surfaces are frequently speckled with a
lighter or darker pattern of green. The surface may
also be short-lineate (Fig. 2) or may present a com-
bination of short lines and speckles (Fig. 3). At
other times the surface may be weakly to promi-
nently striate as in P. anisotomum, P. hederaceum,
and P. wilburii. The stems may be glaucous | as
m,
dii. F «етт the internodes are coarsely short-
costate (Fig. 2) near the apex of the internode, as
P. chiriquense, P. grandipes, and P. schottianum.
These apparently represent areas at which roots
will emerge later.
Stems are commonly smooth but may be minute-
ly wrinkled, as in P. verrucosum, or sparsely warty,
as in P. brunneicaule, or setose, as in P. jacquinii,
or with often branched, trichome-like scales (Fig.
5), as in P. brevispathum. Other species with scaly
stems are: P. hammelii, P. malesevichiae, P. squam-
icaule, P. squamipetiolatum, and verrucosum.
Even stems of species with smooth epidermis are
often weakly fissured in fresh condition (Fig. 2).
Coloration of fresh stems may also be character-
istic; typically, they are medium to dark green when
esh, turning gray-green to brownish, yellow-brown
or reddish brown in age. Often several caras
stages are involved, e.g., medium green
green, to yellowish green and finally ent or
reddish brown. This final stem color is often the
same as the color of the stem of artificially dried
herbarium collections, but at other times the dried
stem color of the herbarium collection is not the
same. Thus it may be important to note the color
of the fresh stem before drying takes place. Natural
aging of the stems often causes a scurfy condition,
which results from numerous close cracks (Figs. 4,
5). Sometimes these minute fissures are ciim
to a "is point of stress yet not visible elsewhere
(Fig. 5
important feature is the extent to which دة
the stem wrinkles or cracks or is otherwise distorted
by the drying process. The results of the drying
Process are usually quite consistent from collection
to collection, such that the dried stem and the fea-
tures it exhibits provide useful recognition char-
acteristics.
еф
imes regular) ribs and intervening sulcae, reflect-
ing shrinkage of the relatively indurate outer sur-
face of the stem compared to the rather soft,
somewhat aerenchymatous stem interior. In some
cases the regularity and severity of this ribbing pro-
vide especially useful characteristics, e.g., in P. sul-
cicaule the stems of which become prominently
ribbed in the course of normal development, and
in P. verapazense where the stems become regularly
and conspicuously ribbed on herbarium collections.
In addition to frequently present longitudinal
ribs, in e.g., P. findens, P. fortunense, and P. helen-
iae, stems may be transversely fissured or checked
with small to large fissures. These may be quite
regular or irregular in severity or spacing. While
sometimes characteristic, these transverse stem fis-
sures tend to be somewhat less consistent than the
longitudinal fissures. However, for some species
such as P. edenudatum, P. ferrugineum, P. iine
P. fortunense, P. ligulatum, P. malesevichiae, P.
mexicanum, and P. wendlandii (among others), the
de-
scribed South American species, i.e., Croat 62785
from the Pacific coast of Colombia
Another feature exhibited by some dried stems
is an exfoliating epidermis, present in P. an
lobum, P. cotonense, P. dodsonii, P. hederaceum, »
schottianum, P. зтићи, P. straminicaule, P. subin-
cisum, and P. tripartitum. In some cases the epi-
dermis not only cracks but may begin to loosen and
fall off or protrude away from the stem (Fig. 8), such
as in P. brevispathum, P. cotonense, P. dodsonii, P.
ligulatum, and P. purpureoviride. Occasionally this
feature is exhibited on fresh stems as well. This is
especially true if the stem is forced to bend by
falling from its support, such as in P. immixtum and
P. sulcicaule. The epidermis appears to be hard and
brittle, while the underlying stem appears to be
green and supple. In some cases the epidermis
seems to be naturally shed and replaced by another
epidermal layer on the fresh stem.
Petiolar scars. Although less conspicuous in gen-
eral than in Anthurium or in P. subg. Meconostig-
ma, the petiolar scars on the stems of P. subg.
Philodendron are nonetheless clearly visible unless
covered with persisting cataphylls. In contrast to
Anthurium the petiolar scars of P. subg. Philoden-
dron are generally much less indented but rather
are more or less flush with the general contour of
the stem surface. They may be moderately incon-
spicuous as in P. rothschuhianum (Fig. 9) or mod-
erately conspicuous as in P. davidsonii (Fig. 10).
346
Annals of the
Missouri Botanical Garden
Philodendron petioles are typically much swollen
at the base, so the scar is usually broader than the
rest of the petiole itself. Petiole scars vary from
(0.5—)1 to 4(—7.5) cm high and (0.7—)1 to 5(-7) ст
diam., but they are rarely more than 3 cm high and
3.5 cm diam. The average height and width for the
32 species studied is 2 cm high and 2.4 cm diam.
The peduncular scar is often quite conspicuous
(Fig. 5) and deep. Of ecological significance is the
fact that these deep holes left when the inflores-
cences fall off are the points of entry for phytoph-
agous insects, especially stem borers, which infect
the older and sometimes the younger portions of the
stems.
Intravaginal squamulae (Dahlgren & Clifford,
1982), so prominent in P. subg. Meconostigma, are
usually present but often inconspicuous in P. subg.
Philodendron (Fig. 11). Mayo (1991) stressed this
as evidence of differences in two contrasting pat-
terns of stem elongation (see discussion under stem
above). In P. subg. Philodendron the intravaginal
squamulae (Fig. 11) are always immediately above
the cataphyll (prophyll of Ray, 1987a), whereas in
P. subg. Meconostigma the intravaginal squammu-
lae are immediately below the cataphyll scars and
often around the scar for the foliage leaf as well.
Roots. Philodendron roots have an anatomically
distinct layer of exodermis beneath the epidermis,
distinguished, among other things, by a long-cell/
short-cell pattern (French, 1987a). “Thick-walled,
pitted sclereids form a cylinder adjacent to the en-
dodermis and similar sclereids also occur singly or
in bands with suberized cork cells in the periderm
of older roots" (French, 1987а). Like those of other
members of tribe Philodendreae, Philodendron
roots have a sclerotic hypodermis. French reported
the scelerotic hypodermis to be distinctive because
of its position next to the exodermis and its occur-
rence in the primary axis. Another distinctive an-
atomical feature of the roots of Philodendron is res-
in canals with sclerotic sheaths (French, 1987a).
All species of Philodendron produce adventitious
roots at some or all nodes. The number of roots
developed seems to have more to do with the en-
vironment than with the species involved. Plants
that are appressed-climbing and in close contact
with the substrate generally produce the largest
number of adventitious roots. Roots may be of two
types, which differ both morphologically and ana-
tomically (Lierau, 1888; Porsch, 1911), either for
anchoring the plant to the substrate or for feeding
(van Tieghem, 1907; Went, 1893). The anchor roots
(Fig. 54) tend to be more numerous and shorter,
often with a dense layer of root hairs (they are
sometimes restricted to the side of the root that
contacts the substrate). They also have а propor-
tionately much smaller central cylinder and more
mechanical tissue to give them strength (Engler &
Krause, 1908) than those of feeder roots. They arise
principally at the nodes but may arise all along the
internodes. Anchoring roots may spread from the
nodes as in P. auriculatum (Fig. 12) or closely ap-
pressed to the surface of the tree on which they are
growing as in P. gigas (Fig. 13). In contrast to the
anchoring roots, the feeder roots (Fig. 14) tend to
be much thicker and longer and usually extend to-
ward the ground. This behavior is to be expected
since Goebel and Sandt (1930) reported that feeder
roots of aroids are negatively heliotropic and posi-
tively hydrotropic. Feeder roots have a much broad-
er central cylinder and broader vessels and sieve
tubes. Feeder roots arise only at the nodes (Gra-
yum, 1990). Normally, the feeder roots are 2-4
times thicker than the anchor roots, and in P. gigas
the feeder roots may be somewhat woody and up to
3.5 cm diam.
Some species, such as P. auriculatum (Fig. 12),
have spine-like branch buds sparsely scattered
along the length of the root, especially near the
stem. Some hemiepiphytes, such as P. solimoesense
A. C. Sm. in South America, have roots that may
become markedly roughened with warty tubercles.
In such cases the only portion of the root that has
root hairs is the apex where the roots branch as
they near the ground. It is unknown whether these
roots are capable of absorbing atmospheric humid-
ity, as is true for some Anthurium species, but cer-
tainly they must be able to take in the free water
that runs down the root.
French (1987a) reported that in P. subg. Philo-
dendron a sclerotic hypodermis is entirely absent
in subterranean roots but present in the aerial roots.
Philodendron subg. Philodendron has elongated,
infrequently anastomosing resin canals that extend
lengthwise through the root cortex (French, 1987c).
They are lined with a layer of epithelial cells that
consist of thin-walled, unlignified cells (рагепсћу-
ma). In P. subg. Philodendron and P. subg. Pter-
omischum the epithelium is surrounded by a sheath
with lignified cell walls. In contrast, P. subg. Me-
conostigma has resin canal sheaths that lack a scle-
renchyma and instead have 2-5 layers of (unlig-
nified) collenchymatous cells, which are easily
distinguished from the ground tissue (French.
1987c
While seldom used taxonomically, roots are уап”
able to some extent from species to species. Fresh
root coloration (ranging from whitish to green Юю
brownish), length, diameter, and surface texture
Volume 84, Number 3 Croat 347
1997 Philodendron Subgenus Philodendron
Table 2. Persistent condition of cataphylls by section.
Deciduous Persisting
% of Sem As 0 of Section
Section Total sect. Intact intact fibers Total sect. total
Calostigma 34 69% 6 6 3 15 31% 49
Macrogynium 1 100% 0 0 0 0 0% 1
Philodendron 10 24% 0 18 14 32 76% 42
Polytomium 100% 0 0 0 0 0% 3
Tritomophyllum "i 8896 0 1 0 1 13% 8
Тога] 55 53% 6 25 17 48 47% 103
(smooth, coarse, or even warty), as well as the dried
color and degree to which they are fissured or fold-
ed, are all features that may be recorded. These
features have not been used extensively since the
roots are generally removed from the stems before
the herbarium specimens are prepared.
Cataphylls. In the taxonomy of Philodendron
probably no morphological feature is more di
nostic than the cataphyll (sylleptic prophyll of йй,
19872). Philodendron exhibits considerable varia-
whether the cataphylls are deciduous or persistent.
This is the single feature that makes the prepara-
tion of keys to species of Philodendron easier than
those for Anthurium. In P. subg. Philodendron 56
taxa have deciduous cataphylls, while 48 have per-
sistent cataphylls. Of the latter, 6 have cataphylls
that persist intact, 25 have cataphylls that persist
semi-intact, and 17 have cataphylls that persist as
fibers. While a few species have cataphylls tardily
deciduous and others have persistent cataphylls
that eventually fall off, relatively few species are
difficult to place in one or the other category. See
Table 2 for a breakdown of persistent condition of
cataphylls by section.
Generally speaking cataphylls are deciduous in
vines and persistent on appressed-climbing plants,
but there are exceptions in both groups. Cataphylls
are bract-like modified leaves which function in the
protection of newly emerging leaves. For vines, the
cataphyll becomes functionless once the leaf has
emerged and generally is promptly deciduous. Typ-
ically it recurls away from the stem on vines and
eventually becomes loosened from the base and
falls free, such as on P. purpureoviride (Fig. 320).
Alternatively, on epiphytes or appressed-climbing
hemiepiphytes with short internodes the cataphylls
cannot easily fall free (Figs. 15, 115, 312). Al-
though they may fall free from the stem and in-
variably promptly dry and usually become withered
or weather into fibers, they are often prevented from
falling by the tight cluster of petioles generally
sleri (Fig. 160), but typically cataphylls persist on
species with short internodes (Fig. 302, P. pirrense;
Fig. 358, P. scalarinerve). Cataphylls often persist
in an organized mass around the apex of the stem
(Fig. 382, P. squamicaule). They may persist intact
in P. grandipes (Fig. , P. jodavisianum (Fig.
248), and P. roseospathum to 341, 343), or vl
a very thin, flaky epidermis that remains intact in
large pieces as in P. dodsonii, P. hebetatum, or P.
strictum (Fig. 398); more frequently they decom-
beneath the epidermis (Fig. 258,
even more frequently the epidermis еа а]-
together (Fig. 202, Р. glanduliferum; Fig. 253, Р.
jodavisianum; Fig. › schottianum). Ecologi-
cally the mass of cataphyll fibers serves a useful
purpose, namely to prevent desiccation of the stem
apex and especially the young roots which emerge
through the moist cataphyll mass. In some species
the layered cluster of cataphylls forms a sodden
mass (Fig. 370, P. schottianum; Fig. 63, P. anto-
nioanum; Fig. 99, P. chiriquense), and in some
cases the youngest cataphylls are protected by a
gelatinous, mucilaginous fluid that fills the inter-
stices of the cataphyll mass. Cataphylls are gener-
ally rigid and firm when fresh, which protects the
young leaf from physical damage. Some are thick
and fleshy with considerable amounts of liquid in
their tissues. As these begin to decompose they
may yield large amounts of watery or even gelati-
nous sap. This fluid may be important as a lubri-
cant for the emerging leaf to prevent damage to the
tender tissues or to prevent desiccation. Since ep-
iphytes are often subject to harsh conditions, this
is probably important to protect the growing point
of the plant and the newly emergent leaves from
Annals of те
Masour! Botanical Garden
persist
ganized network of fibers so they continue to add
protection to the stem apex. Even after the cata-
phylls are reduced to a mass of fibers they retain
proportional
any species, ranging from (5-)10 to 504-70) cm
long and (1.5-)2.5 to 18(-31) cm wide. Since the
is affixed around 1
and may be as much
slender
as 1 cm high, as in P. auriculatum (Fig. 72) and Р.
yellowish epedermas of the cataphslls i one of the
ont РР АСТИ bratures
Croat
Philodendron Subgenus Philodendron
ways less than 30 cm long, 11 less than 20 cm long,
a eis. d mesi, weis A P. brewster-
ense, P. chirri have
sulcate at the base (Fig. 212). In either event they
then И somewhat flattened toward the apex
eS» am raa
edly
as in P. fortunense (Fig. 183) and P.
fines i 11, Sometimes e pts P. lig-
var. heraclioanum also may be weakly
ane а 275, 276). Even petioles not D-sha-
ped often have a slightly thickened, slightly raised
surface. Examples
Канан МИ је the rained. magne of
the petioles, there may be a prominent medial rib,
350
Annals of the
Missouri Botanical Garden
such as in P jodavisianum and occasionally P.
grandipes, giving the petiole a three-ribbed ap-
pearance adaxially. More commonly the medial rib
is broad and obtuse, such as in P. angustilobum, P.
annulatum, P. copense, P. crassispathum, P. ferru-
gineum, P. ligulatum, P. llanense, P. schottianum,
P. smithii, P. tenue, P. thalassicum, and P. triparti-
tum.
Rarely the petioles may be markedly flattened
dorso-ventrally, as in P. platypetiolatum. In this
case the petioles are much broader than thick with
the lateral margins nearly acute.
A few species have petioles markedly thicker
than broad and U-shaped in cross section. Exam-
ples include P. roseospathum, P. jodavisianum, and
sometimes P. davidsonii. Petioles of the latter spe-
cies are more commonly obtusely V-shaped in cross
section.
Petioles of P. subg. Philodendron typically have
very short sheaths when the inflorescence is abort-
ed and much longer sheaths (to accommodate the
emerging inflorescences) when subtending an inflo-
recence (Figs. 38, 67, 93, 150). Generally the
sheath is markedly closed with one margin over-
lapping the other, making it inconspicuous (Figs.
83, 160). Occasionally the petiole sheath may be
conspicuous, such as in P. correae (Fig. 119) and
some members of P. subsect. Glossophyllum.
A geniculum is typically not apparent, but the
genicular area serves the same purpose as it does
in Anthurium. It is involved with inclining or twist-
ing the plane of the blade, presumably optimizing
exposure to sunlight. Usually the genicular area is
somewhat firmer than the remainder of the petiole,
and sometimes it is differently colored, but it is
usually neither swollen nor as conspicuously dis-
tinct as in Anthurium. Typically the genicular area
is the same shape as the remainder of the petiole,
but sometimes the cross-sectional shape is differ-
ent, being more frequently bluntly ribbed and often
rather deeply cracked or scurfy around the circum-
ference (perhaps owing to the bending in response
to light). The geniculum, when apparent, is some-
times thicker than the remainder of the petiole as
in P. brunneicaule, P. ferrugineum, P. heleniae, and
P. tenue. It may be darker than the petiole as some-
times in P. bakeri or P. glanduliferum, or slightly
paler as in P. ferrugineum or P. scalarinerve.
Most species of Philodendron, like Araceae in
general, have glabrous petioles, but there are con-
spicuous exceptions. Several species have conspic-
uously scaly petioles: P. hammelii, P. glanduliferum,
P. malesevichiae (Fig. 283), P. squamipetiolatum (Fig.
387), P. squamicaule (Fig. 382), and P. verrucosum.
The type of scales is also variable. In some species,
such as P. malesevichiae, P. glanduliferum, and P.
squamipetiolatum, the scales are acicular and more
or less terete. The scales of P. glanduliferum and P.
malesevichiae are spreading, while those of Р
squamipetiolatum are sometimes retrorse near the
apex of the petiole. In addition, they are densely
granular-scurfy on the surface. In P. hammelii the
scales are short and broadened laterally, similar to
fish scales, and less than three times longer than
broad. In P. verrucosum and P. squamicaule the
scales are of two distinct types. Philodendron ver-
rucosum has short, broad, and often lacerate scales
interspersed with long-acicular scales. Philodendron
squamicaule has short, purplish, deltoid, broader
than high (about 2 mm high) scales interspersed
among acicular, greenish scales 3-5 mm long.
etiolar scales are not uniformly distributed
. ш Р. squamipetiolatum, P.
squamicaule, and P. verrucosum the scales are usu-
ally scattered throughout the petioles. In P. male-
sevichiae they are present in the upper % to % of
the petioles, becoming increasingly denser toward
the base. In P. glanduliferum they are similar but
more closely aggregated toward the apex. In P.
hammelii the scales are restricted to a small area
near the apex. Philodendron ornatum in South
America is the most extreme example of this re-
duction. In that species the scales are reduced to
stubby protuberances restricted to the apex of the
petiole.
Surface features of both fresh and dried petioles
are often conspicuous and frequently diagnostic.
Though usually solid green, petioles may be irreg-
ularly purplish-spotted as in P. edenudatum (Fig.
163) and P. sagittifolium. This purplish spotting
may continue onto the lower midrib, a feature com-
mon to other types of markings that continue onto
at least the proximal portions of the midrib. The
surfaces of petioles may be minutely speckled but
more commonly bear a light pattern of short dashes
or streaks usually uniformly distributed throughout
the petiole. They may be densely to sparsely
marked with short dashes, which in turn may be
either darker or lighter than the surface. I have
referred to this feature in the descriptions as “short-
lineate” or “lineate.” Though the lineations are
usually short, usually less than 3 mm long, they
may sometimes be longer and sometimes are vari-
able in length with both short and long lines inter-
spersed (Fig. 19). Examples of species with short-
lineate petioles include: P. annulatum, P. ашти"
latum, P. copense, P. cotonense, P. edenudatum, Р
fortunense, P. immixtum, P. ligulatum, P. ритепзе,
P. purulhense, P. smithii, P. subincisum, Ё. that"
sicum, P. warszewiczii, and P. zhuanum. The pet
Volume 84, Number 3
1997
Croat 351
Philodendron Subgenus Philodendron
oles of P. brunneicaule are red short-lineate. The
short lineations may be associated as well with
much longer, continuous, and frequently weakly
raised striations or ribs. Species with petioles de-
scribed as short-lineate to striate include: P. anto-
nioanum, P. clewellii, P. gigas, P. glanduliferum,
and P. panamense. Still other species have more or
less continuous striations throughout: e.g., P. ani-
sotomum, P. davidsonii, P. dressleri, P. findens, P.
madronense, P. rothschuhianum, P. straminicaule,
P. tenue, and P. warszewiczii.
Fresh petioles of P. subsect. Glossophyllum usu-
ally have a distinct purplish (or rarely dark green)
ring (Fig. 20) around the entire petiole just below
its junction with the blade. The species that share
this feature, mostly members of P. subsect. Glos-
sophyllum, include P. annulatum (Fig. 59), P. bak-
eri, P. correae, and P. ligulatum (all three varieties;
see Figs. 271, 275) with a purplish or purple-black
g, and P. auriculatum (Fig. 20), P. immixtum, P.
dolichophyllum, P. pseudauriculatum, and P. wen-
dlandii with a dark green ring at the apex of the
petiole.
Other apparently unrelated species may likewise
have purple rings around the apexes of the petioles.
These include P. brenesii, P. davidsonii, P. dressleri,
P. ferrugineum, P. microstictum, P. NOM P. war-
szewiczii, and sometimes P. grayum
Petioles may produce a cluster of dd droplets
of a sweet, sugary solution on the abaxial surface
at the apex of the youngest leaves. This has been
observed on P. davidsonii subsp. bocatoranum (Fig.
19) and P. megalophyllum Schott, a common South
American species. These droplets appear to have
no function in pollination or in blade orientation,
and I speculate that they act as a food source for
ant guards, which serve to prevent the young blades
from predation by phytophagous insects. This
seems all the more likely because the droplets are
associated with new leaves, which are typically very
tender and thus most easily damaged.
Freshly cut petioles usually form resin droplets
in the same manner as the stems, and may in time
become completely covered with resin. Of rare oc-
currence are slender strands of latex on broken and
eb severed petioles as on P. malesevichiae
(Fig. 21).
Dried petioles yield another suite of character-
istics generally unrelated to those exhibited on
fresh plants
The dried petiole is sometimes diagnostic and in
some cases, such as P. hebetatum and P. schottian-
um, petioles have a dried epidermis that is so con-
spicuously yellow-brown as to be nearly unique.
ough less conspicuous, the petioles of P. schot-
tianum and P. thalassicum also dry quite yellowish
rown.
Petiole firmness is variable in P. subg. Philoden-
dron, but this character is difficult to quantify. Un-
fortunately Engler used this character as one of his
major key characters to separate subsections of P
sect. Cardiobelium, P. subsect. Macrobelium, and P.
Doratophyllum (now P. subsect. Bulaoana) with
rm petioles. This character is difficult to describe
and petioles appear to range from quite firm to
quite spongy without any major discontinuities.
Most species have petioles firm to weakly spongy
when squeezed, and generally they are quite flex-
ible, capable of being bent to a great extent without
breaking. However, spongy petioles, such as those
in P. ligulatum which can be easily crushed by
squeezing, are also more likely to be brittle.
Blades. Blade shape.
of morphological variation in leaf shape in the ge-
nus as a whole is exhibited among the Central
American species of P. subg. Philodendron. The
only blade shapes of Philodendron not shown by
Central American species are those seen on P. goel-
dii G. Barroso in P. subg. Meconostigma, which has
blades reniform in outline and pedately compound,
and members of P. sect. Schizoplacium, which have
blades pinnately lobed with a few, prominently fal-
cate divisions. While P. subg. Pteromischum has
distinctive leaves (even discounting the winged pet-
iole on adult plants, which is definitive), their over-
all shape and size is matched by some other species
of P. subg. Philodendron. Blade shape in P. subg.
Philodendron relates in some cases to sectional dif-
ferences. For example, species with three-lobed
blades are either members of P. sect. Tritomophyl-
lum or P. subsect. Bulaoana. Pinnately lobed spe-
cies are members of P. sect. Polytomium. Some sec-
tions, especially P. sect. Philodendron and P. sect.
Calostigma, are highly variable in blade shape,
with that of P. sect. Philodendron ranging from ob-
long to variously ovate to ovate-sagittate and that
of P. sect. Calostigma ranging from oblong to sub-
cordate, prominently cordate, cordate-sagittate, or
even 3-lobed. Philodendron subsect. Glossophyl-
lum, relatively common in Central America, has
more or less oblong, veces subcordate or cor-
dulate blades. See also section on "Discussion of
Subgeneric Classification" ii additional details.
While lobed leaves in some Araceae, e.g., Mon-
stera, are due to necrotic processes in which growth
stops and tissue rots away (Madison, 1977), the
Virtually the entire range
352
Annals of the
Missouri Botanical Garden
lobed leaves of Philodendron are due to differential
growth of leaf tissue.
In Central America more species have ovate-cor-
date blades than any other shape. In all variations,
38 species have some sort of ovate-cordate leaf
blade. Twenty-two species are described as having
ovate-cordate blades, and 11 as broadly cordate.
Species with ovate to ovate-triangular leaves are
most abundant among P. subg. Philodendron in
Central America. At least 55 species have leaf
blades with well-developed posterior lobes, includ-
ing cordate, sagittate, and hastate. Only 15 species
have non-cordate blades prominently longer than
wide. Eight species have lobed blades, six trilobed,
and two pinnately lobed. Species with trilobed
leaves are: P. anisotomum, P. cotobrusense, P. mad-
ronense, P. rothschuhianum, P. tripartitum, and Р.
angustilobum. Species with pinnately lobed leaves
are: P. radiatum and P. warszewiczii.
Posterior lobes. Philodendron subg. Philodendron
species vary greatly in the shape of posterior lobes:
usually rounded for those species with cordate leaf
bases, e.g., P. glanduliferum (Fig. 197) or P. pla-
typetiolatum (Fig. 306), or cordulate'? leaf bases,
e.g., P. auriculatum (Fig. 69), P. immixtum (Fig.
236), and P. pseudauriculatum (Fig. 314); to more
or less oblong to narrowly triangular and much lon-
ger than wide as in P. verapazense (Fig. 434), or
sometimes in P. sagittifolium (Fig. 346) and P. ten-
ue (Fig. 404). Even in species with well-developed
posterior lobes the lobes may be about as long as
broad, as in P. purulhense (Fig. 322), P. tysonii (Fig.
426), or P. schottianum (Fig. 365). Posterior lobes
may be sagittate as in P. brunneicaule (Fig. 95), P.
hebetatum (Fig. 218), or P. gigas (Fig. 194); to has-
tate as in P. angustilobum (Fig. 49), P. anisotomum
(Fig. 55), and P. mexicanum (Fig. 287); or bluntly
triangular as in P. wilburii var. longipedunculatum
(Fig. 461).
Sinus shape. Philodendron species with posterior
lobes have remarkable variation in the shape of the
n g ort, are usually broadly
rounded with each lobe encompassing about half the width
of the entire blade.
hianum (Fig. 339); parabolic in P. sulcicaule; hip-
pocrepiform in P. hebetatum (Fig. 225), P. lazorii,
P. panamense (Fig. 297), and P. squamicaule (Fig.
374); sometimes closed on live plants as in P. for-
tunense (Fig. 182), P. gigas (Fig. 194), P. pterotum
(Fig. 311), and P. schottianum (Fig. 366). In P. for-
tunense and P. subincisum (Fig. 401) the sinus may
be closed even on pressed plants. The shape of the
sinus varies greatly between live and flattened
dried plants, since the shape of the sinus varies
depending on the angle at which the posterior lobes
are turned up in relation to the midrib. Many spe-
cies have the posterior lobes turned upward at an
angle to the midrib on live plants (see, e.g., Figs.
104, 130, 159, 186, 366, 374), causing the inner
margins of the posterior lobes to become closer to
each other and thus decreasing the apparent size
of the sinus. For example, when the angle of the
posterior lobes is extreme the posterior lobes might
be closed, but when the same blade is flattened and
dried the sinus might become spathulate or hip-
pocrepiform. The description of the sinus as pre-
sented in this work for Central American P. subg.
Philodendron is exclusively that of the flattened si-
nus unless otherwise stated.
The sinus shape may be intraspecifically vari-
able as in P. sousae, for example, with the sinus
ranging from spathulate (Fig. 379) to parabolic (Fig.
380). This is in part related to the age of the plant,
with older plants bearing larger blades that have
larger, more well developed posterior lobes.
Blade size. Blades of P. subg. Philodendron in
Central America as elsewhere are highly variable
in size, but relatively few have blades that are con-
sidered huge. Only 19 Central American species of
P. subg. Philodendron have blades that exceed 75
cm long (although many others approach that size).
Of these, only P. gigas has blades that regularly
exceed 1 m in length, with the maximum record
at 137 cm. The blades of Philodendron ferrugineum
may rarely exceed 1 m in length. By the same token
few species in Central America have small leaves.
Only 37 species have blades less than 25 cm long
on adult plants, and of these only 13 have blades
less than 15 ст long. Philodendron brewsterense,
with blades a maximum of 11 cm long, is the spe
cies with the smallest leaves, and P. chirripoense,
with blades up to 11.6 cm long, has leaves about
as small. The median leaf length for Philodendron
in Central America is 57 cm.
The widest blades belong to P. gigas and Po
diatum (to 90 cm long), but P. pterotum (to 84 cm
wide), P. findens (to 70 cm wide), P. dodsonii (to 66
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
cm), P. schottianum (to 64 cm), and P. warszewiczii
(to 62 cm) also have quite broad blades.
While blade size of Philodendron is often not re-
liable for taxonomic separation and even blade shape
is sometimes unreliable, the length-width ratio is, in
general, more reliable as a taxonomic character.
Blades range from being much broader than long, to
more than eight times longer than broad (in P. creto-
sum), to broader than long. Blades are broader than
long in a number of dap e.g., averaging only 0.62
times as long as broad in P. anisotomum and P. brun-
neicaule and 0.7 times as long as broad in Р. coto-
brusense and P. grayumii. The average blade length-
width ratio for species with simple, unlobed leaves
ranges from 1.6 to 2.5 times longer than wide. Ex-
ceptions include P. cretosum and Р. folsomii (8.3 and
5.2 times, respectively).
Blade margins are typically entire on simple
leaves, never toothed and only rarely sinuate, such
as in P. subincisum. Species with weakly sinuate
blades are: P. basii, P. dwyeri, and P. jefense. On
the other hand, blade margins are frequently un-
dulate to a certain extent, especially on larger
blades. Species that have coarsely undulate leaves
are: P. annulatum, P. antonioanum, P. copense, P.
ferrugineum, P. fortunense, P. grandipes, P. hebeta-
tum, P. panamense, and P. sulcicaule.
Blade margins are frequently hyaline and also
often weakly revolute. Examples of species with hy-
aline blade margins are: P. advena, P. crassispa-
thum, P. davidsonii, P. ferrugineum, P. gigas, P. im-
mixtum, P. hederaceum, P. smithii, P. squamicaule,
P. straminicaule, P. subincisum, and P. wilburii var.
longipedunculatum.
The coloration of leaf blades is highly variable
and largely dependent on the habitat, but some spe-
cies, such as P. roseospathum, P. scalarinerve, and
Р. tysonii, have very dark green blades while others
(e.g., P. lazorii) have lighter green blades. A few
species, such as P. chiriquense, have purple color-
ation on young ye blades but most lose this col-
oration in age. An exception is P. verrucosum,
which has adult ue generally purplish on the
lower surface. A few taxa such as P. ligulatum var.
heraclioanum (Fig. 274) and P. sagittifolium have
leaf blades purplish-spotted on the lower surface,
especially when young. Coloration of lower blade
surfaces is more often restricted to the major veins,
which are frequently reddish (along with the young
petiole) as, for example, in P. antonioanum.
Venation. Midribs. Leaf midribs of P. subg.
Philodendron are more variable in cross-sectional
shape than those of other aroid genera, such as
Stenospermation, Rhodospatha, Spathiphyllum, and
Monstera, yet not as Sra tt significant as in
the related genus Dieffenbac
Midribs on the upper = blade surface of
P. subg. Philodendron are highly variable, being
flattened or variously sunken or raised (sometimes
even in the same species). Fifty-six species have
adaxial midribs at least sometimes raised, 26 have
upper midribs at least sometimes sunken (only P.
cretosum has a deeply sunken midrib), and 54 have
adaxial midribs sometimes flat. Only 12 taxa, Р
antonioanum, P. aromaticum, P. albisuccus, P. co-
loradense, P. dressleri, P. granulare, P. pirrense, P.
purulhense, P. strictum, P. warszewiczii, P. wilburii
var. longipedunculatum, and P. zhuanum, are de-
scribed as having strictly flattened adaxial midribs.
Fifteen species are described as having only convex
midribs, and 15 are described as having only sunk-
en midribs. Only 2 species, P. morii and P. ni-
queanum, are described as having the adaxial mid-
ribs prominently raised.
The adaxial midrib is nearly always to some ex-
tent paler than the surface. In only 12 species are
the midrib and the blade concolorous. An addition-
al 8 species have midribs concolorous to slightly
er.
The lower (abaxial) midrib of Philodendron
blades exhibits less variability. All species have ad-
axial midribs to some extent raised. A few species,
e.g., P. crassispathum, have the abaxial midrib so
broadly convex as to appear nearly flat, but most
species have the midrib noticeably raised. The ab-
axial midrib is generally convex but often narrowly
rounded, thicker than broad, and sometimes bluntly
acute. Species with the abaxial midrib weakly
raised are rare in P. subg. Philodendron. Only sev-
en species have abaxial midribs at least sometimes
described as broadly convex or weakly raised: P.
anisotomum, P. crassispathum, P. fortunense, P. mi-
crostictum, P. platypetiolatum, P. purpureoviride,
and P. straminicaule.
more species have abaxial midribs more
prominently raised. In the largest category, 47 spe-
cies have abaxial midribs more or less convex, i.e.,
more or less hemispherical in cross section. Thirty-
six species have abaxial midribs described as
thicker than broad, narrowly convex, narrowly
rounded or bluntly acute at least part of the time.
A number of these species have abaxial midribs so
prominently raised as to be noticeably thicker than
broad: P. copense, P. cretosum, P. dolichophyllum,
P. hebetatum, P. heleniae, P. jodavisianum, P. pan-
amense, and P. roseospathum. Taxa that have ab-
axial midribs sometimes thicker than broad are P.
scalarinerve and P. wilburii var. longipedunculatum.
The most extreme type of raised adaxial midribs
354
Annals of the
Missouri Botanical Garden
is so raised as to appear to be a cylinder attached
tangentially to the surface of the leaf. This type of
midrib has been referred to as “round-raised.” Ex-
amples of species with such midribs are P. brun-
neicaule, P. dressleri, P. madronense, P. squamipe-
tiolatum, and P. radiatum. In cases where the
midrib is round-raised, the primary lateral veins are
also sometimes round-raised.
While the abaxial midrib is more likely than the
adaxial midrib to be concolorous with the rest of
the blade or darker than the remaining blade sur-
face, 45 species still are reported as having the
midrib paler than the surface. Twenty-four species
have the abaxial midrib described as darker than
the surface.
A number of species, including P. edenudatum,
P. grayumii, P. ligulatum var. ligulatum, and Р
sagittifolium, have one or both midribs maroon- or
purplish-spotted.
Primary lateral veins. The primary lateral veins
(referred to by some authors as “secondary veins”
are those that branch off the midrib and extend to
the margins usually without additional branching.
This use of primary lateral vein is consistent with
that of early aroid workers Engler, Krause, and So-
diro. Engler and later Krause referred to the pri-
mary lateral veins as “nervis primariis.” Sodiro also
referred to the primary lateral veins as “nervis la-
teralibus 1."
On larger cordate blades there is frequent
branching of the primary lateral veins in the lower
part of the blade where primary lateral veins are
widely spaced (Fig. 26) (see also Croat & Bunting,
979). The presence of these veins, referred to here
as “secondary veins,” is not commonly indicated in
this work, but in shape and aspect they are virtually
the same as the primary lateral veins (Fig. 24).
Both surfaces of the blade typically have primary
lateral veins similar to the associated midrib. Usu-
ally they are somewhat less prominently raised than
the midrib. The primary lateral veins on the upper
(adaxial) surface, while usually sunken, may be es-
sentially flat with the surface. Sometimes they are
raised but at the same time contained within a
broad or narrow valley so that the veins appear to
be sunken. Primary lateral veins may sometimes be
“quilted” (Figs. 23, 200, 361), i.e., with the veins
deeply sunken and with the intervening leaf tissue
of the blade that lies between the primary lateral
veins being broadly raised, making the surface ap-
pear like that of a plush quilt.
Primary lateral veins of P. subg. Philodendron
are rarely either absent or so inconspicuous as to
appear to be absent, such as in P. brewsterense,
سد
though they are sometimes not at all conspicuous,
as in P. microstictum (Fig. 289) ог P. sulcicaule (Fig.
403). The number of primary lateral veins varies
from 2 pairs in P. chirripoense to 25 pairs in P.
madronense. Most species have 3 to 6 pairs of pri-
mary lateral veins. These generally arise at an
acute angle with the midrib and, after extending
along or near the midrib, spread at an angle of
generally 40—80° toward the margins, generally
forming a broad arc in the process. The angle of
primary lateral veins for all species may be as little
as 5° to as much as 100°. When species of Central
American P. subg. Philodendron are divided into a
series of range categories based on the angle of
their primary lateral veins, more species (eight)
have primary lateral veins spreading at 50°-60° an-
gle or at 60°—70° (also eight species) than any other
group.
Basal veins. Primary lateral veins that contact the
plexus at the base of the blade and the apex of the
petiole are here referred to as “basal veins” (Fig.
22) (see also Croat & Bunting, 1979). While the
uppermost 1 or 2 pairs usually extend upward and
into the anterior lobe where they join the antemar-
ginal vein, most extend into the posterior lobes.
Generally, basal veins are best developed on plants
that have posterior lobes and are not present, or at
least are few in number, when the blade lacks pos-
terior lobes. Basal veins, like the primary lateral
veins, are typically paired, with usually an equal
number on either side of the midrib.
Seventy-three species (75 taxa) of P. subg. Philo-
dendron in Central America have basal veins. The
number of pairs of basal veins ranges from 1 to 15,
but only eight species (P. angustilobum, P. copense
(to 15), P. findens, P. grandipes, P. thalassicum, P.
pirrense, P. squamicaule, and P. strictum) have 10
or more pairs. Most species have 3—6 pairs of basal
veins, though up to 16 species may have only 2
pairs of basal veins. Only three species, namely P.
lentii, P. mexicanum, and P. scalarinerve, regularly
may have a solitary pair of basal veins.
Posterior ribs. The basal veins are generally to
some extent coalesced near their union with the
petiole. The union or coalescence of basal veins 18
here referred to as the “posterior rib” (Figs. 22, 25)
(see also Croat € Bunting, 1979). Sixty-three spe-
cies (65 taxa) have the basal veins at least in part
united and thus possess posterior ribs, while 34
species (38 taxa) have the basal veins free to the
base (or lack basal veins all together) and thus lack
posterior ribs. Both the number and nature of basal
veins and the length of the posterior rib are 80
characters for distinguishing taxa. The basal vems
Volume 84, Number 3
1997
Croat 355
Philodendron Subgenus Philodendron
may be completely coalesced for a distance d
22) or loosely coalesced. The posterior rib ma
prominently naked for a distance (Figs. 22, 169) or
barely (Figs. 26, 211) or not at all naked (Figs. 161,
163, 287). The basal veins are free or nearly so in
most species with rounded or cordulate leaf bases
(Fig. 20). Such species are: P. auriculatum, P. bak-
eri, P. correae, P. chirripoense, P. cretosum, P. doli-
chophyllum, P. granulare, P. immixtum, P. lentii, P.
ligulatum, P. pseudauriculatum, P. roseospathum, P.
scalarinerve, P. ubigantupense, P. utleyanum, and P.
wendlandii. Even species with short or poorly de-
veloped posterior lobes, such as P. annulatum, P.
clewellii, P. davidsonii, P. microstictum, P. niquean-
um, and P. platypetiolatum usually have the basal
veins free to the base. On the other hand, species
with well-developed posterior lobes usually have
the basal veins coalesced to some degree. Posterior
ribs range from less than 0.5 cm long (in P. stra-
minicaule) and 1 cm long (in P. angustilobum, P.
basii, and P. alticola) to more than 14 cm long in
P. gigas.
Another important taxonomic character is the ex-
tent to which the posterior rib is naked along the
sinus. In general, the longer the posterior rib the
more likely it is to be naked at least to some extent
along the sinus. However, P. tenue is exceptional in
having a well-developed posterior rib and being not
naked along the sinus. The degree to which the
posterior rib is naked along the sinus varies from
as little as 0.5 cm in P. straminicaule to 6 cm in P.
squamicaule. Examples of species with prominently
naked posterior ribs are: P. albisuccus, P. alticola,
P. aromaticum, P. basii, P. brunneicaule, P. copense,
P. dodsonii, P. dressleri, P. findens, P. hebetatum, P.
knappiae, P. panamense, P. purulhense, P. strictum,
P. thalassicum, and P. verapazense.
Lesser order veins. Between successive primary
lateral veins there are possibilities for two addi-
tional orders of veins. Frequently present are in-
аазы ал. primary lateral veins here called “inter-
ary veins” (Figs. 24, 26) (see also Croat &
E 1979). These veins, while decidedly less
conspicuous than the primary lateral veins, are
nevertheless too prominent to be classified as the
smallest-order veins. To qualify as an interprimary,
the vein must extend continuously from the midrib
to very near the margin without major branching.
Generally there is no more than one pair of inter-
primary veins between alternate primary lateral
veins. They are akin to the primary lateral veins in
all aspects except for their reduced size. Like the
primary lateral veins they may bear minor veins,
which may form all along their margins.
The smallest-order veins are here referred to as
"minor veins" (Fig. 24) and may be close, fine, and
conspicuous as in P. sulcicaule, P. tripartitum, and
P. radiatum, to thick, well-spaced, and inconspic-
uous in P gigas, P. granulare, P. grayumii, and P.
ligulatum. The minor veins are not all equally dis-
tinct and sometimes, as in P. dominicalense, the
minor veins are alternately weakly visible and
strongly visible.
The minor veins may arise from the midrib or
from the primary lateral and interprimary veins, but
in either case they form a generally close, uniform,
and parallel array, which extends without interrup-
tion to near the margin of the blade. In most species
the minor veins arise from both the midrib and the
primary lateral veins, but some species have the
minor veins arising from only the midrib. A total of
77 species (81 taxa) have the minor veins arising
from both the midrib and the primary lateral veins
(Fig. 26). In such cases the minor veins are not
always equally arising from one of the two entities
but may, as in the case of P. brenesii, P. davidsonii
subsp. davidsonii, P. ferrugineum, and P. triparti-
tum, be more heavily arising from the midrib rather
than the primary lateral veins. In P. auriculatum,
P. glanduliferum, P. lentii, and P. ligulatum the mi-
nor veins arising from the primary lateral veins are
many fewer than those arising from the midrib. In
P. heleniae the primary lateral veins are only rarely
arising from the midri
In another variation of this venation type, some
species, while having minor veins arising from the
midrib as well as both adjacent primary lateral
veins, have considerably more veins arising from
the distal primary lateral vein than from the prox-
imal vein.
A total of 24 species have the minor veins arising
from only the midrib, and in this case they course
along the primary lateral veins but do not join with
it. Many of the species that have the minor veins
arising only from the midrib are species with oblong
blades, such as: P. bakeri, P. cretosum, P. dolicho-
phyllum, P. granulare, P. heleniae, P. roseospathum,
P. ubigantupense, P. utleyanum, and P. wendlandii.
However, this group also has blades ovate or nearly
so as in P. brewsterense, P. chirripoense, P. cotobru-
sense, P. crassispathum, P. folsomii, P. knappiae, P.
microstictum, P. niqueanum, P. sulcicaule, and P.
verapazense. Interestingly, P. anisotomum, with
deeply 3-lobed leaf blades, also has the minor veins
arising from only the midrib, whereas P. tripartitum,
a similar species with 3-lobed blades, has the mi-
nor veins arising from both the midrib and the pri-
mary lateral veins.
Philodendron dressleri, a species with deeply di-
356
Annals of the
Missouri Botanical Garden
vided leaves, has a more complex venation pattern.
Although the minor veins arise from both the mid-
rib and the primary lateral veins, they also arise
from short secondary veins that regularly branch off
the primary lateral veins. In addition, the minor
veins that arise from the midrib are considerably
fewer in number and weaker than in most species
with this venation pattern. Generally, the confluent
minor veins that arise from the primary lateral veins
and make a broad sweep before continuing to the
margin leave little area for the minor veins, which
arise from the midrib. The latter tend to merge im-
perceptibly with those from the primary lateral
veins. A similar pattern with weak midrib-borne
minor veins is present with P. basii.
At or very near the margin both the primary lat-
eral veins and the minor veins generally turn sharp-
ly toward the apex of the blade. The minor veins
join with other minor veins and with the primary
veins before finally merging into an inconspicuous
and somewhat opaque marginal plexus. This narrow
band is usually chlorophyllous and apparently
veinless. Frequently the outer margin of this chlo-
rophyllous band is a hyaline edge, which is color-
less and typically revolute.
The minor veins are sometimes noticeably inter-
connected by inconspicuous to conspicuous veins,
referred to here as “cross-veins” (Fig. 26). Gener-
ally the cross-veins are markedly perpendicular to
consecutive minor veins where they are sufficiently
prominent to be noticeable, but in some cases the
cross-veins cross transversely from one minor vein
to the next. While P. scalarinerve has cross-veins
so prominent, even on fresh material, as to be easily
visible, other species such as P. chiriquense and P.
copense have cross-veins easily visible only when
the blades are dry.
Secretory ducts and other secretory tissues are
frequently present on Central American members
of P. subg. Philodendron. The contents of the se-
cretory ducts are either latex or tanniniferous com-
pounds (Solereder & Meyer, 1928). No thorough
survey has been made of the nature of the secretory
canals in Central American species, so it is not
always apparent whether these structures are resin
canals or secretory ducts. Secretory canals (also re-
ferred to as secretory files) in Philodendron are al-
ways non-anastomosing and consist of a linear se-
quence of secretion cells, each separated from the
next by cell walls (Solereder & Meyer, 1928). On
fresh and dried leaves these can usually be rec-
ognized by being darker, usually blackened, and in
being intermittent rather than continuous as is gen-
erally true of veins. While the distribution of se-
cretory ducts in Philodendron may be more com-
mon than is apparent from surface examination, not
all species exhibit the secretory ducts clearly. Thus
the presence or absence of distinct secretory ducts
can be useful taxonomically. They are distinctly
visible on P. alticola, P. cotonense, P. grayumii, P.
heleniae, and P. zhuanum, but obscurely visible on
P. antonioanum, and P. bakeri. They are clearly
visible on P. schottianum, somewhat visible on P.
llanense, and obscure on P. findens, even though
these three species have very similar leaf blades.
While perhaps no more reliable than blade
shape, blade size, or other features, blade surfaces
at a magnification of 10X or higher often reveal
another suite of characters, which often yields an-
other degree of confidence (or forewarns of mis-
identification when making determinations. The
surface between the minor veins is frequently
marked by pale sub-surface granulations (perhaps
indicating the presence of druse crystals), short,
pale lineations, gland-like punctiform markings,
reddish or brownish speckling, and also what might
be referred to as "stitching," pale intermittent short
lines appearing on the surface of the blade as
though the blade was loosely sewn with a needle
and thread. Though this phenomenon is much more
common in P. subg. Pteromischum, it is also exhib-
ited in P. subg. Philodendron.
MORPHOLOGY OF REPRODUCTIVE STRUCTURES
INFLORESCENCES
When a plant of P. subg. Philodendron flowers,
the stem apex is terminated by an inflorescence (or
frequently an aborted inflorescence primordium)
(Ritterbusch, 1971; Blanc, 1977c; Madison, 1978)
and new growth is from a bud in the axil of the
penultimate leaf. This new growth pushes aside the
newly developing leaf and the inflorescence that 18
developing in из axil (Ray, 1987a). When the in-
florescences abort, the petiole sheath remains quite
small and unexpanded, but when the inflorescences
develop, the sheath may be much larger. Philoden-
dron inflorescences are sympodial, with each sym-
podium consisting of 1-10 (to 11 in South America)
inflorescences arising typically in what appears to
be the leaf axil of usually one of the upper leaves
(but almost never in the uppermost leaf axil). Often
inflorescences emerge from the mass of cataphyll
fibers, enabling the inflorescence to remain moist
(Figs. 63, 115, 188, 371, 430). The individual in-
florescences are subtended by and enclosed in à
series of moderately coriaceous, whitish to pinkish,
2-ribbed structures (Fig. 195, P. gigas, Fig. 428. Р.
tysonii) called bracteoles (Ray, 1987a). These have
elsewhere been referred to as prophylls (Usher,
AA
Volume 84, Number 3
1997
Croat 357
Philodendron Subgenus Philodendron
1966; Jackson, 1971). At anthesis the bracteoles
are typically deciduous, although sometimes they
persist for a time simply because they are held so
tightly by the developing infructescences.
Typically the inflorescences are borne in the
erect or semi-erect position regardless of the posi-
tion of the stem. Even when the inflorescence 18
borne on the end of a pendent stem, the stem usu-
ally curves upright so that the opening inflores-
сепсе is erect at anthesis (Fig. 273, P. ligulatum
var. ovatum; Fig. 269, P. ligulatum var. ligulatum).
The number of inflorescences per ах! is taxo-
nomically significant. More species (38) have a sol-
itary inflorescence per axil (or are believed to have
a solitary inflorescence) than any other combina-
tion. Many others may be found with only a single
inflorescence if they are seen in the early stages of
flowering or if inflorescences have aborted owing to
lack of pollination, but plants frequently have two
or more inflorescences per axil later in the flower-
ing season. Careful dissection of the leaf axil usu-
ally can provide evidence of the peduncular scar
indicating a lost inflorescence. Taxa known to usu-
ally have only a single inflorescence per axil are:
P. albisuccus, P. anisotomum, P. aromaticum, P.
bakeri, P. basii, P. breedlovei, P. brevispathum, Р.
brewsterense, P. brunneicaule, P. chirripoense, P. cor-
reae, P. cotonense (rarely 2), P. crassispathum, P.
dressleri, P. dwyeri, P. edenudatum, P. folsomii, P.
glanduliferum, P. granulare, P. hammelii, P. hed-
eraceum, P. immixtum, P. jacquinii, P. jefense, P.
knappiae, P. ligulatum var. heraclioanum, P. ligu-
latum var. ligulatum (sometimes 2, rarely to 5), P.
ligulatum var. ovatum, P. madronense, P. mexican-
um, P. microstictum, P. platypetiolatum (rarely 2),
P. purpureoviride, P. purulhense, P. radiatum var.
pseudoradiatum, P. squamipetiolatum, P. utleyan-
um, P. verapazense, and P. warszewiczii (rarely to 3).
Most species with a single inflorescence per axil
are either vines or appressed-climbing hemiepiphy-
tes with internodes longer than broa
e number of upper axils bearing inflores-
cences is also variable. Some species, e.g., P. lentii,
rarely have more than one leaf axil bearing an in-
florescence, while other species, e.g., P. annulatum
and P. fragrantissimum, bear inflorescences in two
or more of the closely spaced upper internodes,
making it appear that the plants have a large num-
ber of inflorescences.
Species that regularly have up to four or more
inflorescences per axil are: P. copense (to 6), P. co-
tobrusense (to 5), P. clewellii (to б), P. dodsonii (2—
5), P. ferrugineum (to 6), P. fortunense, P. gigas (to
7), P. grayumii (to 5), P. heleniae (to 10), P. jodav-
isianum (2-6), P. panamense (to 6), P. ритепзе, Р
squamicaule (to 5), Р. ѕоиѕае, Р. tenue, Р. tysonii (to
5), апа P. verrucosum.
Species with буе or more inflorescences per axil
are rare, totaling only eleven. Only two species, Р.
gigas and P. heleniae, have up to seven inflores-
cences per axil.
PEDUNCLES
Typically the peduncle is terete but it may be
somewhat flattened laterally, especially when there
are clusters of inflorescences per axil. The pedun-
cle is usually whitish or pale green at the base
where it is often hidden by the leaf sheath. The
remaining portion of the peduncle is usually dark
to medium green with the surface often shortly
pale-lineate like the petioles. The peduncle is often
coarsely pale-ribbed near the apex, as in P. copense,
P. dodsonii, and P. findens. Rarely is the peduncle
pinkish red or reddish as in P. malesevichiae, P.
schottianum, and P. roseospathum (or sometimes in
P. sulcicaule) or tinged with red or purple as in P.
chiriquense, P. davidsonii subsp. bocatoranum, P.
heleniae, P. grandipes, P. purpureoviride, and P. ver-
rucosum.
Most peduncles, like other parts of the average
Philodendron, are glabrous, but some species have
peduncles conspicuously covered with trichome-
like glands, e.g., P squamicaule (Fig. 384), P.
squamipetiolatum (Fig. 388), and P. verrucosum
(Fig. 440).
The peduncle is usually much narrower than the
spathe, commonly no more that % to М the width
of the spathe tube, and is almost always broadened
distally, merging almost imperceptibly with the
spathe tube. Usually the color distinction is also
gradual, but in some cases, e.g., P. annulatum, P.
dodsonii, and P. dolichophyllum, there is an abrupt
transition between the green peduncle and the col-
ored spathe. In P. ligulatum there is a purple ring
at the apex of the petiole, much like that at the
apex of the petiole (Fig. 276).
The length of the peduncle, especially relative
to the length of the spathe, may be important tax-
onomically. Some species, e.g., P. antonioanum, P.
crassispathum, P. davidsonii, P. ferrugineum, P. fin-
dens, P. malesevichiae, P. purulhense, P. radiatum,
P. schottianum, and P. zhuanum, have very short
peduncles relative to the length of the spathe. In
contrast, other taxa, e.g., P. advena, P. angustilob-
um, P. dodsonii, P. hebetatum, P. heleniae, P. im-
mixtum, P. mexicanum, P. pterotum, P. rothschu-
hianum, and P. wilburii var. wilburii, have
peduncles as long as or longer than the spathes.
Peduncles range in length from 1 to 25 cm, with
Annals of the
Missouri Botanical Garden
four taxa, P. microstictum, P. pseudauriculatum, P.
wilburii var. longipedunculatum, and P. verrucosum,
all having peduncles that attain the maximum
length of 25 cm. Several additional species have
peduncles that may be more than 20 cm long.
These include P. anisotomum, P. brunneicaule, P.
panamense, P. pseudauriculatum, P. rothschuhian-
um, and P. scalarinerve.
Many species of P. subg. Philodendron have pe-
duncles more than 10 cm long. In addition to those
mentioned above, 37 Central American species
have peduncles more than 10 cm long, while 38
taxa (35 species) have peduncles less than 10 cm
long. Sixteen species have petioles that may be less
than 3 cm long, but only P. knappiae has a pedun-
cle which does not exceed 3 cm in length.
SPATHES
The spathe of P. subg. Philodendron is highly
variable in many regards. The spathe of Philoden-
dron is typically coriaceous and constructed in
such a way that the spathe may open and reclose
without major alteration in its shape. The margins
are invariably much thinner, with the apical portion
of the spathe usually tightly rolled and frequently
acuminate (Fig. 178, P. findens) and the basal por-
tion more conspicuously convolute. Because the
margins of the base are more conspicuously over-
lapped, when the spathe opens the lower portion
remains convolute (Fig. 70, P. auriculatum) where-
as the apical portion separates to very near the tip.
Some species have spathes that open more fully so
that the opening is almost rounded (Fig. 298, P.
panamense) or broadly elliptic (Fig. 288, P. mexi-
canum). In such cases more of the inner tube sur-
face and the pistillate portion of the spadix are ex-
posed. On the other hand, some species have
spathes that open only slightly so the opening is
merely elliptic with the pistillate portion of spadix
not visible without peering down through the top of
the opening (Fig. 63, P. antonioanum).
Species vary greatly in the extent to which the
spathe remains convolute at the base during anthe-
sis. Spathes of some species, e.g., P. angustilobum
(Fig. 51) and P. mexicanum (Fig. 288), unfold to
very near the base. Those of other species, e.g., Р.
antonioanum (Fig. 63), do not open very broadly.
The spathe tubes of P. auriculatum (Fig. 70), P.
panamense (Fig. 298), and P. wendlandii (Fig. 451)
remain convolute only in the lower half. The spathe
of P. davidsonii (Figs. 134, 136) may be convolute
only in the lower % to % in P. squamicaule. That
of P. hebetatum (Fig. 226) is convolute to the upper
% and the spathe tubes of P. aromaticum (Fig. 68),
P. brenesii (Fig. 89), P. copense (Fig. 110), P. sca-
larinerve (Fig. 140), P. findens (Fig. 177), P. roths-
chuhianum (Fig. 340), P. straminicaule (Fig. 360),
and P. schottianum (Fig. 371) remain essentially
closed to the apex of the tube during anthesis.
Mayo (1986) presented a classification of inflo-
rescence types based principally on South Ameri-
can Philodendron species. Eleven inflorescence
types were characterized, and a key was prepared
to separate them. The characters chosen for use in
the key were: (1) presence or absence of resin ducts
in the inner surface of the spathe; (2) solitary vs.
two or more inflorescences per axil; (3) relative
length of the sterile staminate portion of the spadix
(equal to or longer than the staminate portion vs.
shorter than staminate zone); (4) presence or ab-
sence of colorful contrasts on inner surface of
spathe tube; (5) prominently constricted vs. weakly
constricted spathes; (6) presence or absence of res-
in ducts in the spathe; (7) presence of resin ducts
in the sterile staminate portion of spadix vs. with
resin ducts in either the fertile staminate portion or
in both sterile and fertile staminate portions; (8)
presence of non-functional resin canals in the
spathe vs. resin canals lacking in the spathe (usu-
ally correlated with contrasting spathe colors); and
(9) presence or absence of a terminal sterile zone
on the spadix.
While I believe that this classification may be
somewhat artificial, it demonstrates many evolu-
tionarily important morphological features and
shows the complexity of the Philodendron spathe.
Species of P. subg. Philodendron have the spathe
tube relatively well demarcated from the blade ei-
ther by being constricted at the apex or by its con-
trasting color. Some species, e.g., Р. crassispathum
(Fig. 124), lack any constriction above the tube,
thus the spathe is more or less elliptic. Species that
have a weakly constricted spathe include P. advena
(Fig. 40), P. alticola (Fig. 45), P. breedlovei (Fig.
85), P. purulhense (Fig. 323), P. schottianum (Fig.
368), and P. antonioanum (Fig. 63). Despite these
exceptions, most members of P. subg. Philodendron
have spathes to some extent constricted. The sub-
genera Meconostigma and Pteromischum, on the
other hand, have spathes not likely to be constrict-
ed or are only weakly constricted (Mayo. 1986).
Spathe length is relatively variable depending or
the age of the plant and the stage of development.
Elongation of the spathe is typically more or less
arrested after anthesis, but sometimes a consider
able amount of elongation takes place in both the
peduncle and spathe after anthesis, presumably to
accommodate the considerable expansion of the de-
veloping fruits. Spathe length varies from as little
_—- өтт ——ө ө—өөї—- —
Volume 84, Number 3
1997
Croat 359
Philodendron Subgenus Philodendron
as 4.5 cm in P. clewellii and 4.8 cm in P. heleniae
to 29 cm in Р. pterotum and 30 cm in P. warszew-
iczii. Relatively few species have spathes that ever
exceed 25 cm in length even when in fruit.
Relatively few species have spathes less than 10
cm long. These are: P. brewsterense, P. chirripoense,
P. clewellii, P. dwyeri, P. knappiae, P. roseospathum
var. angustilaminatum, and P. ubigantupense.
The shape of the spathe and the relative dispo-
sition of the spadix at anthesis are taxonomically
significant, but seldom described. This is because
many species are still poorly known at anthesis, no
doubt due to the fact that anthesis in Philodendron
lasts only one or two days for each inflorescence.
One of the features that determines the shape of
the spathe at anthesis is the degree to which the
spathe opens. Spathes of some species, such as Р
aromaticum, P. copense, P. dodsonii, P. findens, P.
fragrantissimum, P. grandipes, P. hebetatum, P. jo-
davisianum, P. straminicaule, and P. strictum, open
along the blade portion (see discussion above),
leaving an oblong-elliptic opening (Fig. 110, P. co-
pense; Fig. 140, P. dodsonii; Fig. 378, P. smithü).
In these examples the spadix is either included
within the spathe or is barely exserted, stiffly erect
yet held slightly in front of the top edge of the
spathe. On the other hand, spathes of some species,
such as P. angustilobum, P. mexicanum, and P. tri-
partitum (Fig. 423), may open so broadly as to ex-
pose much of the pistillate portion of spadix. The
pistillate portion of the spadix is usually for the
most part obscured by the prominently convolute
tubular portion of the spathe, as in P. antonioanum,
P. aromaticum, P. copense, and P. dodsonii. Those
species with spathes opening more broadly may
also have the spadix protruding forward somewhat
out of the spathe (see section on Spadix).
The definition of the spathe tube is somewhat
imprecise, in terms of both morphology and color,
but the spathe tube may be defined as the lower
portion of the spathe (that portion which covers the
pistillate portion of the spadix). In reality it usually
extends somewhat above the pistillate portion to in-
clude the lowermost or sterile zone of the staminate
portion of the spadix. In cases where the spathe
has an obvious constriction it is that portion below
the constriction (the balance being the spathe
blade) that is referred to as the spathe tube. Though
the ending of the spathe tube is imprecise, it gen-
erally terminates where the greatest constriction of
the staminate portion of the spadix occurs, usually
immediately above the sterile staminate portion of
the spadix. Functionally, the constriction of the
spathe, coinciding with the narrowest portion of the
staminate portion of the spadix, prevents much of
the pollen from the fertile upper portion of the spa-
dix from falling directly into the chamber of the
pistillate portion of the spathe. This probably serves
more to prevent wastage of the pollen (by contain-
ing the pollen in a place where it can most easily
be carried away by beetles) than it does to prevent
self-pollination, since most species are so markedly
protogynous that no self-pollination is possible, i.e.,
the pistils are probably no longer receptive when
the pollen is shed. Testing for pistil receptivity with
the use of peroxidase paper, Grayum (1996) found
stigmas receptive throughout the period of staminal
dehiscence in P. subg. Pteromischum, but numer-
ous attempts at self-pollinations of members of P.
subg. Philodendron by me and others failed to pro-
duce berries and instead resulted in aborted inflo-
rescences. Not surprisingly those species, e.g., P.
advena, P. crassispathum (Fig. 124), and P. purul-
hense, that have spathes scarcely constricted above
the tube also have spadices not markedly narrowed
above the sterile staminate flowers. It is not known
whether the pollination behaviors of these species
er.
For Philodendron species with a prominent
spathe constriction, the partially closed spathe dur-
ing staminal anthesis, coupled with the swollen
area of the staminate portion of the spadix imme-
diately below it, provides a “pollen well” through
which the beetles must squeeze on their departure.
This probably ensures a greater coverage of pollen
for the departing beetle pollinators.
Spathe tube shape and length are not particularly
useful taxonomically. Tubes range in shape from
virtually oblong (e.g., in P. heleniae, P. ligulatum,
and Р. tripartitum), to oblong-elliptic (e.g., in P
aromaticum, P. copense, P. dodsonii, P. findens, P.
gigas, P. hebetatum, and P. pseudauriculatum) to
elliptic (e.g., P. rothschuhianum and P. smithii), and
range from only 2 cm long in P. clewellii to 14 cm
long in P. davidsonii and P. warszewiczii. Species
with spathe tubes longer than 10 cm are: P. auri-
culatum, P. ferrugineum, P. schottianum, P. sousae,
and P. tysonii. Seventeen species have spathe tubes
less than 5 cm
Even though ЊЕ A tube often remains to-
tally closed at anthesis it is, nonetheless, still some-
what expanded owing to the separation caused by
the flaring of the spathe blade. Generally this pro-
vides considerable room around the spadix to ac-
commodate the pollinating beetles (Figs. 100, 128,
242, 268). At anthesis the spathe tube is consid-
erably more voluminous than before anthesis.
Spathe color. Spathe coloration is generally taxo-
nomically significant though variable depending on
360
Annals of the
Missouri Botanical Garden
the age of the inflorescence. Before anthesis
spathes are frequently green on the outside, but
reddish coloration often develops well in advance
of anthesis. Coloration of the spathe tube and blade
often differs, with the spathe tube more commonly
various shades of green and the spathe blade more
commonly white, whitish, or whitish green. Even
when the spathe is green throughout, the spathe
blade is typically paler green than the tube. The
tube portion of the spathe is commonly colored or
tinged with red, maroon, or purplish violet (some-
times with other shades of red, orange, or purple)
on one or both surfaces. Strong color contrasts (oth-
er than merely green vs. white), so common in
Philodendron, are absent in other ostensibly related
genera, namely Furtadoa, Homalomena, and Anu-
bias (based on the Philodendron Alliance of Mayo,
Bogner & Boyce, 1995) and also Cercestis and Cul-
casia (included in the Philodendron Alliance of
Grayum, 1990). This would also be true of other
presumed relatives including Dieffenbachia, Mon-
trichardia, Nephthytis, Peltandra, Typhonodorum,
and even Zantedeschia.
Generally if the spathe is colored on the outer
surface it is also colored on the inner surface (but
the reverse is not true). The inner surface is typi-
cally much more intensely colored than the outer
surface. The coloration of the inner surface of the
spathe, though commonly more or less restricted to
the tube, may extend well onto the blade, some-
times even to very near the apex, such as in P
breedlovei, P. davidsonii, P. mexicanum, and some-
times P. sagittifolium. In some cases, such as Р
findens and P. schottianum, the color is merely
weakly diffused onto the inside of the blade. At
other times only the very base of the tube is colored
inside, such as in P. ligulatum and P. hederaceum.
The spathe blade is commonly more extensively
colored on the outer than on the inner surface, al-
though blade coloration is highly variable, both in-
ter- and intraspecifically. Nevertheless, most spe-
cies have spathe blades some shade of green or
white on the outer surface, often tinged with red,
pink, purple, or yellow. A few species, such as P.
sagittifolium and P. ferrugineum (Fig. 167), have
purplish spots or blotches throughout much of the
exterior of the spathe surface. Although the external
coloration of the spathe is usually restricted to the
tube portion, it often extends onto the blade, a sit-
uation that happens more frequently on the outer
surface of the spathe than on the inner surface.
Species that have coloration extending well above
the spathe tube are: P. antonioanum, P. breedlovei,
P. dolichophyllum, P. subincisum, and P. roseospa-
thum. Sometimes the coloration of the spathe may
be restricted to the tube, but only near the margins
of the open edge such as in P. grayumii and P.
malesevichiae.
While some species, e.g., P. alticola, P. heleniae,
and P. findens, show little or no distinction in the
coloration of the spathe tube and spathe blade, oth-
er species have distinctly different colors. In some
cases, e.g., P. antonioanum, P. cretosum, P. edenu-
datum, P. findens, P. grandipes, P. lazorii, P. ligu-
latum, P. llanense, and P. roseospathum, the tran-
sition from one color to the next is very gradual. In
other cases, such as P. hebetatum, P. immixtum, P.
jodavisianum, P. panamense, P. pterotum, P. stric-
tum, P. radiatum, P. rothschuhianum, P. scalariner-
ve, P. smithii, P. tenue, P. wendlandii, and P. zhuan-
um, the transition is less gradual but by no means
abrupt. Still other species, e.g., P. ligulatum var.
ovatum, P. dodsonii, P. gigas, P. strictum, and P.
fragrantissimum, show a distinct and abrupt tran-
sition in the spathe tube and blade colors.
The biological significance of the frequently
darker colors on the inside of the spathe tube in
contrast to the paler colors of the spathe blade is
uncertain. It seems unlikely that either the con-
trasting colors or the dark color of the spathe tube
act in attracting pollinators. The presumed polli-
nators, dynastine scarab beetles (see section on
“Pollination Biology”), are believed to orient more
by smell than sight (Faegri & van der Pijl, 1979;
Gottsberger € Silberbauer-Gottsberger, 1991).
Moreover, their normal arrival time, near d "m
would probably preclude their seeing contrasting
colors in any event. In addition, the dark coloration
is usually restricted to the spathe tube and often
not even visible in good light from near the mouth
of an open spathe. It is more likely that the dark-
colored spathe tube acts to encourage these cre-
puscular beetles to stay for an extended period of
time. Mayo (1986) pointed out that, since neither
P. subg. Pteromischum nor P. sect. Meconostigma
have color contrasts in the spathe blade and tube,
P. subg. Philodendron may have different pollina-
tors. Although not enough pollinators have yet been
identified to confirm this possibility, there seems 10
be little evidence in favor of Mayo's hypothesis (see
section on “Pollination Biology"). Some beetle spe
cies, such as Erioscelis proba Sharp, are known t°
visit species of both P. subg. Philodendron and P
subg. Pteromischum (Grayum, 1996) (see also Table
3, “Pollinators of P. subg. Philodendron”). . of
Lenticel-like structures on both the exterior
the spathe tube and the peduncle in some |
dendron species secrete large droplets of a viscous
somewhat sweet substance. Mayo (1986) has shown
that these may consist of clusters of stomata, аһ
PASA
————————————————————————
Volume 84, Number 3
1997
Croat 361
Philodendron Subgenus Philodendron
Table 3. Pollinators of Philodendron subg. Philodendron.
Plant species Voucher
Beetle species
P. anisotomum
P. brenesii Croat 35519
P. brevispathum
P. callosum No voucher
P. correae Croat 66653
P. grandipes Croat 76594
nez 6
nez 6
Croat 43289
P. grayumii Croat 74840
Croat 74840
P. jodavisianum
Croat 35950
P. ptarianum Ramírez 1163
P. pterotum
Croat 10903
P. radiatum
P. rothschuhianum
(Young, 1987)
P. sagittifolium
P. schottianum
P. tripartitum
Thompson 4636
Croat 36110
P. tysonii Croat 66711
*Cyclocephala amblyopsis Bates
*Erioscelis columbica Endródi
Cyclocephala nigerrima Bates
Erioscelis proba Sha
yclocephala rustica (Olivier)
Cyclocephala conspicua Sharp
*Cyclocephala gravis Bates
Cyclocephala sexpunctata Castelnau
Erioscelis columbica Endródi
Cyclocephala rubescens Bates
Cyclocephala sexpunctata Castelnau
Cyclocephala ligyrina Bates
Cyclocephala mafaffa Burmeister
*Erioscelis columbica Endródi
Cyclocephala rustica (Olivier)
*Cyclocephala ampliata Bates
*Cyclocephala cel Bates
Cyclocephala ligyrina
*Cyclocephala ues awe
*Cyclocephala ampliata Bates
*Erioscelis columbica Endródi
*Cyclocephala amblyopsis Bates
*Cyclocephala amblyopsis Bates
Cyclocephala kaszabi Endródi
*Erioscelis columbica Endródi
Cyclocephala sexpunctata Castelnau
Cyclocephala melane
*Cyclocephala amblyopsis Bates
*Cyclocephala kaszabi Endródi
*Erioscelis columbica Endródi
Cyclocephala nigerrima Bates
*Entries were provided by Helen Young, Barnard Coll ege, New York, and were based on unpublished observations
made during 1984—1985 at La Selva (O.T.S. Field Station) in Costa Rica (with the assistance of George Schatz). Entries
without an ا
pointed out their possible ecological significance as
extrafloral nectaries.
The secretion of resin from the inner surface of
the spathe is apparently unique, in Araceae, to
Philodendron (Mayo, 1991). Mayo (1986) enumer-
ated four different types of inflorescence resin ca-
nals, at least two of which occur among Central
American Philodendron. One type, found in P. tri-
partitum, has large-diameter resin canals imbedded
just beneath the epidermis on the inner surface of
the spathe. Resin is secreted directly onto the sur-
сона Another type, represented by P. зтићи, lacks
n canals on the spathe but instead has resin
~ in the staminate zone of the spadix.
were based on determinations made by John Rawlins at Carnegie Museum in ‘ti ar Collections
by S. Thompson were contributed independently to Rawlins.
The two other types of resin canals pertain to P.
subg. Meconostigma and P. subg. Pteromischum. In
the former, exemplified by P. bipinnatifidum Schott
ex Endl., the resin canals аге “J”-shaped and tan-
gential, arising in the aerenchyma of the spathe and
extending to the surface. In the latter, exemplified
by Р. sonderianum Schott, the resin canals аге
evenly distributed between the aerenchyma and
epidermis and are parallel to the surface of the epi-
dermis rather than arising in a J-shaped fashion.
Resin canals usually are present in the proximal
% to % of the spathe blade and sometimes also the
distil portion of the tube. In P. subg. Philodendron,
these resin canals are generally reddish, red-pur-
362
Annals of the
Missouri Botanical Garden
ple, or orange to brownish and generally somewhat
intermittent, as in P. sulcicaule, though sometimes
continuous, as in P. immixtum, P. mexicanum, P.
sagittifolium, and P. wilburii. They are always ori-
ented vertically, paralleling the veins of the spathe.
The resin canals actively secrete a resinous liquid
at anthesis that wells up onto the surface of the
spathe and sometimes runs down the spathe below
the resin ducts. The resin is generally very sticky
and probably functions in causing pollen, some-
times itself not particularly sticky, to adhere to the
bodies of the beetle pollinators as they leave the
spathe.
SPADIX
Philodendron species are monoecious, with the
spadix bearing naked unisexual flowers. The flow-
ers lack a perigon and are closely arranged on a
more or less cylindrical spadix, with the staminate
flowers comprising the upper %-Y, of the spadix
and the pistillate flowers comprising the remaining
basal portion. The average spadix for P subg.
Philodendron in Central America has the staminate
portion 2.2 times longer than the pistillate.
The spadix is usually contained for the most part
inside the spathe at anthesis (see section above on
the Spathe), although in the majority of species the
spadix is weakly exserted in front of the spathe.
The majority of plants at anthesis have their
spathes erect or tipped slightly forward so they can
provide shelter to spadices from water dripping
straight overhead.
Spadices of some species, especially those with
broadly opened spathes (e.g., P. angustilobum (Fig.
51), P. ligulatum (Fig. 266), P. mexicanum, and P.
tripartitum) protrude prominently forward at anthe-
sis. Other species whose spathes remain convolute
to about the middle may also have spadices that
protrude prominently forward; these are: Р. auri-
culatum, P. davidsonii, P. grayumii, P. hebetatum,
P. heleniae (Fig. 229), P. jodavisianum, P. ligula-
tum, P. llanense (Fig. 268), and P. panamense (Fig.
299)
The spadix of Philodendron is usually only
slightly shorter than the spathe. The average length
of the spathe is 16.5 cm, while the average spadix
is 15.8 ст long. Thus on average, the spathe is
nearly 1 cm longer than the spadix. This difference
may be as little as a few millimeters or as much as
The general morphology of the spadix of Philo-
dendron, though highly variable in detail, is more
or less the same for all species in Central America.
The pistillate portion is usually pale green to green-
ish white and obliquely attached to the spathe, and
sometimes markedly stipitate at the base. The flo-
riferous part of the spadix is thus longer on the
adaxial (front) than on the abaxial (back) surface
(Fig. 40). The differences in the length of the ad-
axial and abaxial surfaces of the pistillate portion
of the spadix depend on the angle at which it is
attached to the spathe. These are referred to as
"front side" and *back side" of the spadix in the
descriptions presented in this revision.
The staminate portion of the spadix characteris-
tically has a swollen sterile section at the base.
Typically the staminate portion is broadest at the
sterile section and gradually constricted just above
this. Above the constriction the spadix is fertile and
is gradually broadened usually to a point about %
of its length, then narrowed gradually or abruptly
to the tip.
The staminate portion is usually uniformly white
to creamy white on the outside, although sometimes
with the axis pinkish. Immature spadices are usu-
ally green, though they become white well before
anthesis. The sterile section of the staminate por-
tion is commonly a different color than the fertile
section. Often it is more nearly white at an earlier
stage of development when the fertile flowers are
typically green. At anthesis the sterile staminate
portion is more likely to be tinged with yellow or
tan, reflecting the higher concentration of oil in
these flowers than in the fertile flowers (see below).
However, sometimes at anthesis the sterile and the
fertile staminate flowers are identical in color. The
sterile staminate portion frequently can be distin-
guished easily after anthesis with the sterile portion
maintaining a white color (Fig. 28) while the fertile
portion turns grayish or brownish. The sterile sta-
minate portion can usually be distinguished even
in these cases by the difference in the size of the
spadix (usually larger than the adjacent fertile sta-
minate portion), by the usually larger and more 1t-
regularly shaped flowers, and because the sterile
staminate section typically ends just before the nar-
rowly constricted portion of the fertile staminate
portion (which itself coincides with the constricted
portion of the spathe). The sterile staminate flowers
are often more easily discernible on dried spect
mens than they are on live ones since they tend to
dry a different color, usually somewhat more brown-
-
2
The much higher oil content of the sterile sta-
minate flowers was reported by Pohl (1932), and it
is probably owing to this high oil content that the
sterile staminate flowers are often eaten by a
beetle pollinators (Fig. 29). A high concentration %
а II metn rmm
Volume 84, Number 3
1997
roat 363
Philodendron Subgenus Philodendron
lipids in the mitochondria of the staminodial cells
has been documented (Walker et al., 1983).
The fertile staminate portion of the spadix is typ-
ically more or less clavate and bluntly tapered to-
ward the apex. In a few species, such as P. advena,
P. crassispathum, and P. davidsonii, it is more near-
ly oblong-ellipsoid, often broadest at the base and
bluntly tapered toward the apex. The spadix of P.
errugineum is similarly shaped but is somewhat
more constricted above the sterile staminate por-
tion. Some species, e.g., P. heleniae, have the spa-
dix broadest in the upper % rather than in the ster-
ile staminate portion.
Male flowers. The androecium of P. subg. Philo-
dendron is truncate at the apex, prismatic to ob-
pyramidal and usually irregularly 4—5-sided (Fig.
30). It consists of 2—6 sessile stamens (mostly 3—
4). These are always distinct for most of their length
although often weakly fused at the base. Stamen
number varies within a single spadix and is never
constant (Mayo, 1986). Despite the clear grouping
of stamens it is sometimes difficult to discern clear
floral groupings, especially when the stamens are
distinct. Mayo (1986) reported that some species
have stamens dehiscent laterally rather than ex-
trorsely.
The androecia in Р. subg. Philodendron range
from 2 to 6 mm diam. and show little interspecific
variation. Stamens of all species studied by Mayo
(1986) had druses in their apices. Anthers are ses-
sile to subsessile with a thick connective that is
truncate at the apex and overtops the thecae. The
connective of all species studied by Mayo (1986)
had raphides present. The thecae are ellipsoid, ob-
long or linear, emarginate at the base, and each
opens by a short lateral slit or subapical pore. The
two thecae of a stamen are generally adjacent in P
subg. Philodendron but positioned far apart in P.
subg. Pteromischum an subg. Meconostigma.
Anther thecae lack cell wall thickenings in the en-
Per in P. subg. Philodendron (French,
85).
esee typically emerges in long, slender, some-
what viscid filamentous strands (Fig. 31) (see sec-
tion on “Pollination” for a description of its emer-
gence). These slender strands of pollen do not
persist for many hours, and ultimately the pollen
becomes matted in irregular clusters (Fig. 391, P.
straminicaule).
In P. subg. Philodendron resin canals in the sta-
minate portion of the spadix are situated beneath
the stamens and secrete resin onto the surface of
the stamens. The same is not true of P. subg. Pter-
omischum and P. sect. Meconostigma, where the
resin canals are borne at a deeper level in the axis
and do not secrete resin onto the surface of the
stamens (Mayo, 1986) (Fig. 128, P. crassispathum).
French (1986a) reported that the stamen vascu-
lature of most species of P. subg. Philodendron con-
sists of a single forked bundle with widely divergent
branches. Carvell (1989) reported that a single un-
branched trace supplies each stamen. According to
Mayo (1986) the staminodia have a more distinctive
floral receptacle than do the fertile staminate flow-
ers and often have a multiple-branched vascular
trace, whereas the latter have an indistinct floral
envelope and only a once-branched vascular trace.
In contrast, members of P. subg. Meconostigma
have traces that lack branches or have only short
branches that spread at an acute angle.
Stamens in Р. subg. Philodendron have both
druses and raphide idioblasts with secondarily
thickened walls and tanniniferous idioblasts occur-
ring throughout the ground tissue (Carvell, 1989).
POLLEN
Anthers of Philodendron (Grayum, 1991) have a
two- or more-layer tapetum of a periplasmodial
type. iae mother cell cytokinesis is probably
successive
Philodendron се (Fig. 30) is binucleate (Za-
vada, 1983; Grayum, 1985, 1986, 1992a), inaper-
ы starchy a 4 moderately large size (aver-
aging 40 pm, ranging 28 jum-40 pm), with
h bed polarity (Grayum, 1985, 1991). As in
most aroid genera, it is shed in monads. Pollen is
typically boat-shaped-elliptic to oblong, or occa-
sionally elongate as in P. radiatum. It is usually
round in cross section, but may be very obscurely
keeled in P. hederaceum to moderately keeled in P.
jacquinii, P. jodavisianum, and P. wendlandii, or
prominently keeled in Р. mexicanum. The exine
sculpturing is usually psilate, but sometimes mi-
nutely verruculate, scabrate, or fossulate (P. fra-
grantissimum, P. grandipes, P. jacquinii, P. јодат-
sianum, and P. pterotum) to punctate, subfossulate,
subfoveate, or subverrucate (P. mexicanum and P.
wendlandii).
FEMALE FLOWERS
The pistils of P. subg. Philodendron are closely
aggregated on the spadix in a series of irregular
spirals. Gynoecial characters have long been con-
sidered important in the subgeneric classification
of Philodendron, and lobed stigmas were used as
early as 1832 by Schott. in the recognition of his
greges Meconostigma and Sphincterostigma (the lat-
ter now a synonym of the former). The number of
Annals of the
Missouri Botanical Garden
ovules per locule was used by Engler (1878) in part
to characterize the two largest sections of the genus,
Oligospermum Engl. and Polyspermium (now Ca-
lostigma and Philodendron, respectively) (Mayo,
).
The pistillate flowers consist of a single naked
pistil lacking staminodia. Typically the ovaries are
ovoid to obovoid or elliptic, and terete in cross sec-
tion, or with the sides often somewhat irregularly
angular by compression owing to their close prox-
imity with adjacent pistils. Embryo sac develop-
ment is of the Polygonum-type (Grayum, 1991).
Each ovary for the genus is syncarpous, superior,
and contains 2-47 locules (4-10 in P. subg. Philo-
dendron of Central America). Ovaries range in size
from 0.5 mm (as in P. sousae) to 9.2 mm long (as
in P. advena). The locules are typically oblong, with
thin translucent walls that extend 25—4 the length
of the ovary. The style is barely distinguishable
from the remainder of the ovary, mostly by being
slightly thicker and opaque, rather than translu-
cent. While Dahlgren and Clifford (Dahlgren et al.,
1985) reported no style in Araceae, both Eyde et
al. (1967) and Mayo (1986, 1989) indicated that
the region has a distinct anatomy. Mayo (1989) de-
fined the style of Philodendron as “that portion of
the gynoecium between the base of the stigmatic
epidermis and the ovary locules.” Slender, con-
spicuous styles are rare in Philodendron, but do
exist, as in P. jacquini (Fig. 242).
Each carpel is connected to the stigma apex or
compitum (common stylar canal) by a stylar canal.
The compitum (Endress, 1982) is a cavity or com-
plexly shaped channel into which the pollen may
be inadvertently packed by the beetle pollinators.
This no doubt allows for enough grains to be left to
insure pollination, and to make sure not all are
removed from the stigma by movements of the bee-
tles. The stigmatic epidermis extends into the com-
pitum from the stigma apex (Mayo, 1989). At the
base of the channel or cavity there is a ring of holes
that leads into the stylar canals. Where no compi-
tum is present, such as with “Type B” and “Type
D” styles (see section on “Style Types” below), the
stylar canals lead directly onto the surface of the
style. These canals are readily visible on the dried
stigmas of many species if the preservation is ad-
equate. They are particularly easy to see in fruiting
collections.
The gynoecium has a separate stylar canal for
each carpel (see fig. 1 in Mayo, 1989). Each canal
may open at its upper end into a compitum. A com-
pitum is rare among Central American species (see
section on Style Types below), but is present in P.
correae, P. ligulatum var. heraclioanum, P. smithii,
P. straminicaule, and P. warszewiczii. A compitum
has also been seen on an unusual collection of P
radiatum and some populations of P. tripartitum.
The vascular anatomy of Philodendron ovules
has been studied by French (1986b). Of the five
species he studied, three were South American
while two, P. jacquinii (“P. hederaceum”) and P. in-
concinnum (“P. immixtum"), occur in Central
America. French reported a single vascular trace
for all of these except P. jacquinii, which has mul-
tiple traces. Carvell (1989) reported that the vas-
culature of the gynoecium in Philodendron arises
from a single trace, which divides centrifugally to
yield a single branch trace for each locule. The
carpel traces branch once to form a connection with
the placenta as well as a single dorsal trace. The
placental trace forms individual connections with
each ovule (Carvell, 1989).
Finally, a few miscellaneous anatomical features
of the gynoecium should be mentioned. Both tan-
niniferous idioblasts as well as raphide idioblasts
are lacking in the gynoecium of Philodendron, but
the gynoecium does contain druse idioblasts (Car-
vell, 1989). Like most Araceae, Philodendron has
unicellular ovular and placental trichomes (French,
1987b). These function in secreting mucilage pre-
sumably for the protection of ovules.
Ovules of Philodendron are bitegmic, with the
inner integument forming the micropyle. The integ-
uments are usually completely free from one anoth-
er (Grayum, 1991).
Seemingly the most important morphological fea-
tures of the pistil from the standpoint of systematic
importance are the quantity and distribution of the
ovules and the type of style, discussed in subse-
quent sections.
Placentation. Placentation type for P. subg. Р hilo-
dendron in Central America is usually axile or sub-
basal, with 50 species having sub-basal placenta-
tion and five species with basal placentation. Those
with basal placentation are: P. dwyeri, P. granulare,
P. smithii (also sometimes sub-basal), P. sousae
(also sometimes sub-basal), and P. zhuanum. Forty-
seven species have axile placentation and two ac
ditional species, P. ferrugineum and P. sagittifol-
ium, with mostly sub-basal placentation, sometimes
have axile placentation when they have especially
numerous ovules per locule.
Some general rules regarding placentation type
and number of ovules per locule are the following:
species with basal placentation have only a solitary
ovule per locule and tend to have a few large seeds
per berry. Species with sub-basal placentation ust-
ally have only 1 to a few ovules per locule (aver
وچ ت ج
|
Ми са === олова аса би cm
Volume 84, Number 3
1997
Croat 365
Philodendron Subgenus Philodendron
aging 2.1 to 3.5 respectively for the minimum and
maximum number in the range) and typically fewer
than 6 but rarely to 10 ог to 12 (as in P. brenesii
and P. davidsonii subsp. bocatoranum). Species
with basal and sub-basal ovules tend to have rather
large seeds as well, although sometimes their seeds
are quite small. On the other hand, species with
axile placentation usually have 10 or more ovules
per locule (averaging 14-18 respectively for the
minimum and maximum in the range) but some-
times as few as 3 (P. dressleri and P. warszewiczii),
or 4.
For those species with only a few ovules per loc-
ule, the disposition of the ovules seems to be un-
organized or digitate, but for those with more than
a few ovules, the placentas may be uniseriate or
more generally biseriate or, less frequently, in 1-2
or 2-3 series (see Appendix 2, “Technical Data on
Pistils”). Mayo (1989) reported that, based on his
studies of the gyneocial morphology and anatomy
of 15 mostly South American Philodendron species,
the placenta of even those species whose ovules
appear to be in a single row might actually be bis-
eriate, with the funicles inserted alternately along
the placental ridge. This is also true in Central
American species. Forty Central American taxa of
Philodendron proved to have biseriate placentas,
while much smaller numbers appeared uniseriate
(13), 1-2-seriate (9), 2-3-seriate (11), or digitate
(4). Twenty-six species lacked any appreciable or-
ganization owing to the small number of ovules, and
one species, P. niqueanum, was not studied due to
inadequate material.
Ovary locule number. Mayo (1991) reported that
ovaries in P. subg. Meconostigma range from 3- to
47-locular, and Bunting (1986) reported 2 (rarely
3) locules per ovary as characteristic of P. sect.
Philopsammos.
Though the number of ovules per ovary is highly
significant in separating P. sect. Philodendron and
P. sect. Calostigma, the number of locules is not a
good indicator of relationship. While the average
number of locules per ovary has a slightly lower
range for P. sect. Philodendron (4.6—6.1 vs. 5.5-7.4
for P. sect. Calostigma), the difference is not great.
In P sect. Calostigma the number of locules per
ovary ranges from 1 to 10. In P sect. Philodendron
it ranges from 3 to 10. While P. sect. Philodendron
rarely has more than 8 locules per ovary, and P.
sect. Calostigma rarely has fewer than 5, more spe-
cies (14 in all) have 8 locules per ovary than any
other number.
No species of P. subg. Philodendron in Central
America were found to have bilocular ovaries,
though two species, P. sagittifolium and P. zhuan-
um, may have only a single locule. In the latter
species, the number of locules ranged only from 1
to 3, but in P. sagittifolium the number of locules
ranged up to 8. Often such low locule numbers oc-
cur only near the base of the spadix, where the
ovaries are often somewhat irregular in shape. Fre-
quently the lowermost pistils on the spadix are
larger or smaller than those in the middle of the
adix and are more widely spaced and of irregular
cross-sectional shape. Three-locular ovaries are
are, known only in two species: P. copense and Р.
sagittifolium. Five other species, P. chirripoense, P.
hederaceum, P. immixtum, P. morii, and P. squam-
icaule, may rarely have 3-locular ovaries.
Four-celled ovaries are fairly common in P. subg.
Philodendron. Four species, P. cretosum, P. jacqui-
nii, P. knappiae, and P. subincisum, so far have
demonstrated only 4-locular ovaries. Most species
with 4-locular ovaries also have 5-locular ovaries
on the same spadix. Four species showed exclu-
sively 5-locular ovaries. Only eight species have
locule numbers greater than 10; all but one of these
are in P. sect. Calostigma.
un
Ovules per locule. The number of ovules in each
locule varies from 1 to numerous. Mayo reported
numbers as high as 51 for P. subg. Meconostigma,
but my studies of P. subg. Philodendron show the
highest number found for any Central American
species is 36, in P. fragrantissimum. Only 4 other
species, P. antonioanum, P. panamense, P. squam-
ipetiolatum, and P. verrucosum, have 30 or more
ovules per locule, and 17 species have 20-29 (see
Appendix 2, "Technical Data on Pistils"). Fifty-six
species (62 taxa) have fewer than 10 ovules per
locule.
Along with leaf morphology, the number of
ovules per locule has long been used as a principal
means of assigning sectional affinity in Philoden-
dron (Engler, 1878, 1879, 1899; Krause, 1913;
Mayo, 1989). Central American members of P.
subg. Philodendron generally fall into three cate-
gories in terms of number of ovules per locule.
Many species, including most members of P. sect.
Calostigma, P. sect. Tritomophyllum and P. sect.
Baursia, have one to few (rarely to 4 or 5, but al-
ways with basal or sub-basal placentation). Forty-
four species and four subspecies belong to this cat-
egory. А second group, with axile or sub-basal pla-
centation, generally has 4—10 ovules per locule, but
sometimes up to 14. Nineteen species fall into this
category. The third group has exclusively axile pla-
centation, mostly with many (15+) ovules per loc-
ule. These groups may not be natural, since two
366
Annals of the
Missouri Botanical Garden
species (Р davidsonii and P. roseospathum) have
one variety in each of the aforementioned groups.
It might be expected that, since the pistils of
different species of Philodendron are roughly the
same size (mostly ranging from ] to 4 mm long and
about У to Y as wide), the size of the ovules might
be larger in those species with fewer ovules per
locule than in those with more. Though there are
differences in ovule size (see Appendix 2, “Tech-
nical Data on Pistils”), they are not great. Of 44
species of Philodendron studied that had axile pla-
centation, the ovules ranged from 0.1 to 2.1 mm
long. Alternatively, 48 species with basal or sub-
basal placentation had only slightly larger ovules,
ranging from 0.31 to 0.5 mm long.
Ovules in Philodendron are hemiorthotropous or
orthotropous (Grayum, 1991), rarely hemianatro-
pous (Mayo et al., 1997) and bilaterally symmetri-
al with an eccentric attachment of the funiculus
(French, 1986b). The funicles are usually as long
as or longer than the ovules. The ovules that are
axile usually have funicles of different lengths,
since they are positioned increasingly higher on the
axis. After meeting with the wall of the locule the
funicle is lightly fused to the wall all the way to
the base and can be easily removed intact by pull-
ing it free. This might argue against the assumption
by Mayo (1989) that basal and sub-basal placen-
tation were derived from axile placentation, since
axile placentation could have developed by a
lengthening of the funicular plexus and a fusion to
the wall. While no anatomical study was made of
this phenomenon, the frequent presence of basal
funicles in axile placentas makes it appear that at
least in the case of some Central American species
of P. subg. Philodendron, species with axile pla-
centation may have evolved from species with basal
or sub-basal placentas.
The funicles in P. subg. Philodendron are fre-
quently densely covered with short, usually incon-
spicuous, gland-like trichomes from near the base,
sometimes extending to about half their total length.
The secretory trichomes are continuous with those
of the placentas and lining of the stylar canals
(Mayo, 1989).
Funicles often bear a band of glandular tri-
chomes at or near the base. These were well illus-
trated by Krause (1913) who reported them com-
monly in P. sect. Baursia and P. sect. Calostigma
but with a few in P. sect. Philodendron as well.
These small glands can only be seen under high
magnification and probably secrete mucous into the
ovary, preventing the ovules from drying out.
Funicles are often fused into a thickened, some-
times ramified, more or less translucent placenta.
The entire placenta and its associated funicles and
ovules may be removed, making counting them less
difficult. Sometimes, however, adjacent locules
share a common trunk so that care must be taken
to insure that one is not removing the contents of
two locules.
The free portion of the funicle on species with
axil placentation seems to be proportionately short-
er than the free portion of those with sub-basal pla-
centation. This is perhaps because those species
with axile placentation have somewhat longer loc-
ules allowing greater spacing of the ovules. Those
species with sub-basal placentation and three or
more ovules per locule have the funicles arising
from one small area near the base of the axile wall,
and often have much smaller locules than those
with axile placentation. Ovules need to be separat-
ed for proper development, and funicles of different
lengths allow for this. Thus, in many cases, funicles
are longer in those species with sub-basal placen-
tation. The generally smaller locule size for species
with sub-basal or basal placentation is confirme
by a survey of locule size. Species with basal or
sub-basal placentation have locules ranging in
length from 0.32 mm in P. jefense to 9.5 mm in P
warszewiczii, with the average minimum length 1.9
mm and the average maximum length 3.5 mm. On
the other hand, locules range in length from 0.40
(P. bakeri, P. sousae, P. sulcicaule, and P. wilburü
var. longipedunculatum) to 0.7 mm (P. advena); the
average minimum length is 1.12 mm while the av-
erage maximum is 1.75 mm.
The funicles of a single ovary are usually not of
equal length if the ovules are basal or sub-basal,
since the ovules are usually positioned at slightly
different heights off the floor of the locule. Even
when the ovules are scattered along much of the
length of the axis of the locule wall, the funicles
are of slightly different sizes.
Style and stigma morphology. This treatment fol-
lows the classification of styles and stigmas of Mayo
(1989). Mayo (1986) defined the style in Philoden-
dron as “that portion of the gynoecium between the
ovary locules and the base of the stigmatic ер“
dermis.” Although there is considerable diversity
at the microscopic level, much of the detail of the
style is easily visible only by dissection (see section
entitled “Style Types”). The fresh styles of P. subg:
Philodendron are relatively uniform microscopic?"
ly. Usually they are hemispherical or sometimes
globose or depressed-globose. The microscopically
visible portion is the stigma, a “single continuous
area in which the epidermal cells are greatly elon-
gated into secretory papillae” (Mayo, 1986). This
Volume 84, Number 3
1997
Croat 367
Philodendron Subgenus Philodendron
dense layer of stigmatic papillae is so closely
packed and so engulfed in a gelatinous fluid as to
appear almost solid. Still, a needle can be passed
easily across and through much of its volume in
any direction without disturbing its shape. The stig-
matic papillae are slender, many times longer than
broad, and attached to the surface of the style.
Mayo (1986) reported that the stigmatic papillae
often contain tannin cells.
The stigmatic papillae typically dry to form a
thin, flat, often translucent, wafer-like mantle
these cases the appearance of the stigmas is more
difficult to determine but, regardless of how the
stigma dries, the style apex often can be made eas-
ily visible on a dried specimen by lightly scraping
away the stigmatic papillae. The complexities of the
style types of Philodendron are discussed below.
Style types. The present work adopts the gynoecial
classification of Mayo (1989) in which six distinct
style types were described and illustrated (Fig. 469,
“Style Types in Central America”; see Appendix 2,
"Technical Data on Pistils," which summarizes the
style types for P. subg. Philodendron of Central
America). Only two species, P. niqueanum and P.
utleyanum, have style types not yet known. The ta-
ble also includes the number of locules per ovary,
placentation type, number of ovules per locule, ova-
ry size, disposition of ovary, and the nature of the
ovular sac when present.
Style type is one of the most important gynoecial
features in Philodendron. While the stigma is mod-
erately uniform in its superficial appearance, the
style is highly diverse in morphological features.
Unfortunately, these features are largely hidden by
the stigmatic papillae when the pistil is fresh. De-
spite the shape of the style, the stigmatic papillae,
which cover all or part of the style, may form a
stigma of more or less uniform shape.
Style Type A has a compitum (common fun-
nel into which the pollen may be packed, de-
fined as the space between the upper stigmatic
papillae and the level at which the stylar ca-
nals emerge onto the style surface) with ridged
inner walls and a lobed apex, with each lobe
corresponding to the apex of one carpel. Style
Type A is restricted to P. subg. Meconostigma
and thus will not be considered further here.
Style Type B (Fig. 469A) lacks a compitum.
Instead, the stylar canals open into relatively
broad concavities on the style apex. These
concavities are arranged in a ring with one
hole per locule. Style Type B also has the sty-
lar canal entering directly into the apex of the
locular cavity. Although the style apex may be
completely flattened or broadly concave, it is
sometimes weakly ridged between the aper-
tures of the stylar canals. These ridges meet
in the middle of the style and may even form
a weak central beak. Some styles also have
well-developed stigmatic papillae associated
with the stylar canals causing the surface to
be at least weakly lobed, with a single lobe for
each locule. In dried condition Style Type B
sometimes appears as a button-like structure,
somewhat resembling Style Type D. It is there-
fore important in determining style type to
make comparisons of material at or near an-
thesis. Style Type B seems to be most closely
related to Style Type D and shares with that
type the relatively large stylar canal pores on
the surface of the style apex relatively near its
periphery. Style Type B is the most common
type in Central America, known in at least 70
species. One species, P. tripartitum, though
usually having Type D styles, also has Type B
and E styles in some populations of the spe-
cies (see that species for a discussion of its
style types).
Style Type С (Fig. 4698) is characterized by
being decidedly concave or funnel-shaped at
the apex with no lobes on the margins of the
rim and with the stylar canals arising in a nar-
row cluster at the base of the compitum (fun-
nel). Since the stylar canals are closely clus-
tered there is no central dome (defined as any
stylar tissue that lies above the level at which
the stylar canals emerge onto the style sur-
face). In contrast to style Types B and D, both
of which have rather prominent stylar pores,
the stylar pores of Type С are small, sometimes
barely visible, and in a generally smaller cir-
cle nearer the middle of the style apex. This
style type is rare in Central American P. subg.
Philodendron, and is known primarily in P.
subg. Pteromischum. Only five taxa of P. subg.
Philodendron, (P. correae, P. cotonense, P. lig-
ulatum var. heraclioanum, P. straminicaule,
and Р. warszewiczii) have exclusively Type С
styles. Though their pistils are funnel-shaped
at the apex, these species have funnels gen-
erally not as deep as those of P. subg. Pter-
omischum as illustrated by Mayo (1989). One
other species, P. radiatum, has at least one
collection (Croat & Hannon 63414) with the
368
Annals of the
Missouri Botanical Garden
style type that also has a funnelform apex and
looks like a Type C style. While the dried style
of this collection is distinct and button-like,
the pores are central in a shallow concavity.
Style Type D (Fig. 469C) is similar to Type
B in that it lacks a compitum and has thick
stylar canals emerging in a circle on a flat sty-
lar apex relatively close to the margin of the
stigma apex. It differs from Type B in that the
style is constricted around the circumference
to form a protruded flat “style boss” (defined
as a + domed, circular, stigma-bearing pro-
jection that extends beyond the main part of
the style and is separated from it by a short
neck) that rises above the general level of the
style apex. Thus the style appears to have a
short flat neck at the apex. It is from the “style
boss” that the stylar canals emerge. The stylar
pores are relatively large and borne relatively
near the margin of the style apex. At least 23
species of P. subg. Philodendron in Central
America have Style Type D, the second most
common style type.
It is easy to confuse or misinterpret Style Types
B and D if the specimens are not well preserved,
especially if the material studied is not fresh but
rather rehydrated. Species with Type B styles some-
times have styles that dry with a button-like apex
resembling that of Type D. At least one collection
of P. advena has both Type D and Type B styles.
Some populations of P. tripartitum have not only
Type D styles but Type B and Type E as well (see
the discussion of that species for details).
Style Type E (Fig. 469D) has a slender fun-
nel-shaped to cylindrical compitum with a dis-
tinct raised annulus around the upper rim. The
stylar canals arise in a small cluster at the
base of the compitum just as in Type С styles.
The latter differ, however, in lacking the rim
on the style apex. Type E styles are rare in
Central American P. subg. Philodendron,
found in only P. granulare and perhaps P
smithii. Philodendron granulare has such an
unusual form of the Type E style that it should
perhaps warrant its own status. In P. granulare
the annulus of the stylar funnel actually pro-
trudes well above the surface of the style (at
least in its dried state). Philodendron smithii
was reported by Mayo (1989) as having a Type
E style, but no rim is obvious in fresh material
of the species. It is more appropriately a Type
C style. Philodendron tripartitum, though usu-
ally with Type D styles, has Type E styles in
some parts of its range (see the discussion fol-
lowing that species for details).
Style Type F is narrowly funnel-shaped with
a small dome at the base of the compitum
around which the stylar canals arise. This type
is not known among the Central American
Philodendron. lt is known only from P. burle-
п С. M. Barroso, a member of P. sect.
Baursia (Mayo, 1989) from Amazonian Brazil.
Mayo (1989) has shown that the course of indi-
vidual stylar canals is correlated with the type of
placentation. Those species with basal or sub-basal
ovules, such as P. tripartitum and P. smithii, have
stylar canals that course down the center of the axis
of the pistil and enter the locule near its base.
Those species with axile placentation have stylar
canals that also course down the pistil axis and
enter the locule somewhere above the base of the
locule, but only rarely at the very apex. The stylar
canals are lined with papillose epithelial cells,
which are contiguous with the stigmatic epidermis
(Mayo, 1989). These secrete the gelatinous mucous
that keeps the stigmatic surface moist.
татхи
FRUITS AND SEEDS
In Philodendron the developing pistils remain
within the reclosed spathe after anthesis and pol-
lination until they are fully ripe [except in the rare
case of P. surinamense (Miq.) Engl., a South Amer-
ican member of P. subg. Pteromischum that prompt-
ly loses its spathe after anthesis]. The spathe en-
larges somewhat to accommodate the enlarging
berries. When the berries are mature the spathe
once more begins the process of reopening, but bs
breaks completely free at the base where it 18
obliquely attached to the peduncle. Sometimes the
spathe falls completely free after loosening even
before it opens, but in general it breaks up begin-
ning at the base and falls off (Fig. 33) eventually
falling completely free and leaving a scar just above
the peduncle (Fig. 34). Generally the old, withered
staminate portion of the spadix falls free at this time
as well and the berries are exposed on the remain-
ing pistillate portion of the spadix (Fig. 33, Philo-
dendron hebetatum). In the process of unfolding,
the spathe often develops deep longitudinal fis-
sures, which apparently enable it to unfold. Some-
times the old persistent spathe persists on the ре-
duncle with fragments of the inner surface ex
(Figs. 35, 36).
The berries of Philodendron are cylindrical to
obovoid, generally with a thickened cap-like арех
and not markedly colored, though berry colors from
pale yellow to bright orange or even red to purple
Volume 84, Number 3
1997
Croat 369
Philodendron Subgenus Philodendron
do exist (see section on berry color). Berries in P.
subg. Philodendron are universally soft and fleshy
except for the frequently thickened apex. The seeds
can be seen easily through the sides of the berries.
When fully mature the apical portion of the berry
is easily torn free, and the thin, fragile sides of the
berries are easily ruptured (see section on seeds).
Though little is known about fruit dispersal, the
mesocarp surrounding the seeds contained within
each locule is juicy or gelatinous and is usually
sweet and sticky, making it logically animal dis-
persed. Infructescences that appear to have been
pecked apart by birds (Fig. 36) are frequently seen.
Certainly the sticky seeds, often many per berry,
would logically be easily dispersed on birds’ beaks.
Alternatively, the infructescence is large, and even
faintly scented when fully mature, making it an ap-
pealing meal even for mammals such as monkeys.
Grayum (1996) theorized that those species of P.
subg. Pteromischum with whitish fruits, which may
produce a garlic-like or pepper-like odor at night,
are dispersed by bats. Those with orange fruits may
be dispersed by diurnal animals. Certainly the
manner in which many species of P. subg. Pter-
omischum flower, e.g., on the ends of short, spread-
ing branches some meters above the ground, would
make them superbly positioned as bat fruits. There
are also species of P. subg. Philodendron, such as
P. lentii, which have their branches held in a sim-
ilar manner.
Ants are also probable dispersers of Philoden-
dron seeds. I have seen two different species of ants
carrying away individual seeds of Philodendron.
Ant dispersal is certainly important for those spe-
cies, such as P. megalophyllum in South America,
that live almost exclusively on ant nests. One cul-
tivated individual of that species even set fruit re-
peatedly without being pollinated. Doing so in na-
ture would provide a steady, abundant supply of
berries for its ant dispersers and assure the species
widespread dispersal. Indeed, the species 18 partic-
ularly successful even in areas of white sand soil
where soil nutrients are very low.
Although berry color is known for only a rela-
tively small number of Philodendron species (53
out of 96), some general comments can be made.
onomically as it appears to ђе 1
1983a, 1986a, 1991). Grayum (1996) reported that
for P. subg. Pteromischum the fruits sometimes pro-
vided taxonomically significant characters, with
species related to P. ine having orange
fruits while other species have whitish fruits.
Mature berries of most Philodendron species are
generally described as “white,” but are usually
more nearly ochraceous or somewhat greenish
white. A total of 33 species have either white,
creamy white, or greenish white fruits at least some
of the time. In some species, such as P. advena and
P. smithii, the whitish berries turn somewhat yel-
lowish when fully mature, and the mature fruits of
P. brevispathum are pale yellow. The berries of P.
sagittifolium are usually pale yellowish but rarely
have been reported as orange. While the majority
of Philodendron berries are whitish, 17 species of
Central American P. subg. Philodendron are known
to have berries at least sometimes pale orange to
orange. An additional three species have yellow or
yellowish berries. Philodendron fragrantissimum
has bright red to purple-red berries.
Known species with orange fruits are members
of P. subsect. Glossophyllum. These are: P. auri-
culatum, P. bakeri, P. cotonense, P. pseudauricula-
tum, P. wendlandii, and P. wilburii. Other species
with orange fruits are P. anisotomum, P. brenesii, P.
crassispathum, P. ferrugineum, P. lentii, as well as,
at least sometimes, P. jacquinii and P. sagittifolium.
Three of the above, P. brenesii, P. crassispathum,
and Р. lentii, are members of P. ser. Ecordata. Thus,
it may be significant that most species with decid-
edly orange fruits are members of P. sect. Calostig-
ma. Even P. ferrugineum, with orange berries, and
P. sagittifolium, sometimes with orange berries, are
members of P. sect. Calostigma. Only P. anisoto-
mum (P. sect. Tritomophyllum) and P. jacquinii (P.
sect. Macrogynium) are not members of P. sect. Ca-
lostigma.
The seeds of P. subg. Philodendron are few to
any per berry and are mostly oblong to oblong-
ellipsoid, ellipsoid, ovoid-oblong, or less frequently
ovoid. Typically they are minute (see below). The
seeds are typically more or less terete, though they
may be somewhat flattened, e.g., in P. findens. They
have a rather thick, smooth (e.g., P. granulare) or
striate-costate testa, as well as copious endosperm.
The longitudinally oriented striations may be weak
(P. advena, P. findens, P. cotonense, P. purpureovir-
ide, and P. sagittifolium) to strong (P. ferrugineum
and P. grandipes). Seeds of Philodendron microstic-
tum have both longitudinal striations and much fin-
er cross-etching. A number of species are reported
to have seeds with pale raphide cells on the sur-
face. These include P. hederaceum and P. warszew-
iczii. Seeds of some species, e.g., P. anisotomum
and P. hederaceum, sometimes have a constriction
on the end opposite the funicule.
Philodendron seeds are usually dramatically
smaller than those of Anthurium, which typically
have only two seeds per berry. Seeds of species of
P. subg. Philodendron studied (a total of 48 species)
370
Annals of the
Missouri Botanical Garden
ranged from 0.5 mm to 5 mm and averaged 1.67
mm long; seed diameter ranged from 0.1 to 3 mm,
averaging 0.7 mm. The longest seeds were those of
P. jacquinii and P. hederaceum, at 5 mm. Relatively
few species have seeds longer than 2 mm: Р аа-
vena, P. anisotomum, P. findens, P. dolichophyllum.
P. grayumii, P. purulhense, P. rothschuhianum, P.
sagittifolium, and P. warszewiczii.
The number of seeds in each locule is often
many fewer than the average number of ovules per
locule, presumably owing to the fact that not all are
properly pollinated. A careful examination some-
times shows the aborted undeveloped ovules that
were present at the time of flowering. The percent-
age of ovules in each locule that develops into
seeds varies from species to species. Species of P.
sect. Calostigma that have only one or a few ovules
per locule are more likely to have an equal number
of seeds. On the other hand, species in Р. sect.
Philodendron, especially those with large numbers
of ovules per locule, rarely develop all their ovules.
Because species of P. sect. Calostigma have few-
er ovules per locule and thus fewer seeds, one
might expect them to have larger seeds. Though
admittedly the sample size was small (only 21 spe-
cies studied for P. sect. Calostigma and 17 for P.
sect. Philodendron), the unexpected results were
that the seeds of species of P. sect. Philodendron
in Central America averaged slightly larger than
those of P. sect. Calostigma (to 1.69 mm long for
P. sect. Philodendron and to 1.66 for P. sect. Ca-
lostigma).
Philodendron seeds are largely pale in color,
mostly shades of brown, tan, or white, though for
most species mature seeds have not been observed.
Seeds vary in color from medium green (P. roths-
chuhianum) to whitish (P. roseospathum), tan (P. al-
ticola, P. annulatum, P. granulare, P. morii, P. smi-
thii), yellowish or pale yellow (P brenesii, P.
llanense, and P. strictum), yellow-orange (P. crassis-
pathum, P. purpureoviride, P. wilburii, and P. mex-
icanum), brown (P. ferrugineum), or reddish brown
(P. heleniae). The dried seeds of two species, P.
findens and P. jefense, are described as translucent.
POLLINATION BIOLOGY
Although there are frequent insect visitors to
Philodendron, especially small Hemiptera in the
genus Neelia, which appear not to feed or mate on
the inflorescences (H. Young, pers. comm.), only
the larger beetles are known to be pollinators. The
system of pollination is nearly identical to that of
Dieffenbachia (Croat, 1983b; Young, 1986, 1990).
Pollinators are members of subfamily Dynastinae in
the family Scarabaeidae (Fig. 32). All determined
beetles collected from either Central American or
South American Philodendron are members of the
genera Cyclocephala and Erioscelis. Some species
of beetles are not particularly host-specific, visiting
members of both P subg. Philodendron and P.
subg. Pteromischum; other genera such as Dieffen-
bachia, Homalomena, Syngonium, and Xanthoso-
ma; and sometimes even other families including
some palms (Arecaceae), Cyclanthus bipartitus Poit.
ex A. Rich (Cyclanthaceae), as well as Annona and
Cymbopetalum in the Annonaceae (M. Grayum,
pers. comm.; Schatz, 1990). Cyclocephala negerri-
ma Bates, for example, has been found visiting P.
brenesii and P. tysonii, but also P. standleyi Gra-
yum, a member of P. subg. Pteromischum (see Table
3). Though beetles are not very species-specific
pollinators, individual beetles of some species tend
to be somewhat stratum-specific, visiting only those
species growing at particular height ranges above
the ground (Schatz, 1990; Helen Young, pers.
comm.). These beetles are attracted to the Philo-
dendron inflorescence, usually late in the day or at
dusk. Attractants and/or rewards are apparently a
combination of scent (at least in many cases), а
source of food (oil-bearing sterile staminate flow-
ers), warmth (thermogenesis), and shelter.
Scents produced by Philodendron species are not
always obvious, at least in P. subg. Philodendron.
George Schatz and Helen Young (pers. comm.) have
documented floral odors for species of Philoden-
dron and have identified the principal constituents
of these aromas. Some species have noticeably
sweet aromas in the early evening hours, while oth-
er species have no noticeable scent, at least during
the early evening hours on the first day of anthesis.
Philodendron megalophyllum (in cultivation at the
Missouri Botanical Garden), a South American spe
cies, had a faint spicy aroma detectable directly at
the spathe during the evening, but even this faint
aroma was absent the following morning on day о
of the flowering event. At the same time the stigmas
were soft, juicy, and sticky but without an obvious
flavor or taste both in the evening and the following
morning. Schatz (1990) believed that the pattern of
visitation to Philodendron exhibited by beetles a!
La Selva in Costa Rica was to a great degree ех
plained by odor. He pointed, for example, to the
high degree of specificity exhibited by P hiloden-
dron radiatum and an undescribed species of Cy-
clocephala. The aroma given off by P. radiatum =
made up of compounds unique to that spectes-
At anthesis the open spathe of Philodendron pt
vides ample space at the base in the area surroun
ing the pistillate portion of the spadix. The s
|
Volume 84, Number 3
1997
Croat 371
Philodendron Subgenus Philodendron
blade may provide a certain amount of protection
against rain, as mentioned previously. Beetles typ-
ically spend the first night and most of the following
day inside the spathe where they remove the sticky
exudate from the pistils, eat pollen (Gottsberger &
Silberbauer-Gottsberger, 1991), feed on the sterile
staminate flowers, and mate. Studying P. bipinna-
tifidum, Gottsberger and Silberbauer-Gottsberger
(1991) found that the beetles were active mostly
during the first 10 to 20 minutes after arrival an
during the strongest production of scent. Copulation
was most active immediately after arrival, when the
spadix was warmest; grazing on the sterile male
flowers also subsided by the time the spadix had
cooled off. Old inflorescences that have been pol-
linated often have the sterile staminate portion of
the spadix entirely eaten away.
Typically the number of beetles found in a single
inflorescence is modest, frequently no more than
five and sometimes up to a dozen beetles. However,
sometimes the numbers are simply astounding, with
a
innatifidum. While visiting beetles are often of the
same species, sometimes more than one species of
beetle may be found in the inflorescence. Though
beetles typically do not leave the spathe until about
dusk, they will leave if the inflorescence is suffi-
ciently disturbed. When beetles are disturbed, they
may crawl slowly up the spathe or spadix and ap-
pear at the rim of the open spathe, at which point
they generally fall promptly to the ground where
they disappear in the leaf litter or soil with re-
markable speed.
Thermogenesis, the production of heat in the
spadix by the rapid oxidation of stored starch or
lipids (Walker et al., 1983; Gottsberger, 1990),
plays an important role in the pollination of Philo-
dendron (van Herk, 1937a—c; Van der Pijl, 1937;
Knutsen, 1974; Seymour et al., 1984). Gottsberger
(1984, 1986), studying P. bipinnatifidum, has
shown that, although oxidation of carbohydrates
takes place during preheating of the spadix, lipids
are oxidized thereafter during maximum heating
and are consumed directly, not after conversion to
carbohydrates. This makes the biochemistry of this
species similar to that accompanying heat produc-
tion in some animals. The thermogenic reaction oc-
curs principally in the staminodial region of the
staminate portion of the spadix (James & Beevers,
1950; Henry & Nyns, 1975) involving the inner
surface of mitochondrial membranes (Urdentlich et
al., 1991) and is triggered by an accumulation of
salicylic acid (Meeuse & Buggeln, 1969; Raskin et
al., 1987, 1989; Meeuse, 1975, 1978; Raskin,
1992). The end result of this high rate of respiration
in these plants is the production of heat rather than
ATP as in animals (Meeuse, 1966). To accomplish
this high increase in metabolism the plant's mito-
chondria in the inflorescence switch to an electron
transport pathway commonly referred to as the “cy-
anide-resistant pathwa
e thermogenetic haat rise in Philodendron is
sometimes dramatic, with temperatures rising EM
berger (1991), working with P. bipinnatifidum, re-
ported temperatures of spadices occasionally to
46°C with the highest and most efficient tempera-
tures for the emission of scents being maintained
for 20—40 minutes. Thermogenesis does not create
even or constant temperatures, but rather produces
fluctuations depending on the time of day with def-
inite peaks (Leick, 1910, 1916; Engler, 1920a,
1920b; Foster, 1949; Nagy et al., 1972; Sheridan,
1960; Gottsberger & Amaral, 1984), the greatest
occurring when beetle visitation and odor is most
intense (Gottsberger & Amaral, 1984). Temperature
peaks may occur on two or more successive days.
While the increased temperature is presumably re-
sponsible for the production of scent compounds
(Nagy et al., 1972), and whereas the production of
heat and scent appears closely correlated with
peaks in temperature, there is still controversy over
= exact function of the heat production at least
it pertains to genera that produce foul odors.
Moodie (1976) suggested that heat production and
the higher levels of carbon dioxide production are
components of a carrion, dung, and mammal mim-
icry syndrome and that heat production aids in pro-
viding sufficient warmth in colder climates for the
activity of pollinating organisms. The subject of
thermogenesis and its role in pollination has been
reviewed in great detail by Mayo (1986), Grayum
(1990), and Bay (1995[1996)).
The exact role that thermogenesis plays in the
pollination of Philodendron is still poorly known,
and rather few plants have been studied on an ex-
perimental basis. Despite its probable occurrence
in all Philodendron species, thermogenesis has thus
been documented only for P. selloum K. Koch and
P. bipinnatifidum, now considered by Mayo (1991)
to be synonymous. My own measurements with a
recording thermometer on P. glanduliferum and P.
advena in the field in Chiapas, Mexico, indicated
a definite heating during the early evening hours,
usually peaking between 18:00 and 19:00 hours
The site of the heat production on the spadix is
another possible difference between P. subg. Me-
conostigma and P. subg. Philodendron. Mayo's
372
Annals of the
Missouri Botanical Garden
(1986) investigations on P. subg. Meconostigma in-
dicated that heat production was centered in the
sterile staminate section of the spadix. In P. subg.
Meconostigma, that section is as large as or larger
than the fertile staminate part of the spadix, a sit-
uation unknown in P. subg. Philodendron where the
staminodial segment is always a small percentage
of the total spadix.
Contrary to Mayo, Leick (1916) reported that
heating took place in the “middle and upper part”
of the spadix in P. selloum, presumably implying at
least a part of the fertile staminate portion. Con-
curring with this view, Ron Weeks (pers. comm.)
reported that a perceived temperature rise occurred
in both the sterile and fertile staminate spadix por-
tions of those species of P. subg. Meconostigma that
he studied. Mayo (1986) theorized that, because of
the major morphological differences in the relative
lengths of the sterile and fertile staminate portions
of the spadices, the two subgenera would likely
have different thermogenetic patterns.
In the Central American Philodendron species
observed by the author, the spathe opens broadly
late in the afternoon of the first day of flowering; it
closes slightly the morning after the first night of
anthesis. The evolutionary significance of this slight
closure is uncertain but, even in the most extreme
cases, it occludes only the pistillate portion. The
spathe continues to close on the evening of the sec-
ond day after having been open for about 24 hours
(see discussion below). The beetles, which gener-
ally enter the spathe on the first night of opening,
spend about 24 hours in the spathe tube. This sec-
ond partial closing process usually corresponds
with staminal dehiscence. The now crowded con-
dition of the spathe tube, and the probable desire
on the part of the beetles to seek a new food source
with the onset of dusk, encourages the beetles to
leave the inflorescence. The beetles emerge from
the spathe tube by either climbing up the side of
the spadix or up the inside wall of the spathe. By
the time they emerge from the spathe, it is quite
constricted and they must squeeze through the con-
striction which fits rather tightly around the spadix
just above the sterile staminate portion. In order to
depart the beetles must literally crawl through the
copious strands of pollen that emerge from the api-
cal pores of the stamens. The constriction of the
spathe and its corresponding constricted area on
the spadix help to insure that most of the pollen
that falls into the spathe tube accumulates and is
carried forward and out of the spathe by the de-
parting beetles. Araceae pollen is not very tacky
and probably does not adhere well to the smooth,
hard surfaces of the beetles. However, the beetles
themselves are usually quite sticky from the sugary
secretions of the styles and especially from the res-
in, which arises usually from the inner surface of
the spathe or sometimes from the spadix itself (Fig.
128). Once the beetles have emerged they fly off in
search of another place to spend the night, usually
another open inflorescence. The beetles apparently
have a keen perception of infra-red radiation or of
scent because they are often seen in a “homing-in”
pattern, which is quite direct to the next available
inflorescence (John Rawlins, pers. comm.). Gotts-
berger and Silberbauer-Gottsberger (1991) reported
that beetles flew in a zig-zag pattern toward the
center of fragrance concentration, indicating that
they were very sensitive to the aroma being pro-
duced. They also reported that once the beetles
were within sight of the inflorescence they shifted
to a straight line of flight until they hit the inner
surface of the spathe blade, whence they moved
into the lower portion of the spathe. Gottsberger
and Silberbauer-Gottsberger (1991) have proven
experimentally in the case of P. bipinnatifidum (a
member of P. subg. Meconostigma) that the beetles
use only visual references for location as they near
the inflorescence. Shelter, warmth, food, and cop-
ulation are the driving forces behind this pollina-
tion strategy, and although selectivity is not perfect
in such beetle-pollinated systems (Young, 1986,
1988a) fruit-set in undisturbed populations is high.
The precision and high degree of synchrony of ther-
mogenesis gives evidence of a highly evolved sys-
tem of pollination.
Even though all detailed observations thus far
have been made with P. bipinnatifidum, there 18
considerable confusion regarding the results. Hor-
ticulturist Ron Weeks (Homestead, Florida) report-
ed (pers. comm.) that three members of Р subg
Philodendron, P. bipinnatifidum, P. speciosum
Schott, and P. williamsii Hook. f., showed no vari-
ation in the schedule of spathe opening, the capa-
bility of the plants being hand-pollinated on the
first evening of opening, or in the shedding of pol-
len on the evening of the second day. On the other
hand, he reported that Р. eichleri Engl. showed
great variation in opening periods, temperature
changes, fragrance, and pollen shed, perhaps owing
to weather conditions. Scientific studies carried out
on other plants in P. subg. Meconostigma show
considerable variability. Four separate and conflict-
ing reports were made on material determined as
belonging to P. bipinnatifidum in Brazil. Warming
(1867, 1883) reported a two-day pollination event
with two heat peaks (early evening and late топ“
ing, respectively) with the spathe closing WE
opening during the first night. Gottsberger
Volume 84, Number 3
1997
Croat 373
Philodendron Subgenus Philodendron
Amaral (1984) reported on two plants, one as Р
selloum (now considered to be a synonym of P. bip-
innatifidum) with a three-day pollination event with
two unequal early evening heat peaks, and one as
P. bipinnatifidum with a four-day pollination event
with three unequal early-evening heat peaks. The
spathe was not reported to close during the event.
Confirming the complexity of the thermogenesis
riddle is the fact that Seymour et al. (1983), study-
ing a cultivated but similar plant believed to be the
same species (Mayo, 1986), found both types of
pollination events that had previously been de-
scribed by Gottsberger and Amaral but this time in
a single plant. Clearly more investigation must take
place, at least in P. subg. Meconostigma, to deter-
mine the pollination behavior.
Leick (1916), reviewing work done by Kraus
(1894, 1896) with P. bipinnatifidum, a member of
P. subg. Philodendron from Venezuela, reported a
two-day pollination event with temperature peaks
in the evening of two consecutive days. While it is
not certain that most Central American Philoden-
dron have an elevated temperature on two consec-
utive days, the general pattern of opening and clos-
ing of the spathe and the beetle visitation in P.
bipinnatifidum would appear to match the events of
Central American species observed in the field and
under greenhouse conditions at the Missouri Bo-
tanical Garden. Further detailed studies of this
phenomenon, including a much broader survey of
Central American species, will be carried out by
my student, Jane Whitehill, during graduate studies
at the Missouri Botanical Garden.
Grayum's (1996) observations with Philodendron
subg. Pteromischum showed a similar pattern, with
most species having the spathe beginning to loosen
by early afternoon and being fully open by mid-
afternoon. He reported that for the species of P.
subg. Pteromischum he observed, the pollinators
appeared at the opened inflorescence during a rel-
atively brief time, usually between 19:00 and 19:
15 hours. An important feature in the pollination
story reported by Grayum (1996) for the first time
is that resin secretion from the inner spathe surface
does not begin until 21:00 to 22:00 hours on the
first day of anthesis, and that it then continues until
the end of anthesis.
Once opened, the spathes of P. subg. Philoden-
dron apparently remained open during the night
d were always open the next morning at the be-
ginning of day two, remaining open during the
course of most of the day. During the latter part of
the same day, usually in late afternoon of day two,
the spathe began to close and pollen began to shed
in long filaments. The spathe did not fully close at
this time but remained open near the apex. It re-
mained in this condition into the beginning of the
evening of day two. By the beginning of day three
the spathe was generally fully closed, and the only
evidence that it had ever opened was often some
loose pollen remaining on the closed edges of the
spathe. In addition, the closed spathe is somewhat
less turgid than before anthesis, sometimes allow-
ing it to be forced open without breaking the mar-
gins of the spathe. Doing the same with an uno-
pened spathe is impossible without breaking the
stiff and brittle spathe margins.
The entire pollination episode usually requires
little more than 24 hours, counting just the time
that the beetles are present. The time that the
spathe is to any extent open could be as much as
8 hours longer, since it may open late in the after-
noon and remain open for some hours after the pol-
len has been shed. Grayum (1996) reported that for
the species of P. subg. Pteromischum studied in
Costa Rica the average pollination event required
about 30 hours (i.e., from spathe opening to clos-
ng).
That the intensity of light must play an important
role in flowering behavior is indicated by the fact
that on cloudy days spathes in cultivated collec-
tions open earlier than usual, sometimes as early
as noon on day one of the flowering sequence. This
may support the argument of Buggeln et al. (1971)
that darkness induces opening of the spathe and an
elevated respiration rate in Sauromatum venosum
Kunth.
Armbruster (1984), studying the role of resin in
angiosperm pollination, has questioned the efficacy
of floral resin in the transport of pollen, citing its
possible toxicity and the difficulty of transporting
pollen embedded in resin. While he stressed the
role of resin for other purposes, mainly in nest-
building by bees, it must be pointed out that bees
which use resin for nest building play no role what-
ever in Philodendron pollination. In contrast, the
general availability of resin, its close association
with pollen delivery, the non-tacky nature of Philo-
dendron pollen, and the availability of resin only at
anthesis all point to a strong role for resin in Philo-
dendron pollination. In species with resiniferous
spadices (Fig. 128), the pollen is shed with and
incorporated in the resin from the moment of theca
dehiscence. Alternatively, species that lack stami-
nal resin and instead have resin only on the inner
spathe surface have pollen presented as slender fil-
aments.
Breeding studies (see section on Breeding Be-
havior) have shown that Philodendron species have
few if any genetic barriers to cross-pollination, ow-
—.
374
Annals of the
Missouri Botanical Garden
ing perhaps to the fact that there are other physical
and temporal barriers to self-pollination. Even
when two species of Philodendron are in flower si-
multaneously, there are other parameters that effect
separation. Beetles tend to fly at certain elevations
above the ground (Schatz, 1990), helping to prevent
cross-pollination of species that flower at different
strata. In addition, specific attractants, i.e., species-
specific pheromones, may exist in some species that
attract principally a single beetle species (H.
Young, pers. comm.). The sloppiness in the system,
when it occurs, is owing to opportunistic beetle vis-
itors (С. Schatz & H. Young, pers. comm.), and this
might produce some hybridization.
Perhaps because of the substantial barriers al-
ready present, Philodendron appears to have de-
veloped the ability to cross between sections. In
Anthurium, relatively little cross-breeding was pos-
sible between different sections in the genus (Croat,
1991). In contrast, quite unrelated species of Philo-
dendron, even species in different sections, readily
cross-pollinate and produce intermediate offspring
(Keith Henderson, Cairns, Queensland, Australia,
pers. comm.). For this reason pre-zygotic separation
may be critical to maintaining distinct species
lines. The pollinators of Philodendron, dynastine
scarab beetles, are for the most part not very spe-
cies-specific and frequently switch from one spe-
cies to any other in flower at the same time. Not
only will some species of beetles switch from one
Philodendron species to the next, as is known for
certain in Dieffenbachia (Young, 1986), but some
individuals will also switch to another genus. For
example, beetles that regularly visit D. longispatha
Engl. & K. Krause at La Selva may visit P. gran-
dipes, another species that is terrestrial and about
the same height above the ground (G. Schatz $: H.
Young, pers. comm.). Beetles are also reported to
move from Dieffenbachia longispatha to Xanthoso-
ma undipes (K. Koch & Bouché) K. Koch. Some
individuals of beetle species that regularly visit D.
longispatha will even switch to Cyclanthus bipar-
titus Poit. (Cyclanthaceae). Schatz believes that this
is owing to the fact a small component of the pher-
omone emitted by Cyclanthus Poit. ex A. Rich is
the principal component of the scent given off by
Dieffenbachia longispatha. He believes that during
the end of the flowering season of Dieffenbachia and
the beginning of the flowering season of Cyclanthus
some confusion occurs in the pollinators” behavior.
The unpublished observations of G. Schatz and
H. Young (pers. comm.), and the published results
reported by Helen Young (1986, 1988a, 1988b)
principally for Dieffenbachia, probably are compa-
rable to what is happening in Philodendron. Beetles
that visit any particular species of Dieffenbachia are
often predominantly of one species, but they are
often accompanied by other opportunistic beetles.
Schatz believes that these opportunistic species are
not likely to be effective pollinators, since they are
so catholic in their tastes that they are not likely
to make their next visit to a receptive Dieffenba-
chia. Likewise, Helen Young (Young, 1988a) indi-
cated that the most common species of beetles are
not the most effective pollinators. However, they
may be responsible for the occasional hybrids seen
in Dieffenbachia, at La Selva, where the studies of
both Schatz and Young were carried out. The pol-
lination system described for Dieffenbachia by
Young and Schatz is apparently similar to that of
Philodendron. Schatz (pers. comm.) reported that
while one undescribed beetle species (determined
as Cyclocephala ampliata by H. Young), was found
to visit only Philodendron radiatum, it was accom-
panied occasionally by another more opportunistic
species. Despite the presence of opportunistic
beetle species, some beetle pollinators of Philoden-
dron species are probably much more species-spe-
cific. For example, Grayum (1996), citing unpub-
lished data collected by George Schatz, reported
that two unrelated species of Philodendron subg.
Pteromischum were pollinated by the same species
of beetles, and that both Philodendron species have
floral odors featuring the same two principal com-
ponents. This leads to the conclusion that there is
a degree of specificity among pollinators for certain
species based on their floral odors. In addition, in
the list of pollinators known for P. subg. Philoden-
dron (see Table 3), only three Philodendron species
were observed to have more than a single species
of beetle present at any one time. In each case, two
species of beetles were present. Аз can be seen
from P. grandipes (Table 3), the beetle species need
not always be the same. Though more studies must
be made on pollination biology of Philodendron,
and even though the beetle pollination system 18
somewhat sloppy and imprecise, a combination of
a moderately strong beetle-plant specificity, COU
pled with severe phenological constraints and nar-
row windows of pollination opportunities (perhaps
as little as a few hours per year), works to reduce
interspecific hybridization. Although hybrids. сав
be readily produced under greenhouse conditions,
evidence for hybridization is not usually apparent
among wild populations.
BREEDING BEHAVIOR
In comparison to Anthurium, where breeding
studies were easy to conduct (Croat, 1980, 1983a,
Volume 84, Number 3
Croat 375
Philodendron Subgenus Philodendron
1986a, 1991), Philodendron pollination was diffi-
cult. Cross-pollination attempts were easy in An-
thurium owing to their hermaphroditic flowers and
because plants often had several inflorescences per
plant in different stages of development. In addi-
tion, the plants reached anthesis during the day
when greenhouse personnel and volunteers were
available to make cross-pollinations. In Philoden-
dron, the number of inflorescences available was
always fewer than in Anthurium; Philodendron
sometimes produced only a single inflorescence per
season. In addition, Philodendron was very season-
al in its flowering behavior (unlike Anthurium,
which sometimes flowered all year), making polli-
nation all the more difficult. When flowering did
occur it was often unexpected since it is difficult
to tell when the spathe is ready to open. Moreover,
the spathe generally opens for one day only. Open-
ing usually took place late in the afternoon after
greenhouse personnel left. Even if the opening in-
florescence was found in time, it was generally im-
possible to find another plant with fresh pollen to
use for purposes of experimental crosses. Philoden-
dron pollen does not remain viable very long,
though it can be kept viable for a time in glassine
envelopes. Ron Weeks, a grower from Homestead,
Florida, reports (pers. comm.), that he stores pollen
in film canisters at refrigerator temperatures and
that it remains viable for several weeks. He also
reports that inflorescences cannot be pollinated af-
ter the first evening they open. After the spathe
opens (generally late in the day) it is only during
the evening and night of the first night that the
pistillate flowers are believed to be receptive. At-
tempts to pollinate plants with their own pollen
have always failed if one waits until the pollen
emerges. It should be noted, however, that Grayum
(1996) reported that, based on the use of peridoxase
paper, which purportedly indicates the receptivity
of stigmas (Young, 1986), the stigmas were recep-
tive for up to 24 hours after anthesis. Though it
seems unlikely that pistillate flowers are receptive
after the first evening of anthesis, the pistillate flow-
ers are receptive for an unknown period of time
before the spathe opens, so that most successful
pollinations usually involved cutting a hole in the
spathe after obtaining very fresh pollen from a plant
in the staminate phase of flowering. Though one can
completely remove the spathe then protect the de-
veloping pistils with a plastic bag, it is better to
simply cut a window in the spathe large enough to
see most of the spathe. Then with a small brush
One can spread pollen over as many of the pistils
as possible, again covering the spathe for a time
with a plastic bag to insure that the pollen does not
dry out and fail to germinate. An effective means
of spreading the pollen to insure adequate and uni-
form coverage is to mix the pollen with water.
Failure to remove or at least loosen the bag used
to cover the pollinated spadix later may result in
mold developing in the spathe. Unpollinated inflo-
rescences usually fall off within a week or two. Ron
Weeks (pers. comm.) reported that for Р subg. Me-
conostigma in Florida the unpollinated inflores-
cences may persist for up to a month. Development
time for fruits ranges from only a few weeks or more
generally a few months and sometimes nearly a
whole year. Ron Weeks (pers. comm.) reports that
in P. subg. Meconostigma fruits ripen in South Flor-
ida in 2.5 to 3 months.
PHENOLOGY
Based on a field-oriented study of phenology and
pollination behavior carried out at La Selva in He-
redia Province, Costa Rica, Grayum (1996) report-
ed species of P. subg. Pteromischum flowered for
periods of 4-8 weeks. Obviously, since most mem-
bers of P. subg. Pteromischum have only one or two
inflorescences per axil, vs. sometimes 4 or more for
P. subg. Philodendron, the flowering episodes of the
latter might be longer than two months. The flow-
ering events, however short or long, are not nec-
essarily the same year after year. I suspect that, like
understory vegetation (Croat, 1975), their flowering
рене may be affected by the onset of the rainy
on.
Pasa primarily on a phenological survey of her-
barium collections, flowering (and to a lesser extent
fruiting) behavior has been studied here for Central
American species of P. subg. Philodendron. These
studies resulted in a phenological statement for
each species. This statement follows the description
of each species as a part of the discussion. Some
general comments regarding phenology are impor-
tant.
In studying herbarium material for phenological
variation it is often difficult to determine the exact
state of the inflorescence. However, with experience
it is relatively easy to distinguish inflorescences
that have never opened, i.e., pre-anthesis, from
those which have already opened. Spathes that
have never opened are very tightly closed, whereas
those that have already undergone anthesis are not
so tightly closed. By dissecting the spathe one can
quickly determine if the pollen has emerged. If so,
the spathe has already opened and reclosed over
the spadix.
Spadices at anthesis when they are collected are
usually easy to discern as well because they are
2
376
Annals of the
Missouri Botanical Garden
typically opened when pressed. It is difficult, how-
ever, to easily predict the age of an inflorescence
beyond anthesis and before swelling due to the en-
largement of the pistils. Since an inflorescence at
anthesis is rare (open less than 24 hours), while
every successfully pollinated inflorescence persists
for one to many months, there will always be many
more collections with “post-anthesis” inflorecences
than those described as “in flower.”
Central American members of P. subg. Philoden-
dron fall into several phenological groups. All of
these categories can have variations, and flowering
is rarely consistent throughout any period. Some
categorizations are tentative, as listed in Appendix
, “Phenological Patterns of Central American
Philodendron subg. Philodendron.”
The flowering patterns of Central American
members of P. subg. Philodendron are as follows:
A. FLOWERING IN DRY SEASON AND WET SEASON
This is the largest flowering category among Cen-
tral American P. subg. Philodendron. A total of 47
species fall into this category, including P. fragran-
tissimum, P. panamense, P. sagittifolium, P. gran-
dipes, P. pterotum, and P. radiatum. Although these
species begin flowering during the dry season, the
dry season rarely constitutes the period of greatest
flowering activity (except perhaps in the case of Р
hederaceum, a species which though apparently
flowering all year and thus aseasonal appears to
have more flowering collections made during the
dry season than at any other time of the year). Low
flowering activity is particularly characteristic of
species inhabiting regions of Tropical moist forest
or other areas where marked seasonal changes are
apparent and affect the availability of beetle polli-
nators.
Those species occurring in cloud forest habitats
(usually Premontane rain forest or Lower montane
wet forest) also seem to have more flowering in the
rainy season even though they are more apt to be
in flower in the dry season than their Tropical moist
forest counterparts.
Though perhaps it is merely a matter of poor
sampling, species more common, widespread, or lo-
cally abundant tend to have flowering seasons that
extend from the dry season to the wet season,
whereas rare species seem much more likely to
have flowering restricted to either the dry or the
wet season.
B. FLOWERING ONLY IN WET SEASON
This is the second largest flowering category,
comprising 30 species (32 taxa) believed to flower
exclusively in the wet season (roughly between May
and December in Central America). They represent
largely rare or narrowly distributed species for the
most part and are often species that inhabit the
wettest and frequently the coolest forest types such
as Tropical wet forest, Premontane wet forest, Pre-
montane rain forest, and Lower montane wet forest.
Examples of species in this flowering type are: P.
albisuccus, P. antonioanum, P. chiriquense, P. co-
loradense, P. correae, P. cotobrusense, P. dominica-
lense, P. dodsonii, P. ferrugineum, P. gigas, P. ham-
melii, P. jefense, P. madronense, P. niqueanum, P.
ритепзе, P. purulhense, P. squamicaule, and P. ubi-
gantupense.
Not all species that flower exclusively in the wet
season occur in very wet or cool forest. А few spe-
cies flower only in the wet season because the dry
season in the region where they occur is often too
severe, perhaps so severe as to limit the beetle pol-
linators. These include several Mexican species,
e.g., P. basii, P. breedlovei, P. dressleri, and P. sou-
ae.
ш
С. FLOWERING ONLY IN DRY SEASON
One of the most unusual flowering categories,
and a relatively small one with only 7 species, is à
group believed to flower only during the dry season
(January to April in Central America). This group
is diverse and difficult to characterize. Some of the
species, such as P. bakeri, P. brewsterense, P. chir-
ripoense, P. edenudatum, P. folsomii, and P. knap-
piae, occur in wet to very wet areas, some in areas
where weather conditions are so bad in general dur-
ing the rainy season that it may be more efficient
to compete for pollinators during the dry season
(which would not be very dry in any event). Philo-
dendron dwyeri is unusual in that it flowered at the
beginning of the dry season in an area that in gen-
eral is quite arid in this season. It is known from
only a single individual and may have represented
an unusual, out-of-phase flowering.
D. FLOWERING ALL YEAR
Species that flower aseasonally are usually com
mon species such as P. hederaceum (though it ap”
parently flowers more frequently in the dry season
and Р. jacquinii (which is also often edaphically
versatile or less subject to the pressures of the en-
vironment). Though not as widespread as the two
aformentioned species, P. advena, another member
of this group, is a tough, highly variable and eco-
logically adaptive species. Philodendron purpure”
viride has much the same edaphic preferences as
P. hederaceum, although it is much less widespread 2
Volume 84, Number 3
1997
Croat 377
Philodendron Subgenus Philodendron
Both are highly adaptable, scandent species. Philo-
dendron hederaceum, though flowering all year, is
even more commonly in flower in the dry season
than in the wet season. Philodendron radiatum,
also a member of this group, is as widespread and
even more common than P. hederaceum.
E. FLOWERING BIMODALLY
Based on the historical record of herbarium col-
lections, only a few species are expected to flower
twice per year. This pattern may be much more
common and simply masked by the year-to-year
variation in flowering behavior. Grayum (1996) re-
ported bimodal flowering to be common with Р
subg. Pteromischum. He indicated that the two
modes were quite unequal, that one of them in-
volved far fewer individuals and lasted for a shorter
period of time. Among species of P. subg. Philo-
dendron, P. aromaticum appears to flower in the
mid-dry season and mid-wet season. It is possible
this species is just too poorly known to determine
its phenology. Philodendron morii may also flower
bimodally, with flowering collections seen in March
and November, and with immature fruiting collec-
tions in December, February, and June. Philoden-
dron wilburii appears to flower at the beginning of
the dry season and primarily later at the beginning
of the rainy season.
Several species are poorly known phenologically
because of sparsity of flowering collections: e.g., P
dominicalense, P. niqueanum, and P. ubigantupense
(all seen in flower only once during the early rainy
season). All of these are assumed to be species that
flower entirely during the rainy season.
Fruiting phenology in Philodendron subg. Philo-
dendron is too poorly known to report on here. Most
species appear to develop fruits from between one
and four months after the time of pollination, de-
pending on the size of the infructescence, but too
few mature fruting collections were observed to de-
termine the phenological period. Fruit development
time is affected by the size of the infructescence.
Species with small spadices, such as P. heleniae,
produce mature berries faster than species with
larger spadices.
CYTOLOGY
No karyological studies were made for this revi-
sion, but Petersen (1989) reviewed all chromosomal
literature and made new studies. The chromosomes
of P. subg. Philodendron are small, with counts of
2n = 30, 32, 34, and 36 (rarely 26 and 48). Petersen
(1989) speculated that the base number for the ge-
nus is 18. Very few of the 29 species (other names
were synonyms or hybrids) for which chromosome
counts have been reported are members of the genus
from Central America. Those Central American spe-
cies for which chromosome counts have been re-
ported are: P. radiatum 2n = 32, P. microstictum (as
P. pittieri) 2n = 34, P. verrucosum 2n = 34, and Р.
hederaceum (as P. scandens) 2n — 36. Philodendron
wendlandii was counted at 2n — 54, but Petersen
believed this to be either an error in counting or not
a Philodendron.
GEOGRAPHICAL DISTRIBUTION AND ENDEMISM
The Philodendron subg. Philodendron flora in
Central America is diverse but heavily concentrat-
ed in the southeastern part near South America.
Despite this, relatively few Central American spe-
cies actually enter South America. Most species
that do enter the South American continent range
along the Pacific slope into northwestern Colombia
and Ecuador, with relatively few occurring east of
the Andes and even fewer entering the Amazon ba-
sin.
As was shown for Anthurium (Croat, 1983a,
1986a, 1986b), species diversity of P. subg. Philo-
dendron shows a general diminution from Mexico
to Middle America, followed by a marked increase
approaching South America (Appendix 3, “Section-
al Classification of Philodendron”). Distribution of
Philodendron in Central America is as follow
Mexico has 21 taxa, Guatemala 15, Belize 9, El
Salvador 5, Honduras 13, Nicaragua 18, Costa Rica
48, and Panama 82. Thus species diversity is great-
est in Costa Rica and Panama, with 46% and 79%
of the total Central American species, respectively.
Collectively the number of species in these two
countries comprises 90% of the species total.
Fifty-three species (58 taxa) of P. subg. Philo-
dendron (nearly 59% of the total) are endemic to
Costa Rica and Panama. Endemism is particularly
high in Panama, where 38 taxa (34 species) of 81
47%) are endemic. In Costa Rica 7 of 47 taxa (46
species) (15%) are endemic. Costa Rican endemics
e: P. aromaticum, P. auriculatum, P. chirripoense,
P ка. P. dominicalense, P. microstictum,
and P. wilburii var. wilburii. Mexico has a higher
rate of endemism, with 7 of 21 (20 species) (33%)
endemic. Mexican endemics are: P. basii, P. breed-
lovei, P. dressleri, P. radiatum var. pseudoradiatum,
P. ћедетасешт var. oxycardium, P. sousae, and
subincisum. In Middle America little endemism oc-
curs. With the exception of Belize, which has one
endemic (P dwyeri) no other country in Middle
America has any endemic species.
Annals of the
Missouri Botanical Garden
The distribution of Central American Philoden-
dron reflects the trend for endemism in the genus.
Only 27 species (28 taxa) (a total of 26% of all
Central American species of P. subg. Philodendron)
range into South America, eight (7% of the total)
only to Colombia (Appendix 1). These are: P. ће-
leniae, P. immixtum, P. ligulatum var. ligulatum, P.
malesevichiae, P. mexicanum, P. pseudauriculatum,
P. radiatum var. radiatum, and P. squamicaule.
Sixteen Central American taxa occur in Ecuador
(Appendix 1), all but two (P. brevispathum and P.
jacquinii), ranging along the Pacific slope of the
Andes. These are: P. brunneicaule, P. dodsonii, P.
grandipes, P. hebetatum, P. platypetiolatum, P. pur-
pureoviride, P. hederaceum var. kirkbridei, P. hed-
eraceum var. hederaceum, P. squamipetiolatum, P.
scalarinerve, P. squamicaule, P. tenue, P. triparti-
tum, and P. verrucosum. Of these 14 taxa, 1, P.
hederaceum var. kirkbridei, skips Colombia or has
not yet been collected there. Philodendron dodsonii
is particularly unusual in being absent from Pan-
a.
Eight Central American species, P. brevispathum,
P. fragrantissimum, P. hederaceum, P. jacquinii, P.
jodavisianum, P. sagittifolium, P. strictum, and P.
tenue, range to Venezuela. The ranges of P. brevis-
pathum, P. fragrantissimum, and P. hederaceum
also extend into the Amazon drainage, while the
others occur either on the northern slope of the
coastal cordillera or otherwise in the drainage of
the Orinoco River Basin. Curiously only four spe-
cies, P. brevispathum, P. fragrantissimum, P. hed-
eraceum, and P. verrucosum, occur in the Amazon
drainage. Three additional species, P. jacquinii, P.
strictum, and P. tenue, occur east of the Andes, but
only along the Cordillera de Mérida, the Cordillera
de la Costa or the northern part of the Guiana High-
lands and within the drainage of the Río Orinoco.
The only truly widespread Central American
species of P. subg. Philodendron is P. hederaceum,
which occurs virtually throughout the Neotropics,
and is one of only two species (the other being P.
verrucosum) that occurs on both slopes of the An-
des. Philodendron fragrantissimum is probably the
next most widespread species, ranging from Belize
to the West Indies and into South America to the
Guianas, northern Brazil, and to southern Peru.
Further collecting in Colombia, especially along
the western slope of the Andes, will probably
change these statistics but the figures most likely
reflect the realities of life zone ecology and geologic
history of the area rather than under-collecting.
Since relatively few species of Araceae are known
to occur at lower elevations on both the eastern and
western side of the Andes, it can probably be pre-
sumed that the evolution of the respective Amazo-
nian and Pacific coastal floras occurred indepen-
dently after the Andes began to arise toward the
end of the Cretaceous (Raven & Axelrod, 1974).
The relatively few truly wide-ranging species, i.e.,
those ranging from Mexico to Brazil, appear to at-
test to this isolation. In Central America only one
species, P. hederaceum, really falls into such a cat-
egory, and it is also common in the West Indies
indicating that it may have an ancient origin (or be
easily dispersed). The high rates of endemism in
Costa Rica, Panama, and Mexico perhaps reflect
the isolation of these areas during periods when the
oceans were at much higher levels than they are
today and when the area that is now central Panama
and Costa Rica was disconnected from South Amer-
ica. Much of the present area of Central America
was submerged during early times. At the close of
the Tertiary, 800,000 years ago, sea level was about
100 m higher than today (Holmes, 1969). The land
mass of what is now Central America began to
emerge as a series of islands during the Oligocene
with further uplifting during the Middle Miocene.
It was not until the Upper Miocene and Pliocene
that the final portions of the isthmus of Panama
emerged above sea level (Torre, 1965), and the final
connection of Central and South America was made
about 5.7 million years ago. In order to place these
geological events in relation to the modern aroid
flora, it should be noted that even during this era
precursors to the existing flora probably already ex-
isted, since the angiosperm floras of the Oligocene
were believed to have consisted almost entirely of
existing genera, and the floras of the Oligocene and
Pliocene probably already had existing species
(Takhtajan, 1969). 5
Just as important as geology, from the standpoint
of the isolation of the Central American aroid flora,
are ecological factors that would cause Central
American species to be isolated from those of South
America. Much of eastern Panama consists of broad
expanses of Tropical moist forest with other, gener
ally smaller areas of Premontane wet and Tropic
wet forest. In contrast to Panama, much of the area
of northwestern Colombia in the Department of
Chocó consists of much wetter pluvial forest with
annual precipitation often exceeding 11,700 mm in
some parts of the region (Gentry, 1982). This broad
band of pluvial forest with its own suite of unique
endemic species no doubt acts as a barrier for af“
cies from regions with lesser rainfall amounts.
probably also accounts for the Panamanian of
Costa Rican species that skip the wettest pie
northwest South America but recur in the relative y
drier areas of mesic western Ecuador.
AAA
Volume 84, Number 3
1997
Croat 379
Philodendron Subgenus Philodendron
Just as the Central American P. subg. Philoden-
dron flora is rather isolated from that of South
America, there is a certain amount of isolation
within different parts of Central America. In com-
parison to Mexico and Costa Rica/Panama, Middle
America (Appendix 1) has low species diversity,
with Guatemala having only 15 species, Honduras
13 species, and Nicaragua 18. Most of the species
in Honduras, excepting P. anisotomum, P. mexican-
um, and P. warszewiczii, are shared with Nicaragua
(see Appendix 1). Nicaragua has eight additional
taxa not shared with Honduras: P. brevispathum, P.
grandipes, P. immixtum, P. ligulatum var. ligula-
tum, P. platypetiolatum, P. pterotum, P. tenue, and
P. wendlandii. All of the latter are shared with Cos-
ta Rica and Panama. Guatemala shares only about
half of its species with Honduras and Nicaragua,
namely P. fragrantissimum, P. jacquinii, P. jodavi-
sianum, P. radiatum, P. sagittifolium, P. smithii,
and P. tripartitum. Its other species are shared only
with Mexico (or rarely with Costa Rica and Panama,
e.g., P. mexicanum). These are: P. advena, P. ani-
sotomum, P. glanduliferum, P. mexicanum, P. pu-
rulhense, P. verapazense, and P. warszewiczii.
The low species diversity and the very low en-
demism in Middle America are perhaps explained
by the fact that Central America is rather more re-
mote from existing large land masses to the north
and the south, leaving it isolated from the indepen-
dent evolution that must have been taking place in
both of these larger areas (see below for a discus-
sion of the possible origins of the respective species
in Central American Philodendron). There is strong
evidence, at least based on the distribution of mod-
ern aroid species, that the northwestern part of
Middle America may have been isolated from Costa
Rica by the San Juan Depression. Many of the spe-
cies that occur in Costa Rica or Panama enter into
Nicaragua in only a small area in the southeastern
part of the country. Although the contemporary flora
of Guatemala does not reflect isolation from Mexico
to the same degree, it is possible that the more
elevated portions of Guatemala, Nicaragua, and
Honduras were isolated from major portions of
Mexico at the Isthmus of Tehuantepec
Certainly the Mexican aroid flora appears to be
quite isolated, even when compared to the western
parts of Middle America (here defined as Guate-
mala to Nicaragua). Mexico, in addition to having
one-third of its species endemic, has relatively few
species of Philodendron that range throughout Cen-
tral America. Aside from the aforementioned P.
to northern South America. Two additional species,
P. anisotomum and P. mexicanum, range as far as
Panama. Several taxa, P. advena, P. glanduliferum
var. glanduliferum, and P. verapazense, range only
to Guatemala. Philodendron purulhense and P. war-
szewiczii range to Honduras, and P. smithii ranges
to Nicaragua.
Taken together, Honduras and Nicaragua have
21 species of Philodendron. Of these, 8 species, P.
advena, P. anisotomum, P. mexicanum, P. radiatum,
P. sagittifolium, P. smithii, P. tripartitum, and P.
warszewiczii, appear to be of Mexican origin, or in
the case of the more widespread and variable spe-
cies, namely P. radiatum, P. sagittifolium, and P.
tripartitum, they may have originated in Panama or
Costa Rica and ranged to both Mexico and South
America. Certainly, in terms of morphological vari-
ation, all of these species are much more variable
in Panama and Costa Rica than they are further
north. Philodendron jacquinii has а circum-Carib-
bean distribution, indicating that it may be of West
Indian origin. It is difficult to determine the origin
of P. hederaceum given its extensive distribution.
Three species, P. brevispathum, P. fragrantissimum,
and P. tenue, probably originated in South America
considering their widespread distribution there.
Philodendron platypetiolatum, ranging from Ecua-
dor and barely entering Nicaragua, may be another
South American derivative. The remainder, P. an-
gustilobum, P. grandipes, P. immixtum, P. јодат-
sianum (ranging barely to Chiapas and rare there),
P. ligulatum var. ligulatum, P. pterotum, P. roths-
chuhianum, and P. wendlandii, are probably of
Panamanian or perhaps Costa Rican origin.
The Costa Rican and Panamanian species not
already discussed above appear not to have strong
affinities with South American species, and clearly
did not originate in areas of Middle America. Much
of the flora of adjacent Nicaragua is closely related
to that of Costa Rica. Except for those rather wide-
spread species mentioned above, i.e., P. brevispa-
thum, P. fragrantissimum, and P. tenue, as well as
P. platypetiolatum (already discussed), there are
relatively few species likely to be of South Ameri-
can origin. Philodendron verrucosum is almost cer-
tainly a South American species, since it is rela-
tively widespread there, occurring on both sides of
the Andes. In addition, it seems to have more re-
lated species in parts of South America especially
in the Andes of western Colombia.
Philodendron dodsonii, which occurs in Ecuador
but not Colombia, is just as likely to have origi-
nated in South America as in Central America. The
same is true of P. strictum, which is known from
eastern Venezuela and has relatives in the Andes
of central Colombia, as well as P. hebetatum, which
380
Annals of the
Missouri Botanical Garden
is rather widespread in western South America as
far south as Ecuador. Moreover, the latter appar-
ently does not even reach Costa Rica, making the
case for a South American origin even more likely.
Philodendron grandipes, known from western South
America as far south as Ecuador, might conceivably
be of South American origin despite being very wide-
spread and common as far north as Nicaragua. More
he of South American origin is Р. heleniae,
which is common in Colombia and ranges only to
western Panama. Moreover, it seems to have a close
relative on the eastern slopes of the Andes.
Three species with scaly parts, P. malesevichiae,
P. squamicaule, and P. squamipetiolatum, are mod-
erately rare in Panama, and though all are still
poorly known in Colombia, they are more likely to
have originated in northwestern Colombia where
there are several other relatives with scaly parts.
Some taxa, such as P. ligulatum var. ligulatum,
P. pseudauriculatum, and P. scalarinerve, either
barely enter Colombia or are rare there and are
more likely therefore to have originated in Central
America. All the remaining species of Р. subg.
Philodendron occur in Panama or Costa Rica, with
11 species shared between the two countries.
TAXONOMIC TREATMENT
Philodendron (“Philodendrum”) Schott, Mord
Z. Kunst. 1829: 780. 1829, nom. et orth. c
: P. grandifolium (Jacq.) Schott @
ond Jacq.)
Telipodus Raf., Fl. tellur. 3: 66. 1836[1837]. TYPE: T.
bid (Jacq.) Raf. (Arum grandifolium Jacq.)
Thaumatophyllum Schott, Bonplandia 7: 31.
TYPE | Т. eon Schott
Elopium H. W. t, Oesterr. Bot. Z
34. 1865.
TYPESE surinamense (Miq.) Schott ом sur-
inamense Mi
Baursia Т. Post & Kuntze, Lex. gen. phan. 62. 1903.
PE: Caladium bauersii Rchb.
Appressed-climbing or scandent hemiepiphytes,
sometimes epiphytes or terrestrial herbs rae
stout and arborescent); growth sympodial, an
phyllous, or (more commmonly) diphyllous; cree
chlereids absent, biforines and secretion files pres-
ent. Leaves with spiral phyllotaxy; petioles rarely
geniculate, sheath obsolete or extensive; blades
with parallel-pinnate venation, hypostomatic with
stomates paracytic, simple to trifoliolate, pinnatifid,
bipinnatifid, or spo pedately compound. Inflo-
rescence terminal (appearing axillary), solitary to
multiple in leaf axils; spathe enveloping the spadix
and constricted or not, usually persistent: spadix
monoecious, with sterile male flowers below or both
below and (rarely) above the fertile male region;
flowers naked, unisexual. Male flowers with 2-6
free stamens, connective enlarged; anther dehis-
cence by apical slits or pores; pollen inaperturate,
boat-shaped, subisopolar, bilaterally symmetrical,
shed in monads, medium-sized (mean 40 um; range
28-54 um), the exine coarsely verrucate to subfov-
eolate or (most р psilate; pim at anthe-
sis binucleate and starc ing. Female flowers
virtually always lacking ава ovary (
6(47)-locular, each locule with axile or (rarely) ba-
sal placentation and (1—)4—51 or more orthotropous
or hemiantropous, endospermous ovules. Fruit a
berry, usually white or orange. Chromosome num-
rs 2n = 30, 32, 34, 36. [Description adapted
from Grayum (1996).]
KEY TO PHILODENDRON SUBGENERA
la. Stem of mature flowering plants with a succes-
sion of many leaves terminated by solitary or
ioles with
subg. фон
. Stem of mature flowering plants with a s
—
=
cence(s) 1 to 10 and appearing to be borne in
the leaf axils; petioles of жы) of adult plants
with a short, usually inconspicuous petiole
e stem, not encir-
with conspicuous
leaf scars and iid ae scal
persisting around at least the upper margins
of the petiolar scars; male flowers conspic-
uously elongate, up to 10 times longer than
wide; staminodial zone between staminale
and pistillate zones of the spadix subequ
or longer than fertile zone ___-----------
nostigma
~
=
у
@
5
Ф
~“
d
<
=
Philodendron
Philodendron subg. Philodendron Schott.
Philodendron subg. Euphilodendron Engl.
Bot. Jahrb. Syst. 26: 509. 1899. Baursia 5
Eubaursia EARS in T. Post & Kuntze, Lex
gen. phan. б
Appressed hemiepiphytic climbers or vines with
aerial roots, less frequently terrestrial with creeping :
rhizomatous or deeply rooted stems, rarely >
Volume 84, Number 3
1997
Croat 381
Philodendron Subgenus Philodendron
stemmed true epiphytes, rarely somewhat arborescent
in Central America; sometimes with flagelliform
shoots; sap usually tanniniferous and drying dark
rarely with latex and drying white; stems of mono-
phyllous sympodia with elongated hypopodial inter-
nodes, densely rooted at nodes; internodes often much
longer than broad or about as long as broad, some-
times broader than long at anthesis, sometimes flat-
tened on one side, often coarsely pale-streaked just
below the node, usually green and semiglossy, but
often turning gray-green to brownish or reddish brown
in age; juvenile plants terrestrial or epiphytic and
scandent, the petioles conspicuously sheathed and
subtended by inconspicuous intravaginal squamulae;
cataphylls of mature stems unribbed or variously
ribbed, caducous, marcescent and deciduous or per-
sistent and membranaceous to moderately coriaceous,
remaining intact or more commonly decomposing to
net-like, persistent fibrous reticulum. Leaves usually
long-petiolate; petioles usually with ligulate sheath in
juvenile plants, on adult plants usually sheathed only
at base, variously shaped in cross section, firm or
spongy, usually smooth, frequently densely pale-short-
lineate or pale-striate throughout, sometimes warty or
covered with scale-like processes, rarely geniculate
apically; blades simple and entire, ovate, cordate, has-
tate, sagittate, oblong to elliptic or variously divided,
trifid, trifoliolate, or pinnatifid; midrib raised ог sunk-
en above, raised below; primary lateral veins pinnate,
usually conspicuous, spreading to the margins and
running into an antemarginal collective vein; lower-
most primary lateral veins (basal veins) often coa-
lesced on cordate blades, the posterior rib (coalesced
basal veins) naked along the sinus or not; interprimary
veins sometimes present; secondary lateral and higher
order veins transversely reticulate between the sec-
ondary veins, sometimes all veins slender with no dis-
tinct primary lateral veins; minor veins conspicuous
or obscure, usually fine and closely parallel; cross-
veins (minute veins extending transversely between
the minor veins) sometimes visible; secretory ducts
sometimes appearing like veins, linear, short to long,
obscure to very distinct on lower surface. Inflores-
cences 1-several per axil, usually much shorter than
the petioles; peduncles shorter or longer than the
spathe; spathe erect, usually coriaceous, entirely per-
sistent, often with large superficial resin canals on
inner surface which exude resin, opening widely at
anthesis (usually for about one day), then reclosing
and persisting in fruit, deciduous only on ripening of
fruit, frequently colorful, often bicolorous on outside,
typi somewhat constricted between tube and
blade, convolute at base; tube cylindric to inflated,
uniformly greenish to red ог ol pl т»
blade usually opening widely, becoming +
~
shaped at anthesis, usually white within, sometimes
tinged reddish; spadix sessile to stipitate, divided into
pistillate and staminate portions, each with unisexual
flowers; pistillate zone usually greenish, obliquely
fused at its base to the spathe, free above, usually
much shorter than the staminate portion and separat-
ed from it by a sterile zone of staminodial flowers;
intermediate sterile zone cylindric to ellipsoid, much
shorter than staminate zone in Сеш AERA, usu-
ally thicker than staminat
to clavate, white, usually PON constricted above
the sterile staminate zone; flowers unisexual, naked,
lacking stomial groove, prismatic to obpyramidal; an-
thers tetrasporangiate (with microsporangia embedded
in the abaxial surface of the anther), columnar in
shape, elliptic, ovate to rhombic in cross section, ses-
sile to subsessile; connective thick, apically truncate,
usually irregularly 4—5-sided, overtopping thecae; the-
cae oblong or elliptic, emarginate at the base, dehisc-
ing apically by short, ragged lateral pores; endothecial
thickenings lacking; pollen extruded in strands or
mixed with resin secretion or exuded in amorphous
masses, inaperturate, ellipsoid or oblong or occasion-
ally elongate, medium sized (mean 40 jum, range
28-54 рт), mostly perfectly psilate, sometimes mi-
nutely verruculate, scabrate or fossulate to clearly
punctate, subfossulate, subfoveolate or subverrucate;
sterile staminate flowers naked, usually prismatic,
truncate and usually more irregular than fertile flow-
ers and lacking thecae; pistillate flowers with ovary
syncarpous, ovoid, subcylindric, cylindric or obovoid,
3-9(14)-locular in Central America (2-locular in P
sect. Philopsammos in South America); carpels pre-
sumably equal to number of locules; placentation ax-
ile, sub-basal or basal; ovules 1 to numerous (to ca.
30) per locule, usually hemiorthotropous, rarely hem-
ianatropous, ascending on moderately long or some-
times short funicles; stylar region as broad as or some-
times slightly narrower than ovary; style short,
unlobed, with or without boss (see definition under
Style Type D), funnel, or annulus; central style dome
usually lacking in Central America; stigma sessile,
hemispherical to lobulate. Berries subcylindrical to
obovoid, exposed by the re-opening of the spathe,
white, whitish translucent to red or orange; seeds few
to many per berry, oblong to ellipsoid or ovoid-oblong,
testa rather thick, striate-costate, rarely sarcotestate;
embryo axile, straight, elongate, endosperm copious;
chromosomes: 2n = 30, 32, 34, 36, (26, 48). Species
ca. 700, Central Mexico to Argentina; West Indies
(occurring in all countries of Central and South Amer-
ica except Chile and Uruguay).
382 Annals of the
Missouri Botanical Garden
Кеу а -Враре Drem r Lonbo oR DIVIDED а аа Se ee A
Key 2: BLADES SIMPLE, NON-CORDATE, EITHER OBLONG OR OVATE; SOMETIMES SUBCORDATE TO
3
CORDULATE AT BASE, THE POSTERIOR LOBES UP TO М AS LONG AS ANTERIOR LOBES —
Key 3: CORDATE BLADES WITH PERSISTENT CATAPHYLLS
KEY 4: VINES WITH CORDATE BLADES AND DECIDUOUS CATAPHYLLS
KEY TO PHILODENDRON SUBGENUS PHILODENDRON OF CENTRAL AMERICA
A. Blades deeply divided, either pinnately lobed, or а 2 to trifoliat
A'. Blades not deeply divided, at most a the mar erely prend the lateral margins of the anterior lobe
sometimes deeply concave, but not to such an 5 ert т blade looks decidedly 3-lobed (blades at first
entire, but ea канш». into irregular T in P. ns).
B. В! ades LI t E E. = XU ин 6E. ai.
netimes WCaRILY cordulate or
lobes eb M iu ни nd as the anterior lobe o
KE
B'. Blades жа ба sagittate, ог hastate at deb tum = тын lobes usually more than М as long as the anterior
lobes.
Y 1
C. Cataphylls persistent on stem, either intact or as fibers KEY 3
C'. Cataphylls ultimately deciduous, though sometimes persisting . KEY 4
Key 1: BLADES DEEPLY LOBED or DIVIDED
la. Blades deeply ienie lobed, the divisions extending at least E of the way to the midrib
2a. — bipinnately gu th y lobed, usually divided
e than %-way to b
3a. “Blades lobed ud Ума По the way to the midrib; Mexico (Sinaloa & Nayarit); 0-370
3b. Blade deeply lobed, the divisions extending more than 7, of the way to the m
Blades de
ciduous during the dry season; leaf segments thin, drying pot lit pale yellow-
ellow-brown, with Lai minor veins distinctly visible and darker than surface; lateral
segments with usually 3 o e lobes side, extending to below the middle “of the pinnae;
Mexico (Jalisco & Chispas) ө “El a Guatemala, Honduras, and Nicaragua, 300-1
JR uds G. S. Bunting
500 m
ua E UE A ie ol E E OA i P. warszewiczii K. Koch & Bouché
~
=
. Blades evergreen; leaf blade segments subcoriaceous, drying moderately thick, dark brown or
sometimes reddish brown, the minor veins not distinctly visible on drying, not markedly darker
than qui lateral segments with usually 1-2 lobes e "em аву ко to near the apex
of the pinnae; Mexico to Colombia , 0-700, rarely to 1
if so
lb. RES = 3- ey or trifoli es
Blade a su medial lobe less than 1.7 times longer than the lateral lobes; lateral lobes directed =
owa
6a. ima bes of the blade broadly confluent with the medial lobe by at least 2 ст from the base of
the blade te —— from the depths of the incised area and the apex of the petiole); inflorescences
«gr à 3-9 il.
7a. Medial ie of the blade with 5-12 pairs of primary lateral veins; spathe tube uniformly greenish
to whitish within; Honduras to Panama, usually below я
Tb. ct lobe ob the blade with 18-19 pairs of Pape lara v veins; spathe tube red-violet
e
P
more than cm from the base of the blade; inflorescences usuall
y bra ies to 3 p
8a. Medial сар of blades with more than 18 pairs of primary lateral veins, elliptic: ol mue .
pe
m var. ees
а rothschuhianum (Engl.) Croat & ud
siad dama мы O madronense Croat
pairs of primary lateral veins, usually penis oe to
oblo ES ides aee Mexico to Ecuador.
9a. M нч ms
bro
directed + = toward the apex; minor veins m jai both the midrib and the primary lateral
veins; ripe fruits whitish; Mexico to Ecuador, 0-1300(15 ) m |
Medial lobe mostly 1.5-2 times longer than broad; pri
mostly 2—4, rarely 5 per side, weakly sunken; lateral lobes directed + outward; minor veins
arising only from the midrib; ripe fruits orange; Mexico to Costa Rica, 30-1800 m -----------
о
=
eee erc -—_ - к P. anisotomum Schott
Volume 84, Number 3 Croat 383
1997 Philodendron Subgenus Philodendron
5b. Blades with the medial lobe more than 1.7 times longer than the lateral lobes; lateral lobes directed
efle
inier (spreading) outward or even downward toward the base of the petiole (r
p a "aped the lateral lobes directed downward somewhat toward the base of blad (in direct
iole); M o Panama and South America, 0–1900 m .... Р mexicanum Engl.
10b. Ble Кадык: ihe pe lobes directed outward at ca. 90? angle.
. Latera obes epee with the medial lobe by usually for more ed 4 cm (rarely to 3.3 cm);
basal v 1 side; Honduras to Panama, 0–680 m ___. P. angustilobum Croat & уча
11b. MP po KL eR with the medial lobe E usally less than 3 cm ушга to i cm); basal
eins 4—7 per side; Mexico to Costa Rica, 30-1800 m P. anisotomum Schott
Key 2: BLADES ENTIRE OR SHALLOWLY DIVIDED, NON-CORDATE, EITHER OBLONG OR OVATE; SOMETIMES SUBCORDATE
OR CORDULATE AT BASE, THE POSTERIOR LOBES UP TO %4 AS LONG AS ANTERIOR LOBES
la. Plants vines or at least with internodes much longer than broad.
VINES OR SCANDENT: INTERNODES MUCH LONGER THAN BROAD; BLADES OVATE
2a. Blades = ovate, less than 2 times longer than wide.
За. Blades with primary egos veins И d or fewer or with the primary lateral veins inconspicuous,
а more distinct tha
4a. es more than le cm wide; аб сай more than 10 cm long; Costa Rica, 50-450 m........
us Icones IIT CL Williams
~
=
Blades less than 11 cm wide; ig ag less than 10 с
5a. Internodes less than 10 cm 1 ong; blades less hen) 1. 4 times longer than wide, drying
greenish, licking primary lateral veins; Panama, Cerro Brewster, 850 m ...................
P. brewsterense ^us
In d more than 10 ст long; blades ca. 2 times longer than wide, drying bro
Es to 2 obscure primary lateral veins; Costa Rica, Río ерта del Pacífico, San
Jos 1000 m irripoense Croat & tasen
dim with M пе lateral veins more than 2 and much more (cA з than the mino
{л
E
3b.
ag “Blades ovate to broadly ovate, оу less than 1.7 times longer than wide.
Stems, petioles, and inflorescences densely scaly (petiolar pee Fender and spread-
ing: pus veins on ler phe densely puberulent; Panam uador, 0-1300
US o uamipetiolatum Croat
7b. Stems, petioles, and рака glabrous; major veins оп lower surfac e glabrous;
P. ligul
Panama (Coclé, Veraguas), 770-1200 m ------------+--------+----- igulatum var. ovatum Croat
6b. Blades a ovate to gp ne: яң usually more than 2 times longer than wide.
8a. Blades jer than 12 wide; posterior lobes about as long as broad; Panama to
Colombia, 0-140 m ...... mmixtum Croat
8b. чи ену more than 12 cm mcs posterior lobes much gunk than Гаж Costa
to Panama (Тобто 18000 а Р. lenti Croat & Grayum
Ric )670-1800
2b. Blades + oblong to oblong clips (rarely diam Ys ovate) or oblanceolate, usually more than 2.5 times
longer than wide.
VINES WITH + OBLONG BLADES
9a. Blades lacking distinct primary veins at base of blade; primary lateral veins obscure or
lackin,
10a. Blades cc. gray-green; primary lateral veins not at all apparent; dens = as),
evel .. атирепзе Croat
10b. Blades diving reddish brown or blackened; primary lateral veins 3-6, ieee but still
Па. Blades phan blackened, usually more than 25 cm long; spathe more than 12
m lon ama (Bocas del Toro and Chiriquí), 780-1400 m ....... P. correae Croat
11b. Blades oon reddish brown, usually less than 25 cm ев spathe less than 12
m long; Costa Rica (100-900 m) to Panama, 900-1420 m -..............-.-------
E A nt P bakeri Croat & Grayum
9b. vii with one or more distinct primary veins at base of blade; primary lateral veins
distin
12a. Раше encircled си with a dark purplish (or dark green) ring separating the
iole and the blad
13a. Blades " sei rior lobes usually narrowly rounded and somewhat vengan у
or b r than long, rarely about as long as broad; inflorescences - nius xil;
и 100-970 m ә А ОНИ T. a Croat
13b. Blades with posterior lobes about as broad as long and a i closely near
petiole; inflorescence usually solitary n r EMER
Ta. Blades typically drying red] А eng brown, "rarely n ys tica
cm long; Panama to Colombia, 0-140 m .....................------- pium ‘rot
14b. se brand ot ba едет ted less than 30 cm a
А E P. ligulatum > var. аала
12b. Petioles ш а xul: hears distal ring.
384 Annals of the
Missouri Botanical Garden
15a. Sede less than 10 ст wide; stem minutely pale granular-puberulent at high
gnific
cation; aet pda a funnel-shaped stigma (type E); ovules 1 per locule;
Е ER EA Ба ranulare Croat
15b. Blades afr ip more 15 cm wide (rarely to as little as а sh { in P.
heleniae); stems smooth to mery ridged or folded at high m ен
(sometimes minutely warty but pale gran ia pube; stil with a flat
style apex (type B); ovules 3 or more per loc
6a. oe drying coriaceous, lacking ere ducts; spathes „Яа ка
Аер outside, more than 10 ст long; Costa Rica to ey (210
ааа > lentii Croat & pee
16b. Blades. drying subcoriaceo icuous secretory ducts; spathes red
outside, mostly less than 10 c cm vilium Panama and Ecuador, mar n
1040 m ae Croat
lb. Plants appressed-climbing or sometimes terrestrial, with the internodes typically broader than long or ip
slightly longer than broad.
NON-VINING PLANTS; BLADES OBLONG
17a. Cataphylls deciduo
18a. Blades ja pai tha lobed at base, the lobes typically longer than broad or at least usually
spreading away from the репоје.
19a. снг x thin-drying sinus arcuate with blade tissue nce decurrent on petiole; petiole
dry m diam.; Panama, E of Canal Area, 450-850 m |... P. morii Croat
19b. Blades uk. HET. sinus deco to parabolic or arcuate Ба with blade tissue not at all
decurrent onto petiole; Panama, 100-970 m › annulatum Croat
18b. ae not lobe por or e ا with posterior lobes round, as broad as ee held
c ioles.
20a. Largest leaf blades more than 50 ст basi
21a. Blades usually more than 30 cm wide; petioles usually about E the dena of the
blades or even [ine than the Ызы, Costa Rica and Panama, 0-200 m...
со: P davidsoni Croat
21b. e" less than 30 cm wide; виа typically much shorter than the bis
egens ips reddish brown, 4.4—4.9 times longer than wide; er lacking
tal ring; Panama, Mia o olichophyllum ‘Croat
22b. Blades rias dark gray-green to blackened, preci less than 4.3 times longer
than wide; petiole with a purple or green distal ri
23a. Petioles subterete adaxially, not at ‘all еі) marginally Panama and Co-
lombia, 20-1400 т __ pseudauriculatum Croat
23b. Petioles ери D-shaped with slender wings on the ial m argins; Pan
4, Darién, 50-2009 SS n. P. ligulatum var. heraclioanum Croat
20b. Largest leaf Ниве esu pers wie than 50 em lon
24a. = es usually drying green to yellowish green, thin
. Blades oblong- lanceolate, less than 7 cm wide, obtuse to almost rounded at base
nt along petiole; Panama (Coclé), 700-800 m P. folsomii Croat
25b. Blades оше апи, 11-16 ст wide, зек ie at base; eastern Panama,
morii Croat
24b. Blades dying adie EEE or brownish, gray-brown to reddish brown, moderately
сопасе
26a. Infrescences 2—10 per axil; peduncle less than б ст long; spathe tube reddish
ma outside and within; leaf blades with conspicuous secretory ducts visible
on on the lower dried surface; Panama to Colombia and Ecuador, 20-1 Pes 1450) :
qe гоа
17b. e persisten
ts usually coe and rosulate; petioles deeper than wide; blades usually elliptic to oblong-
di ог acute to narrowly rounded at base; Panama to Colombia, 100-1000 m
NOE A O pu eospathum \ Croat
27b. Murus epiphytic to hemiepiphytic, not rosulate; petioles as deep as wide; TRO шр to oblan
eolate or oblong-elliptic, some ма weakly cordate to cordulate at base.
angle om sinew to oblan eolate, attenuate at base; p rimary lateral v veins aris ing at 25-45
n
angle; cross-veins very conspic d a t colored, or if so
not ue de и Ps to Colombia, oe ong а i suh T Som
29a. ride usually minces t above the middle, acute, obtuse, intus frat le to merely
utely cordulate at
30a. Pris sharply узана: da adaxially (with acute lateral margins); blades acute to
arrowly rounded (never cordulate) at base, mostly more than 2.5 times longer
Volume 84, Number 3 Croat 385
1997 Philodendron Subgenus Philodendron
than petioles; — oe than long; Costa Rica, Nicaragua, ino Panama.
Atlantic slope; 1 P. wendlandii Schott
. Petioles Б ез mein narrowly cordulate at base, less than 1.8 esa longer
ida es often longer than Spa ; sout thwestern Costa oe , Pa-
cific lo se s, 0-1200 res ава аи riculatum Standl. & L. 0. "Williams
29b. Blades broadest s the ae cordate to a ee at base (acute to truncate in
davidsonii subsp. bocator
3la. Blades ovate to proudly жаба less than 60 cm long, drying dark brown; үрти?
lateral veins 5—9 side; petioles terete; Panama (Darién), ca. 1500 m ____
anum Croat
31b. io enc -oblong, ge more than 60 cm long, drying light es нозе
lateral у 18-21 per side; petioles thicker than broad (i.e., with the dichos
и ect ato to the Pune of the blade) and broadly sulcate adaxially;
Costa Rica to Panama, 0—2 P. davidsonii Croat
w
©
c
KEY 3: CORDATE BLADES WITH PERSISTENT CATAPHYLLS
la. Blades with posterior rib (union of basal veins) naked along the edge of the sinus.
2a. Petioles with conspicuous, elongate, hair-like scales.
3a. Pet
iolar scales scatt wel ~ mostly in узе Boo; % of petiole; plants eot Panama and
Co lo mbia, 830—860 m nama, 50-15 n Colombia male. д Croat
3b. Petiole scales dense, c ie the entire кз pem wi hemiepiphytic.
4a. Blades ovate стаг typically more than 1.3 times longer than broa d, semiglossy di
00-1250 m
paler and solid light green to silvery-green Spork Chae Rica to е <1
Pis аше Croat & Grayum
4b. Blades ovate, typically less than 1.3 times longer than broad, cae ind subvelvety above,
paler and tinged with purplish violet between the major "n ei bene не Rica to Peru
(200)500—1500 та SE се О ig Не m L. Mathieu ex Schott
2b. Petioles glabrous or least lacking scales
ба. Petioles steal and sharply flattened adaxially, with lateral margins sharply edged or with
ender w
6a. Petioles merely sharply gnum lacking an actual Mine at the margin.
Ta. Blade
es whitish and m devia cataphylls persisting in semi-intact fragments of thin
ermis; spathe ses green i at most tinged pink within; Costa Rica to Pan-
ama prone a а Р. thalassicum Croat & Grayum
ТЬ. Blades n and semiglossy below Mone persisting in a dense reddish brown,
semi- etd pl 8: ~ ae tube gr bright “ outside, dark red to maroon within,
Fue white; Belize о South is a. 0-900
Blades o ee 1 82. T times diee than wide; cross-veins ome
between minor veins on dried —— dep ge tube green on м Рапата
(Bocas del Toro to Coclé), copense Croat
8b. Blades moiy ovate, 1. 1- Је 7 e Я than wide; cross-veins weal ween minor
veins not a pp Belize to South America,
de pro да (Hook.) G. Don
0-1000 ш...
6b. Petioles with a narrow nal w
9a. Leaf blades pr ib nh ы p slender segments laterally, drying blackened; pet-
iole wing undulate distally; Costa Rica to Panama, Atlantic slope, mos.
Р. ns Croat & Grayum
9b. Leaf blades remaining intact, drying yellow-green; petiole wing usually ME
throughout its entire length; Nicaragua to central Panama, mostly less es
P. pterotum K. Koch EA Мина
5b. Petioles terete to obtusely flattened or U-shaped but not sharply attend adaxially, if sulcate
use
10a. Blades whitish and matte o n lower surface even on е leaves; petioles usually drying
light yellowish brown иа — not in P. s
te ase €— ovate, үе re than 1.8 times nae than wide; plants commonly
miepiphytes; Panama to Ec ndn. — «Mm - — . — atum Croat
11b. Blades ovate to broadly ovate, averaging 13 times longer than wide; typically terres-
cies Costa Rica to western Panama, Colombia, and Venezuela, mostly at 850-1665
чине RA O AS NE P. strictum G. id ning
10b. Blades. мы to yellow-green beneath, usually semiglossy to glossy; petioles desing va
colors, not light yellowish brown (except P. copense, P. schottianum, and P. thalass лага
NARROWLY OVATE BLADES
12a. Blades жщ 1.8 times or more longer than wide.
13a. e tube greenish to yellow-green or whitish inside.
. Leaf blades with lower surface matte, frequently bluish green; style apex prolonged into a
short but distinct neck (style type D); central Costa Rica to western Panama, (775)1000—
2100 m P. thalassicum Croat & Grayum
386
Annals of the
Missouri Botanical Garden
14b. Leaf blades with lower surface glossy ~ semiglossy, not at all bluish green; style apex flat,
not prolonged into a distinct neck (style type B; rarely type C); Panama, except P. jodavi-
sianum (Mexico to Venezuela).
15a. Blades drying yellow-green; basal veins fewer than 5; sap milky white, drying chalky;
inflorescence solitary; cataphylls to 40 ст long; Panama (Darién), a 480 m ___
cid а albisuccus Croat
. Blades drying dark gray-brown to olive-green, typically somewhat iban basal
veins usually more than 5; sap usually clear, drying dark brown to reddish; inflores-
cences 2-6 per axil; cataphylls less than 20 ст long
16a. cas terete to obtusely flattened adaxially, lacki ing prominently pus lateral
margi про преке frequently longer than the spathe; central and eastern Рап-
E nse K. Krause
16b. Pails жые D- shaped or U-shaped, usually flattened adaxially pese promi-
ntly raised margins, often thicker than broad, rarely C-shaped to subterete;
зе usually much shorter than the spathe; S Maio to Panama and Ven-
ezuela (Mérida), 0–1500 m P. jodavisianum G. S. Bunting
13b. Spathe tube red to maroon or violet-purple on inside.
17a. Blades drying алги to ا (sometimes yellow-brownish in P. alticola), lacking
conspicuous cross-v
18a. Pistils with 1 poer per locule; leaf blades with secretory ducts moderately E
oit
—
сл
=
оп;
doughnut-shaped with stylar canals at the bottom of a deep concavity; Panama (Chi-
i P. straminicaule Croat
. Pistils with 12-18 ovules per locule; leaf blades with secretory ducts conspicuous,
abaxial surface smooth on drying; staminate portion of the spadix scarcely constricted
above the sterile portion, the fertile portion stubby and evenly tapered to the apex,
1
Y thicker);
de ed id base raised hut fattened at apex with a Barrow pale ring around its оде
sarph, the stylar canals e
rn Costa Rica pe western yin 800-2500 m ni Р alticola Croat E ue
17b. Blades TE reddish brown with conspicuous cross-vein
isp drying with a conspicuous light reddish жалшы ог yellow-brown epidermis,
smooth and often "ekine cataphylls Е than 25 ст long; primary lateral veins
11-16 per side; basal veins frequently more than 8 per side; Panama үрү del Toro
and Coclé Provinces); 590-930 m P. copense Croat
rown, the epidermis not peeling; cataphylls more than 55 cm
long; primary lateral veins fewer than 8 per side; basal veins up to 8 per rps Panama
(Chiriquf, Bocas del Toro, Coclé, and Veraguas Provinces), 500-1630 m __----
Ile ЕУ O E ao quce dE P. сећа ин Croat
—
со
c
5
т
¿E
EES
7
e
d
i=]
oa
m
4
m
OVATE TO BROADLY OVATE BLADES
12b. MM usually about 1
. Leaf bla
1.5 times longer =“ wide or less (sometimes wider than long).
des with lower surface matte (but never velvety), frequently bluish green, the dried waxy
surface forming an areolate pt central Costa Rica to western Panama, н 1000-2100 m
Р
п.
Mein matte and sc above; central Panama in the region of the isthmus, 300—
а м. P gig as Croat
a uncles ни ually EA s than 10 cm long (rarely longer in fruiting peduncles of Р.
23a. Spathe tube solid reet red on outside; central Panama Mi ger and Coclé),
pat ы or not at all constricted above be.
25a. phylls persisting usually with lar arge e fragments of Lens yellowish
e petioles drying yellowish to yellow nts
as if with a layer of shellac; Costa Rica to Panam m
vu uud uU E pP schot piu d ofer ex Schott
; Cata phylls persisting semi-intact and brow A odiy ellowish;
свет usually drying brown to HETS, init д not yellowish and
1870 rulhense Croat
N
сл
c
ш
= drying ques -brown below. екл ovules mostly
а рег locule; Isthmus of Ране, нони below 500 m —--
р e ы _ P Џапепзе Croat
Volume 84, Number 3 Croat 387
Philodendron Subgenus Philodendron
26b. me blades drying reddish brown, moderately thin; ovules 1-6 p
ig dus P. dodsonii with ca. 20 ovules per locule); Costa Rica
mn tls sually growing over rocks in dry habitats, rarely on
trees; cataphylls eventually deciduous, intact on the older stem;
ules 4—6 per locule; western Mexico (Jalisco & к
350-1250 т P. basii Matuda
. Stems appress sed- -climbing on trees in humid habita ats; cata-
phylls decomposing and breaking up before falling off; ovules
either solitary or ca. 20 per locule; Costa Rica and/or Ecuador.
28a. Spathe tube green; ovules 1 per locule; posterior rib bare-
y or not at all naked along the sinus (up to at most 2.5
cm); southwestern Costa ip surdi ‚о ee
nicalense — & Grayum
28b. Spathe tube dark reddish to МА прасе vules са. 20 per
locule; posterior rib naked to ca. 6 cm along у sinus;
Арчи Rica, principally оп the Atlantic slope at 990-090
and Colombia and Ecuador at 200-1300 m _____
E ib & Grayum
22b. e usually more than 10 cm long (rarely less than 10 cm long) when not at
~
“з
c
esis.
294. = clothed = a dense series of overlapping cataphylls, EA persisting as
reticulum of co bers often overlain with a thin, fragmented epidermis
30a. ma 1 2-1: 5 times longer than wide, drying reddish ivit, ya Pan
ma (Darién), 1000-1560 m rrense Croat
30b. Blades 0.8-1.4 times longer than wide, drying grayish green a 2 ve.
3la. Dried tire with cgi surface semiglossy, sparsely granular, with
moderately conspicuous cross-veins, not speckled, the upper surface
ene raphide edis ovules 6 per locule; Panama, Pan за. Provin
(Cerro Jefe), 550-800 m P. је llenos det
31b. Dried blades with lower surface matte, smooth “ minutely brownish
to whitish speckled, lacking obvious cross-veins, the upper oe
usually with short whitish raphide cells vis ible: ovules 14-20 pe
locule; Panama, Canal Area to Darién Province (Cerro Sapo), 140—
m P. lazorii Croat
29b. Stems with oA a few cataphylls, these usually semi-intact or with a few, thin,
sg tpe
Blades drying и ои to dark yellow-brown; peduncles usually less
than half as the spathe, straight below the spathe; т, d
ees йү» > id aem Costa Rica, Colombia, and Ecua dor
dson ii Croat & а
32Ь. Blades oe dark gray-brown to olive-green, typically RH black-
d above; peduncles usually as long as or longer than the spathe, fre-
quent beni nn below the sé ah о alo ii у оп анана Рап-
тепѕе К. Krause
lb. Blades with posterior rib (union of R А not naked along the sinus, flanked with ade tissue.
33a. Petioles conspicuously scaly distally; terrestrial (except P. verrucosu
34a. Blades less than 25 cm long; petiolar scales close and overlapping broad, usually less than 3
i opé regi а. 800 т P. ћатте Croat
tim
34b. Blades usually more than 30 cm long; petiolar scales more diy scattered, many times longer
han wide.
35a. dum mes stems usually аныны, сазове! across the ground; spathes glabrous; Mex-
and Guatemala, 580-1900 т... P. glanduliferum Matuda subsp. glanduliferum
35b. н Араз не climbers; stems usually climbing trees; spathes pee
ously covered with slender scales, Costa Rica to Peru, 200-1500 m (mostly >
P. verrucosum L. Иа ex Schott
33b. Petioles glabrous and smooth distally, at least not conspicuously scaly; usually — M
Plants consistently terrestrial; petioles D-shaped with raised margins and a medial rib; spathe
tube greenish to whitish within (or usually reddish m Pacific E of eur fica) "elici
(Zelaya) to Panama, Colombia, and Ecuador; mostly 0-750 m ....................--- randipes K. Krause
36b. Plants usually hemiepiphytic (P. tysonii а! terrestrial in some habitats); ‘petioles terete to
potete: spathe tube maroon to red or violet-purple within
a. Spathe not at all constrict ve the tube, the spathe wall usually to ca. 1 cm thick
midway; leaf blades s coriaceous, minor veins etched in the u
central Costa eris to w ern Panama, 1100-2600 m __...... Р с
à mig и со en above the tube (except only weakly r for P. antonioan-
um), the w. ната less than 3 тт thick except at very base; leaf blades not ala
со аи minor veins never etched into surface of fresh leaves.
w
5
c
Annals of th
Missouri а Garden
38a. Primary lateral veins frequently 10 or more per side; spathe и white to pale green
within (sometimes tinged red at very base); Nicaragua to S es or and to Venezuela;
20-1400 m tenue K. Koch & Augustin
38b. depa lateral veins usually ine than 10 per side Мыл na in rs Lysonii); spathe tube
to red-violet, maroon, or violet-purple throughou
30a. Blades СЕЕ sinuate along the margins; rad свећа, Eg than 500 m
» subincisum Schott
39b. 2 with the margins entire, not at all s
. Fresh cataphylls usually red; petioles and midribs lacking conspicuous
purplish spots; basal veins 5-10 p
4la. Blades with primary ciel veins тет КН 6–10 per side, upper sur-
western and а Рапата, “600-1500 II IUS P smit Croat
Alb. ть т with primary lateral veins usually 5—6 per side, upper su
drying dark brown to gray-brown with пене deis cross-veining; pes
aphylls sharply 2-ribbed; Mate ge dark maroon outside; ovaries
wi mi
Veraguas and Coclé Provinces at 8. m .. P antonioanum Croat
Fresh cataphylls green (purplish-spotted in P edenudatum
40b. and P. g
mii); де» and midrib frequently purplish-spotted; a veins wei i
3-5 per side.
=
42a. ii mostly more = 1.8 times longer than wide; petioles 0.85-
.90 times as long as the blade; ur blades Ж! quies cao ts
visible on lower элг шй Рапата, 110—1150 Р. edenudatum Croat
42b. Blades mostly less than 1.8 times longer ndm wide; petioles T 1.
times longer than the blade; dried blades with secretory ducts clearly
visible on the lower surface; Costa Rica to central Panama, 0–1630
Р Р. grayumii Croat
KEY 4: VINES WITH CORDATE BLADES AND DECIDUOUS CATAPHYLLS
[Note: None of the species in this group have the posterior rib of the leaf blades naked along the sinus except Р.
brunneicaule, P. e raceum, P. mexicanum, and sometimes (weakly so) P. coloradense, P. cotonense, P. sulcica ule, P.
wilburii, and P. zhua.
la.
Internodes е elongate, usually much longer than broad; plants usually vines or at least prominently scandent.
Blades with the posterior rib (coalesced basal veins) not naked (i.e., not running on the margin of the
sinus).
Ja. Stems coarsely as perous, covered with branched scales or setose-pubesce
4a. Blades ovate-triangular. a flaring; stems reddi sh
brown, evere ч. trichome-like, frequently branched scales; berries pale уе ellow; styles
very short; See usually in wet habitats; Nicaragua to Panama and South America, 60—
280(13 Р. brevispathum E
4b. Blades vus did thin; posterior devi kon toward one another at maturity; stem
залы covered with elongate setae; pale orange to red or reddi eie се Ben
urring in dry habitats; Mexico to Peer the West Indies, and South America,
. P. jacquinii Schott
1500(250
3b. Stems smooth, not covered with branched scales
5a. i prominently o print босана у with the lateral margins sometimes acute; Nic
0-1500 m > pla typetiolatum Madison
о Ecuador, 1
5b. Petioles — to subterete, not prominently flattened dorsiventrally.
6a. Peduncles usually much shorter than the spathe, usually less prt | ст long at ап-
ар ras 6-10 mm diam.; ovules many — наивне рег
lades narrowly ovate-elliptic or triangular-sagittate or tenti be
usually 1.8 times or more longer than ic iim posterior rib more than 3 cm long.
8a Leaf blades narrowly ovate-elliptie: posterior lobes about as broad as long,
directed toward the base; s + V-shaped; ovules ca. 20 per locule;
Mexico (Chiapas), 0-1000. m мс. P. breedlovei Croat
8b. af blades triangular-sagittate to manglar batat pont r lobes typi-
cally much longer than broad, directed somewhat outward; sinus usually
not V-shaped but agen ii spathulate; ovules atk js per locule;
Mexico to а 0–1900 т mexicanum Engl.
7b. Leaf blades osm usually only slightly longer than wide а to 1.9 times
mais e; wide in P. hederaceum); posterior rib virtually lacking or to at mos
cm
9a аа. drying dark reddish brown, prominently ribbed, usually densely
warty; Costa Rica to Ecuador and Suriname; (250)3 ree PUTAT
sull p M dera ar. kirkbridei Croat
9b. Stems drying either green - we weakly sinis; or light ghee fore brown
and deeply fissured on dry
Volume 84, Number 3 389
Croat
Philodendron Subgenus Philodendron
10a. Stems drying yellowish brown, prominently ridged, smooth es
glossy; spathe tube violet-purple outside; Costa mes to Ecuador.
1600 m purpureoviride | Engl.
10b. Stems drying greenish, weakly striate, matte; spa ж ies od
side; Mexico to West Indies, widespread in South America _____
OES P ni ea ско n аса) Schott
Pe еш» frequently as long as or longer than the spathe, sometimes more
long, usually drying 3—4 mm diam.; ovules 1 or 2 per locule.
po blades us any obvious primary lateral veins; stems prominently sul-
cate opm even when fresh; ovules 1 per locule; SE Costa Rica to central
Panama; 100-700 m Ps com Croat
. Leaf blades with obvious primary lateral veins; stems smooth (or at leas
regularly and prominently sulcate-ribbed longitudinally when — onde
per locule; Panama (Darién); 850-1560 m clewellii сы
е
д
e
—
o
CORDATE VINES WITH DECIDUOUS CATAPHYLLS AND BLADES WITH NAKED POSTERIOR RIBS
2b. Blades with the posterior rib naked and running on the margin of the sinus for usually 1.5 cm or more
Stag meee to as little as 1 cm in P. smit ithi i).
24.5 setose-pubescent; blades be 1 iderabl
space gran ween the inner surface of the spathe n the ydo ЕЕ pistils ram at apex into a
style to ca. 5 mm long; Mexico to Panama, Cuba, and northern South America; 0-1500(2500) m
E jacquinii Schott
. Stems glabrous; blades coriaceous t flated-bulbo
bring uay over spadix; pistils not at g nas de into a ina пој region.
. Stems prominently and regularly sulcate-ribbed when fresh.
ép эмг. less iy 24 cm ín lacking obvious primar 1a lateral veins; Даре to 21 с
Cos i to Isthmus of Panama; 100-700 m .__________ sulci ا Croat
14b. Blades more d 26 cm lo De he ts obvious pairs of primary lateral veins; petioles
more than 29 cm lon ES Mexico to Guatemala; 700-1525 m _____ P. verapazense Croat
13b. Stems smooth or н ~ КЕ not regularly sulcate-ribbed un fresh.
15a. Blades mostly broadly оше, usually more than 25 ст wide; posterior lobes usually
broadly у sue d, about as wide as is stems drying light reddish brown with flaking
epiderm s; Costa Rica is Ecuador, 100-1300 m ______. P. bru. aule Croat & Grayum
156. Blades mostly ovate-triangular or t apg a f to angular state, most less
than 25 cm wide (sometimes ean in mithii); Mexico to Panama; 0-2000
16a. Blades мин to hastate а Е сады lobes 23 times li than beo
posterior rib directed straight toward the apex of the posterior lobe and 1.5-3.5
cm distant from the posterior margin of the blade; Mexico to Panam pee! ro
—
N
c
um Engl.
16b. Blades cordate or sagittate (rarely hastate in some forms of P. wilburii), posterior
lobes usually less than 2 times longer than wide (except sometimes more than
2 times longer in ү; аады m then blades drying reddish brown, not greenish
as in P. mexica = rib mostly curved along and near the m c of
the sinus, irme dins an tt cm from the pos theres I of the bla
17a. e ај и 2-ribbed; blades re more than 35 cm long; ا often
with a ies = Нор. peduncle usually занар "ед ће us Mexico
o Nicaragu mithii Engl.
у Симићи aperte vae 2-ribbed); blades usually less than k cm iis
petioles lacking a purple distal ring; peduncle usually shorter than the
Sache (except often longer in P. wilburii var. longipedunculatum); central
Costa Rica to central Panama, 0-2000 m ._____. P. wilburii Croat & Grayum
—
^J
с"
CORDATE NON-VINES WITH DECIDUOUS CATAPHYLLS AND NON-NAKED POSTERIOR RIBS
1
=
Internodes of mature stems broader than long or sometimes somewhat longer than broad but not elongate
with scandent stems (possibly somewhat scandent in P. breedlovei and P. sousae); plants mostly appressed-
climbing hemiepiphytes
18a. ae oles either daly covered with scales or D-shaped with undulate-winged margin
tha
M ges dom less than 1 ст diam., densely covered with conspicuous, елита ee det
um s broad as long, the major veins on the lower surface densely puberulent; a to
300 n їп Sith аав Ps uamipetiolatum 1 Croat
19b. Petioles sharply D-shaped or sharply flattened with undulate-winged margins, tire than 1
ed ва brous; blades much longer than broad; veins of lower surface glabrous; Panama (Chie
P. fortunense Croat
quí); 1 1300 m ..
18b. Nd Шин, terete to merely ver flattened.
2 or rib naked along the
Pedals ewe less Бик 10 ст long.
Wen s less than 25 cm long, the adaxial surface drying dark brown to yellow-brown.
eere ovate-triangular, 1.8—3 times longer than wide; posterior lobes 1.4—1.8
Annals of the
Missouri Botanical Garden
times longer than wide; inflorescence solitary; Panama (Chiriquí), 1750-2100 m
E E es Р ки Croat
23b. Blades ovate, 1.3-1.6 times longer than wide; posterior lobes about as wide a
long; inflorescences 2-3 per axil; Panama (Chiriquí), 1600 m
а es P. colo dne Croat
22b. een more than 35 ст long, the adaxial surface usually drying green to browni
en to grayish green to dark olive-green (sometimes dark brown in P. zhuanum or
blackish i in P. smithii).
24a. Petioles on drying not markedly flattened, 8-15 mm diam., usually greenish to
dark brown; less than 360 m along the Caribbean ie in Cou Rica: 6
ice maticum Croat & Grayum
24b. Petioles on drying markedly cmi qe to 3—6 cm M^ bere yellowish; 700—
900 m, Coclé Province, Pana P. zhuanum Croat
21b. Peduncle usually 10 or more ст lon
a. Leaf blades with margins convex in lower У of blade.
26a. Stem drying light brown; epidermis often flaking; petioles spongiose, but drying
moderately smooth, flattening to ca. 1 cm wide; blades drying thin and lacking
conspicuous secretory canals; 40-1630 m (mostly less than 600 m), Mexico to
MINE сие A к P. smithii Engl.
26b. Stem drying dark brown; epidermis sulcate or cracked but not flaking; petioles
firm, drying heavily wrinkled, flattening to 3—6 cm wide; blades drying coria-
ceous with conspicuous secretory canals; 700-900 m, Panama (Coclé) ______--
PN P. zhuanum Croat
25b. ag with margins straight to concave in lower % of blade; Costa Rica and Panama.
. Leaf blades drying pale yellow-green, lacking any sign of secretory ducts on
lower surface; ovules 1 per locule; Costa Rica, 1350-1400 m
P. straminicaule Croat
27b. Leaf blades drying dark olive-green or dark brown, with secretory ducts visible
alternating with the minor veins on lower surface; ovules 4—5 per locule; near
Costa Rican—Panamanian border, 1100-1950 m P. cotonense Croat
20b. و rib not naked along the sinus, or weakly and obscurely naked near its base
8a. Blades with posterior lobes turned inward and overlapping; peduncles slender, drying
A mm diam.; 850-1560 m, Serranía de Pirre, Darién Province, Panama .. Р. с lid Croat
28a. Blades with posterior lobes never overlapping (except sometimes in P. үза Б usually
sterile staminate portion of the spadix often conspicuously (50%) broader than the
pistillate portion at anthesis; Mexico to Guatemala, 10-2 WOM E advena Schott
29b. Spathe usually visibly constricted ab hat | the m idle)
Mmadic),
sterile staminate portion of the spadix narrower to s arer broader than the pis-
30a. Inflorescences 1-3 per axil; spathes less than 11 cm long (except 9.5-18 ст
long іп P. brenesii).
3la. Primary ed veins mostly more than 8 pairs; minor veins etched-sunken
on upper blade surface; lower blade surface + bluish green; 800-2200 m.
Costa Кіса and Panama ___ P. brenesii Standl.
31b. Primary lateral veins 4-8 pairs (sometimes to 9 pairs in Р sagittifolium);
minor veins flat or raised on upper blade surface; lower blade surface
medium green to yellow-green, not bluish green; Mexico and Belize.
32a. Leaf blades ca. 1.6 times longer than wide; ovules 1—7 per locule.
33a. Blades drying gray-green below; sap white; ovules 1 per locule;
Belize, dese thau 500 у P. dwyeri Croat
33b. Blades drying dark yellow-brown below; sap clear, turning
rown?; ovules 4—7 per locule; Panama, 1600 m ----------------
HE coloradense Croat
32b. Leaf blades ca. 1.8-2 times longer than wide; ovules 2 —4(5—8) or ca.
20 per locule; Mexico e 1300 m) or widespread, Mexico to
South America, 150-2700
34a. Blades ca. 1.8 times ‘a onger than wide; inflorescence 1 per axil;
ovules 20 per locule; Mexico (Chiapas), 1300 m ----------------
Me CEO: P. breedlovei Croat
34b. Blades 1.85-2 times longer than wide, inflorescences 1-3 per
axil; Mexico (Veracruz) to Colombia, 0-1800 m .........-------
os P. sagittifolium Liebm.
ЗОБ. Inflorescences 4—6 per axil; spathes 15-23 ст long.
P
Volume 84, Number 3
1997
Croat 391
Philodendron Subgenus Philodendron
35a. Cataphylls usually more than 25 ст long; blades usually more than 55 ст
ong, drying coriaceous, reddi
lower surface; Panama, 100-77
sh brown, with minute ig yen ridges о on
70 m poseen Croat
35b. Cataphylls ys than 25 ст long; blades mot a less than 55 € m lon,
drying subcoriaceous, brown to olive-green,
ooth or minutely мања bo
not with interrupted Bea on lower rade Месно E paaie
m
sousae Croat
Philodendron advena Schott, Oesterr. Bot. Woch-
enbl. 5: 289. 1855. TYPE: Mexico. Exact lo-
cation uncertain, described from living mate-
rial, Schott s.n. (lectotype, here designated, L
900230 (89889152)). Figures 37-40, 73.
descri — ene rae Bot. Wochenbl. 5:
5. TYPE: Wes exico: a pecia col-
гаю. еен Ьу "il (holotype, w de-
гне Field Museum Photo 29864. e here
ted; duplicate photo at MO).
аса monticola Matuda, Madroño 10: 170. 1950.
TYPE: Mexico. Chiapas: Sierra Madre, Pacific slope,
Colonia San Juan Panama, 50 km E of Escuintla,
m, Matuda 18169 (holotype, MEXU; isotypes,
DS, NY, UCLA).
биі Јатарапит С. $. ME пач Herb. 9:
6, fig. 242. 1965. TYPE: Mex Veracruz: Cos-
ii dad ec-Huatusco, Route 155 . at Rió Jamapa,
Moore £ Bunting 8872 (holotype, BH).
Terrestrial, epilithic, or hemiepiphytic; stem ap-
pressed-climbing, to 2 m long, sap brownish or-
ange; internodes weakly glossy, 2—4 cm long, 1.5—
2.5(5.5) cm diam., longer than broad, sometimes
obtusely flattened on one side, medium green to
gray-green, drying brown to yellow-brown, epi-
dermis smooth to closely fissured; roots few per
node, drying dark brown to yellow-brown, epi-
dermis sometimes flaking; cataphylls subcoria-
hs tinged reddish, generally deciduous intact,
y emarginate at apex, margins clear. VE
eine to spreading with blades pendent;
petioles 30-45(65) cm long, 7-18 mm diam.,
erect-spreading, subterete to D-shaped, somewhat
spongy to moderately firm, obtusely зеня адах-
ially, sheath 3-11 cm long; blades ovate-cordate to
ci E subcoriaceous, moderately bicolorous,
ualy acuminate at apex, cordate at base,
(32)41-47(64) cm long, 16.5-40 cm wide (1.2-2.5
times longer than wide; 1-1.5 times longer than
petioles), margins hyaline, upper surface dark
green, drying dark brown to blackened, sometimes
yellow-green, lower surface slightly more glossy,
paler, drying yellow-brown to reddish brown, some-
times greenish brown; anterior lobe 20.5-53 cm
long, 12-40 cm wide (3.4-3.8 times longer than
posterior lobes); posterior lobes rounded, 6–14(20)
cm long, 5. 4—15(20) cm wide, directed toward base;
sinus + parabolic to V-shaped or spathulate, 8-15
cm deep; midrib broadly convex to flat, slightly pal-
er than surface above, convex and slightly paler
below; basal veins 3—6 per side, with 0–1(2) free
to base, coalesced (0.4)1.3—4(8.5) cm; posterior rib
not naked or obscurely naked to 1 cm at base; pri-
mary lateral veins (3)4—6 per side, departing midrib
at а (45)50—60(80)° angle, straight or weakly ar-
cuate to the margins, flat to sunken and paler than
surface above, convex below; minor veins weakly
raised, moderately visible, arising from both the
midrib and primary lateral veins. INFLORES-
CENCES spreading-pendent, 1–2(4) per axil; pe-
duncle 5-13.5(17) cm long, 7-11 mm diam.;
spathe coriaceous, (6.5)12-19 cm long ((0.7)1.4—
2.4 times longer than peduncle), weakly or not at
all constricted above the tube, acuminate, usually
greenish throughout, broadest at or below the mid-
dle; spathe blade greenish white outside, spathe red
to maroon or greenish white, glossy inside; spathe
tube sometimes reddish outside, ong,
2.5-3.5(5) cm diam., red to maroon or dark vio-
let-purple, pale-lineate, glossy inside; spadix stip-
itate to 4 mm long; clavate to cylindrical, bluntly
pointed at apex, 11.5—14 cm long, broadest at the
middle or + uniform throughout; pistillate portion
3.3—6.9 cm long, 1.2-2.7 mm diam. at middle, паг-
rowed somewhat at both ends; staminate portion 5—
9.2 cm long; fertile staminate portion tapered to
cylindrical or clavate, usually longer than pistillate
portion, 11-14 mm diam. at base, 8-15 mm diam.
at middle, 5-7 mm diam. ca. 1 cm from apex,
broadest below middle, equal in length to pistillate
portion, narrower than pistillate and sterile por-
tions; sterile staminate portion, 11-30 mm diam.,
usually broader than pistillate portion at anthesis;
pistils 1.7-8.5 mm long, 1. mm diam., ovary
5)6—8-locular, with basal to sub-basal placenta-
tion; ovules 1-4 per locule, arranged digitately in
translucent ovule sac, 0.2-0.6(1.5) mm long, equal
in length to funicle; funicle adnate to lower part of
partition, style 0.9-3 mm long, similar to style type
B; style apex flat; stigma discoid to subdiscoid; the-
cae cylindrical to elliptical, 0.5 mm wide, contig-
uous. INFRUCTESCENCE with spathe green out-
side, dark purple-violet inside, 11.5-14 cm long;
berries white to yellowish; seeds 1(3—4) per locule,
oblong-ellipsoid, sometimes obovoid, 1.7-2.2 mm
quii,
392
Annals of the
Missouri Botanical Garden
long, 0.7-1.8 mm diam., with clear longitudinal
and faint latitudinal striations and speckled with
shiny, silver raphide cells; funicular scar knob-like,
clear, thicker than the seed body.
collected less consistently with July and August the
only consecutive months without fruits reported
Philodendron advena ranges from Central Mexi-
co (Veracruz) along the Atlantic slope to Chiapas
and then along both the Pacific and Atlantic slopes
to southern Guatemala (Santa Rosa and Chiquimula
Departments). It occurs at 70 to 2500 (mostly above
1000) m elevation in a wide variety of vegetation
types, including “Selva Alta Perennifolia,” “Bosque
Caducifolia,” “Selva Mediana Subperennifolia,”
and “Bosque Caducifolia.”
Philodendron advena is a member of Philoden-
dron sect. Calostigma subsect. Macrobelium ser.
Macrobelium. m species is distinguished by in-
ternodes longer than broad, its two-ribbed, gener-
ally decane cataphylls, terete to obtusely flat-
tened, somewhat spongy petioles (about as long as
the blades), and by its generally coriaceous, ovate-
cordate, usually dark brown to blackened-dried to
somewhat sagittate-lobed blades with rounded
lobes with usually four to six primary lateral veins,
and usually one to two green inflorescences with
the inner surface reddish to purplish.
Material from the Pacific slope in Chiapas State,
Mexico, and in adjacent Guatemala often dries
somewhat more greenish. This was described by
Matuda as P. monticola Matuda. Considering the
variability in the species, that taxon is not war-
ranted. Though not closely related to P. purulhense
Croat, P. advena is perhaps most easily confused
with that species, because both may have similar
ovate, black-drying leaves. Philodendron purul-
hense differs from P. advena in its persistent cata-
phylls, a more or less elliptic spathe scarcely con-
stricted above the tube, and ovaries with axile
placentation and 13-20 ovules per locule.
Philodendron advena is probably most closely
related to the eere ecologically versatile P. sag-
ittifolium, which occurs throughout much of its
range. Philodendron irene: is distinguished
by having usually longer, more prominently sagit-
th
broad), which dry typically reddish brown rather
than blackened. In contrast, P advena has blades
averaging 1.5 times longer than broad (ranging from
.3 to 1.8 times longer). Both P advena and
sagittifolium have five to eight locules per pistil
with up to three more or less basal ovules (or they
have the funicle adnate to the lower part of the
axillary wall but extending down to the base).
Two Oaxaca collections are noteworthy. Moore &
from above Vall
drying, there is nothing else out of the ordinary with
the collection (see fig. 235 in Bunting, 1965).
ы specimens examined. GUATEMALA. Alta
41651 (MO); Finca Argentina, 15 mi. W of Telemán, 550-
650 m, 41571 (MO); 24 mi. E of tar 9-11 mi. E
of Hwy. СА-14 to Сођап, 700-800 m, 41498 (MO); near
meto: ca. 1500 m, Standley 70495 1 near Tactic, pe
o Frío, 1400-1500 m, 90484 Chiquimula:
id 3-5 mi. N of [NON P erm
(K, MO, "Us. qe El Pocito, S of San Martín Chile
Verde, on road to Colomba, ca. 2200 m, Standley 85039
(F), 85093 (F); between San Martín Chile Verde and Col-
omba, above Mujuliá, ca. 1800 m, 85723 (F), 85561 (F);
between Finca Pirineos and Finca Patzulín, 1200-1400
m, 86875 (F); Río Samalá, near Santa María de Језиз,
1500-1650 m, 84571 (F), 84669 (F). San Marcos: Can-
а Unión Juárez, near SE portion of Volcán Тасапа,
000 m, Steyermark 36407 (F); Volcan Tajumulco,
1400-1700 e — (F); ca. 3.3 mi. above Finca Ar
afael, 1600 m, е 40963 (MO, NY);
Santa Rosa: Села. 4000 ft., Heyde
N ). Zacapa: summit of Sierra de las Minas, vic. of
Fines Alejandria, 2500 m, Sica 29855 (F) Río
Lima, Sierra de las Minas, below Finca Alejandria, 2000
m 30026 (F). MEXICO. Chiapas: Lagos de Montebello,
3 mi. W о Lagunas, 1460 m, Croat 46636 (MEXU,
MO), 46655 (MO); ca. 6 mi. NW of Pueblo Nuevo Solis-
tahuacán, vic. km 99, 1900-1950 m, 17%07'N, Ks Ww,
Croat & Hannon 65193 eee. MO); 8 mi. NW of
lo Nuevo Solistahuacán, 1900-1950 m, Croat 46429
(MEXU, pu 46430 (MEXU, МО); EE
is IT27N.
ME un 89-90 mi. SW of Pa lenque, 3
m, Croat 40221 (MO); San Juan Panamá, Escuintla, 100
m, Matuda 18488 ded MUR Cascada, i 1800
Ma
oza,
Siltepec, 1000-1300 m, 47462 (MO); Ojo d e AU E
conusco, Nakamura 38 (DS); San Fernando—Morav! -
Моште 84, Митбег 3
1997
393
Philodendron Subgenus Philodendron
mi. NW of San Fernando, 840-940 m, 16%53'N,
93°16'W, Croat & Hannon 65006 (B, MEXU, MO, US);
Mpio. Angel Albino Corzo, above Finca Cuxtepec, 1380
m, Breedlove 52073 (DS); Mpio. Berriozábal, 13 km N of
i 3 DS); m NW of Rizo de
DS); Mpio. Ocos-
ingo, Laguna Ocotal Grande, 3300 ft., Breedlove 15683 (F,
MICH); Mpio. Rayón, Selva Negra, 10 km above Rayón
Mezcalapa, along road to Jitotol, 1700 m, 23291 (DS);
Mpio. Union Juárez, Volcán Tacaná, above Talquian, 2200
m, Breedlove & ua 47712 (CAS); Mpio. Villa Corzo,
base of Cerro s Picos, near Cerro Bola, 1500 m,
16°10'N, 93°15’ e Taks & Thorne 30193 (DS). Oa-
[zs]
o
ке ©
N
py
=
Е
A
ы; MEXU, МО, NY, US); 14 п ті. У of Valle Nacional,
1210 m, Са 39783 (MEXU, МО), 39796 (MO); 21.5
alle Nacional, 1660-1670 m, 48108 (МО);
Teotitlán del Cam ino-Chilchalla: 2.2 mi. beyond vdd
to Huautla de jin 2270 m, 48335 (CM, МО); 3.8 m
a de Jiménez, 2265 m, 48369 (MO)
Ixtlán, Río re bra watershe 5-805 m, 17°37
96°17'W, Boyle et al. 3943 (MO). Tabasco: RI
э, 3.1 mi. E of var ca. 0.25 mi. S of highway, 150
E of Debe din from
este Ж qe idad Autonoma
O). Veracruz: e ME А "Ох апара, т,
17*05'N, 84*35'W, Hammel & Merello 15563 (MO); above
San Jose de Gracia, 1 mi. 5 of hwy. between Córdoba and
Veracruz, 750 m, Croat XR (MO); Conejo—Huatusco, at
m 45, barranca de Santa María, across hwy. from Haci-
EUR El Mirador, Moore & Tuning 8856 un ca. 1200
m, Croat 44013 (MO); Mendoz zaba, Sierra San Cris-
фа, Highway 150-D, ca. 3 km SW of Orizaba, 1260—
m, Croat 39549 (CM, MEXU. МО); Fortín- Huatusco,
Highway 125, 300 m, 19*06'N, 97°02’ W, Croat & Hanno
63107 (B, COL, CR, CM, F, G, INPA, K, LL, M, MO, NY,
aro & Ramarillo 3895 (MO, WIS); Mpio. Yecuatla,
Los Capulines, near Paz de Enríquez, ca. 8 km N of Chi-
conquiaco, 1600 т, 19°47'N, 96°49’ W, Taylor et al.
147 (F, MO, NY).
Philodendron albisuccus Croat, sp. nov. TYPE:
Panama. Darién: Parque Nacional del Darién,
middle slopes on W side of Cerro Pirre, 800—
1500 m, 7%56'N, 77%45'W, 29 June 1988,
Croat 68940 (holotype, MO-3589989; isoty-
pes, B, COL, K, PMA, US). Figures 41-43.
Planta etin ен 2-3 ст longa, 2.8
cm diam.; succus albus cco calcareus; «шерү lev-
iter 1- costata vel acu us 24 Ste persistentia ut fibrae
tenues, pallidae; petiolus usque 51-72 e
lata; inflorescentia 1; pedunculus (3.7)7-12 cm longus;
spatia .9 em longa, extus "quse. intus pallide
ocu 8-20
seminibus; baccae alb.
Hemiepiphytic; чет = appressed-climbing, to
75 ст long, to 2.8 cm diam., sap chalky white;
internodes short, usually sparsely covered with cat-
node, epiderm
pa to flaking and e el gn to
40 cm long, weakly l-ribbed to sharply 2-ribbed,
sparsely short-lineate, persisting as thin, pale fibers
at upper nodes, then de dug. petioles 51—72
cm long, 3—7 mm diam., ing, subterete,
dark green, weakly баа from near base to mid-
dle, obtusely sulcate at the base adaxially, surface
semiglossy; blades narrow ovate, subcoriaceous,
splitting, long-acuminate at apex (the acumen in-
rolled), cordate at base, 50-66 cm long, 24.542
cm wide (1.6–2 times longer than wide), (ca. about
equal in length to petiole), broadest just below
point of petiole attachment, upper surface dark
green, semiglossy, lower surface semiglossy, mod-
erately paler, drying yellow-green; anterior lobe
—48(54 5) ст und 23.1-27.4(42.4) cm wide
posterior lobes 8.5-11.5(16.5
12(18.5) cm wide; sinus hippocrepiform;
flat, concolorous above, bluntly acute, slightly paler
than surface below; basal veins 3—5 per side, with
1 free to base, third and higher order veins coa-
lesced 0.5-2 cm long, the fifth and sixth veins
sometimes coalesced to 5 cm long; posterior rib na-
ked, raised; primary lateral veins (4)6-7(8) per
side, departing midrib at a 60—65^ angle, sunken
above, raised to convex below; interprimary veins
darker than surface; minor veins arising from both
the midrib and primary lateral veins; lesser veins
obscure to visible. INFLORESCENCES 1 per axil;
peduncle (3.7)7-12 cm long, 3-3.5 mm diam., pale
green; spathe semiglossy, 13.5-14.5 cm long (1.2-
1.8 times longer than peduncle), weakly constricted
above the tube, green throughout, pale yel-
low-green (greenish brown post-anthesis) within;
spathe tube densely short pale lineate outside, 5—
7 cm long; spadix 12 cm long at anthesis; pistillate
portion 4.3 cm long in front, 3 cm long in back, 8
mm diam. midway, 7 mm diam. at apex; staminate
portion to 10 cm long; fertile staminate portion to
9 mm diam. toward apex, somewhat narrower just
394
Annals of the
Missouri Botanical Garden
above the sterile portion, to 5 mm diam. 1 cm from
apex; sterile staminate portion 9 mm diam.; pistils
6–6.3 mm long, 1.8–2.5 mm diam., ovary 5—6-loc-
ular, with axile placentation; ovules 2-seriate, style
similar to style type B; style apex flat; stigma sub-
discoid. INFRUCTESCENCE 6.4 cm long, 3 cm
diam.; berries white; seeds 18-20 per locule.
Flowering in Philodendron albisuccus is poorly
known with a flowering collection seen in October,
immature fruits in June, and nearly mature fruits
in July.
Philodendron albisuccus is endemic to Panama,
known only from the Serranía de Pirre in Darién
Province, at ou to 1400 m elevation in Premon-
tane rain fore.
Philodindrod albisuccus is a member of Philo-
short internodes; thin, obscurely one-ribbed cata-
phylls, which persist at the upper nodes as thin,
pale fibers; subterete petioles (about as long as the
blades); the narrowly ovate, moderately cordate
blades with a hippocrepiform sinus; and especially
by the sap in all the cut plant parts, which promptly
turns white and chalky upon exposure to air. Only
one other species in Panama, P. cretosum Croat &
Grayum, has chalky sap. Though both species have
chalky sap (a rare feature for Philodendron) and
18-20 ovules per locule, I do not believe them to
be closely related as they show no other features in
common. Philodendron cretosum differs most par-
ticularly in having slender oblong-lanceolate blades
that are 5.1-8.3 times longer than wide (vs. 1.6-2
times longer than wide for P. albisuccus). Philoden-
dron albisuccus also has the much longer petioles
(51-72 cm long vs. 10-23 cm long for P. cretosum),
more T spreading primary lateral veins (60—
° vs. 25—45° angle for P. cretosum). In addition,
Р Ант: has minor veins more distinct than
those of P. cretosum.
The species is superficially similar to P. alticola
Croat and P. straminicaule Croat & Grayum, both
of which have blades of similar size, shape, and
color upon drying. Both differ by having the inner
surface of the spathe red rather than green.
Additional specimens examined. PANAMA. Dane:
Cerro Sapo, ca. 5 km S of Garachiné, 600-800 m, 7°59'N,
78°25'W, Hamna et al. 14820 (MO, US); Cerro Ба ге-
gion, Altos de Nique—Cerro Рите, са
de Nique, ca. 8 km W of Ca
m, Croat 37851 (MO, PMA); Río Cana-Río Escucha Rui-
do, along ascent of Serranía de Pirre above Cana Gold
vm 1310-1430 m, 37830 (MO, US); Río Cana, SW of
rro Pirre, vic. o
(А 10) arque Nacional Darién, slopes of Cerro Mali,
headwaters of S branch of Rfo Pucuro, ca. 22 km E of
Pucuro, 1300-1400 m, 8?04/30"N, 77°14’ W, Cuadros et
al. 3961 (MO).
Philodendron alticola Croat & Grayum, sp. nov.
TYPE: Panama. Chiriquí: Parque La Amistad,
3.5 mi. W of Cerro Punta, 2 km inside park
along old abandoned roads and trails, 800 m,
8°53'N, 82°35'W, 28 Маг. 1993, Croat 74906
(holotype, M0—4343624—5; isotypes, В, CR, F,
K, PMA, US). Figures 45—48.
Planta epiphytica aut eam desis Bec (1.5-
2)4–10 cm longa, 2-4 c iam., in sicco pallide flavi-
brunnea, subtiliter costata; © аи На à usque ıe 28 cm longa,
incostata, in sicco palli ua; petiolus 42-
70 ст
m lata, sagitta
infra; inflorescentia 1-2; pedun-
m longus; spatha 11.5-14(18) cm longa,
viridis; pistilla (4)5-6(7)-locularia; loculi 12—18-ovulati.
Epiphytic or hemiepiphytic; stem appressed-
climbing; internodes dark green, promptly gray-
green, finally light brown, ане finely ribbed,
(1.5-2)4-10 cm long, 2-4 cm diam., drying pale
yellow-brown, epidermis sometimes loosening and
flaking free in small patches; cataphylls sharply
D-shaped with weakly raised margins, pale green,
weakly glossy, drying thin, to 28 cm long, unribbed,
drying pale brown, deciduous at lower nodes, per-
sisting semi-intact at upper nodes with a few ex-
posed pale fibers, in part persisting as pale fibers
at lower nodes with a few pale fibers and small,
thin fragments of pale brown epidermis. LEAVES
erect-spreading to spreading; petioles 42-70 cm
ong, 7-10 mm diam., subterete, medium green,
weakly glossy, faintly lineate, obtusely flattened
adaxially, weakly spongy, drying greenish to black-
ened, slightly flattened iaid the apex adaxially,
ace drying + matte, sometimes with portions of
epidermis loose and bubbled, tan and translucent;
sheathing subtending an inflorescence, to 6 ст
long; blades narrowly ovate-sagittate, subcoria-
ceous to moderately coriaceous, semiglossy, some-
what bicolorous, weakly and shortly acuminate at
apex, sagittate at base, 44-72 cm long, 21-44 cm
wide (1.6-2 times longer than wide), (about as long
as petioles), broadest somewhat above point of pet-
iole attachment, upper surface dark green, matte,
drying brown to greenish brown and almost matte,
lower surface moderately paler and weakly glossy,
drying yellow-brown; anterior lobe 34-35.5 cm
long, margins convex; posterior lobes 13-26 cm
long, 11-17 cm wide, directed downward; sinus ob-
ovate, 10-19 cm deep; midrib speckled, slightly
paler, drying broadly convex, concolorous above,
narrowly rounded, slightly paler, drying prominent-
ly convex, often flat and irregularly ridged, yellow-
Volume 84, Number 3
1997
Croat 395
Philodendron Subgenus Philodendron
ish brown below; basal veins 5—6 per side, with the
first free to base, the second coalesced 1-5 cm,
third and higher order veins coalesced 4—7.5 cm
long; posterior rib naked for 1-3 cm; primary lat-
eral veins 5-8 per side, those near end of blade
departing midrib at a 50-65” angle, those in the
lower one-half of blade departing at 80-95” angle,
broadly curved to the margins, obtusely sunken and
slightly paler, drying weakly raised above, convex
and paler than surface, splayed out and downturned
at the midrib, drying weakly raised below; inter-
primary veins inconspicuous; minor veins moder-
ately distinct, fine, arising from both the midrib and
primary lateral veins, drying prominulous, alternat-
ing with secretory canals appearing as intermittent
lines, sometimes branched, blackened and minute-
ly sunken. INFLORESCENCES erect, 1-2 per axil;
peduncle + terete, coarsely striate (coarser on
spathe and base of tube, pale short-lineate other-
wise), weaker toward apex, 5-12 cm long, drying
7-10 mm diam., pale green, drying dark brown;
spathe coriaceous, 11.5-14(18) cm long, scarcely
or not at all constricted midway, green to yellowish
green throughout, acuminate; spathe blade dark
green throughout, drying dark brown and un-
marked, smooth outside, pale greenish white to
white on upper two-thirds of blade inside; spathe
tube 2.8-4 cm long, 2-3 cm diam., red to red-violet
to purplish on lower one-third of tube inside; spa-
dix 9-14 cm long; pistillate portion pale green,
0.8-1.2 cm long, 1-1.5 cm diam. throughout; sta-
minate portion 8-12.5 cm long; fertile staminate
portion creamy white, stubby, evenly and bluntly
tapered to apex, 1.5-2 cm diam. throughout, broad-
est at the base, much broader than the pistillate
portion; sterile staminate portion 1.5 cm diam.; pis-
tils 3-4 mm long, 1.7-1.8 mm diam., ovary (4)5—
6(7)-locular, with axile placentation; ovules 12-18
per locule, 2-seriate, weakly translucent, 0.4-0.5
mm long, longer than funicle; funicle 0.1-0.2 mm
long, adnate to lower part of partition, style similar
to style type B; central style dome sometimes weak-
ly developed; dried style base raised but flattened
apically with a narrow, pale ring around its outer
margin; stylar canals emerging as tiny funnels at
base of small apical depressions and arranged sep-
arately in a ring, drying as minute funnels extend-
ing above the surface of the style boss; style apex
weakly concave; central domes and small depres-
Sions surrounding stylar canal exits; stigma cover-
Ing entire style apex; androecium truncate, oblong
| 1 mm long; thecae + oblong, ca. 0.3
Pee wide; sterile staminate flowers in part prismat-
1C, in part weakly clavate or irregularly 4—6-sided,
1.2-1.8 mm wide. INFRUCTESCENCE in early
fruit to 14 cm long; spathe becoming dark brown
with paler lines in tube; berries white with dried
style bases ca. 2 mm long, brown, with a narrow
smooth ring around its margin; seeds tan, ovoid,
weakly mammiliform, 1.4—1.8 mm long, 1-1.2 mm
diam.
Flowering in Philodendron alticola is document-
ed by just a few collections and might be bimodal.
Specimens in Panama were collected both in bud
and immature fruit in February (nearly mid-dry
season in Panama), but post-anthesis collections
were also made in August and September. No ma-
ture fruits were seen. The region where P. alticola
occurs is decidedly less seasonal (at least in terms
of totally rainless days) than is much of the rest of
anama. This might induce a less seasonal flow-
ering in Philodendron.
Philodendron alticola is known only from eastern
Costa Rica and western Panama in the region ad-
jacent to the frontier at 800 to 2500 m elevation in
Tropical Lower Montane wet forest or possibly Trop-
ical Lower Montane rain forest. It is one of the most
high-ranging species of Philodendron subg. Philo-
dendron Schott in Central America, hence the name
“alticola” (meaning high-dweller).
Philodendron alticola is a member of P. sect.
Philodendron subsect. Philodendron ser. Fibrosa.
In addition to the high elevations at which it occurs,
the species is characterized by having the inter-
nodes longer than broad and drying yellowish
brown and finely ribbed; by its subterete petioles
about twice as long as the blades, drying darkened
and sometimes with bubbly epidermis; the ovate-
sagittate blades, which dry mostly dark brown
above and yellowish green below with conspicuous
blackened, interrupted and sometimes branched
secretory ducts; and by the usually paired, short-
pedunculate inflorescences with the spathe scarcely
constricted above the tube, green outside, white in-
side on the blade and red to purple on the tube
within. The species is most similar to P. stramini-
caule, which is also subscandent and has similarly
colored blades. That species differs in lacking
prominent intermittent laticifers and the fertile sta-
minate portion of the spadix constricted above the
sterile portion; the sterile staminate portion of the
spadix only slightly thicker than the pistillate por-
tion; and a doughnut-shaped style with the style
tubes in the bottom of a concavity upon drying. In
addition, it has only 1 ovule per locule (vs. 12-18
per locule for P. alticola). In contrast, P. alticola
has conspicuous laticifers, the staminate spadix
scarcely or not at all constricted above the sterile
396
Annals of the
Missouri Botanical Garden
portion, and the sterile portion of the spadix much
thicker than the sterile staminate portion. The dried
style base of P. alticola is flat at the apex with erect,
tubular-extensioned stylar pores.
The species is also similar to P. schottianum H.
Wendl. ex Schott, which also shares the conspicu-
ous latex canals on the lower blade surface and a
similarly stubby spadix, but that species differs in
having shorter internodes, cataphylls that persist as
a dense mass of fibers (vs. semi-intact or as only a
few pale fibers interspersed with small fragments of
epidermis in P. alticola), by its petioles, which dry
smoother and more matte, and by its more broadly
ovate blades, which often have conspicuous cross-
veins.
Additional specimens examined. COSTA RICA. Li-
món: Cordillera de Talamanca, Atlantic slope, Valle de
Silencio, along Río Terbi, 0.5-1.5 airline km W of Costa
Rican-Panamanian border, 00— m, 9?8'N,
82°57'W, Davidse et al. 28735 (CR, МО); S side of un-
named cordillera between Río Terbi and Río Siní, 2-4
airline km W of Costa Rican—Panamanian border, 2300—
2500 m, 9°9-11'N, 82%57-58'W, Davidse et al. 28931
(MO). PANAMA. Chiriquí: vic. of Las Nubes, 2.7 mi. NW
of Río Chiriquí Viejo, W of Cerro Punta, 2200 m, Liesner
323 (MO, US), 325 (MO, US); Cerro Punta, Las Nubes,
2000 m, Croat 26492 (MO).
Philodendron angustilobum Croat & Grayum,
sp. nov. TYPE: Costa Rica. Heredia: Estación
Biológica La Selva, confluence of Río Sarapi-
quí and Río Puerto Viejo, 10°26’N, 84%01'W,
50-80 m, Grayum & Chavarría 8302 (holo-
type, MO). Figures 49-52, 74.
Planta hemiepiphytica; internodia 3-6(15) cm longa,
5-3 cm diam.; cataphylla 12-33 cm longa, acute 2-cos-
5-10 mm diam.;
А с
el atribrunnea supra, flavibrunnea
vel atricanibrunnea infra; lobis lateralibus manifeste pa-
tentibus, in sicco denigratis infra; inflorescentia 1-3; ре-
dunculus (4.5)9-11(15) cm longus; враћа (11)13-18 ст
longa, lamina spathae extus interdum suffusa alba; tubo
spathae pallide viridi, extus interdum suffusa marronino
basi; pistilla 7-10-locularia; loculi (1)2-ovulati.
Hemiepiphytic; stem appressed-climbing, grow-
ing to 2.5—6 m high in trees; internodes matte to
weakly glossy, 3-6(15) cm long, 1.5-3 cm diam.,
dark green, drying gray (within 4-6 internodes),
epidermis drying light brown, semiglossy, often
peeling (underlying stem blackened), fissured
transversely, longitudinally ridged; cataphylls soft
(especially near base), 12-33 cm long, bluntly to
sharply 2-ribbed or sharply 1-ribbed to near the
apex then 2-ribbed below, green, sometimes tinged
red, deciduous, persisting deciduous, intact; peti-
oles 17-54 cm long, 5-10 mm diam., subterete,
somewhat spongy, dark green, weakly flattened to-
ward apex with an obscure medial rib adaxially,
with adaxial margins rounded to weakly angular,
surface weakly glossy, drying blackened; sheathing
3-9 cm long; blades deeply 3-lobed, subcoria-
ceous, moderately to conspicuously bicolorous, 30-
48 cm long, 20-48 cm wide (1-1.5 times longer
than wide), (0.8—1.8 times longer than petioles), up-
per surface dark green, drying dark gray to dark
brown, semiglossy, lower surface drying dark yel-
low-brown to dark gray-brown, weakly glossy, mod-
erately paler; anterior lobe 28-39 cm long, 8.5-
14(17) cm wide (1.8—1.9 times longer than lateral
lobes) rarely broadest at the base, usually nar-
rowed toward the base (4-12 cm wide just above
the base); sinus arcuate to broadly V-shaped; lat-
eral lobes (7.5)15-22(27) cm long, 3-8(12) cm
wide, broadest usually toward the apex, directed
outward, broadly spreading (90—127? angle), broad-
ly confluent with medial lobe 3.3—6.5 (rarely to 14)
cm, acute; midrib broadly convex to flat-sunken
above, convex, slightly paler than surface, drying
blackened below; basal veins 8—10 per side, pin-
nately arranged along a stout medial rib; posterior
rib not at all naked or rarely naked for 1–1.5 cm;
primary lateral veins 5-6 per side (45—75° angle
on median lobe), gradually curved to the margins,
sunken above, convex, drying blackened below; in-
terprimary veins weakly raised and darker than sur-
face below; minor veins distinct, darker than sur-
face below, arising from both the midrib and
primary lateral veins. INFLORESCENCES 1-3 per
axil; peduncle (4.5)9-11(15) cm long; spathe
(11)13-18 cm long, 8-10 mm diam. (1.4-1.6(2.4)
times longer than peduncles), greenish white out-
side, greenish white to white within; spathe opening
broadly at anthesis, convolute in only lower one-
half of tube, barely or not at all constricted above
the tube; spathe blade sometimes tinged white out-
side; spathe tube light green, sometimes tinged ma-
roon at base outside; spadix held erect, not pro”
truded forward, oblong to oblong-tapered, rounded
to bluntly acute at apex, (9.5)11-15.5 cm long,
broadest near the middle, constricted weakly above
sterile staminate portion; pistillate portion pale
green, oblong to oblong-tapered, 4.5—5.7 cm long,
8-10 mm diam. at apex, 8-12 mm diam. а! middle,
(5)11-13 mm wide at base; staminate portion 8-11
cm long; fertile staminate portion oblong-tapered,
8-13 mm diam. at base, (6)12-13 mm diam. а!
middle, 8-9 mm diam. ca. 1 cm from apex, usually
broader than the pistillate portion; sterile staminate
portion ca. 1 cm diam.; pistils 1.5-3 mm long, 11-
1.8 mm diam., ovary 7-10-locular, (0.8)1.1 mm
diam., with sub-basal placentation; ovules 1(2) Рег
Volume 84, Number 3
1997
Croat 397
Philodendron Subgenus Philodendron
locule, contained within transparent ovule sac, 0.6
mm long, usually shorter than funicle, style similar
to style type B; style apex rounded; stigma button-
like, subdiscoid, 1 mm diam., 0.2-0.4 mm high,
covering center of style apex; thecae oblong to ob-
long-ovate, contiguous, sometimes divaricate. JU-
ENILE plants with petioles 8-10 cm long; blades
oblong, ченин to weakly subcordate.
Flowering in Philodendron angustilobum has
been ا د the early dry season through
the mid-wet season, including Februrary, March,
May (the greatest number), and August, but too few
collections exist to be certain of phenology.
Philodendron angustilobum ranges from Hondu-
ras (Olancho) to Panama, apparently being restrict-
ed to the Atlantic slope from Honduras to Costa
Rica and ranging from near sea level to 680 m
elevation. In Panama it has been collected on both
slopes near the Continental Divide from 800 to
1430 m elevation, but a juvenile collection from
near sea level in the Canal Area is probably also
this species. It is known from Tropical wet forest
and Premontane wet forest life zones.
Philodendron angustilobum is apparently a
member of P. sect. Tritomophyllum [though M. Gra-
yum (pers. PNE ^g a it is closest to P. ligu-
latum, a member t. Calostigma subsect.
Glossophyllum ser. m This : species is
characterized by its moderately long internodes,
spongy petioles, and 3-lobed blades, which
blackened with broadly spreading lateral lobes that
rter than the medial
Engl., which has sagittate blades that dry green to
brown and occurs typically in much drier habitats.
The sole Honduran collection might prove to rep-
resent another species or perhaps even a hybrid. It
iffers from material collected from Nicaragua,
Costa Rica, and Panama in having the medial lobe
broader, more triangular, and not at all constricted
at the bas
Two и (Croat & Grayum 59933 and
Croat 67616) from the same plant from southwest-
em Costa Rica near Golfito differ in having the
posterior lobes less narrowed; they may prove to
represent another species.
Additional specimens examined. COSTA RICA. Сағ
tago: CATIE, near ban Molina, SE of Florencia 7
Turrialba, са. 680 m, 9"52'30"N, 83°40'W, Grayum 3877
R, MO). Puntarenas:
Coto Brus, Las Cruces d poit near San Vito
de Java, ca. 4000 ft., Croat 32960 (MO); 6 km W of San
ips de Java, 1200 m, 849'N, 82°58’ W, 57230 (MO); ca.
1 km NW of Golfito, 11 km SW of Interamerican Highwa
«100 m, 8°11'N, 83*12'W, Croat & Grayum im (CR,
F, K, MEXU, MO, US); Golfito- Villa Bricefio, 3.1 mi. NW
of center of Golfito, 30 m, 8?11'N, 83?12'W, Croat 67616
border with Colón Dept., along Río Grande, 35 m,
15°31'N, 8542", Croat & Hannon 64522 (CAS, CR,
EAP, HNMN, K, MO, PMA, US). NICARAGUA. Zelaya:
Río Sucio, 2 km E of Bonanza, 140 m, Neill 4024 (MO).
PANAMA. Canal Area: ca. 1 mi. E of Fort Sherman, <25
m, 9197, 79°57'30"W, Croat & Zhu 76283 (MO). Chi-
i: Cerro Colorado, 15.6 mi. ab Río San
m 1330 m, Croat 48439 (MO), 24 mi. above bridge
over Río San Félix, 1430-1500 m, 48489 (MO
, 33180 ( 1
Lago Fortuna, trail to Rfo , 845'N, 82°18'W,
76372A (MO). Coclé: vic. of La Мы, N of El Valle de
Antón, 800-900 m, 8”38'N, 80709", 67209 (CM, MO).
Panamá: Cerro Campana, above Su Lin Motel, 14759
(MO, NY, SCZ).
Philodendron anisotomum Schott, Oesterr. Bot.
Z. 8: 179. 1858. Philodendron fenzlii Schott
var. anisotomum (Schott) Engl., in A. DC. &
. DC., Monogr. Phan. 2: 1879. TYPE:
Guatemala. Las Nubes, Wendland 321 (lecto-
type, here designated, GOET). Figures 44, 53—
6:75
узы tao ne aie Hemsl., Diagn. Pl. Nov. Mexic.: 37.
TYPE: Gua ш Barranca Honda, Volcán
ucgo, 3800 ft., Salvin s.n. (holotype x
Philodendron дара Schott, ge res Z 179.
1858. TYPE: Costa Rica. Cart and еа
vi oM s.n. (holotype, destroyed. вод ic. 2592
, Publ. Field. Mus. Nat
ist., n 18:137. 1937. TYPE: Costa Rios.
Мајне n pts de San Ramón, 1050 m, Brenes
5762 (holotype, F).
Usually a hemiepiphytic vine or sometimes on
rocks; stem appressed-climbing, green as juvenile,
graying with maturity, minutely and densely striate,
sap watery, unscented, leaf scars conspicuous, 8—
10 mm long, 7-9 mm wide; internodes smooth, se-
miglossy, to 2.5 cm long, 4—10 mm diam., usually
longer than broad, medium green to olive-green,
epidermis thin, tan, peeling; roots olive green,
smooth, few per node; cataphylls thin, semispongy,
to 10 cm long, unribbed, bluntly or sharply
1-ribbed, green, drying pale yellow-green, decidu-
ous. LEAVES erect to spreading; petioles 21-57
ст long, (2)3-9 mm diam., terete, moderately
spongy, whitish toward apex, slightly flattened to-
ward apex adaxially, surface dark green striate at
base; blades triangular in outline, deeply 3-lobed,
subcoriaceous, moderately bicolorous, long-acumi-
nate at apex (the acumen apiculate, to 3 mm long),
hastate at base, 20-35 cm long, 20-42 cm wide
(0.8-1 times longer than wide), (0.6-1 times the
398
Annals of the
Missouri Botanical Garden
petiole length), upper surface dark green, semi-
glossy to glossy, drying dark brown to dark gray-
green, lower surface semiglossy, paler, drying yel-
low-brown to yellow-green; anterior lobe
oblong-lanceolate to oblanceolate, almost elliptic,
16-30 ст long, 7-15 cm wide (1-1.3(1.5) times
longer than lateral lobes); lateral lobes broadly con-
fluent 1-3(8) cm with medial lobe, 10-25 cm long,
3-8.7 cm wide, directed outward (90° angle from
midrib), acute to bluntly acute; sinus arcuate; mid-
rib + flat to sunken, paler than surface above,
broadly convex below; basal veins 4—7 per side,
sometimes with last vein free to base, most veins
coalesced 1.5-13 cm, 2 veins coalesced to 17 cm,
drying reddish brown to yellowish brown; posterior
rib naked; primary lateral veins 4-5 per side, de-
parting midrib at a 50—60° angle, narrowly sunken
above, convex below; interprimary veins weakly
sunken and concolorous above, weakly raised and
darker than surface below; tertiary veins visible,
darker than surface below; minor veins fine, close,
weakly visible to distinct below, arising from both
the midrib and primary lateral veins but mostly
from the midrib. INFLORESCENCES erect-spread-
mm diam.; spathe 7.4—16.6 cm long (0.8-1.4(1.7)
times longer than peduncle); spathe blade green to
greenish white to creamy yellow, tinged with violet-
purple outside, tinged with violet-purple, at least
sometimes, with yellowish resin canals visible in-
side; spathe tube green outside, 4-6 cm long, 2—
3.5 cm diam., dark violet-purple inside; spadix
sessile; white, drying golden-yellow throughout, ta-
pered, + rounded at apex, 7.5-10.8 cm long,
broadest near the base; pistillate portion pale green
(post-anthesis), ellipsoid, 3.6 cm long, 1.3 cm diam.
at apex, 1.2 cm diam. at middle, 6 mm wide at
base; staminate portion to 8.4 cm long; fertile sta-
minate portion white, ellipsoid, tapered at apex, 8
mm diam. at base, 5 mm diam. at middle, 5 mm
diam. ca. 1 cm from apex, broadest at base, much
narrower than the pistillate portion; sterile stami-
nate portion tannish, 8 mm diam.; pistils (1.5)2.9—
4.4 mm long, 1.7-2.3 mm diam., ovary (6)7-8-loc-
ular, (1)2.2-3.2 mm long, ovule sac 0.7 mm long,
with sub-basal placentation; ovules 3 per locule,
contained within translucent, gelatinous ovule sac,
ca. 0.4 mm long, longer than funicle; funicle 0.2—
0.3 mm long (can be pulled free to base), style
similar to style type B; style apex flat; stigma dis-
coid, 5—6 mm diam., 0.1 mm high, covering center
of style apex; the androecium = prismatic, margins
irregularly 4—6-sided, 0.7—1.1 mm long; thecae ob-
long, 0.3 mm wide, + divaricate; sterile staminate
flowers irregularly 5—6-sided, 1 mm wide. IN-
FRUCTESCENCE 9.5-16 cm long, peduncle to 14
cm long; spadix to 5.3 cm long, to 3 ст wide; ber-
ries orange, rhomboid; seeds 1-2(3) per locule,
(16)21–22(29) per berry, light brown, 1.5-2 mm
long, 0.5 mm diam., with weak constriction (nipple)
opposite funicular end of seed.
Flowering in Philodendron anisotomum appears
to be restricted to the rainy season in Central
America. One collection was seen in flower in May
but most are from July through December (the latter
month is sometimes the beginning of the dry season
in parts of Middle America). Immature fruits were
collected from December through July, with mature
fruits known only from March, April, and July.
Philodendron anisotomum ranges from Mexico to
Costa Rica, at 30 to 1800 m elevation. In Mexico,
the species ranges from Nayarit to Oaxaca and
Chiapas mostly along the Pacific coast, but also oc-
curs in Puebla (Ajenjibre) and Morelos (Cuerna-
vaca). One collection, Moore & Bunting 8874, from
near Córdoba in Veracruz state, appears out of
range for the species.
Collections from Guatemala are few, but all are
from the Pacific slope except one collection from
Baja Verapaz between El Chol and Rabinal (Croat
& Hannon 63670). In Honduras and Costa Rica,
the species is nearly restricted to the Pacific slope.
In Costa Rica the species occurs in Premontane
moist forest. Philodendron anisotomum is a member
of P. sect. Tritomophyllum.
This species is distinguished by its deeply three-
lobed blades with frequently much smaller, falcate
lateral lobes broadly confluent with the medial
lobe.
Philodendron anisotomum is easily confused
with P. tripartitum (Jacq.) Schott, which differs ш
having proportionately narrower medial lobes
(mostly 3-3.5, rarely to 1.7 times longer than
broad) with 4—9 prominently sunken primary lateral
veins and lateral lobes typically directly more ог
less toward the apex. It has only one ovule per
locule and white fruits. In contrast, P. anisotomum
has medial lobes less than 1.5-2.8 times longer
than broad with 2-4(5) weakly sunken primary lat-
eral veins and usually much smaller lateral lobes
typically directed outward, as well as 3 ovules per
locule, and orange fruits.
Material from Nayarit (McVaugh 13363, Moore
& Bunting 8703) is not only geographically isolated
from populations in Puebla, Cuernavaca, an Oa-
xaca, but also differs morphologically by having
much larger blades (medial lobe >23 cm wide)
with the lateral lobes narrowly rounded rather than
pointed.
Volume 84, Number 3
1997
399
Philodendron Subgenus Philodendron
Costa Rican collections differ in sometimes ћау-
ing two inflorescences per axil, rather than solitary
inflorescences, which is more typica
Philodendron dagilla Schott, ваһиди on the
same page as P. anisotomum, was considered а syn-
onym of P. tripartitum by Krause (1913), but it is
clearly synonymous with P. anisotomum. Schott dis-
tinguished it from P. anisotomum by its longer, less
spreading lateral lobes, but they are well within the
Additional specimens examined. COSTA RICA. 1911,
without locality, Pittier & Durand 3090 (BR); Worthen s.n.
(MO). Alajuela: San Pedro de San b engl 1075 m, Bre-
nes 4874/33 ) (F); San Ramón, 1500-1600 m, Tonduz
17719 (BM, K, P); San Ramón- Aw 2.3 km N of Río
Balsa, 1050-1150 m, 10°11'N, 84%30'W, Stevens 14193
(MO); ca. 5.7 km N of Quebrada Volio, 1100-1150 m, ca.
0°08'N, 84°29'W, 14165 (CR, MO); San Ramón-Balsa,
5.7 mi. N of San Ramón, 1200 m, Croat 46837 (MO); San
1-3 km E of San Ramón, ca. m,
Liesner 14198 (B, CR, MO); above Río San ix 15 km
of San Ramón, 800 m, 10%14'N, 83°31'W, Lent 3108
(F); Finca Los Ensayos, ca. 11 mi. NW of Zarcero, 900
m, Croat 43522 (MO); Zarcero mam ca. 100 m, Smith
A447 ); ca. 7.5 mi. N of Zarcero, ca. 1000 m, Croat
43499 (MO); ca. 15 km N of у Pasta 1350 т, Williams
m, Cavan 3085 (DUKE); Catilón San Carlos a
So 975 m, A. Smith H1669 (F, US); San Isidro de San
Ramón, 1259 m, 10 °04'46"N, 84?26'30"W, Herrera 58
(AAU, M, MEXU, MO); Monteverde Biological Reserve,
Río Peñas Blancas, 900 m, 10°18’N, 84°45'%, Haber &
Bello 7181 (MO); D I. m, 9?17'N, 84^84'W, Burger
et al. 10770 (CHAPA, 89 Rio Реје, 4 Кт 5 of Ciudad
uesada, 960 m, Lent 1274 (BM, CR, F, GH); Río Trojas,
2 km N of La Luisa, pec m, 1674 (CR, US). Cartago:
Juan Viñas, 1300 m, Carpenter 608 (US); 5 slope of Vol-
cán Irazú, Standley 36638 (US); Stevens 48 (US). Guan-
acaste: La Cruz de Abangares, 1400 m, Haber & Bello
07'N, 83°04'W, Davidse et al. 25635 (MO); Cantón de
sg Aires, Quebrada Dorora (tributary of Río Kuiyé),
| "№, 84*18'W, Grayum et al. 6096 MO); San José,
landley 47358 (US), са. 1130 m, 41210 (US), 47334
(US), 1150 m, 33262 (US); 1.4 km NW of Brazil de Santa
Ana, 800 m, Taylor 17378 (NY, US); San tt S of
San José, 1160 m, Standley 49293 (US); San Pedro Mon-
tes de Oca-Curridabat, ca. 1200 m, 32830 (О 5), 1250 т,
41307 (US); Rfo Мапа Aguilà ar, near San José, 1200 m
38952 (US); Las Pavas, 1070 m, 36085 (US); camino de
ape near San José, 1200 m, 32171 (US); Acosta, Z.P.
Cerros de Escazú, Río Tabarcia, 1600-1
9*50' 52"N. 84%04'40"W, маре ae (CR, MO); Valle
del Candelaria, 1000-1 m, Morales E rios 4596
(CR, INB); Santiago de Puriscal, Ec 8 (F). EL
SALVADOR. Ahuachapán: Padilla piles pe 1922,
pe (US), 60 (05); 2-3 mi. NE of Bridge Imposible,
1250 m, Croat 42162 (MO). San Salvador: Tone-
pete Calderón 200 (US); 650-850 m, Standley
€ €: eis US). ise Alta Verapaz: Cob-
m, Standley 90275 (F); Semococh, TE km
mnis ies on Cobá n Road, се 4714
N of Cobán, along Highway C 4, 13
(MO); 5 mi. 5 of Cobán, nh ли а СА-14, 1300 m,
41364 (MO); San Juan Chamelco, Wilson 41008 (F). Baja
Verapaz: Mpio. Rabinal, El Chol-Rabinal, Highway 6,
8.7 mi. N of El Chol, 1330 m, 15%03'N, 90?29'W, Croat
& Hannon 63670 (GH, LL, MO, TEX). Chiquimula: Que-
a: Es
[жге 63390 п, NE of Escuintla, 89605
(F, MEXU). а Los СЕ along Rfo Pinule, 1 mi.
W of San Pedro Pinula, 1400 m, Steyermark 32931 (F).
Jutiapa: San José Acatempo-Río de los Esclanos, Cuesta
ei la Conora, gue m, Standley 60616 (F). m
enango: Reserve INDE, “Зата María," km 199, 1200—
1300 m, 14745' N T W, Croat & Hannon 63435 (B,
M, US); 1 1020 m, Standley
67137 (Е);
Volume 84, Number 3
1997
oat 405
Philodendron Subgenus Philodendron
primary lateral veins; minor veins arising mostly
from the midrib, fewer from the primary lateral
veins; lesser veins obscure above, fine, moderately
conspicuous and darker than surface below. IN-
FLORESCENCES = erect, 1-2 per axil; peduncle
7-11.3 cm long, 9-11 mm diam., medium to pale
green, unmarked, semiglossy; spathe erect-spread-
ing, subcoriaceous, 13—14.5 cm long (1.2-1.8 times
longer than peduncle), weakly constricted midway
(6 cm above base); spathe blade broadly flattened,
curved weakly forward, pale greenish outside, to 9
cm long (opening elliptic in face view, 5.5 cm
wide), pale green, heavily suffused with red (В &
K Red-Purple 3/10) throughout inside; spathe tube
oblong-ellipsoid, medium to dark green outside,
densely short lineate throughout outside, 6 cm long,
3.5 cm diam., red to maroon (B & K Red-Purple
3/10) at base, weakly so toward apex inside; spadix
oblong (weakly tapered or weakly clavate), 11-24
cm long, broadest usually at the middle; pistillate
portion medium to dark green, tapered toward the
apex, 2.3-4.2 cm long, 2.7-4.3 cm long in front,
2.4-2.7 cm long in back, 8-12 mm diam. at apex,
9-15 mm diam. at middle, 5-7 mm wide at base;
staminate portion 9.5-12 cm long; fertile staminate
portion weakly tapered or weakly ellipsoid, 8-16
mm diam. at base, 10-16 mm diam. at middle, 6–
10 mm diam. ca. 1 cm from apex, broadest at the
middle or + uniform, broader than or as broad as
the pistillate portion, as broad as the sterile portion;
sterile staminate portion broader than or as broad
as the pistillate portion; sterile staminate portion
5—7(8—9)-locular, 0.7-1.2(2.1)
mm long, (0.7)1.1-1.3 mm diam., with sub-basal
placentation; ovules (3)4 per locule, 1-seriate (or in
2 series of 2), contained within translucent ovule
sac, if present, 0.2-0.4 mm long, usually longer
than funicle, style ыш, to style type В; style
crown usually as broad as ovary; stigma discoid,
0.7-1 mm diam., 0.1-0.3 mm high, covering entire
style apex; the androecium margins 4—6-sided and
mm diam. at apex; thecae oblong
pen, slightly obovate); sterile staminate flow-
rregularly rounded to bluntly 5-sided, 2.3-3
mm E 1-1.5 mm wide, white. INFRUCTES-
CENCE with berries orange.
Flowering phenology in Philodendron auricula-
tum is unclear, but possibly flowering is initiated
in the late rainy season. Only a single flowering
collection exists (October), but there are a modest
number of collections collected post-anthesis dur-
ing the dry season and early rainy season (January
through June). A single mature fruiting collection
was made in January.
Philodendron auriculatum occurs only in south-
western Costa Rica on the Pacific slope from San
José and Puntarenas Provinces (as far west as Car-
ara), ranging from near sea level to 1200 m ele-
vation in Tropical wet forest life zones.
Philodendron auriculatum is a member of P.
sect. Calostigma subsect. Glossophyllum ser. Glos-
sophyllum. This species is characterized by short
internodes, sharply two-ribbed, deciduous cata-
phylls, moderately long, markedly spongy, some-
what flattened petioles (averaging slightly shorter
than the blades), oblong-elliptic to oblong-oblan-
ceolate, pale yellow-green-drying blades, which are
usually narrowly cordulate at base, and one to two
greenish inflorescences, which are red to maroon
within at base.
Philodendron auriculatum is probably most
closely related to P. pseudauriculatum Croat, which
ranges along the Atlantic slope of Central America
Pirre). That species differs in its usually darker
gray-green-drying leaf blades with a dark green
ring at the apex of the petiole, and in having the
leaf base acute, rounded, or broadly subcordate at
base (lacking the narrow auriculate lobes so com-
mon for P. auriculatum). In addition, P. pseudaur-
iculatum has the whitish spathe clearly demarcated
from the contrasting green peduncle. In contrast, Р.
auriculatum has a yellowish green spathe that is
not at all demarcated from the peduncle.
Additional specimens examined. COSTA RICA. Pun-
are ear Inter-American Highway in vic. of Piedras
edes Croat 32956 (CR, MO); hills N of Palmar Norte,
along trail to Jalisco, 50—700 m, I 35205 50 соуе
at NE base of e m 3 Fen erto Quepo
924'N, 84°10'W, Fire 6612 (CR, К. мо,
US); El General Valley, pees Río Sonador, near Pan-
American Highway, 600 m, Williams et al. 28808 (F);
along the Río Cacao above Pan-American Highway, 900
m, Williams et al. 28687 (Е); - Cruces Tropical нк:
ical Garden, Сапібп Coto Bru m W of San Vito de
Java, 1200 m, 8°49’N, 82°58'W, Croat 57268 (MO); 8.8
mi. N of Villa Neily, 1010 m, 66171 (МО); Osa Península,
vic. Boscosa, Croat annon 79297 (INB, MO); Esqui-
nas Ridge, 150-250 m, Gómez 19677 (MO, NY, RSA, US);
Rincón de Osa, vei 1819 (F, MO); ca. 5 km W of
Rincón de Osa m, 842'N, 83°31' * Sar a &
Liesner 7298 (F, зе MA 7306 (CR, F); 50-
8°42'N, 83°31'W, Burger & Gentry 8978 (CR, F); 0 N.
83°31’ W, Aguilar 1507 (INB, MO); Piedras Blancas—Rin-
; of Рап i Highway, 90-105 m
8°46'N, 83°18’ W, e 67691 (MO); Quebrada Aparicio-
оше, Aguabuena, Rincón de Osa, 200—400
42'N, 83°31'W, E а et al. 4013 (MO); аы
Pes cón de Osa, ca. 6 km W of Inter-American eene
at Chacarita, 100 m, 8°45'N, 83^18'W, Croat & Grayu
59730 (CM, CR, K, MO, SAR); Parque Nacional Cuni;
406
Annals of the
Missouri Botanical Garden
vado, Sirena, 0-150 m, 8?27—30'N, 83°33-38' W, Kernan
25 (CR, MO); Estación Sirena, 5 of Río Sirena along Río
Camaronal, 0 m, 8”28'N, 83°35'W, Knapp 2188 (CR, L,
MO, US); Talamanca Range, Pacific slope, forested foot-
hills of mountains E of Quepos, 150-250 m, 929'N,
84°03'W, Burger et al. 10603 (F, MO). San José: Puriscal,
Z.P. La Cangreja, 300 m, Morales et al. 3239 (CR, INB);
Carara—E] Sur de Turrubares, 280—370 m, 9?45'30"N,
84732", Grayum 10445 (CR, INB, MO).
Philodendron bakeri Croat & Grayum, sp. nov.
TYPE: Costa Rica. Guanacaste: W slope of
Cerro Nubes, ca. 2 km E of Silencio de Tilar-
án, large patch of remnant primary forest, 900
m, 10?28'N, 84°53'W, Grayum, Herrera &
Sleeper 4992 (holotype, MO-3392250; isoty-
pes, CR, DUKE). Figures 77, 78.
Planta hemiepiphytica; internodia pleurumque longiora
quam lata, (2)7—9 ст longa, (0.7)1—1.5 ст lata; cataphylla
5-10 cm
longa vel oblanceolata, basi cuneata vel rotundata, 12—
25.5 em longus; inflorescentia 1; pedunculus 2.6—5.2 cm
longus, 2-5 mm diam.; spatha
spathae extus pallide viridi vel crema sufi
marronina, intus viridi vel albida suffusa rubra; tubo spa-
thae intus atrimarr а t carmesino; pistilla 5—
8(10)-locularia; loculi 1(2)-ovulati; baccae aurantiaceae.
Hemiepiphytic; stem scandent, slender, to 1.5 m
long; internodes (2)7-9 cm long, (0.7)1-1.5 cm
diam., usually longer than broad, medium green, +
matte, becoming light brown, epidermis fissured
minutely longitudinally upon drying; roots dark
brown, thin; cataphylls subcoriaceous, 5-10 cm
long, unribbed, bluntly to sharply 1-ribbed or rarely
sharply 2-ribbed, green to reddish or maroon, de-
ciduous, narrowly rounded at apex, margins clear.
LEAVES spreading; petioles 4-12 cm long, 2-4(6)
mm diam., spreading, subterete to C-shaped, some-
times tinged reddish, sometimes maroon at base,
obtusely somewhat flattened or bluntly sulcate
adaxially, rounded abaxially, surface semiglossy,
with narrow purple ring at apex; sheathing 2.7-3.2
cm long, sheathing to ca. 2 cm but for Y to % of
its petiole length when subtending inflorescences;
geniculum subterete, maroon, 2.7-4 cm ong,
darker than petiole; blades + oblong or narrowly
oblong to oblanceolate, moderately coriaceous, acu-
minate to long acuminate, sometimes acute at apex
(the acumen sometimes short apiculate, 1-3 mm
long), cuneate to rounded at base, 12-25.5 cm long,
(2.74—10 cm wide (2.5-3 times longer than wide),
(2-3 times longer than petiole), margins thin, nar-
row, reddish, upper surface drying reddish brown,
semiglossy, lower surface weakly glossy; midrib flat
to sunken, slightly paler than surface above, nar-
rowly convex, reddish violet below; basal veins
lacking; posterior rib lacking; primary lateral veins
3—4 per side, departing midrib at а 45-55” angle,
ascending to the apex, inconspicuous or slightly
sunken above, not distinct below; tertiary veins vis-
ible, darker than surface; minor veins obsurely vis-
ible to moderately distinct, fine, close, arising from
midrib only; secretory ducts usually obscurely vis-
ible on lower surface of dried blade. INFLORES-
CENCES 1 per axil; peduncle 2.6-5.2 cm long, 2-
5 mm diam., obscured by petiole sheath; spathe
moderately coriaceous, 7-11.5 cm long (2.2-2.7
times longer than peduncle), scarcely or not at all
constricted, pale green to cream, often tinged light-
ly or heavily with reddish lineations, sometimes sol-
id red or maroon, or with red speckles or lines
(rarely seen in Panamanian specimens), green to
whitish, suffused with red inside; resin canals ap-
pearing medially, especially near the constriction;
spathe tube cylindrical, semiglossy outside, 4-5 cm
long, deep maroon, red, or crimson inside; spadix
sessile; cylindrical, sometimes clavate or weakly ta-
pered, 7.2-8.5(9.5) cm long, + uniform throughout;
pistillate portion pale greenish, cylindrical, 1.8-2.4
cm long, 5.8-8 mm diam. at apex, 6-9 mm diam.
at middle, 5-6 mm wide at base; staminate portion
4.8-8.9 cm long; fertile staminate portion white,
cylindrical, clavate or weakly tapered, 6-11 mm
diam. at base, 8-9 mm diam. at middle, 7 mm
diam. ca. 1 cm from apex, as broad as or sometimes
broader than the pistillate portion, if detectable,
narrower than the sterile portion; sterile staminate
portion broader than the pistillate portion, white, 6
mm diam.; pistils 1-2.2 mm long, 0.6-1.4 mm
diam.; ovary 5-8(10)-locular, 0.4-1 mm long, 0.8-
1 mm diam., with sub-basal placentation; ovules
1(2) per locule, sometimes contained within trans-
parent, gelatinous ovule sac, 0.2-0.5 mm long,
equal in length to or longer than funicle; funicle
0.2 mm long, style 0.7 mm diam., similar to style
type B; style apex with depressions surrounding
stylar canal exits; stigma brush-like, discoid,
0.5(1.5) mm diam., 0.2–0.6 mm high, covering en-
tire style apex; the androecium truncate, margins
irregularly 4—5-sided, 1.3 mm long, 0.3-1.4 mm
diam. at apex; thecae oblong, sometimes oblong-
elliptical; pollen spheroidal to ellipsoidal or elon-
gate, less than 0.2 mm long, 0.1 mm diam.; sterile
staminate flowers 3—6-sided or irregularly 4-sided,
0.9-3.9 mm long, 0.4-1.3 mm wide. INFRUC-
TESCENCE pendent; seeds 1(2) per locule. JU-
VENILE leaves and spathes tinged red.
Flowering in Philodendron bakeri occurs from
the early dry season, December through April, with
post-anthesis collections from January through May.
Volume 84, Number 3
1997
Croat 407
Philodendron Subgenus Philodendron
Immature fruiting collections were made in May
and June
Philodendron bakeri is known along the Atlantic
slope of Costa Rica from 100 to 900 m elevation
in Premontane wet forest and Tropical wet forest life
zones and also occurs in Central Panama at 900 to
1420 m in Tropical wet forest and Premontane rain
forest life zones. It is expected to be found in in-
tervening ar
Philodendron bakeri is a member of P. sect. Ca-
lostigma subsect. Glossophyllum ser. Glossophyl-
lum. The species is characterized by its scandent
habit, slender stems (frequently pendulous on flow-
ering portions), reddish, deciduous cataphylls, sub-
terete petioles (one-fourth to one-third as long as
the blades), more or less oblong blades, and by the
solitary inflorescence with the spathe green, suf-
fused with red or maroon outside, and heavily suf-
fused with red within, especially at the base. Also
characteristic are the bright orange fruits.
Philodendron bakeri is most similar to P. immix-
tum, also a vine with more or less elongate blades.
That species differs in having thicker internodes
drying with broad fissures and an often loose, flak-
ing epidermis, and thinner blades that dry greenish
or greenish brown and are more typically subcor-
date at the base. In contrast, P. bakeri has minutely
fissured stems that lack a loose, flaking epidermis
and blades that dry typically reddish brown, mod-
erately coriaceous, and are merely rounded to ob-
tuse at the base.
Though both species have spathe tubes that are
reddish on the inner surface, P. immixtum differs
in that the tube is not reddish on the outside.
Both P. bakeri and P. immixtum are members of
P. sect. Calostigma subsect. Glossophyllum, with
one ovule per locule, but locules of P. immixtum
differ in being longer, with the transparent envelope
enclosing the ovules being about twice as long as
the ovules, whereas those of Р. bakeri have the
ovule(s) embedded in a gelatinous matrix complete-
ly filling the envelope.
Philodendron bakeri may also be confused with
Precociously flowering shoots of Р. sagittifolium.
oth species have petioles that may be tinged pur-
plish violet on both ends. Philodendron sagittifol-
ium differs in having pistils with 7-8 locules with
24 ovules per locule in a transparent envelope (vs.
1 ovule per locule in a gelatinous matrix).
Croat 44312 from La Selva, Costa Rica, differs
in drying dark gray-brown above, dark brown be-
low, and in lacking ува kii It is perhaps a
Juvenile of some other speci
The species is named in bu of Richard Baker,
an aroid specialist previously at the Field Museum,
who made the first collection in 1974.
ae ional «вм ехапипеа. Ages m Ala-
: Cariblanco-Laguna Hule, c m W of Costa
eg Highway 9. 117 N, 84°13’ W. er et al. 227 (Е,
MO); Monteverde Biological Reserve, Río Pefias Blancas,
900 m, 10%19'N, 84°44'W, Haber & Bello 6932 (MO),
7907 (CR); M. Uie gie W of Bijagua, near Hío
Zapote, ca. 600 m, 10°44’N, 8505", Burger et al. 11628
(F. MO); ves e 4 km NNE of Bijagua, on slopes
Rosita: 800 m, Bello i (CR); Bello et al. 4537 (CR,
NB) Reserva Forest e San Ramón,
10912'40"М, 84°36’ 20"W. Herrera 6748 (CR, MO). Gude-
acaste: N side of Lake Arenal, 1 km NW of dam, 650 m,
of Las Кем. 450-550 m
et al. 6543 (МО); Rio Peje-Río Sardinalito, Volcán Barva,
700—750 m, 10°17'30"N, 84%04'30"W, Grayum & Jerm
6784 (MO); La Zona Protectora, Río Peje-Río Guacimo,
northern slopes of Volcán Barva, along ~ = main
road across Quebrada pe виши to Rfo Guacimo, 250 m,
Grayum & Schatz 3219 (DUKE); La Selva Field. Station,
100 m, Hammel 7805 (DUKE, MO); 100-150 m, Croat
44312 (MO); 6 km by road from Río Peje crossing, 5 km
SSE of Magsasay, 10?21'N, 84°03-04’W, Schatz & Gra-
yum 667 (DUKE); Magsasay, 700 m, 1. ر 182 (MO).
ón: Hacienda Tapezco-Hacienda La rte, 29 air
k of Tortuguero, 'N, 47W Davidson
et al. 6982 (LAM); rtuguero National Park, 0 m,
10°34’N, 83°31'W, Cro б 61210 (МО); 2 km W of Rfo
Toro Amarillo on road W from Guápiles, 275 m, 10^13'N,
83750", Thompson & Rawlins 1225 (CM). PANAMA.
Penonomé, Llano Grande—Coclecito, 4.3 mi.
Grande, 410 m, 8°42'N, 80%26'W, Croat 67456 (CM, Es
MO, PMA); El Copé region, Alto Calvario, 5.2 mi. above
El Copé, 930 m, 49159 (MO); above Alto Calvario,
m и m, Sisina ma & Andersson 4546 (MO); El Valle
esa, a 860-900 m
Cao 3 7419 > ш 7508 (МО). Panama: El Lláne Ски
Road, Km 19. m, 919'N, 78%55'W, de Nevers &
Herrera 5875 E NA. Е q іа pt Nusagandi,
325-350 m, Croat 67395 (MO); Km 10, 33710 (MO); 0.5
= E of El Llano, ад 5202 (МО).
Philodendron basii Matuda, Anales Inst. Biol.
Univ. Nac. Méxic. 32: 153, Fig. 8. 1961.
TYPE: Mexico. México: Barranca de Malinal-
tenango, NE (published as SE) of Zacualpan,
1200 m, 18?43'N, 99°36’W, Matuda 37244
(holotype, MEXU). Figures 76, 81–84.
Almost always terrestrial, or trailing over rocks;
stem green, stout, succulent, leaf scars conspicu-
ous, 2—4 cm long, 4-5 cm wide; internodes 1—4 cm
long, 2.5-7 cm diam., broader than long, drying
olive-green, epidermis peeling, light brownish,
408
Annals of the
Missouri Botanical Garden
roots long, 5 mm diam., light reddish tan; cata-
phylls 20-28 cm long, weakly to sharply 2-ribbed,
medium green with darker ribs and dark green
speckles in lower one-half of cataphyll, drying tan-
nish, persisting intact, eventually deciduous; peti-
oles 22-70 cm long, 8-15 mm diam., terete, me-
dium green, dark striate; blades
ovate-cordate, drying moderately thin, weakly bi-
colorous, semiglossy, acute at apex, cordate at base,
16-56 cm long, 9-38 cm wide (1.5-1.8 times lon-
ger than wide), (0.7—0.9 times the petiole length),
broadest at or above middle; upper surface medium
green, weakly glossy, drying medium yellow-brown;
lower surface slightly paler, semiglossy, drying yel-
low-green to yellow-brown; margins sinuate or
weakly undulate; anterior lobe 35—48 cm long,
37.546 cm wide (2.5—8.5 times longer than pos-
terior lobes); posterior lobes 4–19 cm long, 18-23
cm wide, directed inward, rounded to obtuse at
apex; sinus spathulate, 10-13 cm deep; midrib
weakly sunken to flat, heavily dark green striate,
paler than surface above, convex, sparsely dark-
lineate, paler than surface below; basal veins 3-6
per side, with 1 pair free to base, the third through
fifth coalesced 3—3.5(4) cm; posterior rib naked for
1-2.5 em; primary lateral veins 4—7 per side, de-
parting midrib at a 40—60° angle, + straight to the
margins, sunken and paler than surface above, con-
vex and paler than surface below; interprimary
veins sunken and concolorous above, flat and
darker than surface below; branches of primary lat-
eral veins + ruffled-sunken and concolorous above,
raised and paler than surface below; minor veins
predominately arising from the primary lateral
veins, those arising from the midrib seem to dis-
appear soon after leaving the midrib. INFLORES-
CENCES probably 1 per axil; peduncle 7-8 cm
long, 7-8 mm diam., green; spathe 12.6-13.7 cm
long (1.7-1.8 times longer than peduncle), apicu-
late at apex; spathe blade pale green or white,
tinged red near base outside, 8 cm long, white
tinged with red near base, pale-speckled inside;
spathe tube ellipsoid, dark green outside, 6 cm
long, 3 cm diam., maroon-red (B € K Red-Purple
10/3) inside; spadix sessile; tapered, 11.8-12 cm
long, broadest at the base; pistillate portion cylin-
drical, 2.5-3.8 cm long, 1.5-1.9 cm diam. midway,
slightly narrower on both ends; staminate portion
7.6–13 cm long; fertile staminate portion clavate,
9-15 mm diam. at base, 11-15 mm diam. at mid-
dle, 5-10 mm diam. ca. 1 cm from apex, broadest
above the middle, narrower than the pistillate por-
tion, broader than the sterile portion; sterile sta-
minate portion narrower than the pistillate portion,
1-1.8 cm diam.; pistils 2.9-3.9(5.7) mm long,
surface
1.4—2 mm diam.; ovary 4—6-locular, 2 mm long, 1.4
mm diam., with sub-basal placentation; ovules 4—6
per locule, 2—3-seriate, contained within transpar-
ent ovule sac, 0.3 mm long, equal in length to fu-
nicle, style similar to style type D; style apex
rounded, with style boss and with depressions sur-
rounding stylar canal exits; style boss narrow but
pronounced; stigma slightly discoid to hemispher-
oid, 1.6-2.2 mm diam., 0.6-1.4 mm high, covering
almost entire style apex; the androecium 4—6-sided;
thecae oblong, 0.3 mm wide; sterile staminate flow-
ers 4—6-sided, 1.7 mm long, 0.9-1.6 mm wide. JU-
VENILE leaves + sagittate at base.
Flowering in Philodendron basii is recorded dur-
ing the early wet season with post-anthesis collec-
tions made during September and in January, al-
though too few collections overall have been seen.
Philodendron basii is endemic to Mexico, rang-
ing from western Jalisco, south and east to the
states of Colima, México, and Guerrero, at 350 to
1200 m elevation in *Bosque Pino-Encino" and
*Selva Baja Caducifolia." Matuda (1962) reported
it from as low as 200 m elevation.
Philodendron basii is a member of P. sect. Ca-
lostigma subsect. Macrobelium ser. Pachycaulia.
The species is characterized by its very stout,
green, succulent stems, which usually trail over
rocks; weakly two-ribbed cataphylls, which remain
intact and are usually soon deciduous; long-petio-
late, moderately thin, ovate cordate blades with a
sinuate margin; and by the green spathes with the
tube reddish purple within.
Although Philodendron basii is not easily con-
fused with any other species, it is ecologically very
similar to P. warszewiczii, which also has thick, suc-
culent, bare stems and occurs in very dry habitats.
The latter is distinguished by its deeply dissected
leaf blades. Matuda compared P. basii with P. smi-
thii, to which it bears only a superficial resem-
blance. That species differs in having more slender
internodes, which are usually longer than broad;
more narrowly ovate blades; and (usually) two 1m-
florescences per axil with longer peduncles.
Though no specimens have been seen from Co-
lima State in Mexico, Matuda (1962) reported hav-
ing seen this species there.
tween bridge over s
pagayo with Río Omitlan, 850 m, Croat 45766 (MO). J б
isco: Highway 200, 10-13 km SE of El Тино, valley 9
Río las Juntas, 250-330 m, McVaugh 25396 (MICH): €
б mi. S of Tuito, 350 m, Croat 45442 (COL, MEXU, MO.
US
Volume 84, Number 3
1997
Croat 409
Philodendron Subgenus Philodendron
Philodendron breedlovei Croat, sp. nov. TYPE:
Mexico. Chiapas: Mun. La Trinitaria, Monte
Bello National Park, E of Lago Tzikaw, 13 May
1973, Breedlove 35181 (holotype, DS; isotype,
MEXU). Figures 79, 80, 85.
Planta hemiepiphytica; internodia usque plus quam, 7
em, ca. 1.5 cm diam.; cataphylla decidua; petiolus sub-
longus, 7 mm di am.; lamina anguste ovato-
que 37 cm lon паун й іп ѕіссо ai
brunnea; ен ga 1; pedu noci 6 cm longus,
10.5 ст longa; lamina spathae љета и a
alba; tubo dra extus virenti, intus rubro; pistilla 5—6-
locularia; loculi 20-ovulati.
Hemiepiphytic; internodes to 7 cm or more long,
ca. 1.5 cm diam., usually longer than broad, drying
light brown, semiglossy, weakly and irregularly
ribbed; cataphylls deciduous; petioles 34 cm long
(slightly longer than blades), 7 mm diam., subter-
ete; blades narrowly ovate-cordate, acuminate and
slightly inequilateral at apex, sagittate at base, 37
cm long, 20 cm wide (1.8 times longer than wide;
about equal in length to petiole), semiglossy, upper
surface drying dark olive-green, lower surface dry-
ing yellowish brown; anterior lobe 31.5 cm long, 21
cm wide (4 times longer than posterior lobes); pos-
terior lobes 7.5 cm long, 8.7 cm wide, narrowly
rounded, directed toward base; sinus + V-shaped,
5-6 cm deep; midrib drying narrowly raised and
darker below; basal veins 3—4 per side, with 0 free
to base, 1 pair coalesced for 2.8 cm; posterior rib
to 3 cm long, not naked; primary lateral veins 5—6
per side, arising initially at an acute angle then
forming a gradual arch to margin, departing midrib
at a 55^ angle toward apex, 60—70? angle midway,
to 80" angle near the base, weakly arcuate to the
margins, drying weakly raised and paler below; mi-
nor veins arising from both the midrib and primary
lateral veins. INFLORESCENCES 1 per axil; pe-
duncle 6 cm long, 6 mm diam.; spathe 10.5 cm
long (1.75 times longer than peduncle), slightly
constricted midway, densely speckled-lineate, es-
pecially on the tube, red within, darker red on the
tube within, weakly acuminate at apex, convolute
to half its length at base; spathe blade greenish
White to white, paler along the margins outside;
spathe tube medium green outside, 5 cm long; spa-
dix sessile; 6.5-8 cm long, broadest above the mid-
dle; pistillate portion 2.9 cm long, 1.7-1.9 cm
iam., broadest at the middle; staminate portion 7.2
em SÉ fertile staminate portion to 2.2 cm diam.
midway, usually broader than the pistillate portion;
sterile staminate portion 1.7 cm diam.; pistils
3.3-3.8 mm long, 1.7-1.8 mm diam.; ovary dera
locular, with axile placentation; ovules 20 per loc-
ule, 0.3-0.4 mm long, equal in length or slightly
longer than funicle; style similar to style type B,
1.3-1.5 mm diam.; style apex truncate, minutely
warty, irregularly 3—5-sided; stigma 2.3-2.5 mm
diam., depressed with 5—6 small stylar canals; ster-
ile staminate flowers 2.7-3.1 mm long, 0.9-1.6 mm
wide.
Flowering in Philodendron breedlovei is based on
a single collection in post-anthesis condition made
in May. Flowering is probably in the early wet sea-
son, which begins in May in Mexico.
Philodendron breedlovei is known only from the
type locality in Mexico (Chiapas) in “Bosque Pino-
Encino” or “Bosque Caducifolio,” at 1300 m ele-
vation. Since its type locality is very near Guate-
mala, it is expected to be found there as gy
Philodendron breedlovei is a member sect.
Philodendron subsect. Philodendron ser. rs таш
The species is characterized by its long internodes;
subterete petioles about equaling the blades; nar-
rowly ovate-cordate, yellow-brown drying blades;
solitary, short-pedunculate spathes with the outer
surface white on the blade and reddish on the tube;
and especially by its ovaries, which have parietal
placentation and about 20 ovules per locule.
Philodendron breedlovei is most easily confused
with P. sousae Croat, which has similarly shaped
blades with weakly coalesced basal veins that often
dry a similar yellow-brown color. That species dif-
fers in having only 1—3 basal ovules per locule and
persistent stigmas bearing a conspicuous rim. The
old stigmas of P. breedlovei instead are truncate and
smooth with up to six more or less equally spaced
stylar canals in a ring around the center.
Philodendron brenesii Standl., Publ. Field Mus.
Nat. Hist., Bot. Ser. 18: 140. 1937. TYPE:
Costa Rica. Alajuela: La Palma de San Ra-
món, 1050-1100 m, Brenes 5110 (holotype, F).
Figures 36, 86—89.
Hemiepiphytic, sometimes terrestrial; stem gray-
en, to 2 m long, glaucous; sap watery, spicy-
scented; internodes weakly glossy, becoming matte,
2.5 ст long, 2.5-5 cm diam., about as long as
broad, medium to dark green, weakly glossy to se-
miglossy, drying gray, epidermis brown, crisp; cat-
aphylls to 24 cm long, sharply 1-ribbed to sharply
2-ribbed, sharply and broadly sulcate, pale green
to greenish brown to reddish or weakly tinged red
near base, densely short dark lineate, drying tan-
nish brown, promptly deciduous, obtuse at apex,
margins clear to pale; petioles 20—53 cm long,
0.6-1.7 cm diam., subterete, somewhat spongy, yel-
lowish green, very broadly convex to weakly flat-
tened to obtusely flattened with obtuse medial rib
410
Annals of the
Missouri Botanical Garden
adaxially, rounded to convex abaxially, with adaxial
margins rounded, surface sparsely short, dark green
or reddish lineate, sometimes with green to reddish
ring around apex; blades narrowly ovate, subcor-
iaceous, short- to long-acuminate at apex, + sag-
ittate at base, 28-79 cm long, 8-38 cm wide,
(1.6)1.8-2(2.4) times longer than wide, 1.1-1.4
times longer than petioles, upper surface medium
to dark green, semiglossy to glossy, lower surface
pale green to bluish green, matte to glaucous; an-
terior lobe 30-52 cm long, 20-30 cm wide (4.3-
5.5 times longer than posterior lobes); posterior
lobes 5.5-12 cm long, 8.8-13 cm wide, rounded,
directed toward base; sinus narrowly V-shaped;
midrib very broadly convex to flat, whitish to pale
green to pale reddish green, sometimes sparsely
short red-lineate, at least near base above, narrowly
convex to convex, pale green to reddish, matte,
sometimes dark red-lineate below; basal veins 5-6
per side, 2 free to base, the third and fourth some-
times coalesced to 3.5 cm; posterior rib not naked;
primary lateral veins (5-6)8-12(18) per side, de-
parting midrib at a 60-95” angle, sunken above,
convex and paler than surface below; minor veins
visible, etched-sunken and paler above, slightly
raised and slightly darker than surface below, aris-
ing from both the midrib and primary lateral veins.
INFLORESCENCES erect, 1-2 per axil; peduncle
4.5—10.5 cm long, 0.5-1 cm diam., convex adaxi-
ally, rounded to angular abaxially, medium green,
semiglossy; spathe thin, 9.5-18 cm long (1.7-2.1
times longer than peduncle), constricted + at the
middle, pale yellowish green throughout, spathe
blade cream inside; spathe tube 4-6 cm long, se-
miglossy outside, deep magenta with resin canals
inside; spadix sessile, + tapered, 11.5-17.3 cm
long, broadest at the base; pistillate portion раје
yellow, cylindrical, 2.8-8 cm long, 0.8-1.4 cm
diam., tapered toward base; staminate portion 8.6—
10.8 cm long, only slightly broader than pistillate
portion; fertile staminate portion cream, generally
tapered, 9-14 mm diam. at base, 1.1-1.2 cm diam.
at middle, 4-6 mm diam. ca. 1 cm from apex,
broadest at base, usually narrower than the pistil-
late portion; sterile staminate portion 1-1.5 cm
diam.; pistils 3.1—4.7 mm long, 1.7-2.8 mm diam.;
ovary 5-8-locular, 1.8 mm long, 1.7 mm diam., with
sub-basal placentation; ovules 6-12 per locule, ar-
ranged in 2 series of 6 ovules, contained within
gelatinous matrix (no true envelope), 0.4 mm long,
+ equal in length to funicle, style 1 mm diam.,
similar to style type B; style apex flat to weakly
rounded; stigma subdiscoid, 1.5-2.1 mm diam.,
0.3-1.7 mm high, inserted on center of style apex,
shallowly depressed medially; the androecium trun-
cate, 4—6-sided; thecae oblong to ovate, 1.3 mm
wide, contiguous; pollen cream, elongate to sub-
spheroidal, 0.1—0.2 mm long, to 0.1 mm diam. IN-
FRUCTESCENCE with berries yellow maturing to
orange; seeds 2 per locule, very pale yellow, 1.4—
1.7 mm long, 0.7 mm diam. JUVENILE plants
creeping, appressed-climbing; internodes 8-15 mm
long, 2-2.5 cm wide; petioles 7-8 mm diam, D-sha-
ped, flattened adaxially, weakly striate; blades nar-
rowly ovate, 19–23.3 cm long, 9-12 mm wide; ba-
sal veins 1-2; posterior rib not naked; primary
lateral veins 5-10 per side.
Flowering in Philodendron brenesii may be asea-
sonal. Flowering collections have been made in
July and August, and many post-anthesis collec-
tions have been made between March and Novem-
ber. Fruiting collections have been made mostly
during what is the dry season and early wet season
in Costa Rica, December through May. Mature
fruiting collections have been made in December,
January, February, and May.
Philodendron brenesii ranges from Costa Rica to
central Panama, at 800 to 2200 m elevation in Pre-
montane rain forest and Tropical Lower Montane
rain forest life zones.
Philodendron brenesii is a member of P. sect. Ca-
lostigma subsect. Macrobelium ser. Ecordata. The
species is distinguished by its mid-elevation habi-
tat, bright green internodes about as long as broad,
sharply two-ribbed, promptly deciduous cataphylls,
and narrowly ovate blades with a narrow V-shaped
sinus, more or less free basal veins, and a more ог
less bluish green lower surface.
Philodendron brenesii is most easily confused
with P. crassispathum Croat & Grayum, which has
similar venation but more broadly ovate blades and
a nearly ellipsoid spathe with very thickened walls
(usually more than 1 cm thick) and no sign of a
constriction between the blade and tube portions.
The peduncle on the latter is sometimes so short
that it appears to be lacking. In contrast, the spathe
of P. brenesii is of normal thickness and shape (e-&
the spathe is divided into a tube and blade portion).
Philodendron straminicaule can sometimes be con-
fused with the more narrow-leaved forms of P. bre-
nesii, but that species differs in having a hippocre-
piform, rather than a narrowly V-shaped, sinus.
The leaf blades of Philodendron brenesii closely
resemble those of P. validinervium Engl. from
coastal Ecuador between Nanegal and Gualea. That
species differs in having proportionately longer and
narrower posterior lobes, more closely spaced and
prominently raised primary lateral veins, a long-
pedunculate spathe (1.2 times longer than the pe
Volume 84, Number 3
1997
Croat 411
Philodendron Subgenus Philodendron
duncle vs. 1.6–3.1 times longer), and а 4-locular
ovary with about 4 ovules per locule (vs. a 5-7-
locular ovary with up to 12 ovules per locule).
Philodendron brenesii is one of the most com-
monly cultivated species throughout the Meseta
Central in Costa Rica. A specimen collected in Co-
clé Province of Panama (Croat 67578) is somewhat
disjunct from the nearest populations in Chiriquf
Province, but perhaps belongs here as well. The
material has juvenile blades more broadly ovate
than in other populations of the species. It also
lacks the bluish green coloration on the lower blade
surface and has the dried midrib dark rather than
pale. The adult blades of the Coclé collections are
remarkably similar to those of the Chiriquí collec-
tions, and the other differences may be due to the
fact that these plants occur near the lower part of
the elevational range. A number of differences in
the pistil argue that this collection may represent a
different species. Pistil differences in the Coclé col-
lection include, among other things, the presence
of a style funnel and a style dome (lacking in the
Chiriquí populations). The Coclé collection also has
eight locules per ovary and two = per locule,
perhaps another important differe
A Costa Rican collection усе et al. 8789)
from Tarrazú in San José Province is unusual in
having smaller leaves with indistinct primary lat-
eral veins and petioles drying minutely wrinkled.
The dried blade color, minor venation, and dried
stem characters otherwise match P. brenesii. While
this might represent a new species, more informa-
tion is needed. Herrera et al. 8789 also is similar
in stature and blade shape with P. knappiae from
Chiriquí Province in Panama. The collection differs
from P. knappiae in its pale gray-green drying (vs.
dark brown) lower blade surface, lack of secretory
ducts between the minor veins, and a deeply sunk-
en style into the apex of the pistil on drying (vs. a
style held above the apex of the pistil on drying).
Additional specimens examined. COSTA RICA. Ala-
Tw San Ramón-Balsa, ca. 5.7 km N of Quebrada Volio,
1100-1150 m, 10%08'N, 84°29'W, Stevens 14124 4. Б
MO, NY, ae Cerro el Chayote-Zarcero, Poveda 1
(CR, F, MO); 8.9 mi. NW of San Ramón, Tx m,
10*10'30N, 84°30'W, Croat 68084 (B, CM, DUKE, F, K,
5 MEXU, MO, NY, US); 1. am mi N of Angeles Norte,
1200 m, Croat 46879 (MO); ca. 11 mi. NW of
900 m, Croat 43521 (CR, MO); ny past Zarcero, 1800
m, m, Hoover 1350 (CR, MO); Volcán Poás—Volcán Barba, 7
i. N of Carrizal, 1850 m, Croat 35491 (F, MO). Cartago:
са. 7.3 km NE of Pacayas, 5200 ft., Wilbur et al. 16086
(MO); road to Moravia, ca. 20 km E of Rfo Pacuare, 1150
m, 9°50'N, 83%24'W, Thompson & Rawlins 1230 (CM);
hp de la Muerte, between summit € Empalme, aes
m, Croat 35415 (CR, L, MEXU, MO); La Cangreja,
km 5 of El Tejar, 1850 m, Williams et al. 24191 (F, NY:
e
Río Grande de Orosi, 15 km S of Tapantí, E slope above
río, 1500 m, Burger & Liesner 6713 (F); Río Naranjo, 3.5
km E of Cachí, 1360 m, Lent 1431 (F); Tapantí Hydro-
electric Reserve, “ч Río Grande de Orosi, 4.5 km ђе-
yond small bridge, 1500-1700 m, Croat 36111 (MO); ca.
ет S of jet. of А — & Río Grande de Orosi,
00-1800 m, 9?43'N 7'W, Croat & Grayum
petet (MO); 1200 m, ES 990 (F). Guanacaste:
Guachipelín—El Volcán de la Vieja, Brenes 15565 (Е, NY);
SW slopes of Volcán Rincón de la Vieja and Volcán Santa
María, trail from Hacienda Guachipelín, 1400 m,
10%48'N, 85%21'W, Burger & Pohl 7771 (CR, F, MO,
PMA); Fila del Volcán Cacao, 1400-1520 m, Chacón &
ight 2302 (MO); 1 km N i
of Monteverde, 1200
|ui 9519 (CR, INB, MO, МУ); Río agro 820
m, 10746'30"N, 85?20'35"W, Rivera 660 (CR, MO). He-
redia: 2 km S of Vara Blanco, 1900 m, Wilbur et al. 15711
(DUKE); NW slopes of Volcán Barba, Río San Rafael, Lent
1299 (CR, F, US); не Poás—Vara Blanca, 1.5 km past
divide in road, 1930 m
0 = Protectora Río vien fo Guácimo, N
slopes Volcán Barba, Стауит & Schatz 3232
(DUKE); Río e Rafael, уверена = of Volcán Barva,
1500 m, 10?13'N, 84?05'W, Grayum et al. 7750 (MO).
Limón: Moravia, 1300 m, Williams 161 a (EAP); Cantón
de Talamanca, Bratsi, Amubri, А
==
> *
Lori, 1700 m,
Monteverde Cloud Forest Reserve, 1450-1650 m,
10°18'N, 84°47'W, Burger & Baker 9767 (CR, F); 1700
m, 10°20'N, 84750", Haber & Bello 4115 (MO); 1700
m, Haber 2413 (MO); me ки m, Haber & =e mage
9827 (INB, MO); Zarcero region, Palmira, 5700
eee 143 e Osa, vic. Boscosa, re 78806 (CR, TNB,
МО). 5 : vic. of Vara Blanca, 1880 т, Croat 35519
Me р Pará m Cerros de! Zurquí; 1
84701", Burger et al. 10242 (Е, МО); SW бой
of Ра T Blanco, lower slopes of Cerro Zurquí, таме
& Utley 1270 (F); La Palma-San €— Utley
532 (F); 2 km N of на 12. С m W о ites
American Highway, 2 ш = (СЕ, МО);
along CA-2, Cerro de p rales N of turn off for road
aiaia pe m, Croat 32857 (МО); Patarrá, Cerro El Es-
9°53'N, 84°02'W, Chacón & do
DS R, МО); pen Cerros de Escazú, 1950-2100
Morales 1305 (CR, INB); Rincón de la Vieja, Boucler 233
(CR); Tarrazú, Herrera et al. 8789 (MO). PANAMA. Chi-
riquí: Callejón Seco, Volcán de Chiriquí, 1700 m, Wood-
son & Schery 510 (F, SH MON 4 km past divide in road
to Alto Quiel from Boquete, 1600 m, 8°49'N, 82°28'W,
e 1337 ја байса. Сп Grande, 5. 9 пи. ђе-
ond Los Plan it 5 m, 845'N, 82°14’ W,
asi 67793 (AAU Д
ОМ, QCA, SAR, TEX, US); 5.5 mi. NW of Los
Planes de Hornito, 1320
74914 (МО); Сето. Colorado, ca.
Félix bridge, 800-1200 m, Croat ж (МО); 24 mi. N
of Río San Félix, 1430-1500 m, Croat 48487 (MO); above
San ява 33184 (MO); Los Planes de Hornito beyon
Gualaca, 900 m, Croat 48879 (MO, SAR, » ne
quete sars W of Río Caldera, ca. 2 ks NW o
Mono, 1700 m, 849'N, td Grayum el T
> = ті. W of Chame, 1300 m, 8°35'N, 81750",
Cr 9090 (AAU, CR, MO, NY, PMA, VDB); near Con-
oad Divide, 1500 m, Antonio 1497 (MO, NY); Cerro
412
Annals of the
Missouri Botanical Garden
Hornito, S slope approached from Los Planes de Hornito,
640 m, 8°41'N, 82°11’ W, Croat 67937 (CAS, MO); Cerro
Небе, 1650 т, Croat 27000 (CM, MO). Coclé: Alto
Calvario along summit of Continental Divide 5.5 mi. N of
El Copé, 3.5 mi. N of Escuela Barrigón, 850 m, 8”39'N,
80°36'W, Croat 67578 (CM, CR, K, мо, NY, риа, US);
1.5 mi. N of El Copé, ca. 900 m, Croat 44577 (MO).
Philodendron brevispathum Schott, Bonplandia
7: 29. 1859. TYPE: Panama. Canal Zone: Cha-
gres River, Fendler 431 (holotype, MO; iso-
type, K). Figures 90, 91.
Philodendron arcuatum K. Krause, in Engl. & K. Krause,
Pflanzenr. at se (Heft 60): ia 1913. TYPE: Bo-
livia. Pan Acre, Cobija (on Brazilian border.
SW ^ Rio Banc: 1102: 68°44’ W, Ule 8819 ње
lotype, B).
Philodendron holmquistii G. S. Bunting, Acta Bot. Venez.
1975. Philodendron brevispathum subsp.
ria = S. Bunting) G. S. Bunting, Phytologia
64: TYPE: Venezuela. Amazonas: Pueblo
età onn zone between the Río Pacimoni and the
orest, 1°50'N, 66730", Steyermark & Bun-
ting 1 22 быры. ae isotype, MY).
Terrestrial or hemiepiphytic; stem scandent,
coarsely scabrous, densely covered with trichome-
Li often branched scales; internodes elongate,
5-14(23) cm long, 8-15 mm diam., longer than
Бол: epidermis reddish brown, loosening and
flaking, without fissures; roots drying tan to dark
brown, few per node; cataphylls 6–9 cm long, soft,
unribbed, green, drying dark reddish brown, even-
tually deciduous; petioles 10-43 cm long, 3-7 mm
diam., subterete to bluntly C-shaped, somewhat
spongy; blades ovate to ovate-triangular, concol-
orous, semiglossy, thin, acuminate at apex, cordate
at base, 16-36 cm long, 11-26 cm wide (1.4-2
times longer than wide), (0.8—1.6 times longer than
petiole), broadest at or above middle, upper surface
semiglossy, lower surface semiglossy; anterior lobe
12-30 cm long, 10.8-24 cm wide (1.3-2.4 times
longer than posterior lobes); posterior lobes some-
what triangular to narrowly triangular to rounded,
or rounded to with hastate or flaring lobes, 5-15
cm long, 4.9-11.5 cm wide, subacute; sinus narrow
or sometimes V-shaped, 9-11 cm deep; midrib
sunken above, convex to raised, drying slightly pal-
er below; basal veins 3—4 per side, with 1 free to
base, coalesced 4-10 mm, or with 2-3 veins co-
laesced to 3 or 4 cm; posterior rib not naked or
obscurely and briefly naked to 0.5 cm long; primary
gres veins (3)4—5 per side, departing midrib at a
0° angle, gradually ascending to the margins
edd sunken above, convex and drying digity
darker below; interprimary veins almost as con-
spicuous as primary lateral veins; minor veins aris-
ing from both the midrib and primary lateral veins;
tertiary veins visible and darker than surface below.
INFLORESCENCES erect, 1 per axil; peduncle 1–
7 cm long, 2.5-5 mm diam., subterete; spathe 6-
10.5 cm long (1.4—4.6(9) times longer than pedun-
cle); spathe blade green outside, red inside; spathe
tube green to greenish white outside, 4—4.5 cm
long, 2-2.5 cm diam., green to white inside; spadix
sessile; 7-9.5 cm long, whitish to yellowish; pistil-
late portion 2-3 cm long, 3.5 cm diam. throughout;
staminate portion 5-7 cm long; fertile staminate
portion 5-10 mm diam.; sterile staminate portion
mm diam.; ovary ca. 6-locular, with axile pla-
centation; ovules 6-14 per locule, 0.3-0.4 mm
long, longer than funicle; funicle 0.2 mm long, ad-
nate to lower part of partition, style similar to style
type B; stylar canals emerging at base of pro-
nounced apical depressions; style apex drying dark
brown with a pale undulate margin and a central
solitary stigmatic pad; the androecium truncate,
prismatic, margins distinctly scalloped, ens
4—5-sided, 0.9 mm long; thecae oblong, 0
wide, nearly contiguous. INFRUCTE е ke
peduncle 4 cm long, spathe 5—6 cm long; spadix,
3.54 cm long, 2.5 cm wide; berries pale yellow, 6
mm long, 3 mm diam.; seeds 4—6 per berry, oblong
to weakly ovoid, 1.4 mm long, 0.4-0.5 mm diam
Flowering in Philodendron brevispathum is ap-
parently aseasonal, perhaps owing to its habitat
along and near streams. Flowering collections have
been made in February through April, July, August.
and November. Immature fruiting collections were
also made throughout most of the year, January
through April, July, September, and November.
Though long known only from the Caribbean
coast of Panama, Philodendron brevispathum is now
known to be a locally rare but more widespread
species with special ecological requirements. It
ranges from Nicaragua to Panama at 0-280 m wi
disjunct populations in South America, there rang
ing from Venezuela to Brazil (Amazonas), Colombia
(Vichada, Meta), Ecuador (Napo), Peru (Loreto),
and Bolivia, at elevations of 60 to 230 m.
Philodendron brevispathum is a member of P
sect. Philodendron subsect. Solenosterigma. lt
grows as a low hemiepiphyte or in drier areas as à
terrestrial herb, usually along streams, on river
banks, or in swamps. Maguire 36157 is unusual in
being reported as a vine growing to 13 m in trees.
The species is recognized by its scandent habit, its
thin ovate to ovate-triangular leaf blades with
rounded to hastate posterior lobes and a qu
narrow (sometimes V-shaped) sinus, and especi
by the stems that are densely covered with не
chome-like, often branched scales and dry with а
Volume 84, Number 3
1997
Croat 413
Philodendron Subgenus Philodendron
flaky brown periderm. While the posterior leaf
lobes are commonly somewhat triangular (and may
even be narrowly triangular and subacute at apex),
they may be nearly pea on the same collection
(Davidse & Gonzalez 1
Bunting (1988) i eR the South Ameri-
an material as P. brevispathum subsp. holmquistii,
У osos PM by having 7-14 ovules per locule (vs.
14—18 for the typical material in Central America)
and pistils with the apex convex with 3—4 stigmatic
pads. Yet my observations indicate that Central
American material also has 6–14 ovules per locule.
This species is closest to P. muricatum Willd. ex
Schott (an older name) and may not be separable
from it. The latter species is distinguished by hav-
ing densely verrucose-warty petioles and usually
smaller blades with more rounded lobes. However,
Delta Amacuro, RIAM pd and Krukoff 7250 from
southern Amazonas State, Brazil (the type of P. am-
plectans А. C. Sm., a synonym of P. muricatum),
have narrow, more or less triangular blades like
those of P. brevispathum. On the other hand, not all
specimens of P. brevispathum have similar lobes.
Central American material commonly has more
rounded or elongate lobes that are turned somewhat
inward, but some sheets (e.g., Burger & Antonio
11236) have blades identical to those of South
American plants. In addition, some South Ameri-
can collections (Croat 58586, Davidse 4294) have
the posterior lobes noticeably rounded and scarcely
longer than broad. Despite this variation, the ver-
rucose petiole character is adequate to separate Р.
brevispathum from P. muricatum.
Philodendron brevispathum has also been con-
fused with Р. jacquinii, but that species has merely
puberulent stems (with trichomes simple and un-
branched), thinner, more broadly ovate leaves (of-
ten also puberulent on the petiole and lower mid-
rib), a conspicuously bulging spathe tube, and more
elongate styles
Additional specimens examined. COSTA RICA. He-
redia: Puerto Viejo-Guápiles, along Río Puerto Viejo, 7
km N of Buenos Aires, 10?23'30"N, 83?48'30"W, Croat
684 MO); La Selva Field Station, Grayum m
(DUKE, MO); Grayum 2642 (DUKE, F, MO),
Hammel & Trainer 10810 (DUKE); Río F Мо де E
between Río Sucio and railroad tracks, SW of Finca Zona
Ocho, 110 m, 10°18’N, 83%52'30"W, Grayum & Hammel
5568 (MO). Limón: Río Bananito-Cahuita, near Punta
Vargas ca. 4 km S of Cahuita, 0-10 m, Burger et al. 10493
(F, MO); 16 airline km SW of Barra del Colorado, 10-120
m, 10°39’N, 83°40’ 40" W, Davidse & Herrera 31254 (MO);
Refugio Barra del фе ан area between Río Chirripocito
and Río Sardina, 12 m, 10%38'N, 83°45'W, Grayum et al.
9742 (AAU, CR, MO); Fines Tapezco-La Suerte, 29 air
km W of Tortuguero, 40 m, 10?30'N, 83747", Davidson
& Donahue 8842 (МО); 40 m, 10°30’N, 83^47' W, 8970
(RSA, MO); Gandoca (slightly to N of trail from Mata de
Limón), a m, 9*36'N, 82?36'30"W, Grayum et al. 8024
hes K, M, MO); Cerro Coronel, 10-40 m, 10?40'N,
40'W, Бави et al. voti A MO); Parque Tortu-
= National Раг m, Robles 1410 (CR,
MO); 4 m, 10°32’N, ' 83°30 W, "Robles з 1877 (CR, F, MO);
near Boca de las Lagunas de Tortuguero, 0-3
10°34'N, 83°32'W, Burger & Antonio 11236 (F, MO, U)
NICARAGUA. Matagalpa: Rancherfa, 11 km а! МЕ de
Muy Muy, 280 m, 12°46'N, 85°31'W, Moreno 24433 (MO).
Río San Juan: Quebrada Santa Crucita, 50 m, 11%02'N,
Sábalos, cerca de “La Toboba,” 70-90 m, 11%03-04'N,
84°28-29'W, Robleto 1833 (MO, US). Rivas: Isla de Ome-
tepe, La Argentina m, 11%27'N, 85°32’W, Mo-
reno 22112 (CM, MO). Zelaya: T Barra de Punta Gorda,
MO); Santa Marta, `5 m, 14°18'N, 8337'W, Stevens & Mo-
reno 19623 (MO); SW of Bluefields, 10—40 m, 11?59'N,
83°46'W, Stevens 19736 (CAS, L, MO, UWL). PANAMA.
Colón: пое de la Borda, Croat 10012 (MO, SCZ). San
Blas: Playón Chico, road to Isisukun, 0-10 m, 9°20'N,
78°13'W, ыш 596 (МО).
Philodendron brewsterense Croat, sp. nov.
TYPE: Panama. Comarca de San Blas: Cerro
Brewster, 850 m, 9°18'N, 79°16'W, de Nevers,
Herrera, Hammel & Charnley 5545 (holotype,
MO-3 ADS nom 92.
Planta
4
longa, € sein leniter ¡ga acute 2- -cos-
tata, decidas petiolus 9-11 cm longus, m
m. et obtuse o: lamina pss leniter
subcordata, 95:1 cm longa, 7-8 cm lata, in sicco fla-
vibrunnea; nervis lateralibus 1 Sad sag DE
ls mh s 4. à cm longus, 3.5 mm latus; spatha o
nino in superfice mbabus maronnina, 9-9, 30 ст vic
pistilla 5- x адна oak 2-ovulati.
Epiphytic; stem scandent; leaf scars inconspic-
uous; internodes terete, 5-6.5 cm long, 4-5 mm
diam., much longer than broad, drying yellowish
brown, epidermis fissured narrowly; roots dark
brown, drying moderately fuzzy, slender, 1 mm
diam., very few per node; cataphylls weakly a
sharply 2-ribbed, glossy deciduous; petioles 9—
cm long, to 4 mm diam owly and поћи
sulcate adaxially, vidi reir wrinkled: sheath
conspicuous, the пр free-ending (2-3 mm long);
lades ovate, acuminate at apex (acumen inrolled,
1-2 mm long), weakly subcordate at base, 9.8-11
cm long, 7-8 cm wide (1–1.4 times longer than
wide; about equal in length to petiole), broadest in
lower one-third, margins revolute, drying reddish
brown, upper surface medium green, drying grayish
414
Annals of the
Missouri Botanical Garden
green, semiglossy, lower surface weakly glossy,
moderately paler, drying yellowish green; anterior
9–10.5 cm long, 7-8 cm wide (5.9-7.1 times
longer than posterior lobes), broadest in lower one-
third; posterior lobes 2-2.4 cm long, 2.8 cm wide,
rounded; sinus obtusely angular, 5 mm deep; pos-
terior rib lacking; midrib concave above, moderate-
ly raised, drying paler than surface below; basal
veins numerous on each side but none of them out-
standing, primary lateral veins lacking or indistin-
guishable from minor veins; minor veins close, aris-
roon throughout on both surfaces; spathe blade 6.9
cm long; spathe tube 4.5 cm long; spadix
throughout (probably juvenile), narrowly indst не at
apex, 7.3 cm long, constricted below sterile sta-
minate portion; pistillate portion drying grayish,
slightly tapered toward the apex, 2.1–2.5 cm long
in front, 1.2 cm long in back, 4.5—5 mm diam. at
apex, 5—6.8 mm diam. at middle; staminate portion
4.5-6.1 cm long; fertile staminate portion white,
gradually tapered towards apex, 6.5—6.8 mm diam.;
pistils 1.1 mm long; ovary 5-locular, 0.9 mm diam.,
ovule sac ca. 0.5 mm long, with sub-basal placen-
tation; ovules 2 per locule, contained within trans-
parent ovule sac, ca. 0.2-0.3 mm long, longer than
funicle; funicle 0.1-0.2 mm long (can be pulled
free to base), style 0.7–0.9 mm diam., similar to
style type D; style apex rounded; style boss broadly
and gently rounded; stigma button-like, distinctly
lobed, 1 mm diam., 0.3 mm high, covering entire
style apex, inserted on style boss; the androecium
truncate, prismatic, margins irregularly 4—6-sided,
0.9-1.1 mm long, 3-5 mm diam. at apex; thecae
oblong to weakly ellipical, 0.4 mm wide, + parallel
to one another; sterile staminate flowers subround-
ed, 5-9 mm wide, grayish to pale orange.
Flowering in Philodendron brewsterense is based
on a single flowering collection made in April in
an area that is somewhat aseasonal.
Philodendron brewsterense is endemic to Panama,
where it is known only from the type specimen col-
lected on Cerro Brewster in Comarca de San Blas
Province, at 850 m elevation in a Premontane rain
forest life zone.
ilodendron brewsterense is a member of P. sect.
Calostigma subsect. Glossophyllum ser. Glossophyl-
um. The species is recognized by its scandent hab-
it with internodes much longer than broad, drying
yellowish brown and narrowly fissured; small (<8
cm diam., «11 cm long), coriaceous, ovate, yellow-
ish brown blades lacking primary lateral veins; and
solitary inflorescences with the outer surface sur-
face таг
Philodendron brewsterense is apparently unrelat-
ed to any other species but is most similar to P.
blades usually more than 20 cm long and with four
to six pairs of primary lateral veins.
Philodendron brunneicaule Croat & Grayum,
sp. nov. TYPE: Panama. Coclé: Alto Calvario,
6 mi. N of El Copé, Atlantic slope along Con-
tinental Divide, along trail which heads off old
lumber trail and leads down to Las Ricas, Li-
món, and San Juan, 710—800 m, 8?39'N,
80*36'W, 22 June 1988, Croat 68713 (holo-
type, M0—3591332). Figures 93-97.
Planta —€— internodia 6–20 cm longa, 1-
1.2 cm diam., rubrobrunnea; cataphylla 20—40 cm longa,
incostata, o petiolus teres vel C-formatus, leviter
и adaxialiter 21-63 cm longus, 4-11 mm
; lamina ovata, cordata aut interdum sagittata br,
25-62 cm longa, 16—52 cm lata; inflorescentia 1; pedun
culus 4—20 cm longus, 8-9 mm т par = € pé
longa; lamina spathae extus cremea, intus virid
spathae extus viridi, intus cerasino; рыша ere pre
laria; loculi (13)15—18-оушан; baccae albae.
Hemiepiphytic; stem light reddish brown to rusty
red, leaf scars conspicuous, 8 mm long, 12 mm
wide; internodes sparsely short dark-lineate, some-
times weakly warty, semiglossy, 6-20 cm long, 1-
1.2 cm diam., longer than broad, dark green to ol-
ive-green to brown, epidermis loosening and flak-
кч reddish brown; cataphylls soft, 20-40 cm long,
]-ribbed, greenish
dish, deciduous; petioles 21-63 с
mm diam., erect-spreading to рибне terete, to
C-shaped, somewhat flattened adaxially, medium
green, surface matte, sometimes striate and dark
green- to red-lineate; geniculum 3 cm long, 2.5-3
cm diam., thinner and paler than petiole; blades
ovate, subcoriaceous, bicolorous, acuminate at
than wide), (0.7-1.6 times longer than petiole), up-
per surface dark green, glossy to semiglossy, lower
surface glossy to semiglossy, paler; anterior lobe
23-65 cm long, 1 1.5 times
longer than wide); posterior lobes rounded to ob-
tuse, 8-20 cm long, 9-26 cm wide; sinus spathulate
to hippocrepiform; midrib flat to sunken, paler than
Volume 84, Number 3
Croat 415
Philodendron Subgenus Philodendron
surface above, narrowly convex to round-raised,
paler than surface, sometimes tinged maroon below;
basal veins 5-9 per side, with 0-2 free to base,
(2)3-4 coalesced 1.9—4.5 cm; posterior rib naked
for 1—4.5 cm; primary lateral veins 4—6 per side,
departing midrib at a 45-55” angle, conspicuously
sunken above, narrowly convex and tinged maroon
below; interprimary veins weakly raised and darker
than surface below; minor veins arising from both
the midrib and primary lateral veins; tertiary veins
isible and sometimes darker than surface below.
INFLORESC ENCES erect, 1 per axil; peduncle 4—
20 cm long, 8-9 mm diam., medium green, whitish
at base, short dark green lineate; spathe glossy,
11-21 ст long (1.05-2.85 times longer than pe-
duncle), constricted midway above the tube; spathe
blade cream outside, greenish white (at anthesis)
inside; spathe tube 6.5-9 cm long, green outside,
cherry-red inside; spadix 9—17 cm long, broadest
above the middle or = uniform throughout; pistil-
late portion cylindrical to obovoid, 3.8-7 cm long,
7-13 mm diam. at apex, 8-14 mm diam. at middle,
10-14 mm wide at base; staminate portion 4.6—
17.5 cm long; fertile staminate portion clavate, 9—
16 mm diam. at base, 12-20 mm diam. at middle,
7-10 mm diam. ca. 1 cm from apex, broadest at
middle, broader than the pistillate portion, narrow-
er than the sterile portion; sterile staminate portion
as broad as or narrower than the pistillate portion,
0.9-1.6 cm diam.; pistils 1.3-3.2 mm long, 1-1.8
mm diam.; ovary 5(6)7-locular, 0.9-2.1 mm long,
0.9 m Eng with axile placentation; ovules
(1315-18 per locule, 2-series, 0.1—1.7 mm long,
longer than or equal in length to funicle, style 1.2
mm diam., similar to style type B; style apex round-
ed; stigma subdiscoid, lobed, 1.5 mm diam., 0.3
mm high, covering not quite entire style apex; the
androecium truncate, 3—6-sided, 2.2 mm long; the-
cae oblong, 0.2-0.5 mm wide; sterile —
flowers blunt with one side scallope
long, 0.7-1.9 mm wide. INFRUCTESCENCE with
berries white. JUVENILE plants with internodes to
10 cm long, 4-10 mm diam.; cataphylls reddish,
persisting at upper nodes, sharply C-shaped in
Cross section.
Flowering in Philodendron brunneicaule proba-
bly occurs in the early rainy season, but a single
(post-anthesis) collection was made in January.
Most post-flowering collections have been made be-
tween April and July, with immature fruits collected
in August and October.
Philodendron brunneicaule ranges from Costa
Rica to Panama, Colombia (Valle), and Ecuador
(Esmeraldas), at 50 to 1300 m elevation in Tropical
neicaule is a member of P.
sect. Philodendron subsect. Platypodium. The spe-
cies is characterized by its appressed-climbing
habit, long internodes with thin, flaking reddish
brown epidermis (hence the name “brunneicaule,”
from “brunneus” meaning brown, and “caulis”
meaning stem), white, unribbed, deciduous cata-
phylls, somewhat adaxially flattened and red-line-
ate petioles, ovate blades with reddish-brown-dry-
ing veins, large, solitary inflorescences borne at
several of the uppermost internodes, and green
spathes colored cherry-red inside on the tube.
ilodendron brunneicaule is probably related to
P. ernestii Engl. from Amazonian Ecuador and Peru.
That species shares long internodes with flaking
rown epidermis and similar, solitary inflores-
cences. It differs, however, in having stems that are
often warty and petioles that are undulate-winged
vs. more nearly terete for P. brunneicaule. It is note-
worthy, however, that a single collection from Ama-
zonas Department, Peru (Vásquez & Apanu 19051,
МО), appears to lack a ен wing. This may
prove to be P. brunneicaule, but if so it would be
the first collection from east of the Andes.
In Central America Philodendron brunneicaule is
most easily confused with P. copense. See that spe-
cies for the differences.
Additional specimens examined. a RICA. Ala-
juela: San Ramón- peu. ca. 0 7 km N of bridge over
pipe е and са. 7.5 km N of Río Balsa, 700-800
'N, 84° ar W, aed 13859 (CR, F, MO);
000 m, 10°12’N,
84732"М, те e Judziewicz 14667 (CR,
Мобнйоніе Río Peñas Bi 900 m Bello 369 (CR,
Soco
5517 (МО); 830 т, 10°16'N, 8411’ үу, Croat 68302 (СМ,
G, M, MO, NY). Heredia: Puerto Viejo-Guápiles, 7 km
N of Buenos Aires, <100 m, 10?23'30"N, 83"48'30'
Croat 68402 (MEXU, MO); La oet Field Station, Gra-
eraldas
yum 2790 (F, MO). ECUADOR. E : Quinin чн
Bilsa Biological Station, Montañas de Mache, 35 km
of Quinindé, 5 km W of Santa Isabela,
10?21'N, 79%44'W, Pitman & Bass 1085 (MEXU, MO;
NY, QCNE). PANAMA. Bocas del Toro: Fortuna Dam
area, Fortuna Dam-Chiriquí Grande, 2.8 road mi. N of
Divide, 850-950 m, 8°45'N, tee W, McPherson 9661
(MO); 1.2 mi. N of PUN. E 3m of bri
Pru Eod 60468 (AAU,
tinental Divide, 1170 m, 8°44’N, 81°17'W, Croat 66655
(L, MO). Chiriquí: Gualaca-Fortuna Dam, 10.1 mi. NW
of Los Planos de Hornito, 1300 m, 82?17'W, 8'45'N,
Croat 49836 (COL, K, MO, NY); 9.4 km above El Copé,
900 m, Croat 44733 (MO). Coclé: El Copé, Alto
416
Annals of the
Missouri Botanical Garden
Calvario above El Copé, 4.5 mi. N of El Copé, 580-740
m, 8°38’N, 80° d W, шен a (AAU, MO, W); 930
m, 49183 (MO, US). Pan ano—Cartf road, 10
m from Inteis American = near El Llano, 330 m,
Croat 33823 (MO); Km 14, 350-500 m, Folsom et al.
1483 (MO); 10.1 mi. above highway, 325-350 m, Croat
67365 (MO). San Blas: El Llano-Cartí Road, km 19, 350
m, 9?19'N, T8'55' W, de Nevers et al. 5599 (B, K, MO,
COLOMBIA. Valle: Bajo Calma region, Buenaventu-
ra-Málaga, Km 28, 50-150 m, 3°59'N, 77°03'W, Bay 240
(CUVC, MO).
Philodendron chiriquense Croat, sp. nov. TYPE:
Panama. Bocas del Toro: Cerro Colorado, 9.2
km W of Chame, along trail E of road leading
down to stream, 1450-1480 m, 8°35'N,
81°50’W, 6 July 1988, Croat 69068 (holotype,
MO-3599857; isotypes, B, CAS, CM, COL,
CR, F, GH, К, MEXU, PMA, US). Figures 98—
100, 107.
Planta ces: internodia 1—4 ст longa, (1.5—
2)3—4(6-10) cm diam.; cataphylla (20)55-70 ст longa,
acute pa sabes Eie uh in sicco rubrobrunnea, per-
sistentia semi-intacta; petiolus subteres vel D-formatus,
27-108 ст longus, in sicco 9-15 mm diam.; lamina trian-
gulari- -sagittata, ейн basi, 32.5-91 cm longa, 11-44
diam., albus suffus patha 14.2 cm longa,
lamina spathae extus alba, intus albida, suffusa marronina
usque ad dimidium; tubo spathae extus pallide v iridi, 1 in-
tus marronino; pistilla Block lan loculi cirea 14-ovulat
Hemiepiphytic; stem appressed-climbing to 4 m
high or creeping, to 30 cm long; internodes short,
very thick, semiglossy, 1-4 cm long, (1.5-2)3—4(6—
10) cm diam., much broader than long, medium to
pale green, coarse white-ribbed at upper edge, com-
pletely hidden by cataphyll fibers, drying brown;
roots several per node, to 4 mm diam., light reddish
brown to dede brown with flaky lighter brown epi-
dermis, weakly glossy; cataphylls (20)55-70 cm
long, sharply 2-ribbed, sharply sulcate with acute
brown, with two low ribs, persisting semi-intact at
upper nodes, as pale fibers at base, eventually as
dark brown to reddish brown fibers; petioles 27—
108 cm long, 9-15 mm diam., subterete to obtusely
D-shaped, firm and flexible, obtusely to weakly flat-
tened adaxially, with adaxial margins sharp to blunt-
ly rounded, dark green, base reddish, weakly glossy,
surface faintly and densely white striate to coarsely
pale striate, drying dark brown; blades triangular-
sagittate, moderately coriaceous to subcoriaceous,
short- to long-acuminate at apex (the acumen some-
times inrolled), cordate at base, 32.5-91 cm long,
11-44 cm wide (1.34-3.28 times longer than wide),
(0.58-1.05 times longer than petiole), about equal
in length to petiole, broadest at or near point of pet-
iole attachment, upper surface dark green, with vel-
vety sheen, drying gray-green to yellow-brown above,
lower surface yellow-green, semiglossy, paler, drying
reddish brown below; anterior lobe 39.5—68 cm long,
22-53 cm wide (1.2-1.8 times longer than posterior
lobes); posterior lobes 6.2-29.6 cm long, 10.3-26.5
cm wide, directed inward, obtuse to rounded; sinus
parabolic, hippocrepiform or spathulate; midrib nar-
rowly rounded to almost flat to weakly sunken, dry-
ing gray-brown, paler than surface above, narrow-
rounded to convex, concolorous or paler than sur-
face, drying reddish brown below; basal veins (3-
4)5-8 per side, with 0-1 free to base, (1)2—3 coa-
lesced 0.9-7.5(11.5) cm, posterior rib naked 24
cm, sometimes not naked but with the rib sometimes
running very near the margin; primary lateral veins
7-8 per side, departing midrib at a 60—70° angle,
nearly straight to the margins, obtusely sunken, con-
colorous or paler than surface above, prominently
convex to narrowly rounded, concolorus or paler than
surface below; interprimary veins many, distinct; mi-
nor veins fine, moderately distinct; “cross-veins” vis-
ible on lower surface, sometimes weakly visible
above. INFLORESCENCES + erect, 3 per axil; pe-
duncle 9 cm long, 8-9 mm diam., white tinged red,
pale white striate toward apex, turned at 100° angle
to spathe at anthesis; spathe 14.2 cm long (I. 57
times longer than peduncle), weakly constricted
above the tube; spathe blade white outside, whitish,
suffused maroon to about midway inside; spathe t
globose, 6 cm long, pale green, tinged red on front
outside, densely short, pale streaked throughout out-
side, maroon inside, spadix stipitate to 5 mm long;
ovate, 11.6–12.5 cm long, broadest below the mid-
dle; pistillate portion greenish white, clavate, 2.7 cm
long, 1.6 cm diam. at apex, 1.5 cm diam. at mi middle,
1.1 cm wide at base; staminate portion 10.9 cm long;
fertile staminate portion weakly tapered, 1.8 cm
diam. at base, 1.6 cm diam. at middle, 1 cm diam.
ca. 1 cm from apex, broadest at base, broader than
the pistillate portion, narrower than the sterile por-
tion; sterile staminate portion broader than the pis-
tillate portion, 1.8 cm diam.; pistils 4.4 cm long, 18
mm diam.; ovary 5-locular, with axile placentation;
ovules ca. 14 per locule, 2-seriate, contained wi
gelatinous matrix (no true envelope), 0.6 mm long.
longer than funicle, style 2 mm diam., similar to
style type B; style apex rounded; stigma subdiscoid,
lobed, 1.5 mm diam., 0.3 mm high, drying lobed,
covering center of style apex; the androecium trun-
cate, margins 4-6-sided; thecae oblong, 0.2 mm
Volume 84, Number 3
1997
Croat 417
Philodendron Subgenus Philodendron
wide; sterile staminate flowers blunt, 4 mm long, 1.6
mm wide. JUVENILE blades with lower surface dark
maroon.
Flowering in Philodendron chiriquense is known
only during July. While this is, in general, the early
wet season in Panama, the region around Fortuna
where the species is common is somewhat less sea-
sonal than other areas of Panama.
Philodendron chiriquense is endemic to Panama,
known from Bocas del Toro, Chiriquí, and Coclé
Provinces in Tropical Lower Montane rain forest and
Premontane rain forest life zones at 500 to 1630 m
elevation. It is frequent in the cloud forests near
the Continental Divide in the Fortuna Dam area.
ilodendron chiriquense is a member of P. sect.
Philodendron subsect. Philodendron ser. Fibrosa.
The species is distinguished by its short, very thick
internodes; sharply two-ribbed, red-tinged cata-
phylls, which persist as semi-intact, dark brown fi-
bers; obtusely flattened to D-shaped, densely
pale-striate petioles; huge, triangular-sagittate
blades; and up to three inflorescences per axil with
the spathe tube pale green outside and maroon
within.
Philodendron chiriquense is similar to P. с
which also has persistent reddish brown EA
fibers and reddish-brown-drying, more or less tri-
angular blades, but that species differs in having
petioles drying reddish brown, smooth and matte,
with peeling periderm (usually blackened, minutely
fissured and semiglossy in P. chiriquense), primary
lateral leaf veins drying reddish brown and darker
than the surface (pale and lighter than the surface
in this species), and minor veins and “cross-veins”
both distinct (minor not distinct and “cross-veins”
scarcely or not at all visible in P. chiriquense).
The only collection from Coclé, Croat 44566,
may represent another species. This collection dif-
fers from Chiriquí material in having the primary
lateral veins scarcely paler than the surface and
more prominent *cross-veins."
Additional specimens examined. PANAMA. Bocas
del Toro: Cerro Colorado, 8.6 mi. W of Chame, 1450-
1480 m, 8?35' N, 81750", Croat 69133 (CM, МО); 7 mi.
W of Chame, 1500 m, 8°35’N, 81°50" W, 69215 (MO, NY).
hiri San Félix,
485 m, Croat 75011 (CM, MO, PMA); Fortuna Dam area,
Gualaca-Chiriquí Grande, 1 km S of Continental Divide,
1075 m, 8°45'N, 82°18'W, Croat 66865 (MO); Los Planes
0
of Pla
m, 8°44’N, 82 mow Croat 49905 (K, MO, US); 8.3 mi.
NW of Las Planes de Hornito, 1260 m, 8°44'№, 82716",
Croat 49940 (MO); 10 mi. NW of Los Planes de Hornito,
1260 m, 82717", 8'45'N, Croat 50101 (MO); Fortuna-
pce Grande, 1170 m, 8°44’N, 81°17W, Croat 66666
"S
2.
s
Ax
©з A
Ro
ш
>
=
=>
т]
8 x
| ==
2
HE
2
со
N
о
—
ЈЕ
=
Crit & Zhu 76345 (MO); Fortuna Lake area, 3.4 km N
of Quebrada Chorro, 1.6 mi. N of center of bridge over
lake, 1205 m, 843'N, 82714", Croat 74956 (MO). Co-
clé: El Copé region, near Continental Divide, ca. 1.5 mi.
N of El Copé, 900 m, Croat 44566 (MO); Alo: Calvario
along summit of Continental Divide 5.5 mi. N of El Copé,
N, 80
850 m, 8°39’ *36'W, Croat 67573 (MO, NY); ca. 6
mi. N of El Copé, 770 m, "№, 80735'W, Croat & Zhu
76780 (MO, PMA). Veraguas: Santa Fe region, ca. 15 k
past Escuela Agrícola Alto Piedra above Santa Fe, Río
Caloveborita, Atlantic watershed, 500 m, Sytsma & An-
dersson 4772 (MO).
Philodendron chirripoense Croat & Grayum, sp.
nov. TYPE: Costa Rica. San José: Canaán-
Chirimol, above Río Chirripó del Pacífico,
Burger & Liesner 7139 (holotype, F; isotype,
CR). Figure 101.
Planta hemiepiphytica; M scandens; internodia
m longa, minus quam iam.; cataphylla cete
ак еј teres, 7-7.4 cm ана 1 mm diam., obtuse com-
planatus adaxialiter; lamina anguste ovata, subcordata
5 ст lata; nervis basalibus
1
gus,
rubella; pistilla (3)5-6(8)- -locularia; loculi 8-12-ovulati.
Hemiepiphytic; stem scandent; internodes very
long and slender, 20 cm long, less than 3 mm
diam.; roots drying dark brown; cataphylls un-
known, probably <10 cm long, lanceolate, decid-
uous; petioles 7-7.4 cm long, 1 mm diam.,
obtusely flattened abaxially; blades narrowly ovate,
very long-acuminate at apex (the acumen 2.4—2.6
ст long), subcordate at base, 11-11.6 cm long, 5—
5.5 cm wide (ca. 2 times longer than wide), (ca. 1.5
times longer than petiole), about equal in length to
petiole, drying brown; posterior lobes rounded to
obtuse, 2-3 mm long, 1.4-1.7 cm wide; sinus ar-
cuate with blade decurrent on petiole, 2-3 mm
deep; midrib drying with up to 5 ribs above; basal
veins 2-3 per side, with all free to base, obscure;
primary lateral veins 2 per side, departing midrib
at a 40—45? angle, straight to the margins; minor
veins arising from the midrib only. INFLORES-
CENCES nearly as long as leaves, probably 1 per
axil; peduncle 6 cm long, 4 mm diam., 0.83 times
the petiole length; spathe 7.4 cm long (1.2 times
longer than peduncle), green to reddish within;
spadix pinkish throughout (fide field notes, in part
= pistils 2.2 mm long, 1.1-1.2 mm diam.; ovary
3)5-6(8)-locular, with axile placentation; ovules
Annals of the
Missouri Botanical Garden
8-12 per locule, mostly 1-seriate, 0.1-0.2 mm long,
slightly longer than funicle; funicle ca. 0.1 mm
long, adnate to lower part of partition, style similar
to style type B; style apex domed; the androecium
truncate, oblong, prismatic, margins irregularly 4—
5-sided, 0.7-1 mm long; thecae oblong, 0.3 mm
wide, + parallel to one another; sterile staminate
flowers irregularly 4—5-sided, 0.8-1.2 mm wide.
Flowering in Philodendron chirripoense is based
on a single fertile collection made in December,
which is the early dry season on the western slope
of Costa Rica.
Philodendron chirripoense is known only from the
type specimen from the Pacific slope of southern
Costa Rica, in Premontane wet forest at 1000 m
elevation.
Philodendron chirripoense is tentatively placed in
P. sect. Philodendron subsect. Canniphyllum. The
species is characterized by its scandent habit, very
long and slender internodes (drying <3 mm diam.),
slender petioles about as long as the small, narrow-
ly ovate, subcordate blades, and especially by the
inflorescence being longer than the petioles.
Philodendron chirripoense is not similar to any
other species in either Central or South America,
but it might be confused with P. microstictum, the
only other species that has the inflorescence longer
than the leaves. The latter has larger blades (more
than 13 em wide), which are often broader than
long.
Philodendron clewellii Croat, sp. nov. TYPE:
Panama. Darién: middle slopes of W side of
Cerro Pirre, cloud forest, 850-1050 m, 29
June 1988, Croat 68945 (holotype, MO-
3589994; isotypes, F, K, PMA, US). Figure
102.
ata, in
runnea, nervis basalibus liberis (aut leniter coalatis); in-
florescentia usque 6; pedunculus 3.5-16 ст longus, 3-6
m diam.; spatha 4.5-7 cm longa; lamina spathae extus
cremea, intus purpurascente cum margine albido; tubo
spathae in superficiebus ambabus purpurascenti; pistilla
6(7)-locularia; loculi 2-ovulati.
3
E
S
Epiphytic; stem scandent, smooth, thick, mod-
erately glossy; internodes moderately glossy, 12-18
cm long, 3.5 cm diam., longer than broad, medium
green to dark gray-green, epidermis fissured weakly
longitudinally; cataphylls 18-25 cm
sharply 2-ribbed, green, caducous; petioles 24.5-
53 cm long, (2)7-14 mm diam., subterete, spongy,
dark green, obtusely flattened adaxially, surface
green or white streaked-lineate; blades broadly
ovate-cordate, subcoriaceous, moderately bicolo-
rous, acuminate at apex (the acumen 1-2 cm long),
cordate at base, 30-59 ст long, 20—45 ст wide
(1.25-1.83 times longer than wide), (0.95-1.65
times longer than petiole), upper surface dark
green, drying brown, semiglossy, lower surface
slightly paler, drying brown, sometimes red-brown
ог green-brown, semiglossy, paler; anterior lobe
26.7-42.2 cm long, cm wide (2.36-
3.95(5.28) times longer than posterior lobes); pos-
terior lobes broadly rounded, often overlapping, 7-
18 cm long, 12-21 cm wide, obtuse to rounded;
sinus narrowly closed; midrib weakly raised to al-
most flat, purple-spotted, paler than surface above,
convex, darker than surface below; basal veins 3-
5 per side, all free to base (or weakly fused); pos-
terior rib absent or to 1.5 cm long, not at all naked;
primary lateral veins 4—7 per side, departing mid-
rib at a 45-60° angle, straight to the margins,
weakly sunken, concolorous above, raised below,
secondary veins weakly visible; minor veins visible,
sparse, discontinuous below, arising from both the
midrib and primary lateral veins. INFLORES-
CENCES to 6 per axil; peduncle 3.5-16 cm long,
3-6 mm diam., drying 2-4 mm diam., green, heavi-
ly red-striped; spathe 4.5—7 cm long ((0.62)1.28-
2.33 times longer than peduncle); spathe blade
cream outside, purplish, with margins whitish in-
side; spathe tube purple outside, 2.5—3 cm long, 6-
0 mm diam., purple inside; spadix sessile; cla-
vate, 6 cm long; pistillate portion green, tapered
toward the apex, 2.5 cm long, 6 mm diam. at apex,
7 mm diam. at middle, 8 mm wide at base; stami-
nate portion 3.7 cm long; fertile staminate portion
white, clavate, 7 mm diam. at base, 9 mm diam. at
middle, 8 mm diam. ca. 1 cm from apex, broader
than the pistillate or sterile portions; sterile sta-
minate portion 7 mm diam.; pistils 1.6 mm long,
1.1 mm diam.; ovary 6(7)-locular, with sub-basal
placentation; ovules 2 per locule, contained within
translucent ovule sac, 0.2 mm long, equal in length
to funicle, style similar to style type B; style spes
concave; stigma discoid, unlobed, 0.6 mm diam.,
0.1 mm high, covering interior faces of stylar fun-
nel; the androecium truncate, 4—6-sided; thecae ob-
long to elliptical, 0.1-0.2 mm wide; sterile stam
nate flowers irregularly shaped, 2.1 mm long, 9 PM
wide, the margins blunt. JUVENILE plants with in-
ternodes 5-6 cm long, 8 mm diam., dark green;
petioles terete, sheathing to midway. 1
Flowering phenology in Philodendron clewellii is
unclear because there are too few fertile collec-
tions. It has been collected in flower in December
Volume 84, Number 3
1997
419
Philodendron Subgenus Philodendron
and post-anthesis in June, giving a hint of a bi-
modal flowering. Though Cerro Pirre is not mark-
edly seasonal, these two months in general would
mark extremes, with December the general begin-
ning of the dry season and June the beginning of
the rainy season in Panama.
Philodendron clewellii is endemic to Panama,
known only from the Serranfa de Pirre, from 850
to 1560 m elevation in Premontane wet forest.
Philodendron clewellii is a member of P. sect.
Calostigma subsect. Oligocarpidium. The species is
distinguished by its thick, smooth stems; usually
long internodes; sharply two-ribbed, deciduous cat-
aphylls; subterete petioles; ovate-cordate, dark-
brown-drying blades with free or weakly fused ba-
sal veins and narrow, closed sinus with often
overlapping posterior lobes; and especially by its
clusters of up to six small inflorescences per axil
with long peduncles, and spathe purple on the out-
side of the tube and cream on the blade.
In fertile condition, Philodendron clewellii does
not resemble any other described species, but the
leaf blades are similar to those of P. tysonii Croat.
That species differs in having no more than three,
much larger inflorescences at anthesis (to 12 cm or
more long). In addition, the leaves of P. tysonii usu-
ally dry darker. In the blade shape and lack of a
naked posterior rib, it is similar to P. grayumii
Croat. That species differs in having generally more
reddish brown blades with a broader sinus and
much stouter peduncles.
e species is named in honor of Andrew Clew-
ell, who made several expeditions to Panama while
working for Florida State University and who
helped collect the type specimen.
Additional specimens examined. PAN
Cerro Pirre region, 1000—1400 vilae Gentry ^ "usd 7028
(K, MO); Río Ca pares Esc oe Croat 37831
(MO); ca. 12 km N of Alto de Nique, 15 560 m, Croat
37915 (F, MO); ca. 16 N of Alto de Nique, 1530-1550
m, Croat 37946 (MO); middle slopes on W side, 800-
1050 m, 7°56'N, 77°45'W, Croat 68957 (MO).
Philodendron coloradense Croat, sp. nov.
TYPE: Panama. Chiriquí: Cerro Colorado,
along mining road 31.6 km beyond bridge over
Rfo San Félix (10.6 km beyond the turnoff to
Escopeta), 1690 m, ca. 8*30'N, 81°45'W, 15
July 1976, Croat 37168 (holotype, MO-
2395435). Figures 103-106
Planta pawiepiphytipa; internodia usque 10 cm longa,
m dia ue 20 cm lon-
gus, 1.5 cm ; lamina ovato-cordata, 26-33 cm
longa, 17.5-25 cm тй ا 2-3; pedunculus 8
cm longus, ad angulum ca. 140° infra spatham flexus; spa-
tha usque 10 cm a, in tubo viridis, in lamina extus
prera intus viridis; pistilla 4—5-locularia; loculi 4—7-
ovulat
Hemiepiphytic; appressed-climbing; internodes
smooth, sparsely cracked but only weakly or not at
all ribbed, somewhat flattened on one side (at least
at the upper nodes), to 10 cm long, 3 cm diam.,
medium green, soon turning gray, drying light
brown; cataphylls to 20 cm long, unribbed, green,
deciduous; petioles 35-37 cm long, 1.5 cm diam.,
subterete, firm, drying reddish brown, obtusely flat-
tened adaxially, surface drying finely and irregu-
larly striate; sheathing at base, 4—5 cm long;
lades ovate-cordate, subcoriaceous, semiglossy,
weakly bicolorous, acuminate at apex, prominently
cordate at base, 26-33 cm long, 17.5-25 cm wide,
1.3-1.6 times longer than broad, upper surface dry-
ing dark brown, lower surface drying dark yellow-
brown; anterior lobe 20—23.5 cm long, margins con-
vex; posterior lobes 9-12 cm long, directed
somewhat upward at an angle to the midrib, drying
directed toward the base; sinus obovate, 6.5-9 cm
deep; midrib flat, pale green above, convex, paler
than surface below; basal veins 4—6 per side, and
with the first free to base, third and fourth veins
coalesced 1.5—3.5 cm, pale green; posterior rib not
at all naked or naked up to 2 cm, only weakly
curved; primary lateral veins 4—5 per side, depart-
ing midrib at a 45-55” angle, weakly curved to the
margins, pale green, weakly sunken above, weakly
raised below, drying paler than surface, flattened,
with acute margins below; interprimary veins only
occasionally present; minor veins easily visible on
oth surfaces, arising from both the midrib and pri-
mary lateral veins, drying weak and paler than sur-
face and usually alternating with blackened secre-
ory ducts, surface minutely granular upon
magnification. “INFLORESCENCES 2-3 per axil;
peduncle to 8 cm long, drying dark reddish brown,
minutely striate, bent at ca. 140° angle beneath the
spathe; spathe to 10 cm long, to 5 cm wide when
flattened, drying dark reddish brown throughout
within, spathe blade pale green outside, drying red-
dish brown with prominent resin canals extending
from the blade well into the tube within; spathe
tube green, finely striate outside; spadix bluntly
pointed at apex, 9 cm long; pistillate portion 2.5—
2.7 cm long in front, 1–1.1 ст long in back, 10
mm diam. at apex, 9 mm diam. at middle; staminate
portion 8.2 cm long; fertile staminate portion 1 cm
diam. at base, 1.2 cm diam. at middle, 9 mm diam.
ca. 1 cm from apex, sterile staminate portion 9 mm
diam.; pistils 2.5 mm long, 1.4 mm diam.; ovary 4—
5-locular, with axile placentation; ovules 4—7 per
[mad
Annals of the
Missouri Botanical Garden
locule, 2-seriate, 0.2 mm long, contained within
trans matrix; funicle 0.1-0.2 mm long, ad-
nate to lower part of partition, style similar to style
type B; style apex flat; stigma discoid, covering
most of style apex except for center, drying irreg-
ularly 5-lobed, 0.9-1.3 mm diam.; the androecium
truncate, weakly oblong, probably prismatic, mar-
gins irregularly 4—5-sided, mostly 4-sided, 1.3-1.6
mm diam. at apex; sterile staminate flowers irreg-
ularly 4—6-sided, prismatic, 1.4 mm wide. IN-
FRUCTESCENCE with seeds 5—7 per locule, 0.4—
0.5 mm long.
Flowering in Philodendron coloradense is docu-
mented by a single collection made in July, but the
plant has three inflorescences, probably all of
which opened after the onset of the rainy season in
May. The region where the collection was made,
though at a high elevation, is on the western slope
of the Continental Divide, usually much affected by
the dry season.
Philodendron coloradense is endemic to Panama,
known only from near the Continental Divide at Cerro
Colorado in Chiriquí Province at 1600 m elevation.
Philodendron coloradense is a member of P. sect.
Calostigma subsect. Macrobelium ser. Macrobel-
ium. It is characterized by having grayish inter-
nodes longer than broad; green, unribbed, decidu-
ous cataphylls; obtusely flattened petioles;
ovate-cordate reddish-brown-drying blades with a
spathulate sinus and a weakly developed posterior
rib that is barely or not at all naked; and paired,
green inflorescences bent somewhat at the apex of
the peduncle.
The species is perhaps closest to P. grayumii,
which differs in having 7-8 (vs. 4-5) locules per
ovary and 3-4 (vs. 5-7) ovules per locule. In ad-
dition, the lower leaf surface of P. grayumii dries
glossy and is epunctate with dense secretory ducts
at higher magnifications. In contrast, the blades of
P. coloradense dry more or less matte on the lower
surface and are minutely speckled with only sparse
secretory ducts at higher magnification.
Additional specimen examined. PANAMA. Chiriqui:
Cerro Colorado, along mining road 20.5 mi. N of bridge
over Río San Félix, 8.3 mi. beyond Chame and turnoff to
Escopeta, 1630 m, Croat 75039 (MEXU, MO. PMA. US).
Philodendron copense Croat, sp. nov. TYPE: Pan-
ama. Coclé: Alto Calvario, 6 mi. N of El Copé,
Atlantic slope, along Continental Divide, along
trail which heads off old lumber road and leads
down to Las Ricas, Limón, and San Juan, 710—
800 m, 8739'N, 80°36'%, 22 June 1988, Croat
68765 (holotype, MO-3584056-8; isotypes,
AAU, B, CAS, CM, COL, CR, K, PMA, US).
Figures 109-111, 113-115.
Planta hemiepiphytica; internodia 1.5-2.5 ст longa; 2-
7 ст фат.; cataphylla 7-25 cm longa, plerumque 1-cos-
tata, in sicco flavibrunnea vel atribrunnea, persistentia
semi-intacta; petiolus acute D-formatus, 53-109 ст lon-
gus, 8-20 mm diam., epidermide interdum dense
brevi-lineata, in sicco atriflavibrunneus; — ovato-
56-84.5 ст longa, 22—47 ст lata,
nnea; inflorescentia usque 6; "MEO
7-7.5 em longus, 7-15 mm diam.; spatha 13-18.5 с
longa, lamina spathae snnt — Јави ёзге ext
uso purpureoviolaceo, intus
marronino; pistilla S ame rari Tali 20-22-ovulati:
baccae albae.
Hemiepiphytic; stem appressed-climbing, scan-
dent; leaf scars inconspicuous; internodes short,
thick, 1.5-2.5 cm long, 2-7 cm diam., broader than
long, drying yellow-brown, epidermis moderately
glossy, finely and acutely ridged on younger stems,
lighter and more matte on older stems; roots drying
brown, 15-75 cm long, 1-5 mm diam., numerous
per node, epidermis peeling; cataphylls drying
chartaceous, 7-25 cm long, usually 1-ribbed, pale
red to reddish brown, drying yellowish brown to
dark brown, persisting semi-intact as fibers.
broad medial rib adaxially, surface sometimes
densely short-lineate, drying smooth and matte to
semiglossy, light reddish brown or dark yellow-
brown, the epidermis smooth and often flaking;
blades ovate-triangular, moderately bicolorous to
concolorous, abruptly acuminate at apex (the acu-
men (1)1.7-2.5 ст long), cordate at base, 56-84.5
cm long, 22-47 cm wide (1.8-2.7 times longer than
wide), (0.77—1.1 times longer than petiole), broad-
est below point of petiole attachment, margins
broadly undulate, occasionally downturned, upper
surface dark green, glossy to rigo drying
reddish brown, gray-green or olive-green, matte to
weakly semiglossy, lower surface drying ne sei
yellow-brown to reddish yellow-green; anterior lobe
32—64 cm long, 20-46 cm wide (2.6-3.3 times lon-
ger than posterior lobes), broadest at or near base,
margins concave; posterior lobes 14. 5-21.5 cm
long, 22-47 cm wide, broadest at or near base, di-
rected downward and inward to downward and out-
ward, obtuse to semirounded; sinus spathulate to
widely hippocrepiform; midrib weakly raised to flat
above, thicker than broad, paler than surface, dry-
ing concolorous or darker than surface below;
veins 6–10(15) per side, with 1 free to base, num-
bers 4—7 coalesced 6-9 cm, prominently raised be-
low; posterior rib naked for 3.5—4 cm; primary lat-
eral veins 11-16 per side, departing midrib at 2
90° angle, straight to weakly arcuate, drying
reddish brown, sunken and paler than surface
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
above, raised, drying slightly darker than surface
below; interprimary veins as conspicuous as pri-
mary lateral veins; minor veins arising from both
the midrib and primary lateral veins; lesser veins
drying prominulous; “cross-veins” moderately con-
spicuous on both surfaces. INFLORESCENCES
erect, to 6 per axil; peduncle 7–7.5 cm long, 7-15
, green, coarsely lineate; spathe сопа-
ceous, 13-185 cm long (1.8-2.6 times longer than
peduncle), + constricted midway on the tube, acute
at apex; spathe blade green, short pale-lineate to-
ward base, striate near apex outside, 10.5 cm long
(opening broadly oblong-ovate in face view); spathe
tube oblong-ellipsoid, green, tinged purple-violet,
paler along margin outside, 8.5-9 cm long, 3.5 cm
diam., maroon, conspicuously short-lineate inside;
spadix sessile; oblong, 12.5 cm long, constricted
at base of fertile staminate portion and tapering at
both ends; pistillate portion white, cylindrical to
ellipsoid, 6.5—7.5 cm long, 2.2-2.5 cm diam.
throughout; staminate portion creamy white, 9 cm
long, 9 mm diam. throughout, broadest in lower
one-third; fertile portion 9-11 mm diam.; sterile
portion narrower than the pistillate portion, 9-15
mm diam.; pistils 1.9-2.3 mm long, 1.9-2.3 mm
diam.; ovary 3—5-locular, drying irregularly angled,
walls embedded with granular, crystal-like parti-
cles, with axile placentation; ovules 20-22 per loc-
ule, style similar to style type B; style apex broadly
and shallowly concave; stigma button-like, 0.5-0.7
mm long; sterile staminate flowers 2-2.4 mm long,
1-1.5 mm wide. INFRUCTESCENCE with berries
white; seeds 0.8-0.9 mm long, 0.3-0.4 mm diam.
Specimens of Philodendron copense hint at bi-
modal flowering. A single flowering collection in
post-anthesis condition was made in June, imma-
ture fruits were collected in January, and mature
fruits were collected in December. The mature
fruits in December could be the result of a flow-
ering in the early rainy season, but immature fruits
in January probably were the result of flowering in
the late rainy season or early dry season. More flow-
ering specimens are needed for more conclusive
statements on phenology.
Philodendron copense is endemic to Panama (but
should be expected in eastern Costa Rica) and is
known from Bocas del Toro and Coclé (El Copé) at
390 to 930 m elevation in Premontane rain forest.
Philodendron copense is a member of P. sect.
Philodendron subsect. Plat ium. This species is
characterized by its appressed climbing habit;
short, thick internodes (2-7 cm diam.); usually one-
ribbed cataphylls which persist semi-intact; D-sha-
ped petioles with flaky, yellow to yellow-brown epi-
dermis upon drying; large, ovate-triangular blades
drying reddish brown with many conspicuous
“cross-veins”; and up to six inflorescences per axil
with the spathes green or reddish tinged outside
and maroon within on the t
Philodendron copense is most easily confused
with P. brunneicaule, with which it occurs. Though
the latter species has blades that dry a similar col-
or, it is distinguished by having long internodes;
brown, rem epidermis; ovate blades that lack
the minute “cross-veins” upo ing; and stouter
банди that are solitary in each axil.
Philodendron copense is also similar to P. chiri-
quense, another species with an ovate-triangular
leaf that dries reddish brown. Philodendron chiri-
quense differs by having much shorter, thicker in-
ternodes, conspicuous reddish brown persistent fi-
bers (rather than semi-intact), and dark brown to
blackened striate petioles, which lack an exfoliat-
ing smooth epidermis.
Additional specimens examined. PANAMA. Bocas
del Toro: Fortuna Dam area, Dc inq Grande,
4.3 km N of the Continental Divide, 590 m, 8°46'N,
Coclé: near Continental Divide, along lum
i С т, Croat 44563 (MO, RSA, U);
44593 (COL, MO); 6 2 mi. above El Соре, 930 т, Croat
49156 (L, MO, SAR, US); 5.6 mi. N of El Copé, 800 m,
8°39'N, 80%36'W, Croat 75064 (M, MEXU, МО, aM
SCZ); 75091 V eris ca. 5.6 > N of El Соре,
75091 (MO); 9.4 km س 1 Copé, 750-900 m, 44729
(В, СЕ, К, МО, РМА, ТЕХ).
Philodendron correae Croat, sp. nov. TYPE:
Panama. Chiriquí: Gualaca—Chiriquí Grande,
vic. Lago Fortuna, along trail to meteorological
station on Rio Hornito departing from N side
of highway, ca. 0.5 km S of Centro de Cientif-
cos, 8745'М, 82°18’ W, 24 July 1994, Croat &
Zhu 76395 (holotype, MO-4619416; isotypes,
B, CAS, COL, CR, F, K, NY, PMA, US, VEN).
Figures 116-119.
СЕЎ и aut raro terrestris; ee ob-
ata uno latere, usque 7 cm m 1.5°m
dam pora (8)13-16 cm longa, шы decidua;
iolus D-formatus, 7—16.5 cm longus, plus minusve
ni пене lamina plus minusve oblonga, rotunda vel len
iter subcordata Pm mah m longa, 5-12 cm lata; ner-
vis lateralibus I 4
inflorescentia 1; йкы 10.5-12 cm lon
18.4 cm longa, extus flavialba, intus leviter palidiori;
interdum viridis basi, tubo intus suffuso obscure rubro;
pistilla 5-6-locularia; loculi 1-ovulati.
Hemiepiphytic or rarely terrestrial; stem ap-
pressed-climbing or spreading; sap viscous, un-
scented; internodes obtusely flattened on one side,
422
Annals of the
Missouri Botanical Garden
moderately glossy, to 7 cm long, to 1.5 cm diam.,
usually much longer than broad, medium green to
tannish, drying gray, epidermis becoming grayish
brown and peeling; roots mostly 15-30 cm long,
drying 2-3 mm diam.; cataphylls (8)13-16 cm long,
unribbed, green, deciduous, turning mushy; peti-
oles 7—16.5 cm long, (3)6-15 cm diam., D-shaped,
obtusely flattened (drying usually sulcate) adaxially,
+ spongy, narrowly rounded abaxially, with adaxial
margins raised, surface with a conspicuous purple-
black ring around apex; sheathing usually to near
the apex, with margins erect and incurled, the tip
free-ending; blades + oblong, subcoriaceous, se-
miglossy, conspicuously to moderately bicolorous,
abruptly long-acuminate or sometimes acute at
apex (the acumen inrolled if present), rounded to
weakly subcordate at base, 21-46 cm long, 5-12
cm wide (3-5 times wider than long), (ca. 2-4 times
longer than petiole); upper surface semiglossy, dark
green to medium green, drying dark brown to
blackish brown, lower surface glossy and slightly
to moderately paler; sinus 5-12 mm deep; midrib
flat to broadly convex, paler than surface above,
convex or thicker than broad and bluntly acute,
darker than surface below; basal veins lacking; pri-
mary lateral veins weak, 4–6 per side when present
but sometimes not apparent, departing midrib at a
60—70° angle, arcuate to the margins, slightly sunk-
en, weakly visible above, convex, slightly darker
than surface, weakly visible below; minor veins
fine, moderately visible to distinct, arising from
both the midrib and primary lateral veins. INFLO-
RESCENCES 1 per axil; peduncle 10.5-12 cm
long, 5-6(14) mm diam., subterete; spathe coria-
ceous, 15-18.4 cm long (1.43-1.75 times longer
than peduncle), weakly or not at all constricted,
spathe blade yellow-white (B & К yellow-red
9/7.5-10 (at anthesis)) throughout, slightly paler in-
side; spathe tube sometimes green outside, 6.5—7.5
cm long, 1.6-2 cm diam., slightly paler and tinged
dull red inside; spadix 8-19 cm long; stipitate 7—
10 mm long; pistillate portion green to pale yellow,
cylindrical, 7.9 cm long, 1.3-1.4 cm diam. through-
out; staminate portion 4—8.4 cm long; fertile sta-
minate portion tapered; pistils glossy, 3.3-3.8 mm
long, 2.3-2.6 mm diam.; ovary 5-6-locular, with
sub-basal placentation; ovules 1 per locule, style
similar to style type C; style funnel shallow, some-
times deep subcylindric; style apex rounded, some-
what flat, with simple funnel; stigma + discoid,
covering interior faces of stylar funnel. INFRUC-
TESCENCE (post-anthesis) with seeds 1 per locule,
1.5-1.7 mm long, 7-8 mm diam
Flowering in Philodendron correae has been re-
corded only during the early rainy season, June
through September, but relatively few fertile collec-
tons have been seen overall.
Philodendron correae is endemic to western Pan-
ama in Chiriquí and Bocas del Toro Provinces at
180 to 1400 m in Premontane rain forest and Trop-
ical Lower Montane rain forest.
Philodendron correae is a member of P. sect. Ca-
lostigma subsect. Glossophyllum ser. Glossophyl-
lum. This species is distinguished by its internodes
generally longer than broad; more or less oblong,
dark brown- to black-drying blades with rounded
to weakly cordulate bases; primary lateral veins
only weakly visible; and solitary inflorescences with
the spathe blade yellowish white on both inner and
outer surfaces.
Philodendron correae is closest to P. ligulatum,
which differs in having rather prominent primary
lateral veins and by occurring at generally lower
elevations (sea level to 900 m). Folsom et al. 5471
is noteworthy in having blades that dry greener
than is usual for the species. It also has a Type B
style type; the only other specimen studied for
ovules (Croat 66748) had a Type C style. The dif-
ference may have been due to the age of the flow-
ers, but this situation warrants further investigation.
Philodendron correae is named in honor of Mir-
eya D. Correa A., among the first scientists to col-
lect in the Fortuna area, and the first to collect this
species.
Additional specimens examined. PANAMA. Bocas
del Toro: Fortuna Dam area, Chiriquí Grande—Fortuna,
1.2 mi. N of Continental Divide, 910 m, 'N, 82717,
Croat 60436 (CAS, L, MO); 6.6 mi. N of middle of bridge
over Fortuna Lake, 780 m, 8°45’N, 82°18'W, Croat 66779
(MO); along gravel road near Continental Divide, 1170 m,
8°44’N, 82°17'W, Croat 66653 (MO, PMA, US). Chiriquí:
Cerro Colorado, above San Félix, along mining road 18-
27 mi. N of Pan-American Highway, 1200-1500 m, Croat
33151 (MO); 13-14.6 km N of Chame, 1390-1410 m,
Croat 37217 (MO); Fortuna Dam area, N of Gualaca, 11.8
mi. N of Los Planes de Hornito, 1400 m, Croat 48676
MO, US); Gualaca—Chiriquí Grande, at junction of road
to IRHE headquarters, 1200 m, 8°45'N, 82°18'W, Croat
~
(PMA); ca. 10.7 mi. from Planes de Hornito, 4000 ft.,
Antonio 5148 (MEXU, MO); along aquaduct to IRHE pie’
r source near dam, 1200-1300 m, 8^45'N, 82 p
Croat 66626 (MO); 1400-1500 m, Folsom et al. 547
(MO).
=
I]
Philodendron cotobrusense Croat & Grayum,
sp. nov. TYPE: Costa Rica. Puntarenas: along
road about halfway between Flor del Roble
and Las Alturas de Coto Brus, 1250 m.
8°54'N, 82°51'W, 13 July 1985, Grayum &
Hammel 5689 (holotype, MO-3446320; 150-
types, CR, K, US). Figure 120.
Volume 84, Number 3
1997
Croat 423
Philodendron Subgenus Philodendron
Planta амер рун; internodia 1 1-3 ст longa, 1.5—2
dia pider
idua; petiolus
32.5-35 cm longus; lamina profunde 3- lobata, 37-38 ст
longa, 46-52 ст lata, venis confertis; nervis lateralibus I
18-19 utroque; inflorescentia usque 5; pedunculus 6–9
cm longus; Hun 6-14 ст longe, lamina spathae extus
cremeoflava, intus eburnea; tu pathae extus viridi, in-
tus rubro- EIR pistilla 6-7- Ua: loculi 3-ovulati.
Hemiepiphyte; stem appressed-climbing; inter-
nodes short, thick, 1-3 cm long, 1.5-2 ст diam.,
mostly longer than ~ epidermis silvery gray;
cataphylls to 14 cm long, sharply 2-ribbed, yel-
low-green, tinged pinkish, deciduous; petioles
32.5-35 cm long, 4-9 mm diam., subterete, firm,
obtusely flattened adaxially, surface matte; blades
deeply 3-lobed, margins incised to within 1.5-2 cm
of base, subcoriaceous, weakly bicolorous, long-
acuminate at apex (the acumen probably down-
turned), + cordate at base, 37-39 cm long, 46-52
cm wide (0.7-0.8 times longer than wide), (1-1.2
times longer than petiole), usually about equal to
petiole, upper surface dark green, semiglossy, lower
surface weakly glossy, somewhat paler; median
obes 30.5-32 cm long, 13-17 cm wide; lateral
lobes 24-27 cm long, 9-12.6 cm wide (2.14-2.88
times longer than wide), directed + upward, broad-
ly confluent to 2.5 cm with medial lobes; midrib
broadly convex, closely veined above, convex be-
low; basal veins lacking; primary lateral veins 18-
19 per side, departing midrib at a 50-65” angle,
broadly curved to the margins, sunken above, con-
vex below; interprimary veins almost as conspicu-
ous as primary lateral veins; tertiary veins visible,
slightly darker than surface; minor veins arising
from both the midrib and primary lateral veins but
mostly from midrib in upper half of blade. INFLO-
ESCENCES to 5 per axil; peduncle 6–9 cm long,
3-4 mm diam., terete; spathe 6-14 cm long (1.8—
3.5 times longer than peduncle); spathe blade
creamy yellow outside, creamy white inside; spathe
tube green outside, red-violet to purplish inside;
spadix 9.5-12 cm long; pistillate portion yellow-
green, 7-10 mm diam.; staminate portion 5-8 cm
long; fertile staminate portion white, 8-12 mm
iam.; ovary 6-7-locular, with sub-basal placenta-
tion; ovule sac 1 mm long; ovules 3 per locule,
digitately arranged in series of 2-3, contained with-
in transparent ovule sac, 0.2-0.3 mm long, slightly
shorter than funicle; funicle ca. 0.3 mm long (can
be pulled free to base), style similar to style type
B; style apex flat; stigma subdiscoid, 1 mm diam
0.2-0.3 mm high, covering entire style арех; the
ci truncate, + oblong, prismatic, irregu-
larly 4—6-sided, 0.7-1.1 mm long; thecae oblong,
0.3 mm wide, + parallel to one another.
Flowering in Philodendron cotobrusense has been
recorded only during the early rainy season in July.
Philodendron cotobrusense is known only from
eastern Costa Rica in the Cordillera de Talamanca,
at 1000 to 1250 m elevation in a Premontane wet
der transition to rain forest life zone.
odendron cotobrusense is a member of P.
sect. PAG hyllum. This species is characterized
y its т. climbing habit; sharply two-
ribbed, deciduous cataphylls; short, thick inter-
nodes with a silvery-gray epidermis; petioles about
equaling the deeply three-lobed, closely veined
blades (with more than 16 pairs of primary lateral
veins); and especially by the inflorescences in clus-
ters of up to five per leaf axil with the spathe tube
green outside, red-violet within, and the blade
creamy yellow on the outside and creamy white
within.
Philodendron cotobrusense is most similar to P.
madronense Croat. That species differs in having
the medial segment even more closely veined (to
18 or more) and not at all confluent with the lateral
lobes. In addition, it has solitary inflorescences.
Philodendron cotobrusense is also similar to P.
tripartitum, but that species differs in usually hav-
ing narrower, oblanceolate to oblanceolate-elliptic
blades with more widely spaced primary lateral
veins (usually 4—10, rarely to 12), and fewer inflo-
rescences per axil (up to three) with longer pedun-
cles. In addition, P. cotobrusense has 3 ovules per
locule whereas P. tripartitum has 1.
Davidse 24202, tentatively placed in P. triparti-
tum, seems to be somewhat intermediate with P.
cotobrusense. See the discussion of that species.
Additional specimen examined. COSTA RICA. San
José: General Valley, between Canaán and Chimirol, Río
Chirripó del Pacífico, 1000 m, 9°27'N, 8337" W, Burger
& Liesner 7119 (F)
Philodendron cotonense Croat & Grayum, sp.
nov. TYPE: Panama. Chiriquí: vic. of El Hato
del Volcán, at ~ Lagunas, 3 mi. from Nueva
California, 1.7 mi. past new airstrip, 1380 m,
8°46’N, 8 82°40, 17 Sep. 1987, Croat 67708
(holotype, MO-3584981; isotypes, AAU, B,
CAS, CM, COL, CR, DUKE, F, K, L, G, NY,
OOM, PMA, QCA, RSA, SAR, TEX, U, US,
VEN). Figures 108, 121-123.
Planta plerumque hemiepiphytica; internodia 1-3 cm
longa, 2.5-3 cm diam.; cataphylla 18-31 cm longa, in-
costata vel acute 1-costata, decidua intacta; petiolus sub-
teretes, 17.5-53.5 ст longus, 5-8 mm
guste ovato-triangularis, a on ст longa, 11.5-20.5 ст
lata; inflorescentia 1-2; cu 6.
longus, spatha 11-17 cm inen liis spathae extus di-
Annals of the
Missouri Botanical Garden
lute viridi vel cremeoflava vel alba, intus pallide viridi vel
rronino v u
atripurpureo; pistilla 56 locularia; loculi 4—5-ovulatii;
baccae vivide aurantiacae
Hemiepiphytic vine or sometimes terrestrial;
stem appressed-climbing but with ends often di-
vergent, fertile to at least 5 m; leaf scars conspic-
uous, 1-1.3 cm long, 1.2-1.3 cm wide; internodes
terete, semiglossy, sometimes matte, drying irregu-
larly ridged, 1-3 cm long, 2.5-3 cm diam., usually
broader than long, green, soon becoming gray to
silvery-gray, drying yellowish brown, epidermis
sometimes flaking free; roots short, stubby, light-
colored at tips; cataphylls 18-31 cm long, unribbed
to sharply 1-ribbed, yellow-green to green, prompt-
ly deciduous intact; petioles 17.5-53.5 cm long
(averaging 34.4 cm long), 5-8 mm diam., subterete,
weakly spongy, dark green, obtusely flattened, es-
pecially toward the apex adaxially, with abaxial
margins broadly rounded, surface pale greenish li-
neate, semiglossy; blades narrowly ovate-triangu-
lar, subcoriaceous, usually gradually acuminate at
apex (the acumen 1-3.5 cm long), prominently
lobed at base, 23-45 cm
(0.7—1.4 times as long as petiole), broadest near
point of petiole attachment, moderately bicolorous
to almost concolorous, upper surface drying dar
brown to dark olive-green, semiglossy, lower sur-
face weakly glossy, usually yellow-brown, some-
times yellow-green; anterior lobe 16-31 cm long,
margins weakly convex to flat or concave; posterior
lobes 6–16.5 cm long, 3-8 cm wide, broadest at or
near the middle, sometimes narrower at point of
petiole attachment; sinus parabolic to obovate or
almost oblong, 5—7.5 cm deep; midrib flat to weakly
sunken, concolorous to paler than surface above,
convex, paler than surface, often tinged purple
(with coloration continuing onto petiole apex) be-
low; basal veins 3—4 per side, the first often free to
base (sometimes all free to base), the second and
third coalesced 1–2.5(4.5) cm; posterior rib usually
not naked except sometimes for up to 5 mm at the
base; primary lateral veins (3)4-5(6) per side, de-
parting midrib at a 30-65” angle (70-90” angle
near base), weakly curved to the margins, some-
times drying undulate, often prominently down-
turned at midrib, sometimes tinged purplish,
scarcely to weakly sunken above, narrowly raised
below; minor veins obscure to clearly visible, aris-
ing from both the midrib and primary lateral veins,
close and almost indistinguishable from the secre-
tory canals, secretory canals appearing as intermit-
tent lines, usually more prominently downturned at
the midrib and appearing to cross over the minor
veins, drying blackened. INFLORESCENCES 1-2
(usually solitary, sometimes 2) per axil; peduncle
5)10-17.5 cm long, subterete, pale-striate;
spathe 11-17 cm long (0.94—1.6 times longer than
peduncle), somewhat constricted above the tube;
spathe blade light green to cream-yellow to white
outside, pale green to cream-white to white inside;
spathe tube dark to medium green, sometimes pale
striate-lineate outside, 3-5.5 cm long, 1.3-2 cm
diam., maroon to crimson pale red-violet or dark
purple inside; spadix sessile, (6.7)10.5-16.5 cm
long; pistillate portion green, cylindrical to ovoid,
2 cm long, 4-7 mm diam.; staminate portion 4.2-
7.1 cm long; fertile staminate portion to 5 mm diam.
throughout; sterile staminate portion whitish to
cream-white, to 5 mm diam.; pistils 0.9 mm long;
ovary 5-6-locular, 0.6 mm diam., with sub-basal
placentation; ovules 4 or 5 per locule, arranged dig-
itately, «0.1 mm long, longer than funicle; funicle
«0.1 mm long, style 0.9 mm long, 0.5 mm diam.,
similar to style type C; style apex flat, drying with
5—6 depressions alternating with ribs from a raised
center; stigma hemispherical, inserted on stylar
funnel; the androecium irregularly 3—6-sided; the-
cae ovate to cylindrical; sterile staminate flowers
0.8 mm long. INFRUCTESCENCE with berries
bright orange; seeds drying pale yellow-brown, nar-
rowly ellipsoid, 1.3-1.5 mm long, 0
diam., with weak striations, appendages absent. JU-
VENILE plants with petioles 4.7—7.7 cm long;
blades lanceolate (acumen 6.5—10.5 cm long), 1.7-
2.5 cm wide.
Flowering in Philodendron cotonense has been
recorded in both Costa Rica and Panama during
June, and post-anthesis or early fruiting collections
have been made from July through December (ex-
cept October). Mature fruits have been collected
only in March. However, several post-anthesis col-
lections have been made during March, indicating
perhaps a bimodal flowering period. Too few data
are available to be certain.
Philodendron cotonense is known only from east-
ern Costa Rica and western Panama in Premontane
wet forest and Premontane rain forest, Tropical Low-
er Montane wet forest, and Tropical Lower Montane
rain forest life zones, as well as the transition zone
between these zones, at 1100 to 1950 m Sie
Philodendron cotonense is a member of P. sec à;
Calostigma subsect. Glossophyllum ser. Ovata. This
species is characterized by having internodes usu-
ally longer than broad (except distally), subterete
petioles, and more or less triangular blades aver
aging over twice as long as broad, with posterior
Volume 84, Number 3
1997
Croat 425
Philodendron Subgenus Philodendron
lobes typically much longer than broad and often
directed somewhat outward. Also characteristic are
the conspicuous secretory canals visible on the
dried blade surface and the solitary long-pedun-
culate inflorescences in each leaf axil.
Philodendron cotonense is most easily confused
with P. wilburii var. longipedunculatum, which dif-
fers in having longer and more slender internodes
(3-8 ст long and 8-15(20) mm diam.); sharply
2-ribbed cataphylls; and more typically hastate
leaves ranging from 1.03 to 1.4 (averaging 1.29)
times longer than wide, with proportionately nar-
rower lobes directed outward at a broader angle
(averaging 91? vs. 121? for P. cotonense). Philoden-
dron wilburii var. longipedunculatum also differs in
having mostly two inflorescences per axil with pro-
portionately much longer peduncles, averaging 1.35
times longer than the spathe (vs. rarely longer than
the spathe in P. cotonense).
hilodendron cotonense might be confused with
smaller plants of P. alticola, which also have se-
cretory canals visible on the blades. The latter spe-
cies differs in typically having much larger blades
(minimum 44 cm X 21 cm) that dry greener and
thicker; persistent cataphyll fibers; and a short-pe-
dunculate inflorescence with the spathe usually not
at all constricted.
Noteworthy is Davidse et al. 28367, which has a
narrower sinus and the basal veins all free to the
base with the uppermost primary lateral veins more
or less obscure. Otherwise, it fits well into P. co-
tonense.
Additional specimens examined. COSTA RICA. Pun-
tarenas: vic. of San Vito de Java, ca. 1 km S of San Vito,
on road to Villa Neily, 1100 m, Croat 66169 (CR, K, MO,
NY, US); 4000 ft., Croat 32905 (MO); Zona Protectora Las
Tablas, San Vito Coto Brus-Sabalito, Finca Neblinas,
m, 8°54’N, 82°51'W, Grayum & Hammel 5690
(MO); Cerro Pando, ridges а Ко Cotón and Río oe
; Laguna fenetre San Vito, Coto Brus, 1200
m, Gómez-Laurito 10774 (F); Las Alturas, n Río Co-
tón, 1340 т, 8%56'30"N, 82°50'W, Davidse 24093 (CR,
MEXU, M MO); Cordillera de Talamanca, area pop Río
Canasta, 9.5 airline km NW of Agua Caliente, between
Cerro ся and Cerro Pittier, 1500-1600 m, 9°02'N,
8259'W, Davidse et al. siae (Мој ea Cotón-Mellizas,
1300-1450 m, ca. 8°54'N, 8 avidse et al. 25562
(CR, K, MO); N of Santa osi on ки Cotón, 5 of Agua
Caliente, 1100 m, 8"57"М, 82°56’ W, Davidse et al. 28236
(COL, CR, MO); N of Pies 1400-1500 m, 8%57'N,
82°50'W, Davidse 24166 (CR, MO, Use. канй ‘Tres Col-
inas, 1800-1850 m, 9°07'N, 83°04'W, Davidse et al.
25606 (CR, MO); Le ruces dod Garden, Coto
Brus, ca. 4 km SE of San Vito, 1150-1200 m, 847'30'N,
82°58'W, Grayum et al. 8113 (CM, CR, L, MO, QCA); Río
Coto Brus, near Cotón, 23 km N of La Unión, Croat 26692
(MO); Río yid vic. of first large concrete culvert before
Finca Las s at Cotón, ca ran , Croat 44349
(MO); Río Pieds Blancas, Cern Мавы Fila Cos
tefia, Fila Cruces, 950-1150 m "8-49 18"N, 83°11’ 15"W,
Grayum 10646 (CR, INB, MO, VEN). PANAMA. Chiri-
quí: Volcán-Río Sereno, 7 mi. N of Volcán, 8°50'N,
8238 'W, nd 66226 (CM, MO, PMA, US); 13.7 mi. W
of Volcán, 1200 m, 8°51'N, 82°43'W, Croat 66343 (МО);
S of Boquete, Cerro Pate оа upper NE slopes and
ummit, 1900-2000 m, 8°46'N, 82%25'W, Croat 66504
(CAS, L, MEXU, MO, PMA).
Philodendron crassispathum Croat & Grayum,
sp. nov. TYPE: Panama. Chiriquf: Cerro Col-
orado, above San Félix along mining road, 18-
27 mi. off Pan-American Hwy., above Chame
(turnoff to Escopeta), 1200-1500 m, 8°35’N,
81%50'W, 12 Mar. 1976, Croat 33150 (holo-
type, МО–2395067; isotypes, В, F, K, PMA,
US). Figures 112, 124, 127, 128, 149.
Planta hemiepiphytica, interdum terrestris; internodia
1.5-8 ст longa, (1-1.8)2.5-3.5 cm diam.; cataphylla car-
nosa, 8.5-24 cm longa, persistentia semi-intacta, denique
decidua; petiolus subteres, obtuse complanatus, 20—
ina ovato-cordata,
violaceopurpureo vel rubro; pistilla
7-10-ovulati; bacca virides, aurantiacentes vel albidae
Hemiepiphytic, sometimes terrestrial; stem ap-
pressed-climbing or scandent, creeping, becoming
matte and gray, smooth when fresh, sap watery,
spicy-scented, leaf scars conspicuous, 9-20 mm
long, 2-3.5 ст wide; internodes drying conspicu-
ously wrinkled, epidermis weakly glossy, 1.5-8 cm
long, (1-1.8)2.5-3.5 cm diam., usually broader
than long, dark green, epidermis brownish and
cracking; roots dark brown, smooth upon drying,
elongate, 3-50 cm long, 1-3 mm diam., few per
= cataphylls fleshy, 8.5-24 cm long, sharply
-ribbed, green to dark green, dark green short-
ism drying yellowish brown to pale green, per-
sisting semi-intact at upper nodes then eventually
fibrous then deciduous. LEAVES erect-spreading to
spreading; petioles 20—31(48) cm long, 5-19 mm
diam., subterete, somewhat spongy, sometimes
tinged brown near apex, obtusely flattened with ob-
tuse medial rib adaxially, convexly rounded abax-
ially; blades broadly ovate-cordate, very coria-
ceous, acuminate at apex (the acumen inrolled, 2—
5 mm long), cordate at base, 14—29 cm long, 11—
24 cm wide (1—1.56 times longer than wide), broad-
est at or near the middle, margins hyaline, whitish
or reddish, tightly curled under when dried, upper
surface dark green, glossy, drying semiglossy to
Annals of the
Missouri Botanical Garden
matte, lower surface paler, drying greenish gray to
yellowish green to brownish, drying semiglossy to
matte; anterior lobe 10–25 cm long, 11–33 cm wide
(1.6-2.5 times longer than than posterior lobes);
posterior lobes 4.5-10 cm long, 5-16.5 cm wide,
obtuse to rounded; sinus spathulate to hippocrepi-
form; midrib flat to very broadly convex, drying yel-
lowish green, paler than surface above, weakly con-
vexly raised, paler than surface below; basal veins
3-4(6) per side, with 0-1 free to base, most of the
remainder coalesced 1-2.5(4.4) cm, flat above,
weakly raised below; posterior rib not naked if
present; primary lateral veins 4-6 per side, de-
parting midrib at a 40—70° angle, + straight to the
margins, flat, paler than surface, drying obscure
above, raised and paler below; interprimary veins
obscure above, weakly sen pt minor veins
etched above, visible and r than surface be-
low, arising from midrib ni INFLORESCENCES
shorter than leaves, 1 per axil; peduncle 3.5-7 с
long, 1-2.5 cm diam. (dried), terete, drying TER
spathe fleshy, (6.8)10-14 cm long, 4.5-10 cm
diam., not at all constricted, = ellipsoid, bluntly
acute to rounded at apex, the walls to 1 cm or more
thick midway; spathe blade red outside, red to
white inside; spathe tube green, sometimes yellow-
ish to orange-red outside, violet-purple to red in-
side; spadix (6.5)9-14 cm long; pistillate portion
grayish to golden-yellow, cylindrical to slightly el-
lipsoid, 1.5-7 cm long, 10-25 mm diam. through-
out, with 13—15 flowers visible per spiral; staminate
portion 5—9 cm long; fertile staminate portion white
with orange-brown droplets, cylindrical, weakly
constricted above sterile portion, tapered toward
apex, 15-23 mm diam. throughout, broader than
pistillate and sterile portions, 22-27 flowers per
spiral; sterile staminate portion 17-24 mm diam.;
pistils 4—4.5 mm long, 4—4.5 mm diam.; ovary 6—
7-locular, with axile placentation; ovules 7-10 per
locule, 0.6-1 mm long, 0.3 mm diam.; funicles with
free portion ca. 1 mm long, the remainder loosely
fused to the wall of the locule; style similar to style
type D, button-like and concave on drying, 1.3-2
mm diam., the margins pale; androecium margins
— 0.6-1.3 mm long; sterile staminate flow-
= globose, 1.4-2.2 mm long. INFRUCTES-
CENCE erect; pistillate spadix 4.5-5 cm long, 4 cm
diam.; berries green becoming orange to whitish;
pericarp white; mesocarp yellowish white; seeds yel-
low-orange, 2.9-3.5 mm long, 0.7-1 mm diam
Flowering in Philodendron ован ар-
pears to occur during the dry season and early rainy
season based on flowering collections made during
February through April and in August. Post-anthe-
sis collections have been made from January
through October. Mature fruiting collections have
been made only from January through March, in-
dicating that they may take up to almost one year
to develop.
Philodendron crassispathum ranges from central
Costa Rica to western Panama, at 1100 to 2600 m
elevation in Premontane rain forest and Tropical
dec
ribbed cataphylls, subterete, EY flattened,
somewhat spongy petioles, coriaceous ovate-cordate
blades with the minor veins etched above, and ез-
pecially by the very fleshy more or less ellipsoid,
een spathe with walls typically 1 cm or more
thick, colored white within on the blade and red in
the tu
Philodendron crassispathum is most closely re-
lated to P. brenesii, which differs in having narrowly
ovate blades with a narrow V-shaped sinus and
mostly free basal veins, and a comparatively thin
ا о constricted above the tube portion.
ast, P. crassispathum has blades with a typ-
ау Баве to e sinus and a
sually obvious posterior ri
Additional specimens examined. COSTA RICA. Ala-
juela: Río San Pedro, Cerro Azahar, 15 km NW of San
Ramón by air, 1400-1500 m, 10%9'30"М, 84%34-35'W,
Liesner et al. 15502 (CR, K, MEXU, MO, WIS); Monte-
verde Reserve, Cerro omogo, 1600-1 m, Dryer
1470 (CR, F). Cartago: Río Dos Amigos-Río Villegas.
narrow ridge W of R m, 9°42'N,
83°47'W, Grayum et al. 3762
past town of Río Macho, along road to Hu
9*1 7'N, 83%45'W, Hoover 1346 (CR, MO); Тарап Ке-
serve, Quebrada Salto-Río Grande de Orosi, ca. 1 km 5
of jct., 1500-1800 т, 9°43'N, 83747 "1, Croat & Grayum
68226 (CR, MO), Grayum et al. 6301 (MO); 1380 m,
Croat 79077 SpA INB, MO); Tapantí sae Preserve,
O mi. SW of Paraíso, Croat 47045 (CM, MO), 47043
(MO, SCZ). Heredia: Río "4: Sete Río Nuevo, E
slopes of Ser Barba, 2000 m, 10°6’N, 84°03’ W, Burger
& Baker 9500 (CR, F, ISC, MO, NY, PMA); 9 km SE of
Virgen dd Socorro, 9 km E of Isla Bonita, 1530 m.
10°14’N, 84°05’ ier Loiselle 228 (MO); Cerro Chompipe. N
of San Rafael, 2000 m, Lems s.n. (Е, NY, US); Cerro се
las Caricias, N га sane Iid, 2000-2400 m, Standley &
Valsti 51910 (US); San sapri Las Vueltas, N -—
rro Chompipe, 2100-2200 'N, 84 04"
Sas 13989 (MO); Volcán Barva, Geen Chompipe, near
: Río Las Vueltas, 12
=
ps
de Talis. Bratsi, A
Rio Lari and Río Dapari, Tal-
83°06’ 15"W, Herrera 5504 (INB; Mor Cordillera de
S РБ "ГП" РБ РНЕ РИИ
Volume 84, Number 3
Croat 427
Philodendron Subgenus Philodendron
manca, Cerro Biricuacua, 2600 т, 9?23'55"N,
83^10'10"W, Herrera 6219 в, INB, MO); 920'20'N,
8313'33"W, Bittner 1864 (INB, MO). tarenas: Mon-
tev , 84°48'W, Croat
61195 (MO); Cerro Negro, 1500—1600 m, Haber & Bello
2843 (MO); Brillante Trail to Veracruz, river valley along
Continental Divide, 1600 m, 10?20'N, 84°50’W, Haber et
al. 4584 (MO). San José: 5 slopes of Cerro Zurquf, са.
5 km М of San Isidro de San José, 1500-1800 m,
Utley & Utley 408 (DUKE); 4 km N of Cascajal, 7 km N
ubes, on CR-216, 1500-1600
o: C
, Kress et al. 86-1949
л SEL); 9.2 mi. W of ae 1450-1480 m, 8°35'N,
81750", Croat 69069 (L, MEXU, MO, P, US); near Con-
tinental Divide, 9.4 mi. from Chame, ca. 1700 m, ca.
oe N, 81%45'W, McPherson 8918 (CAS, MO); Fortuna
Dam area, along trail on Continental Divide, ca. 1200 m,
McPherson 9031 (MO). Bocas del Toro-Chiriquí: Chi-
riquicito—Calderas, Elfin forest, at Divide, on trail, Kirk-
bride & Duke 975 (MO); С ue Colorado, 1 1 m,
8°35'N, 8150", McPherso (МО,
Cerro Colorado, 20 mi. N ^ Eo San Félix, 1660 m,
8730'N, 81°46’ W, Croat 74987 (MO, US); 1110-1750 m,
8°35'N, 81%54'W, Hammel & Trainer 14932 (MO); Bo-
quete region, SW slope of Cerro Pate Macho, 1630-1780
m, 8°46'N, 82°25'W, Croat 66382 (MO, QCA, SCZ); For-
Lake-Chiriquí Grande, 4.5-5 km
tuna Dam area, Fortuna
. N of bridge over Вауапо
s 846'N, 82°16'W, ас 68027 (COL, С, MEXU,
Philodendron cretosum Croat & Grayum, sp.
nov. TYPE: Panama. Coclé: vicinity El Valle
de Antón, at forested flat area near Finca Ma-
MO-04619350-51; isotypes, AAU, B, CAS,
CM, COL, CR, F, K, L, M, MEXU, NY, OOM,
P, PMA, QCA, SEL, TEX, US, VEN). Figures
125, 126, 144.
, Planta epiphytica aut hemiepiphytica; internodia brev-
la, 1-3 cm diam.; succus calcareus, in sicco mox calca-
reus; cataphylla 23 42 cm longa, obtuse 1-costata, in sic-
co тк flavibrunnea, persistentia ut fibrae pallidae;
tiolus obtuse et late aut profunde et anguste alias
savas 10. 5-23 3.5 с cm i s, 3-6 mm diam.; lamina
line 29-84.5 cm longa, 5-12 ст lata;
ботан 1-3; рейси (2.5)4–8.9 ст Па са.
mm diam.; зраћа 9-13.5 cm longa; lamina spathae
ee pallide vitii ш viridialba, intus viridialba: tubo
"hae extus viridi, intus albo; pistilla 4-locularia; loculi
m 18-20-seminibus
Epiphytic or hemiepiphytic; stem appressed-
climbing, elon ngate, sap chalk-white; internodes
short, 4-9 cm lo ong on lower stems, short on flow-
ering plants, 1-3 cm diam., as broad as long or
slightly longer than broad, gray-green; roots drying
brown, few per node; cataphylls thin, mem-
branous, 23-42 cm long, bluntly 1-ribbed, green,
drying pale yellowish brown, persisting as pale fi-
bers; petioles 10.5-23.5 cm long, 3-6 mm diam.,
medium green, semiglossy, bluntly and broadly or
deeply and narrowly sulcate adaxially, with adaxial
margins obtuse or sometimes acute; blades linear
to oblanceolate, chartaceous to weakly subcoria-
ceous, weakly bicolorous, drying greenish to yel-
lowish brown on both surfaces, matte and paler be-
low, acuminate to weakly acute at apex (the acumen
"peo attenuate at base, 29-84.5 cm long, 5—
12 cm wide (5.1-8.3 times longer than wide), (2.6—
4.7 times longer than petiole), much longer than
petioles, broadest in upper one-third; midrib prom-
inently and narrowly sunken above, thicker than
broad, bluntly acute, glossy and darker than surface
below; basal veins lacking; primary lateral veins 6—
9 per side, departing midrib at a 25—45° angle,
straight to weakly arcuate to the margins, sunken
above, convex below; interprimary veins sunken
above, slightly raised below; minor veins arising
from midrib only; tertiary veins visible, darker than
surface below. INFLORESCENCES 1-3 per axil;
peduncle (2.5)4-8.9 cm long, са. 10 mm diam.,
drying 3—5 mm diam., shorter than petiole, subter-
ete or irregularly angled, white-striate; spathe sub-
coriaceous, semiglossy, 9-13.5 cm long (1.8-3.3
times longer than peduncle), moderately constrict-
ed midway or just below the middle, abruptly acu-
minate at apex; spathe blade lanceolate, pale green
to greenish white outside, 5.5-8 cm long, greenish
white inside; spathe tube ellipsoid, green, short
white lineate outside, 3.5—5.5 cm long, 1.8-2 cm
diam., white (at anthesis) inside; spadix stipitate
to 5 mm long; 11-15 cm long; pistillate portion
white to pale greenish, weakly ellipsoid, 3—3.5 cm
long; staminate portion 4.5—7.8 cm long; fertile sta-
minate portion white; sterile staminate portion 1.2
cm diam.; pistils 0.6-0.9 mm long, 3 mm diam.,
drying blackish; ovary 4-locular, with axile placen-
tation; ovules ca. 20 per locule, 2—3-seriate, 0.3—
0.4 mm long; funicle ca. 0.4 mm long, adnate to
lower part of partition, style similar to style type D;
style apex flat, with a small style boss; stigma hemi-
spherical; the androecium oblong, 3—5-sided, 0.8-
1.2 mm long, 0.4—0.6 mm diam. at apex; sterile
staminate flowers irregularly 4—6-sided, 1.5-2.1
mm long, 0.7-1 mm wide. INFRUCTESCENCE 6
ст long, 2.5 cm diam.; seeds 18-20 per locule,
obovoid to weakly ellipsoid, 0.8-1 mm long, 0.4—
iam.
Flowering in Philodendron cretosum occurs in
both Costa Rica and Panama from March through
July (except May), mid-dry season to early rainy
season. It probably flowers over a broader period
428
Annals of the
Missouri Botanical Garden
because post-anthesis collections have been made
in January, April, June, July, and November. The
post-anthesis collection in January means that the
species may have flowered later than July or that
there is some bimodality in the flowering. Immature
fruits have been collected in November.
Philodendron cretosum ranges from Costa Rica
to Panama, from near sea level to 900 m elevation
in Tropical wet forest and Premontane rain forest life
zones. The species is perhaps restricted to the At-
lantic drainage but has been collected principally
from areas along the Continental Divide in Panama.
Philodendron cretosum is a member of P. sect.
Philodendron subsect. Canniphyllum. This species
is recognized by its short internodes, white chalky
sap (hence the name “cretosum,” meaning “full of
chalk”), persistent, thin, pale cataphyll fibers,
bluntly sulcate petioles much shorter than the
blades, and especially its linear to oblanceolate
blades. Few other species in Central America have
white chalky sap, and no other species with white
sap have narrow, non-cordate blades.
Philodendron cretosum is not easily confused
with any other species in Central America. It re-
sembles P. tenuipes Engl. from Ecuador, which also
has elongated blades with acute bases and persis-
tent cataphyll fibers, but that species differs in hav-
ing blades drying darker brown with the primary
lateral veins closer and more numerous (more than
15 pairs) and smaller spathes (usually less than 7.5
cm long).
One collection, Gémez et al. 19121, reportedly
from the slopes of Volcán Miravalles along the
Guanacaste—Alajuela border at about 1500 m,
would be well above the c
ed elevational
range and in a Lower Minas rain forest life zone.
It is doubtful if it was collected in this life zone or
at that elevation. Neither Mike Grayum nor I have
encountered this species above 600 m in Costa
ica.
Additional specimens examined. COSTA RICA. Ala-
m, Bello 1070 (CR, MO); 950 m, 1092 (INB,
MO); Naranjo—Aguas Zarcas, along Highway 15, 8.5 km
NE of Villa Quesada, 600 m, Croat emen (MO); Cafias—
Upala, 4 km NNE of Bijagua, Río Zapote, 400 m, Croat
36295 (MO); San Ramón, Bittner &
= 19121 E MO, US
N
Field Station, 50-100 m, 10°26'N, 84%01'W, Grayum
1894 (MO), 8546 (MO); Croat 61212 e Limón: Barra
de Colorado Refugio, Rio Chirri o—Río Sardina, 10—
15 m, 10%38'N, 83°45'W, Grayum 9022 (CR, MO); Río
Segundo, Asunción, Estribaciones Fila Matama, Cerro
Matama, 300 m, Gómez et al. 23424 (MO). PANAMA.
Coclé: 9.4 km above El Copé, 750-900 m, Croat 44743
(MO); El Valle region, La Mesa, above El Valle de Antón,
m, 8°38'N, 80709", Croat 37344 (F, МО); 800-
900 m, 67123 FO MO, US); 13344 (MO); 775 m, Croat
74782 (M, MO); 900-1000 m, 8”40'N, 80%07'W, Knapp
5799 (MO, NY). Colón: Nuevo pd baas ca. 3
m, Croat 33555 (MO). Panamá: Cerro
Hutchison & Dressler 2974 (BH, F, M, US: ye 17164
(F, MO, PMA); 150 m, 35985 (MO); 780-875 m, 25254
(М 0); 800 m, sb "№, 79°56'W, 74775 (MO); Cerro Jefe
region, 3-3.5 mi. МЕ of Altos de Pacora, 700-750 m,
9°15'N, 7925 X Croat 68697 (MO); along road to sum-
mit, 750—800 m, 9°14’N, 79*22'W, Croat 67090 (MO); El
Llano-Cartí, 6.8 mi. from the highway, 350 m, Croat
49123 (MO); Campamento de los guardabosques de IN-
RENARE, 800-900 m, 8?40'N, 79%55'W, Correa et al.
9516 (STRI). San Blas: Nusagandi, km 19.1, de Nevers
& Herrera 7950 (MO). Veraguas: Santa = region, Es-
cuela Agricola Alto Piedra - Rar Ds Boc km from
school, 730—770 m, Croat 25910A (MO).
Philodendron davidsonii Croat, Aroideana 6:
9-41. 1983. TYPE: Costa Rica. Limón: N of
Siquirres, originally collected by Jim Tally of
Miami, Florida, Croat 52232A (holotype, MO-
3000000-1; isotypes, B, COL, CR, K, MEXU,
NY, US). Figures 10, 129, 130, 136.
Epiphytic climber, often occurring high in canopy;
sap clear; stem appressed-climbing, thick, creeping,
leaf scars conspicuous, 1.5-2 cm long, 1-3 cm wide;
internodes short, thick, broader than long, 1.5-5 cm
long, 3-9 cm diam., olive-green to gray-green, be-
coming tannish white to brownish, scurfy; epidermis
peeling and cracking with age; roots few per a
to 7 mm diam., light reddish brown, drying dark
reddish brown, smooth, epidermis peeling; саја-
phylls soft, 28-54 cm long, unribbed to ws
ribbed to sharply 2-ribbed near apex, green
dish, sometimes dark green-striate, promptly deci
uous; petioles 50—85 cm long, 1-2 cm diam., thick-
er than broad, obtusely V-shaped, broadly sulcate
adaxially, rounded abaxially, with adaxial margins
sharp, surface medium to dark green-striate with a
weak maroon to dark green ring around apex; blades
ovate-oblong, coriaceous, acute to + acuminate а!
apex (the acumen inrolled), subcordate to cordate àt
base, 50-76 cm long, 21-43 cm wide (1.7-2.6 times
longer than wide), upper surface dark green. semi-
glossy to glossy, lower surface matte, much paler (of-
ten tinged purplish m when young); margins ns hy-
aline to yellowish gr te; anterior
lobe 48-77.5 ст oni 20-50 cm wide (3.463
times longer than posterior lobes); posterior lobes
9-19 cm long, 9.6-17.8(24) cm wide, rounded; mid-
rib flat to weakly raised, yellowish green 10 cream,
paler than surface above, convex, weakly striate, pal-
== A = = === O EE ИНОНИ» NEED RR un —————
Volume 84, Number 3
1997
Croat 429
Philodendron Subgenus Philodendron
er than surface below; basal veins (1)3–6, mostly 4
per side, with 1-2 (or sometimes all) free to base,
4th to 6th veins coalesced 3—4 cm; posterior rib 0.8—
4 cm long, not at all naked or naked to 4 cm; pri-
mary gutem ones (10)18-21 per side, departing
midrib a 5" angle, straight or weakly arcuate
to the margins, etie to weakly quilted and paler
than surface above, convex to raised and paler than
surface below; interprimary veins weakly sunken
and paler than surface above, weakly raised and
darker than surface below; minor veins darker than
surface below, arising from both the midrib and pri-
mary lateral veins. INFLORESCENCES + erect,
1-3 per axil; peduncle (3.5)5-9 cm long, 7-27 mm
diam., flattened adaxially, pale green to reddish
green, or white, often dark green-striate; spathe co-
riaceous, 15-27 cm long ((2.1)2.9—4.6 times longer
than peduncle), weakly constricted near the middle
(opening 16.5 cm long); spathe blade at anthesis
pale green, pale speckled, with margins reddish pink
outside, 10-19 cm long, 2-6 cm diam. (the opening
to 9 cm long), tinged reddish to maroon (B & K red-
purple 2/7) inside; spathe tube pale green, minutely
white-short-lineate outside, 5.9-14 ст long, 3-5.5
cm diam., reddish to maroon (B & K red-purple 10/3
to 2/10) inside; spadix sessile or weakly stipitate;
tapered, 14.5-25 cm long, tapered, broadest near the
base; pistillate portion green to yellow-green, slightly
red toward the apex to cylindrical, 3.3-10 cm
long, 1.1-2.5 cm diam. at apex, 1.4-1.7 cm diam.
at middle, 1.5-3 cm wide at base, with 19-25 flow-
ers per spiral; staminate portion 9.2-20 cm long;
fertile staminate portion creamy white, becoming or-
ange-red (post-anthesis), tapered, 1.6-2.8 cm diam.
at base, 1.5-2 cm diam. at middle, 7-12 mm diam.
ca. 1 cm from apex, broadest at the base, narrower
than the pistillate or sterile portions; sterile stami-
nate portion narrower than the pistillate portion,
white, 17-28 mm diam.; pistils 3.3-4.7 mm long,
1.6-2.5 mm diam.; ovary 8-14-locular, 3.7-4 mm
long, 2-2.6 mm diam., with sub-basal placentation;
ovules 4-8(12) per locule, striate, 2-seriate, usually
contained within translucent ovule sac, 0.2-0.6 mm
long, about equal in length or shorter than funicle;
funicle 0.2 mm long, style 1.2-2.3 mm long, 1.6-
2.1 mm diam., similar to style type B or D; style
apex usually flat; style apex flat or with a broad boss,
pronounced (with divot in center); stigma subdiscoid
to ш unlobed to weakly lobed, 1.8-1.9 mm
, 0.3-0.7 mm high, covering entire style apex,
eee medially; the androecium truncate, mar-
gins 4—6-sided; thecae elliptical to oblong-obovate,
0.1 mm wide; sterile staminate flowers blunt, 4—5-
sided, 3.5-4.5 mm long, 1.5 mm wide. Berries
greenish (nearly ripe).
Philodendron davidsonii ranges from Costa Rica
(no doubt into adjacent southeastern Nicaragua as
well) to Panama on the Atlantic slope, from near
sea level to about 200 m elevation in Premontane
wet forest and Tropical wet forest life zones.
Philodendron davidsonii is a member of P. sect.
Calostigma subsect. Macrobelium ser. Ecordata.
This species is characterized by its generally short-
creeping habit often high in the canopy (reportedly
to 30 m); short, thick internodes; sharply two-
ribbed cataphylls, which are deciduous intact; long,
stout, broadly sulcate petioles; large ovate-oblong
blades with many sunken primary lateral veins; and
one to three inflorescences per axil with the cori-
aceous spathes green outside and reddish to ma-
roon within.
Philodendron davidsonii has two subspecies; the
typical subspecies is found throughout the range of
the species, and P. davidsonii subsp. bocatoranum
is known only from the type locality in Bocas del
Toro Province.
KEY TO THE SUBSPECIES OF PHILODENDRON DAVIDSONII
Base of blade cordate to subcordate; primary lateral
veins 16-21 per side, arising а! 65-75” angle;
Costa Rica to Panama, 0-100 m ree GS
sp. davidsonii
Base of blade acute; primary — veins s 10-1 per
side, arising at 50-55” angle; ma (Bocas del
Toro), 190-220 m iio зз aa bocatoranum
Philodendron davidsonii Croat subsp. david-
sonii
Internodes 1-5 cm long, 3-8 cm diam.; cata-
phylls 38-54 cm long, sharply 2-ribbed; petioles
56-85 cm long, obtusely V-shaped, broadly sulcate
adaxially, with adaxial margins sharp, with a dark
ring at apex; blades 49.5-67 cm long, 21-43 cm
wide; basal veins 3-6, mostly 4 per side, with 1-2
free to base, 4th to 6th veins coalesced 3—4 cm;
posterior rib 0.8—4 cm long, not at all naked to 4
cm; primary lateral veins (16)18-21 per side, de-
parting midrib at 65-75” angle. INFLORES-
CENCES 1-3 per axil; peduncle (3.5)5—9 cm long,
flowers with style similar to style type D; style apex
usually flat with a broad style boss bearing a medial
divot; stigma subdiscoid to globular, weakly lobed.
Berries greenish, nearly ripe.
Flowering in Philodendron davidsonii subsp.
davidsonii occurs in the late dry season and early
wet season with post-anthesis collections from
March through August (except April) and immature
fruiting collections from June and November. Cul-
tivated collections at the Missouri Botanical Garden
flowered in April, May, June, and July.
430
Annals of the
Missouri Botanical Garden
Philodendron davidsonii subsp. davidsonii rang-
elevation in Premontane wet forest and Tropical wet
forest life zones. No specimens have been seen in
Costa Rica above 100 m elevation.
Additional specimens examined for P. davidsonii subsp.
davidsonii. COSTA RICA. Heredia: La Selva Field Sta-
tion, ca. 100 = Jacobs 2708 (MO), Grayum 2931 (MO),
2829 (M D Limón: pri comis
Suerte, 29 air km W of Tort
83°47'W, Deter 3956 (CM, E MO); 7097 (L, 54
RSA), 8782 (MO); Manzanillo de Talamanca, са. 5 т,
38'N, 82739'W, Grayum & Burton 4326 (MO); Refugio
Barra del Colorado, between Río Chirripocito and Río Sar-
dina, 12 m, 10738'N, 83°45'W, Grayum 9807 (CR, MO).
PANAMA. Bocas del Toro: Fortuna Dam-Chiriquí
Grande, 10 mi. below the divide off highway 1.2 mi., 300—
400 ft., Kress et al. 86-1992 (MO, SEL)
Philodendron davidsonii subsp. bocatoranum
Croat, subsp. nov. TYPE: Panama. Bocas del
Toro: Ojo de Agua, 7 km W of Almirante, 190—
220 m, 9°16'N, 82°28’W, 4 Aug. 1976, Croat
56853 (holotype, MO-3636098-99). Type live
at MO. Type plant is a re-collection of a sterile
live collection vouchered as Croat 38177 on 8
July 1983. Figures 19, 131-135
m diam.; cataphylla obtuse vel acute
ioribus, tum de-
76.5 em longa, 25-35 cm lata; inflorescentia 2; pedun-
culus 5.5—6 cm longus; spatha 23-25 cm longa.
Internodes 1-2 cm long, 4—9 cm diam.; cata-
phylls 28—45 cm long, ca. 6 cm wide, unribbed to
bluntly ribbed to sharply 2-ribbed near apex, bat
sisting semi-intact at base; petioles 50.5—67 c
long, flattened to sharply sulcate adaxially with ч.
axial margins conspicuous and erect, maroon ring
at apex weakly apparent; blades 58–76.5 cm long,
25-35 cm wide; basal veins 1-4, prominently as-
cending, all free to base or weakly coalesced at
base to 1 cm; primary lateral veins 10-16 per side,
departing midrib at 50-55” angle. INFLORES-
CENCES 2 per axil; peduncle 5.5-6 cm long;
spathe 23-25 cm long; flowers with style similar
to style type B, lacking a boss; style apex flat; stig-
ma subdiscoid, unlobed. Berries not seen.
Flowering in Philodendron davidsonii subsp. bo-
catoranum occurs during July (based on only two
collections).
Philodendron davidsonii subsp. bocatoranum is
endemic to Panama, known only from the type lo-
cality in Bocas del Toro, at 190 to 220 m elevation
in Premontane wet forest.
The taxon is distinguished by its thick, short in-
ternodes, two-ribbed, deciduous cataphylls, long,
sharply sulcate petioles, and ovate-oblong blades
acute at the base.
Philodendron davidsonii subsp. bocatoranum dif-
fers from the typical subspecies in having blades
acute rather than decidedly lobed at the base and
11 or fewer primary lateral veins (vs. about 16 or
ore) arising at more acute angles with the midrib
(50-55* vs. 65-757).
Additional Ades examined. PANAMA. Bocas del
Toro: vic. of Ojo de Agua, 7 km W of Almirante, 190-
22 m, 9°06'N, 22728 W, Croat 38177 (CAS, HUA, М,
MO, SCZ, VEN).
Philodendron dodsonii Croat & Grayum, sp. nov.
TYPE: Ecuador. Pichincha: along Río Blanco
across from Villa Hermosa on road 1.9 km N
of main Sto. Domingo de Los Colorados-Es-
meraldas е е departing main highway 25
km NW of Santo Domingo, 410 m, 0°5'S,
79°15'W, Г Маг. 1992, Стоа 72982 (ћојо-
C
QCNE, US). eins 137-143, 1 150.
anta oe internodia brevia 2.5-3 с
am.; cataphylla usque 20-33 cm longa, leniter vel os
2-costata, interdum in жайы persistentia, demum fibrosa;
petiolus subteres, 52-93 cm longus; lamin peso 36-87
cm longa, 28—66 ст lata, esr supra, eere pal-
lidior et oU: infra, in sicco atriflavibrunnea et impolita
vel leniter nitida; costa postica manifeste
pistilla (4)5-locularia; loculi ca. 20-ovulati.
Hemiepiphytic; stem appressed-climbing, leaf
scars conspicuous, 1-2.7 cm long, 1.5-3.5 cm
wide; internodes short on ae stout, matte to
glaucous, 2.5-3 cm diam., frequently longer than
broad on nonflowering plants, gray-green, becoming
whitish to grayish with age, epidermis flaking; cat-
aphylls thin, spongy, to 20-33 cm long, weakly to
sharply 2-ribbed, sometimes unribbed,
dark green-striate, persisting in parch
mats, eventually fibrous, rarely deciduous intact,
margins clear; petioles 52-93 cm long, 9-25 mm
diam., subterete to obtusely flattened abaxially, soft,
drying black, surface pale, dull whitish-streaked,
thinly dark green-striate, drying black blades
ovate, subcoriaceous, semiglossy, slightly bicolo-
rous, very short acuminate to + acute а! «p
weakly cordate to sagittate at base, 36-87 cm long,
Мојите 84, Митбег 3
1997
Croat 431
Philodendron Subgenus Philodendron
28-66 cm wide (1.3—1.6 times longer than wide),
(0.7–1.2 times longer than petiole), about equal in
length to petiole, upper surface dark green, drying
dark brown to dark yellow-brown, lower surface
moderately paler and glossy, drying yellow-brown
and matte to weakly glossy, margins weakly undu-
late and upturned, hyaline; anterior lobe 31—76 cm
long, 39-70 cm wide (1.65—2.9(4) times longer than
posterior lobes); posterior lobes 11-29 cm long, 9—
31 cm wide, broadly rounded to broadly obtuse;
midrib flat to sunken, slightly paler than surface
above, convex, concolorous below; basal veins 7-8
per side, and with the first free to base, numbers
3-7 coalesced 4—12 cm; posterior rib prominently
naked to 6 cm along the sinus; primary lateral veins
(4)7-8 per side, departing midrib at a
gle, to the margins, sunken and paler than surface
above, convex to round-raised, usually darker than
surface, sometimes paler than surface near base be-
low; interprimary veins sunken and concolorous
above, raised and concolorous below; tertiary veins
visible, slightly darker than surface below; minor
veins darker than surface, drying smooth below,
arising from both the midrib and primary lateral
veins. INFLORESCENCES 2-5 per axil; peduncles
5-9.5(14) ст long, 6-10 mm diam., subterete, pale
green, white-streaked; spathe 16-18 cm long
((1)1.8-3.5 times longer than peduncle), moderate-
ly constricted above the tube; spathe blade white
outside, reddish inside; spathe tube red-purple to
dark reddish (B & K red-purple 3/7.5) outside, red-
purple to dark reddish inside, sap mango-scented;
spadix 13-15(18) cm long; pistillate portion 6 cm
long in front, 5 cm long in back, 2.5 cm diam. at
middle, 2.2 cm wide at base; staminate portion 11
cm long; fertile staminate portion ca. 1 cm diam.;
sterile staminate portion 2.5-3 cm diam.; pistils 4
mm long, 1.7-2.3 mm diam.; ovary (4)5-locular,
with axile placentation; ovules ca. 20 per locule,
2-seriate, 0.3-0.4 mm long, slightly longer than fu-
nicle; funicle 0.2-0.4 mm long, adnate to lower part
of partition, style similar to style type B; central
style dome fairly well developed; style apex broadly
domed; stigma inserted on entire style apex; the
androecium truncate, + prismatic, oblong, margins
irregularly 4-5-sided, 1-2.5 mm long; thecae ob-
long, 0.3 mm wide, contiguous, + parallel to one
another; sterile staminate flowers clavate, irregular-
ly 3-5-sided, 2.3-3.7 mm wide.
Flowering in Philodendron dodsonii probably oc-
curs during the rainy season, June-September, in
Central America. It is known to flower only during
September there, with post-anthesis collections also
made from July through September and mature
lts in March. In South America, flowering col-
lections have been made in June and July, post-
anthesis from May through September, immature
fruiting from February through November (except
May, August, and October), and mature fruiting in
September.
Philodendron dodsonii is known from Costa Rica
and Ecuador and is expected to be found on the
Caribbean slope of Panama and probably on the
Pacific slope of Colombia. In Costa Rica, it occurs
at 240 to 1300 m elevation, principally on the At-
lantic slope but also on the Pacific slope on the
Fila Costefia in Tropical wet forest and especially
in Premontane rain forest life zones. In Ecuador,
this species is known from Tropical wet forest life
zones at 20 to 1750 m in Pichincha Province.
Philodendron dodsonii is a member of P. sect.
Philodendron subsect. Philodendron ser. Fibrosa.
This species is distinguished by its stout, short in-
ternodes, weakly two-ribbed cataphylls that persist
in parchment-like mats and are eventually fibrous;
more or less terete petioles; large ovate, yellowish-
brown-drying blades with naked posterior ribs; and
short-pedunculate spathes with the tube dark green
to reddish outside and red within. Another feature
useful for determination is the presence of tiny
granulations or protuberances on the lower leaf sur-
face. Juvenile blades often have minute pale dots
on the surface.
Philodendron dodsonii is most easily confused
with P. dominicalense, a species with very similar
features, which differs in having a single ovule per
locule (vs. ca. 20 for P. dodsonii) and an outer green
spathe tube (vs. red for P. dodsonii).
Philodendron dodsonii also appears close to Р.
schottianum, but that species differs in having cat-
aphylls drying with a much thicker, yellowish epi-
dermis and coarser fibers; petioles drying yellowish
brown (vs. almost black for P. dodsonii); blades dry-
ing with raised minor veins (vs. essentially smooth
for P. dodsonii); and inflorescences usually scarcely
constricted above the tube.
Another species that might be confused with P
dodsonii is P. grayumii. It has blades similar in
shape, size, and color upon drying but which differ
in having the upper surface glossy upon drying and
in usually having laticifers visible on the lower sur-
face, and especially in having mostly free basal
veins with the posterior rib (when present) rarely
naked along the sinus.
A collection near San Vito and the only wild-
collected specimen from the Pacific slope in Costa
Rica differs in having longer internodes. It perhaps
represents another species, but in other respects it
matches well with P. dodsonii.
432
Annals of the
Missouri Botanical Garden
This species was recognized as distinct by Sodiro
(in herb.), under the name P. robustum Sodiro ined.,
previously published for a different species by
Schott (1
Ра аи вија dodsonii is named in honor of Cal-
away Dodson, Director of the Centro Científico Río
Palenque and staff member of the Missouri Botan-
1 en, who first studied the species for the
о
fide herbarium specimens), there is little likelihood
that it could be the same as a species occurrin
the western slopes of the Andes. The stalls
appears to show leaf blades with basically free ba-
sal veins and little or no posterior ribs, whereas P.
dodsonii has well-developed posterior ribs. A col-
lection from Ecuador in Sucumbios (Croat 50317)
may prove to be this species. Though the inflores-
cence is immature its blades match those of P. dod-
sonit. Two other collections, Gentry 71004 from Bo-
livia (La Paz Province, 1500-1550 m) and Croat
55455 from Colombia (Cundinamarca Dept., 2000
m), may prove to be P. dodsonii. If so, the range
statement could be affected.
Additional specimens examined. COSTA RICA. Ala-
juela: Río Sarapiquí, at bridge on road to Colonia Virgen
del Socorro, 830 m, 10°16'N, Am Croat 68318 (CR,
МО); 17 km NW of San Ramón, 785 m, 10?14'15"N,
84°33'W, Croat 68138 (MO); Río nM. ca. 20 km
NW of San Ramón, 850 m, 10?12'30"N, 84^32' W, Crayum
et al. 6327 (MO). bo pc Río Reventazón, CATIE, Tur
rialba, ca. 600 m, 9°54’N, 83*39' W, Grayum et al. 3820
(МО); along Camino Raiz de Hule, SE of Platanillo (Tsi-
400 m, Croat 36796 (MO, US). Heredia:
Di ios, 240 m 10°03'N,
W, Grayum et al. 8662 (MO); Rio Blanco, Rio Frio-
Limón, d of Guápiles, Quebrada Dan
highway, 360 m, 1012'N, 83%49'W, Per | 68422 (СМ,
МО). Mos: Las illl Botanical Garden, 1300
m, Croat 44420 (MO); 64 (MO); ca. 4 km SE of San
Vito, 1150 m, 8°47'30"N, 8258", Grayum 8112 (INB,
MO); Las Cruces Botanical Garden-Río Jaba, ca. 3.5 km
SE of San Vito de Coto Brus, ca.
847'30"N, 82°58'W, Grayum 5980 (MO). San José: San
Isidro del General-Dominical, SW of San Isidro, 4.8 mi.
from Río Pacuare, 1000 m, Croat 35260 (MO); 990—
m, Croat & Hannon 79114 (CR, INB, MO); Tarrazá, vic.
Hormiguero, 1100-1200 m, Croat 78974 (CR, INB, MO n
Bore BIA. Valle: Cali-Buenaventura Highway, 1
Е of Cisneros, 220-260 m, Croat 62829 (COL, из.
NY). ECUADOR. Cotopaxi: Quevedo-Latacunga, 3 km
E of El Palmar, 800 m, Dodson & Gentry 10253 (MO); La
Mana, 1 km N of Pucayacu, 750 m, Croat 73276 Mos 2
m N of Pucayacu near Río San Francisco, 690 m, Croat
57080 Faxon! QCA); Rfo Guapara, ca. 20 km NW El Cor-
6 m, Sparre 17106 (S), 17164 (S), 17150 б),
171 08 de. 1: 71 45 e Río Pilaló, Tenefuerste, km 52-5.
п & Dode i 4 a: ) Copan
Cañar- razo— var 000-1250 ^
Camp E E-3714 (MO, NY, 9. E- Bs. (MO, NY S). Can
Azoques—El Triumfo Road,
:
a
m Machala-Loja Road, 890 |
ОСА). Esmeraldas: Quinindé, Bilsa Biological Station,
Montafias de Mache, 35 ce ы of Quinindé, 5 km
Sta. Isabela, 400-600 m, 0?21'N, 79744", Рита
Marsh 1146 (MO, QCA); Viche, e 16514 i Fila
de Bilsa, 7 km E of San José de Bilsa, ca. 80
Esmeraldas, 12 km SE of El Salto on jl
Road, 280 m, 0°37'N, 79°51'W, Gentry et al. 72977 (MO);
Rio Esmeraldas, opposite мај larer ma e
6361 (S); near San Mateo , Croa
A); Río Lita, near Lit tis а т,
Croat 38944 (MO). Guayas: Teresita, 2 km W of Bucay,
270 m, Hitchcock 20537 (NY, US); 3 km W of Bucay, 270
m, Hitchcock 20489 (NY, US), 20440 is Cordillera
Chongon-Colonche, 600 m, 1%48'S, 80
Bonifaz 5237 (GUAY, MO); ин) idi 160 m,
Croat 61593 (F, MO, QCA). Imbabura: 13-15 km E of
Lita, 800 m, Croat 38918 MOX Cachaco, 9 km E of Lita,
630 m, Croat 39000 (MO). Los Ríos: Babahoyo-Montal-
ve, Hacienda Clementina, 20n m, Sparre 17966 (S); Centro
Científico Río жу ји 230 m, Croat 38651 (MO); 250
m, 50658 (МО 4 220 m, 0%35'S, 79°12'W, 73803
ichincha: Resna ENDESA, Río Ca-
negal, 13 km
ант pek 13 (С); 35 km NW of Santa Domingo, Río
Blanco, 250 m, Gentry 961 ds (MO); Quito, Parroquia
Nan egal: ака ng Río Umachaca near Hacienda El Carmen,
1250 m, 0*07—7.5'N, 78°38’ W, рог et al. 28781 (DAV,
MO); Reserve Maquipucuna, Hacienda Esparragos—Cerro
de Sosa, ca. 6 km airline SE of Nanegal, 1500-1600 m,
0°7'N, 78°38'W, Webster & Bainard 27500 (DAV, MO);
nc quin 210 m, Croat pie (MEXU, MO, ب
QCA, WIS); Santo Domingo de orado
Zaracay, 500 m, Sparre 15190 G 1519 (S), Г oe
Rancho Brahman, ca. 10 km nto Dom
los Colorados on road to Esmeraldas 400 m , Sparre
(5); El Paraíso-Saguangal m from El Paraíso.
00 m, 78°46'W, 0^10'N, poss et al. |. 37799 (ОСА).
Philodendron dolichophyllum Croat, sp. 10“
TYPE: Panama. San Blas: Nusagandi, E El Lla-
no—Carti ману 9 mi. N of main highway, Ner-
gan Igar (Nergan "e 350 m, 9°20'N, 79W
2 July 1994, u 76569 (holotype:
MO-4619523-25; = В, СОГ, СЕ, F,
NY, PMA, US, VEN). Figures 145-148, 151.
Volume 84, Number 3
1997
Croat 433
Philodendron Subgenus Philodendron
Planta plerumque hemiepiphytica; internodia 4–6 ст
longa, 2.5-4 cm diam.; cataphylla usque 20-60 cm longa,
acute 2-costata, decidua; petiolus subteres, 28-41 ст lon-
gus, 7-20 cm diam.; lamina oblongo-lanceolata, anguste
rotunda vel leniter subcordata basi, 53-75 cm longa, 12—
15.2 cm lata, in sicco rubrobrunnea; nervis lateralibus 1
17-20 utroque; inflorescentia 2-3; pedunculus 9-12.5 ст
longus, 5-17 mm diam.; spatha 14-20 cm longa; lamina
spathae extus viridi vel purpurescenti, intus albida; tubo
spathae extus purpureo aut marronino, intus purpureo vel
rubello; pistilla 6—8(9)-locularia; loculi 3—7-ovulati; bac-
cae aurantiacae.
Hemiepiphytic, rarely terrestrial on steep banks;
stem appressed-climbing, to 3 m long, elongate,
leaf scars conspicuous, 1–1.4 cm long, 1-1.3 cm
wide; roots few, dark yellow-green, faintly ridged,
becoming reddish brown, semiglossy, to ca. 30 cm
long, 3—4 mm diam.; internodes short, semiglossy,
somewhat scurfy, 4—6 cm long, 2.5—4 cm diam., as
long as broad or slightly longer than broad, dark
green, drying brown; cataphylls coriaceous, 20—60
ст long, sharply 2-ribbed, pale greenish white,
sometimes with reddish base, drying red-brown, de-
ciduous. LEAVES erect-spreading to spreading, the
lowermost somewhat pendent; petioles 28-41 cm
long, 7-20 mm diam., subterete, somewhat obtusely
flattened adaxially, weakly glossy, moderately
spongy, dark green to gray-green with a moderately
conspicuous dark ring at apex, drying brownish;
blades large oblong-lanceolate, subcoriaceous to
weakly coriaceous, acuminate to gradually acumi-
nate at apex, gradually tapered toward base or nar-
rowly rounded to weakly subcordate at base, 53—
75 cm long, 12-15.2 cm wide (4.4—4.9 times longer
than wide), (1.6-2.3 times longer than petiole),
broadest at or near middle, upper surface medium
green, moderately bicolorous, drying reddish
brown, semiglossy, lower surface semiglossy, paler;
midrib raised to slightly convexly raised to broadly
convex, paler than surface when fresh, drying
darker than surface above, rounded, somewhat
thicker than broad to acute, paler than surface be-
low; basal veins lacking; primary lateral veins 17—
20 per side, departing midrib at а 70-80” angle,
slightly arcuate to margin, weakly sunken to mod-
erately obscure above, weakly raised and paler than
surface below; interprimary veins slightly less con-
Spicuous than primary lateral veins; minor veins
distinct, prominently visible below upon drying,
arising from the midrib only. INFLORESCENCES
2-3 per axil; peduncle 9-12.5 cm long, 5-17 mm
diam., medium green, minutely white striate, clear-
ly demarcated from the colored spathe; spathe co-
Flaceous, 14-20 cm long (1.6-1.85 times longer
uncle), acute at apex; spathe blade green
to purplish outside, whitish inside; spathe tube ob-
long-ellipsoid, purple or maroon outside, purple to
reddish inside; spadix sessile, (6)9.5-19 cm long,
broadest near the base or above the middle; pistil-
late portion cylindrical to ellipsoid, 3.2-5.8 cm
long, 1.1-1.5 cm diam. at apex, 1.1-1.7 cm diam.
at middle, 1.1-1.2 cm wide at base, with 15-17
flowers visible per spiral; staminate portion 6-9.8
cm long; fertile staminate portion white, tapered to
clavate, 8-11 mm diam. at base, 7-10 mm diam.
at middle, 5-18 mm diam. ca. 1 cm from apex,
narrower than or as broad as the pistillate portion;
sterile staminate portion as broad as the pistillate
portion, 8-11 mm diam.; pistils 1.4-3 mm long, 8—
5 mm diam.; ovary 6-8(9)-locular, 9-25 mm
diam., with sub-basal, sometimes basal placenta-
tion; ovule sac 0.5 mm long; ovules 3-7 per locule,
contained within transparent ovule sac, 0.1–0.9 mm
long, almost equal in length to funicle, style similar
to style type B; style apex flat; stigma discoid, un-
lobed, 0.6-1.3 mm diam., 0.1-0.4 mm high, cov-
ering center of style apex; the androecium truncate,
irregularly 4—6-sided, 0.4—0.8 mm diam. at apex;
thecae ovate or oblong, 0.2 mm wide; sterile sta-
minate flowers irregularly blunt, 1-2 mm long, 1
mm wide. INFRUCTESCENCE with berries or-
ange, narrowly ovoid-ellipsoid, tapered and ob-
scurely beaked at apex, 1.2 mm long, 9 mm diam.;
seeds 2-3 per locule, 0.8-2.8 mm long, 0.2-2.5
mm diam.
Flowering and phenology in Philodendron doli-
chophyllum are unclear. Flowering collections have
een made in February, June, and August, with
post-anthesis collections from March, July, August,
and October, and immature fruiting collections from
March, June, July, and September.
Philodendron dolichophyllum is endemic to Pan-
ama (though to be expected in eastern Costa Rica),
from 325 to 650 m elevation in Tropical wet forest
life zones.
Philodendron dolichophyllum is a member of P.
sect. Calostigma subsect. Glossophyllum ser. Glos-
sophyllum. This species is distinguished by its ap-
pressed-climbing habit with at least the apical in-
ternodes short; sharply two-ribbed, deciduous
cataphylls; subterete petioles (about half as long as
the blade); and especially by its long, slender
spreading leaf blades (hence the name “dolicho-
phyllum”), which dry typically reddish brown and
have weakly sunken primary lateral veins.
Philodendron dolichophyllum is not easily con-
fused with any other species in Central America,
but bears a superficial resemblance to P. pseudaur-
iculatum with which it may occur. Both species
have petioles and blades of comparatively equal
Annals of the
Missouri Botanical Garden
lengths and spathes clearly demarcated from the
peduncle. Philodendron pseudauriculatum differs
in having proportionately much shorter leaves with
blades that dry gray-green and have prominently
sunken primary lateral veins. In addition, the outer
surface of the spathe tube in P. dolichophyllum is
typically maroon or purple, while the blade is green
to purplish. Philodendron pseudauriculatum has the
spathe tube white to pinkish white at anthesis.
Philodendron dolichophyllum might also be con-
with Р. auriculatum, which differs in having
proportionately longer petioles and blades drying
ostly yellow-green and with a lower blade length/
petiole ratio (1.4-1.6 vs. 1.6-2.3 times longer than
iole) than P dolichophyllum. In addition, the
spathe of P. auriculatum is yellowish green outside
and not clearly demarcated from the peduncle (clearly
demarcated from peduncle in P. dolichophyllum).
Additional specimens examined. PANAMA, Bocas
m, 8°45'N, 82%15'W, McPher-
son 11136 (MO). Coclé: El Copé region, Alto Calvario,
200—400 m, 8?45'N, 80%35'W, enis & Davidse 2638
(MO). Panamá: El Llano—Cartí, 10.1 mi. N highway, 325—
350 m, 3°20'N, 7858 W, Codi So (Е МО, ВЗА, кы
Mile 12, 200—500 т, Croat 22908 (F, DUKE, MO, NY);
Mile 7, 460 m, 9°1 9'N, 79759", Croat 75112 (MO); Km
18, 900-1000 ft., 9^16'N, 78°58’W, Sytsma 1068 (MO);
ca. 16-18 km N, 2n m, Tyson & Nee 7359 (L, MO, SpA;
El Llano—Cartí ar Nusagandi, 300—400 m, 9°2
126 (BM, мо); El Llano—Cartf, 23-29
ican Hwy, 300—400 m, 9°22'N, 78°69’ W. Knapp 1878 iy
MO, NY); Km 22, 350 m, 9°19'N, 18°55'W, de Neve
Herrera 7849 (MO). Veraguas: Santa Fe region, ا Jd
Río San X ceni Escuela Agrícola Alto de Piedra, at
Río Segundo Brazo, 480 m, 8?33'N, 81%08'W, Croat
66895 (CM. MO. PMA. SAR).
Philodendron dominicalense Croat & Grayum,
sp. nov. TYPE: Costa Rica. San José: along
road between San Isidro del General and Do-
е са. 4.8 mi. E of Río Pacuar, ca. 1000
ca. 919'N, 83746", 22 May 1976, Croat
35268, (holotype, MO-2395109, MO-
2395111). Figures 153, 154.
Ply рата internodia 2. 5-10 ст longa, 2—
cm diam
sistenta semi-intacta; petiolus subteres, 37—74 cm longus,
.5-2 em diam.; lamina ovato-cordata, 41-46 cm longa,
19–37.5 ст lata | in sicco atribrunnea; costa postica haud
nuda aut usque 2.5 cm; inflorescentia 2; pedunculus 8-9
cm longus; spatha 14—14.5 cm longa, omnino viridis; pis-
tilla 5—6-locularia; loculi 1-ovulati.
Hemiepiphytic; internodes coarsely white-striate
beneath each node, somewhat soft, drying semi-
glossy, 2.5-10 cm long, 2-5 cm diam., gray-green,
drying yellow-brown, epidermis fissured closely;
roots moderately few per node, to ca. 30 cm long,
drying 2-3 mm diam., reddish brown, enc
sharply ridged; cataphylls soft, 16-29 cm long,
weakly to sharply 2-ribbed, green to pale per: dry-
ing thin, brown, persisting semi-intact at upper
nodes; petioles 37-74 cm long (averaging 51 cm
long), 1.5-2 cm diam., subterete, weakly spongy,
medium green, weakly flattened near apex adaxi-
ally, surface light green streaked; blades ovate-cor-
date, acuminate at apex, prominently lobed at base,
41—46 cm long, 19-37.5 cm wide (1.1-1.7 times
longer than wide), upper surface dark green, se-
miglossy to weakly glossy, drying dark brown, lower
surface semiglossy, much paler, drying yellow-
brown; posterior lobes rounded, 12.5-15 cm long,
about as broad as long; sinus hippocrepiform, 8-
11 cm deep; midrib flat to raised, paler than surface
above; basal veins 56 per side, with the uppermost
free to base, second basal vein coalesced no more
than 1 cm, (2)34 coalesced 2.5-5 cm; posterior
rib not at all naked or naked up to 2.5 cm; primary
lateral veins 4—6 per side, raised or sunken, darker
than surface, drying dark brown below; interpri-
mary veins in part sunken, the remainder flat but
visible below; minor veins alternately strongly or
weakly visible, with the more prominent veins
weakly stitched below, arising from both the midrib
and primary lateral veins. INFLORESCENCES 2
per axil; peduncle 8-9 cm long, 1–1.5 cm diam.,
fleshy, drying dark brown; spathe 14—14.5 cm long,
weakly constricted above the tube, green through-
out, narrowly acuminate at apex, drying dark brown
outside; spathe tube 5.5-6.5 cm long, 2-2.3 cm
diam.; spadix sessile; to 11.7 cm long; pistillate
portion cylindrical, broadest midway, 1.2-1.3 cm
diam., weakly tapered in both directions; staminate
portion 8.5-9 cm long; fertile staminate spadix
Neuen at sterile portion, constricted to 9-10 mm
a. 1.5 cm above base, then clavate upward, bluntly
едн at apex, 10-11 mm diam. in upper one
third, 7-8 mm diam. 1 cm а apex; sterile sta-
minate portion 12-13 mm diam.; pistils 1.6 mm
long; ovary 5-6-locular, 0.9 mm diam., with
sub-basal placentation; ovules 1 рег locule, con-
nicle; funicle 0.2—0.3 mm long, wi
chomes near base, style 0.7 mm diam., similar to
style type B; style apex flat to concave; the androe-
cium truncate, + prismatic, margins mostly irreg-
ularly 5-sided, 0.7-1.1 mm long; thecae oblong, 0.4
mm wide, + parallel to one another, contiguous
Volume 84, Number 3
1997
roa 435
Philodendron Subgenus Philodendron
PREADULT leaves with petioles 17-20 cm long;
blades to 31 cm long and 15 cm wide.
Flowering phenology of Philodendron dominica-
lense is unclear owing to its rarity, but since it was
found in flower in May, during the early wet season,
it probably flowers and fruits in the wet season.
Philodendron dominicalense is endemic to south-
west Costa Rica on the Pacific slope in the vicinity
of the type locality along San Isidro—Dominical
road, at ca. 1000 m, in Premontane rain forest.
Philodendron dominicalense is a member of P.
sect. Calostigma subsect. Glossophyllum ser. Ovata.
This species is distinguished by its appressed epi-
phytic habit; thick grayish green stems with inter-
nodes mostly longer than broad; soft, ly
2-ribbed, semi-persistent, intact cataphylls; subter-
ete petioles about as long as the blades; ovate-cor-
date brown-drying blades with the posterior ribs na-
ked for a short distance to the sinus; and paired
short-pedunculate inflorescences with green outer
spathe surfaces and one ovule per locule.
Philodendron dominicalense is vegetatively al-
most identical to P. dodsonii, but that species dif-
fers in having the spathe red-purple to dark reddish
outside and ovaries with axile placentation and
about 20 ovules per locule (vs. spathe tube green
outside, ovaries with basal to sub-basal placenta-
tion, and 1 ovule per locule). Philod. dodsonii
also differs in usually having persistent fibrous cat-
aphylls, a sunken upper midrib (vs. flat to raised),
and posterior ribs usually prominently naked for 6
cm (vs. not at all naked or naked to only 2.5 cm).
This species is also similar to P. schottianum,
especially in shape and color of the dried blades.
The latter species differs in having shorter inter-
nodes with thicker cataphylls, which dry yellow and
weather promptly into a coarse network of fibers
(vs. persisting semi-intact at upper nodes then de-
ciduous).
Additional specimen examined. COSTA RICA. San
José: San Isidro del Сем DO iod: SW of San Isi-
dro, 6 mi. from Río Pacuare, 1000 m, Croat 35454 (MO).
Philodendron dressleri G. S. Bunting, Ann. Mis-
souri Bot. Gard. 50: 25. Fig. 2. 1963. TYPE:
Mexico. Nayarit: Tepic-San Blas, along Hwy.
24, 14-16 mi. SW of junction with Hwy. 15
(Tepic-Mazatlán), 75 m, 5 Sep. 1961, Moore
& Bunting 8688 (holotype, BH; isotypes, K,
MO, US). Figures 152, 157-160.
Hemiepiphytic; stem scandent, 3—3.5 cm за
44.6 cm diam., leaf scars conspicuous, 3—3.5 c
long, 44.6 cm wide; internodes short, stout, suc-
culent, scurfy, 3—6.5 cm long, 4—7 cm diam., broad-
er than long, pale olive-green to gray-green, becom-
ing grayish white with age, epidermis peeling; roots
few per node, brownish, smooth, long, to 7 mm
diam.; cataphylls 16-22 cm long, sharply 2-ribbed,
drying weakly 2-ribbed, densely short dark striate,
sometimes deciduous in dry season; petioles 34.5—
70 cm long, 1-1.8 cm diam., terete, somewhat
spongy, surface dark green-striate, dark green ring
around apex; blades ovate in outline, deeply in-
cised-lobate, bipinnatifid, weakly bicolorous, matte
to semiglossy, acute to weakly obtuse at apex (the
acumen apiculate), cordate at base, 30—46.5 cm
long, 27—40.5 cm wide (0.99-1.2 times longer than
wide), (0.9-1.2 times longer than petiole), broadest
near the middle; anterior lobe 20.5—30 cm long
with up to 6 segments, 12-19 cm long and each 3—
5-lobed; posterior lobes rounded in outline, turned
up at an angle to midrib with ca. 5 similarly lobed
segments; sinus closed or nearly so; segments pin-
natifid, 12.3-27 cm long, divided to within 2-15
cm from the midrib; the interlobal sinus divided
0.4—0.7 the length of the lobe; midrib flat, dark
green-striate, paler than surface above, round-
raised, pale striate, paler than surface below; basal
veins 4 per side, 3—4 coalesced 5-10 mm; posterior
rib naked for 2.5 cm; primary lateral veins 5—6 per
side, departing midrib at a 45-60” angle, +
straight, eventually branching to the margins, flat
to weakly raised above, round-raised and paler than
surface below; reticulate veins visible, darker than
surface below; minor veins arising from both the
midrib and primary lateral veins; tertiary veins
sunken and paler than surface above, raised and
paler than surface below. INFLORESCENCES 1
per axil; peduncle 9-16 cm long, 7-9 mm diam.
dried), green; spathe 12-21 cm long (1.3 times
longer than редипсје), weakly constricted above
the middle, = obtuse at apex; spathe blade dark
green outside, pinkish, with darker punctations in-
side; spathe tube green outside, to 4.5 cm long,
purplish violet to wine-red or crimson inside; spa-
dix 10-15 cm long; pistillate portion to 3.5 ст
long, 2.2 cm diam.; staminate portion to 14 cm
long; fertile staminate portion clavate, to 1.9 cm
diam. at base, 2.4 cm diam. ca. 1 cm from apex;
sterile staminate portion 2.4 cm diam.; pistils 3.7(7)
mm long; ovary 4—5-locular, with axile placenta-
tion; ovule sac 2.5 mm long; ovules 3-4 per locule,
2-seriate, contained within translucent envelope,
0.4 mm long, longer than funicle; funicle 0.2—0.3
mm long, adnate to lower part of axillary wall, style
similar to style type B; central style dome some-
times present; style apex flat; stigma subdiscoid to
hemispheroid, 1.8 mm diam., 0.7 mm high, cover-
ing entire style apex; the androecium truncate, +
436
Annals of the
Missouri Botanical Garden
prismatic, irregularly 5—6-sided, ca. 1.6 mm long;
thecae = oblong, 0.4 mm wide, nearly contiguous,
t allel to one another. INFRUCTESCENCE
with spathe 10—13 cm long, pistillate spadix 4—6.5
cm long, 3.5—4 cm diam.; seeds 3-4 per locule, 2
mm long, 0.8 mm diam.
Flowering in Philodendron dressleri is probably
during the rainy season. Post-anthesis collections
are known from July and September with immature
fruiting collections known from December and Jan-
uary.
Philodendron dressleri is endemic to west-central
Mexico from coastal Nayarit including Tres Marías
Islands (off the western coast of Mexico) to southern
Sinaloa, from sea level to 370 m elevation in *Selva
Baja Caducifolia."
Philodendron dressleri is a member of P. sect.
Polytomium. This species is characterized by its
stout, succulent stems with short internodes, weakly
two-ribbed, deciduous cataphylls, terete, somewhat
spongy petioles (about as long as the blades), bipin-
natifid blades divided about midway to the midrib,
and solitary green inflorescences with the spathe
tube purplish violet to wine-red within.
Philodendron dressleri is the most northerly-
ranging species of Philodendron, extending almost
to the Tropic of Cancer. It is probably most closely
related to P. warszewiczii, but is also similar to P
radiatum, both of which differ in having gradually
much more deeply divided (almost to the midrib)
blades (vs. pinnatifid 0.4—0.7 the way to the midrib
in P. dressleri). Philodendron warszewiczii ranges
from Honduras to western Mexico, but no further
north than the state of Jalisco. Philodendron radia-
tum ranges no further north than Chiapas on the
Pacific slope.
The species is superficially most similar to P
radiatum var. pseudoradiatum, which also has
blades divided less than halfway to the midrib.
However, that taxon differs in comprising more
scandent plants with slender stems having inter-
nodes longer than broad (2-12 x 1-2.5 cm for P
radiatum var. pseudoradiatum vs. 3-3.5 X 44.6
cm for P. dressleri).
For an additional photo of this species see Bun-
ting (1965: 332).
Additional specimens examined.
Я
ХІСО. Nayarit:
near Sangaite, Е of San Blas, Philbrick 414 (BH); Tepic—
Puerto Vallarta, along Hwy. 200, 33 mi i x
N of Las Varas, 370 m, Croat 45360 (CM, MO); 6 mi. S
of Mazatan, Dressler & Wirth 2732 (UC, US); W of Jal-
cocotán, Dressler 1051 (UC); 5-6 mi. E of San Blas al
eng Gentry et al. 19477 (US); Tres Marías Islands
. Sinal
lez-Ortega 6632 (US); Mpio. Concordia, Sindicatura Mes-
illas, Саћада La Calera, 200 m, Trejo 1112 (US); Sindi-
catura Panuco, La Calera, Gónzalez-Ortega 271 (MEXU);
Mpio. Mazatlán, Gónzalez-Ortega 7393 (US).
Philodendron dwyeri Croat, sp. nov. TYPE: Be-
lize. Cayo: Macal (Macaw) River, Guacamallo
Bridge, 16%52'N, 89%05'W, Dwyer & Liesner
12334 (holotype, MO-2179389). Figure 155.
cm diam.; cataphy
cm longus, 4 mm
diam., leniter longior quam laminae; lamina ovata, 33 cm
onga, 21 cm lata, in sicco cana-viridis supra, flaviviridis
7-8 cm profundus; inflorescentia 1; pedun-
culus 7 ст longus, 5 mm Фат.; spatha 9.5 cm longa,
pistilla 7-8-
e
5
Ф
<
=
E
5
2,
б
б
©
=
e
Е
Ф
=
S
З
>
5
Ф
£
locularia; loculi l-ovulati.
Epiphytic or epilithic; sap white; internodes
slightly longer than broad, ca. 2 cm diam., semi-
glossy, epidermis light brown, drying conspicuously
wrinkled and i
prominent ridges, semiglossy, with thin broad
scales; cataphylls not seen, probably deciduous;
petioles 36.5 cm long, 4 mm diam., terete; blades
ovate, acute at apex (the acumen + inrolled, 5 mm
long), cordate at base, 33 cm long, 21 cm wide (1.5
times longer than wide), (0.9 times the petiole
length), slightly shorter petioles, margins sinuate,
upper surface drying gray-green, lower surface pal-
er, drying yellow-green; anterior lobe 24.6 cm long,
21 cm wide (1.2 times longer than wide); posterior
lobes 9 cm long, 9.5 cm wide, rounded, directed
toward base; sinus 7-8 cm deep; midrib convex and
concolorous above, convex and paler, drying yel-
lowish, closely fissured and minutely warty below;
basal veins 3 per side, with 1 free to base, 1 coa-
lesced, second and third veins coalesced 1.9 cm;
posterior rib not naked; primary lateral veins 5 per
side, departing midrib at a 50-60(75)° angle; minor
veins moderately distinct, arising from both the
midrib and primary lateral veins. INF LORES-
CENCES 1 per axil; peduncle 7 cm long, 5 mm
diam.; spathe 9.5 cm long (1.3 times longer than
peduncle), visibly constricted above the tube, ellip-
tic and to 5.5 ст wide when flattened, pne
throughout, green, drying reddish brown within;
spathe tube ca. 4 cm long; spadix sessile; 7.2 cm
long, broadest + uniform throughout; pistillate por-
tion cylindrical, 1.3 cm long, 1 cm diam.; staminale
portion 6 cm long; fertile staminate portion cylin-
drical, 1 cm diam., broader than the рне
tion, sterile staminate portion not detectable; ae
1.1 mm long, 0.6 mm diam.; ovary 7-8-locular, 2
mm diam., with basal placentation; ovules 1 pe
———— MÀ ——
—
Volume 84, Number 3
1997 A
Croat 437
Philodendron Subgenus Philodendron
locule, 0.1 mm long, + equal in length to funicle,
style 0.6-0.8 mm diam., similar to style type D;
style apex + flat, with style boss; stigma subdis-
coid, unlobed, 0.7-1.1 mm diam., 0.1 mm high,
covering interior faces of stylar funnel; the androe-
cium truncate, margins 4—6-sided; thecae oblong,
very elongated, 0.2 mm wide.
Flowering in Philodendron dwyeri is documented
by a single post-anthesis collection from January,
early in the dry season.
Philodendron dwyeri is endemic to Belize, known
only from the type collection in Cayo District of
Belize at the Macal (Macaw) River near the Gua-
camallo Bridge at less than 500 m elevation.
Philodendron dwyeri is a member of P. sect. Ca-
lostigma subsect. Macrobelium ser. Macrobelium.
This species is characterized by having internodes
longer than broad; white sap; terete petioles (slight-
ly longer than blades); ovate blades drying gray-
green above and much paler gray-green below with
a very narrow sinus; solitary inflorescences with a
green spathe; and ovaries with seven to eight loc-
ules and one ovule per locule.
Philodendron dwyeri is probably most easily con-
fused with Р. breedlovei from adjacent Chiapas,
which is also a vine with blades of similar size and
shape. That species differs, however, in having
blades that dry dark yellow-brown with a more or
less V-shaped sinus and ovaries with about 20
ovules per locule.
Additional specimen examined. BELIZE. Cayo: S of
сетат Bridge, Whitefoord 2837A (ВМ).
Philodendron edenudatum Croat, sp. nov.
PE: Panama. Veraguas: along road from
Santa Fe to Rfo Calovébora, vic. of Alto Pied-
ra, on Atlantic slope, 0.6 mi. N of Escuela
Agricola Alto Piedra (now Escuela Primer
Agricola Alto Piedra), 735 m, 4 Apr. 1976,
Croat & Folsom 33988 (holotype, MO-
2389069; isotypes, B, COL, CR, F, K, NY,
US). Figures 156, 161-164.
Planta hemiepiphytica aut aha internodia 2.5-7.5
cm longa, 2-5 cm diam., viridia vel cana, in sicco pallide
favibrunneas cataphylla 20-31 cm bus acute 2-costata, de-
cidua; petiolus subtere aut obtuse D-formatus, 34-41 cm lon-
gus, ]- 1. 5 cm diam., virens, guttatus purpureus; lamina ovata
vel awe ovata, lux basi, 34-63 cm longa, 17-33 ст
2 mm diam
ina spathae extus en pes pude viridi; tubo spa-
7 extus atriviolaceopu atrimarronino; pistilla
docularia loculi (3)4-ovulati.
| Hemiepiphytic to epiphytic; serais omn
eaf scars inconspicuous, 2 cm long, 2-2.5 cm
wide; internodes moderately smooth and glossy,
loosely and irregularly ribbed and grooved, 2.5-7.5
cm long, 2-5 cm diam., green to gray, soon brown-
ish and + densely transversed-fissured to scurfy
and glossy to semiglossy, drying pale yellowish
brown; epidermis flaking; roots l-few per node,
reddish brown; cataphylls 20-31 cm long, D-
shaped to sharply 2-ribbed, green, spotted with
purple, persisting intact at upper nodes, eventually
deciduous; petioles 34—41 cm long, 1-1.5 ст
diam., subterete to obtusely flattened or obtusely
D-shaped adaxially, especially toward apex, medi-
um to dark green, surface densely pale lineate, pur-
ple-spotted; sheathing up to % its length; blades
ovate to narrowly ovate, subcoriaceous, semiglossy,
moderately bicolorous, abruptly acuminate at apex,
cordate to subcordate at base, 34—63 cm long, 17—
33 cm wide (1.8-2.2 times longer than wide), lon-
ger than petiole, upper surface drying gray to gray-
brown, lower surface drying yellow-brown; anterior
lobe 29-51 cm long, margins convex; posterior
lobes 6-17 cm long, 6–13 cm wide, directed down-
ward; sinus arcuate to V-shaped or rarely parabolic,
(46-12 cm deep; midrib flat to concave, paler than
surface above, convex to narrowly rounded, ma-
roon-spotted and darker than surface, drying paler
than surface below; basal veins 3—4 per side, sec-
ond and higher (or sometimes only third and fourth)
coalesced 0.5—4.5 cm; posterior rib not at all na-
ked; primary lateral veins 5-6 per side, departing
midrib at a 60—75? angle, weakly curved to grad-
ually curved downward just before reaching the
midrib, weakly sunken and paler above, convex
and darker below; minor veins moderately distinct
to + obscure, arising from both the midrib and pri-
mary lateral veins, drying minutely undulate. IN-
FLORESCENCES 1 per axil; peduncle 7-9 mm
long, 9-12 mm diam., + terete, light green, some-
times faintly tinged reddish medially on one side,
clearly demarcated from spathe; spathe semiglossy,
10-13.5 cm long, 2-2.5 cm diam. midway, weakly
constricted above the tube, pce ngu
abruptly acuminate at apex (the acumen ca. 8 m
long); spathe blade greenish yellow outside, т
green, faintly striate inside, resin droplets forming
on blade surface within; spathe tube 5 cm long,
dark violet-purple with thin greenish margin (ca. 5
mm wide) and along a narrow band adaxially, faint-
ly pale striate-speckled to faintly pale lineate out-
side, dark maroon inside; spadix sessile; gradually
tapered to a blunt apex, 7. 3-11.5 cm long; pistillate
portion greenish, 4 cm long in front, 3.2 cm long
in back, 1.2 cm diam.; staminate portion 7.3 cm
long; fertile staminate portion 1.4 cm diam. at base,
1.1 cm diam. at middle, 8 mm diam. ca. 1 cm from
438
Annals of the
Missouri Botanical Garden
apex, broadest at the base; sterile staminate portion
1.4 cm diam.; pistils 2.3 mm long; ovary 7-locular,
1.7 mm diam., with sub-basal placentation; ovules
3(4) per locule, contained within gelatinous matrix
(no true envelope), 0.5 mm long; funicle 0.3 mm
long (can be pulled free to base), style 1 mm diam.,
similar to style type B; style apex flat; stigma lobed,
1 mm diam., 0.2 mm high, covering apical depres-
sions (forming a ring without papillae in the center
of the apex), depressed medially; the androecium
truncate, prismatic, irregularly 4—6-sided, mostly
4—5-sided, 0.9-1.2 mm long; sterile staminate flow-
ers irregularly 4—5-sided, prismatic, 1.3-1.8 mm
ong.
Flowering in Philodendron edenudatum possibly
occurs in the dry season, based on a single, post-
anthesis collection made in April.
Philodendron edenudatum is endemic to Pana-
ma, known in Premontane rain forest life zones at
110 to 1150 m elevation. It perhaps occurs in the
adjacent Сћосб of Colombia.
Philodendron edenudatum is a member of P.
sect. Calostigma subsect. Macrobelium ser. Macro-
belium. This species is characterized by its slightly
elongate internodes; sharply 2-ribbed cataphylls;
somewhat D-shaped, usually purple-spotted peti-
oles shorter than the blades; and ovate leaf blades
with maroon-spotted midribs and posterior ribs
which are never naked (hence the epithet *edenu-
datum," meaning not naked). Also characteristic
are the solitary greenish inflorescences with the
spathe tube dark purple-violet inside.
Philodendron edenudatum was first collected in
1979 at Alto de Piedra in Veraguas and more re-
cently on Cerro Pirre in the Darién.
The species appears closest to P. grayumii and
is perhaps only subspecifically distinct from that
species. The latter differs in having more regularly
and conspicuously ridged dried stems with a glossy,
brownish yellow epidermis; petioles always longer
than the blades (1.09-1.44 times longer); usually
larger, more broadly ovate leaf blades (1.2-1.5
times longer than wide) with secretory ducts easily
visible on the lower surface; and much larger inflo-
rescences with ovaries having axile placentation
(vs. sub-basal in P. edenudatum).
Additional specimens examined. PANAMA. Darién:
Parque National Cerro Pirre, W side, 550—760 m
°46'
ico, 17 km N of E А
& Zhu 77087 (MO, NY, US); 100 m, 77185 (CAS, CM,
MEXU, MO, VEN); vic. Cerro Pirre, trail to Rancho Frio
on slopes of Cerro Pirre, 200-450 m, 7°58'N, 77°43’ W,
Croat & Zhu 77157 (AAU, CM, MO, US); Serranfa de
Pirre above Cana Gold Mine, Río Cana—Río Escucha Rui-
0, 000 m, Croat 37741 (MO). Ver : Santa
Fe-Río Calovébora, 1.7 mi. past Escuela Agrícola Alto
Piedra, 570 m, 8°38'N, 81?08'W, Croat & Zhu 76861
(CM, МО); trail to top of Cerro Tute, 1050-1150 m, Croat
48904 (MO); 48906 (MO, US).
Philodendron ferrugineum Croat, sp. nov.
PE: Panama. Panamá: along El Llano-Cartí
road, 8.3 mi. above Inter-American Hwy.,
m, 17 July 1987, Croat 67400 (holotype, MO-
3582221-22; isotypes, AAU, B, A 3
CR, F, K, L, MEXU, PMA, US). Figures 165-
168, 173, 174, 185.
Planta hemiepiphytica; internodia brevia, 3-5 mm
diam.; cataphylla (15)26-48 cm longa, incostata aut ob-
tuse 2-costata prope apicem; petiolus teres aut semiteres,
41-67 cm longus, 1—1.3 cm diam., circa tam longus quam
amina; lamina an e ovata, manifeste cordata i
(38)56-85(102) cm longa, 17-56(62) ст lata, in sicco
rubribrunnea; venis minoribus obscuris, arcte dispositis,
ut videtur intermittentibus; inflorescentia 4-6; pedunculus
longus, 1-2 ст diam.; spatha 16-23 cm longa:
lamina spathae extus pallide viridi, intus albida; tubo spa-
1 eo-purpureo, intus
purpu
cularia; Јосић 4—7-ovulati; baccae aurantiacae.
Hemiepiphytic; stem appressed-climbing, pale to
dark green, soon brown, to 1 m long, leaf scars
conspicuous, 2.5-3 cm long, 2-5 cm wide; inter-
nodes semiglossy, short on adult plants, 3-5 cm
diam., sometimes longer than broad, pale to dark
green, becoming gray, finally brown, somewhat
scurfy, sometimes transversly fissured; roots dark
rown, to ca. ст long, 1-2 cm diam., drying
reddish brown; cataphylls (15)26—48 cm long, un-
ribbed, except bluntly or sharply 2-ribbed near
apex, green, blotched with purple-violet, to paler
green or whitish to densely dark green speckled,
maroon spotted or tinged, sulcate between ribs, dry-
ing reddish brown, deciduous, persisting weakly at
upper nodes, rounded at apex margins clear to hy-
aline. LEAVES spreading-pendent, scattered even-
ly along stem, clustered at or near stem apex; pet-
ioles 41-67 cm long, 1-1.3 cm diam. (about as
long as the blade), erect-spreading, terete ог sub-
terete, dark green, firm, weakly and obscurely sul-
cate adaxially, often obtusely flattened and obtusely
ribbed toward apex, to bluntly and broadly sulcate
near base adaxially, surface weakly glossy to matte,
weakly and densely light green-lineate or striate,
purplish red ring around apex; sheathing to 6 cm
long; geniculum thicker than petiole, 2.5-3.5 cm
long, slightly paler than petiole; blades pendent,
narrowly ovate, moderately coriaceous, bicolorous,
gradually to strongly acuminate at apex (the acu-
men tightly inrolled), cordate at base,
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
85(102) ст long, 17-56(62) cm wide (1.3-2.3
times longer than wide), (0.8—1.3 times longer than
petiole), about equal in length to petiole, broadest
somewhat above point of petiole attachment, or at
lower one-third, margins hyaline, weakly to strongly
undulate; upper surface dark to medium green, dry-
ing coriaceous, reddish brown, semiglossy to glossy,
weakly arched along midrib; lower surface very
weakly glossy to matte, much paler, drying with
minute uninterrupted ridges; anterior lobe 42—70
cm long, 9-22.2 cm wide ((7.5)12-27 times longer
than posterior lobes), margins broadly convex; pos-
terior lobes 14-27 cm long, directed downward or
inward, sometimes overlapping, obtuse to rounded;
sinus strongly spathulate to hippocrepiform; midrib
flat to weakly convex, paler than surface above,
convex to narrowly rounded, lineate, sometimes
maroon-spotted, dark green, paler than surface,
drying dark yellowish brown below; basal veins
2-3-5 per side, with 1(2) free to base, 2-5 coa-
lesced 2-4.5 cm, convex and paler than surface
above, bluntly acute to convex below; posterior rib
not naked, straight to weakly curved; primary lat-
eral veins 5-8 per side, departing midrib at a 45—
70° angle, + straight, slightly curved toward apex,
pale green to whitish, usually obtusely sunken,
sometimes to weakly and bluntly raised above, con-
vex and darker than surface below; interprimary
veins weakly visible to inconspicuous above and
below; secondary veins drying inconspicuous; mi-
nor veins moderately obscure, close and apparently
intermittent, giving veins a bumpy look, arising
from both the midrib and primary lateral veins. IN-
FLORESCENCES semi-erect to erect, 4–6 per axil;
peduncle 2.5-11 cm long, 1-2 cm diam., whitish
at base; spathe coriaceous, 16-23 cm long (1.6–
4.2 times longer than peduncle), slightly constrict-
ed above the tube, acuminate (the acumen inrol-
led), dark green outside; spathe blade with lateral
margins rolled back, sometimes pale green outside,
whitish inside; spathe tube green, heavily spotted
with purple-violet outside, 6–8(12) cm long, pur-
ple-violet to light maroon, at least the lower ¥,, oth-
erwise greenish white inside; spadix sessile; cla-
vate, weakly protruding forward at anthesis, 12-20
cm long, broadest above the middle, constricted
weakly above sterile staminate portion; pistillate
portion pale green to yellow, cylindrical or tapered
toward the apex to narrowly ellipsoid, 3.7—6.5(8)
em long, 1.3-1.5 cm diam. at apex, 1.6 cm diam.
at middle, 1.1-1.8 ст wide at base; staminate por-
tion 8.6–16 ст long; fertile staminate portion white
lo yellowish, + ellipsoid to clavate, 1.1-1.5 cm
diam. at base, 1.3-1.9 cm diam. at middle, 1.1-1.5
em diam. ca. 1 cm from apex, broadest at the mid-
dle or at the base, broader than the pistillate por-
tion, broader than the sterile portion; sterile sta-
minate portion narrower than the pistillate portion,
white, 1.5 cm diam.; pistils 2-3.4 mm long, (0.7—
0.8)1.6-2 mm diam.; ovary 8-10-locular, 1.6 mm
diam., with sub-basal (axile) placentation; ovules
4-1 per locule, 1-seriate, 0.4—0.5 mm long, longer
than (or equal in length to) funicle; funicle adnate
to lower part of axile wall; style 1.1—1.6 mm diam.,
similar to style type D; style apex flat, with raised
annulus; style boss broad; stigma button-like with
medial depression, 1.3 mm diam., 0.2-0.5 mm
high, covering entire style apex except (including
annulus) in center, medially and shallowly de-
pressed; the androecium truncate, margins 4—6-sid-
ed, irregularly scalloped or lobed on at least one
margin; thecae oblong, 0.4-0.5 mm wide,
contiguous; sterile staminate flowers rounded or
blunt, 2.2-2.5(4) mm long, (1.9)2.2-3.2 mm wide,
2.2-2.5(4) mm long. INFRUCTESCENCE erect to
semi-erect; berries turning orange, obovoid-ellip-
soid, apex blunt, 0.8-1.2 mm long, 0.5 mm diam.;
mesocarp pale yellow to orange; seeds 3-7 per loc-
ule, pale and striate or brown and smooth, strongly
sulcate, 1-1.5 mm long, 0.5-0.6 mm diam., en-
closed in a translucent envelope. JUVENILE
blades narrowly ovate, rounded at base, gradually
acuminate at apex, + shorter than petiole.
Flowering in Philodendron ferrugineum is re-
corded by only one collection in August, although
post-anthesis collections range from May through
Sa tea indicating that the species probably
flowers in the rainy season.
Philodendron ferrugineum is currently known
only from Panama, from 0 to 770 m elevation in
Tropical wet forest and Premontane wet forest life
ones.
Philodendron ferrugineum is a member of P. sect.
Calostigma subsect. Macrobelium ser. Macrobel-
ium. This species is distinguished by its short in-
ternodes; unribbed, deciduous cataphylls; terete to
subterete petioles about equaling the blades; and
large, thick, cordate blades drying reddish brown
with the minor veins obscure, close, and apparently
intermittent with a bumpy look. Also characteristic
are the 4—6 inflorescences per axil with the spathe
tube green outside and maroon inside.
Philodendron ferrugineum can be confused with
P. llanense Croat, with which it occurs in both Bo-
cas del Toro and on the El Llano Cartí Road and
Cerro Jefe in Panamá Province and which has sim-
ilar blades. The latter species differs in having cat-
aphylls becoming fibrous and blades drying green-
ish brown to somewhat blackened, never
440
Annals of the
Missouri Botanical Garden
conspicuously reddish brown. Philodendron ferru-
gineum has also been confused with P. grayumii.
See the discussion of that species for differences.
A similar, apparently undescribed species from
Colombia resembles P. ferrugineum in its large
ovate, thick blades (especially in living condition)
and in having three to four inflorescences per axil.
Examples include Croat 56246, 56708, and Mon-
salve 911 from Bajo Calima near Buenaventura in
Valle Province. These differ, however, in drying
grayish brown above and yellowish brown beneath
and in having interrupted secretory ducts between
the minor veins. In addition, they do not have the
undulated and puckered minor veins on drying that
are so characteristic of specimens of P. ferrugi-
neum.
А noteworthy collection is McPherson 11479,
which is somewhat intermediate between P. ferru-
gineum and P. grayumii. That collection has blades
that dry more reddish brown and have a bumpy
surface but also secretory ducts. It may prove to be
a new species.
Additional specimens examined. PANAMA. Bocas
del Toro: Chiriquí Lagoon, on Cayo Agua, 5 m, 910'N,
82°W, McPherson 11479 (AAU, MO); Escudo de Veraguas
Island, 5 m, 9*05'N, 8135", McPherson 11410 (MO,
PMA). Colón: Santa Rita Ridge Road, along trail to Río
Indio, 10.6 km from Transisthmian Highway, 380 m, Croat
34349 (MO); along route between Sabanitas and Porto-
bello, 3.9 mi. from highway, 250 m, 9%22'30"N,
7974130" W, Croat 75155 (AAU, BR, СМ, L, MO, PMA);
Río Agustín, Río Guanche, ca. 0 m, 9°30'N, 79?40' W,
Churchill et al. 6018 (MO); Río Iguanita, ca. 3 km abov
bridge on Portobelo road, <100 m, 9°27’N, 79?40'W,
Croat 49749 (MO). Panamá: + Llano—Carti Sn Croat
33145A (MO); Km 7-12 km m, 8 (MO);
5-6 mi. N of highway, Be TR ‚ 34787 o Mile 3.4,
1000 ft., 49098 (MO); Mile 33732 (CAS, F, че, т
Km 7-12, 360-400 т, 251 MO); Km ca. 17,
915'N, 78°50'W, oe o (MEXU, MO); Cerro Jefe
region, 9°15'N, 79*30'W, Croat & Zhu 76216 (CM, MO);
21 km above Pan-American Hi way, 600 m, Croat 35886
(MO); 750-800 m, 9*14'N, 79°22’W, Croat 67092A (CM,
MO, PMA); 4.6 km beyond peak on road to Altos de in
cora, 26.3 km from Inter-American Highway, 600 m, Croa.
35923 (MO); at Altos de Pacora, 750 m, 9715'М, 79°29’ W.
Croat & Zhu 76607 (CM, MO); 0.8 mi. Бе turnoff to
Altos de Pacora, 770 m, 9°15'N, 79°29'W, Croat €: Zhu
76612 (CR, MO, NY). San Blas: Nusagandi, along El
Llano—Cartí гед 0.7 mi. beyond Сипа Headquarters, lo-
cated 10.9 mi. N of Pan-American a 450 m,
9°18'N, sw. oe 75116 (CM, MO, PMA, TEX);
3 m, 92 79W, Hamilton & Stockwell 1073
(MO); 1-2 mi. N jh age on road to Cartf, 250-275
m, 9°20'N, 79°W, Croat & Zhu 76577, (CAS, L, MO,
PMA, US); 76580 (CM, MO, SEL, WIS); Mile 10.1, 300
m, 9°20'N, 79°W, Croat & Zhu 77029 (COL, CR, MEXU,
MO); 77030 (MO, PMA).
Philodendron findens Croat & Grayum, sp. nov.
TYPE: Panama. Chiriquf: Fortuna Dam area,
Fortuna-Chiriquí Grande, 1.8 mi. NW of cen-
ter of dam, 1080 m, 8?45'N, 82*18'W, 27 June
1994, Croat & Zhu 76502 (holotype, MO-
4619581–84; isotypes, B, COL, CR, F, K, NY,
PMA, US, VEN). Figures 7, 34, 169-171, 175,
177, 178.
Planta plerumque hemiepiphytica, varius terrestris; in-
ternodia 4-9 cm longa, 24 cm diam.; cataphylla usque
35 ст longa, acute 2-costata, persistentia semi-intacta no-
dis superioribus, fibrosentia, demum decidua; petiolus
D-formatus, ا 19 cm longus, 0. e EM cm diam.,
cum ala marginali tenui, erecta; lamina ovato-cordata,
25)40-76 cm v longa, (20. 5)28-70 cm diam., "fin dens inter
nervos later. videtur fere pinnata, in sicco deni-
grata; se i А 2: pedunculus 4—12(15) em longus,
0.3-1.4 ст diam.; spatha 13-22.8 cm longa; lamina spa-
en extus virenti vel pallide viridi; intus viridialba; tubo
athae extus viridi, intus rubro; pistilla 5—6-locularia;
loculi 15-20(22)-ovulati.
"tm,
Usually к rarely terrestrial on steep
anks; stem appressed-climbing, medium green,
ossy, sap wate, svi sented leaf scars con-
spicuous, 3. m long, 5 cm wide; internodes
short, thick, ab pee 49 cm long, 24 cm
iam., longer than broad, dark olive-green, epi-
dermis cracking, yellowish, fissured longitudinally
and transversely; roots moderately few per node,
drying dark brown, semiglossy, acutely ridged; cat-
aphylls thi
broadly sulcate abaxially, persisting semi-intact at
upper nodes, becoming fibrous and eventually de-
ciduous, blunt to acute at apex, margins weakly
пена oo у; m: (2543-119 cm v"
1.1-1.5 times longer than wide), (0. 6-1.1 times
longer than petiole), margins usually promptly
diae into segments by dividing between the pri-
ary lateral veins, making blades appear almost
ind e, upper surface dark green, semiglossy, Фу"
ing blackened to dark reddish brown, nearly matte,
lower surface semiglossy to matte, paler; anterior
kin de 8-57.5 cm long, 15.5-55(79) cm wide (2-2
mes longer than posterior lobes); een
es (7.2)11.5-32 cm long, 8.6-28(37.5) cm w1 ips
obtuse to rounded; sinus + parabolic; midrib we :
ly to deeply sunken, paler than surface above. con
Volume 84, Number 3
1997
Стоа!
Philodendron Subgenus Philodendron
vex, weakly striate, slightly paler than surface be-
low; basal veins 5-12 per side, with 0—
to base, in part coalesced 5—
naked for 1-7 cm; primary lateral veins 5—7 per
side, departing midrib at a 40—50° angle, deeply
sunken and paler than surface above, convex, usu-
ally concolorous below; interprimary veins visible,
few near apex above and below; tertiary veins vis-
ible, darker than surface below; minor veins weakly
raised and darker than surface, arising from both
the midrib and primary lateral veins; secretory ca-
nals obscurely visible. INFLORESCENCES erect,
2 per axil; peduncle 4—12(15) cm long, 0.3-1.4 cm
diam., pale to medium green, prominently, densely
white-streaked; spathe 13-22.8 cm long (1.4—2.4
times longer than peduncle), constricted midway
slightly above the tube; spathe blade medium to
pale green outside, greenish white inside; spathe
tube oblong-ellipsoid, green, densely and minutely
lineate-speckled outside, 7-9 cm long, red (B & K
red-purple 4/10, 2/10) inside; spadix sessile, ta-
pered, 13-23 cm long, broadest just below the mid-
dle; pistillate portion gray-white, tapered toward the
apex, slightly curved, 4—8.5 ст long, 1.4-2.1 ст
diam. at apex, 1.6 cm diam. at middle, 1.9 cm wide
at base; staminate portion 14.3-17.5 cm long; fer-
Ше staminate portion creamy white, tapered, 1.8—
2 cm diam. at base, 1.3 cm diam. at middle, 9 mm
diam. ca. 1 cm from apex, broadest at base, nar-
rower than the pistillate portion, narrower than the
sterile portion; sterile staminate portion broader
than the pistillate portion, 1.6 cm diam.; pistils
(3)7.7-8.4 mm long, 2-2.5 mm diam.; ovary 5—6-
locular, locules (1.6)5.8-6.1 mm long, with axile
placentation; ovules 15-20(22) per locule, 2-seri-
ate, 0.3-0.5 mm long, longer than funicle, style
similar to style type B; style apex rounded; stigma
subdiscoid to discoid, + lobed, 1.2-2 mm diam.,
0.1-0.5 mm high, covering entire style apex; the
androecium truncate, 4—6-sided; thecae oblong, 0.5
mm wide, contiguous; sterile staminate flowers
blunt, 3.1 mm long, 1.2 mm wide. INFRUCTES-
CENCE 12 cm long, 3.7 cm diam., 19.5 cm long;
stipe 4.5 cm diam.; berries irregularly quadrangular
: bluntly 4-5-dded; with a kiwi-fruit scent, 3.5—
mm diam.; seeds 1-2 per locule, ca. 17 per
beny, ч somewhat flattened, 1.5-2.2 mm long, 0.3-
diam., translucent and with fine striations.
ант stems dark green to yellow-brown,
semi-glossy to glossy, 2.4 cm long, 5 mm diam.;
blades thin, = PRE-ADULT petioles broadly
convex adaxially, the margins spreading winged, in-
curled, erect; res moderately bicolorous.
Flowering i in Philodendron findens occurs in May
and July, but post-anthesis and immature fruiting
collections from March through November (except
October) suggest a broader flowering period, with
plants flowering earlier in the dry season or much
later in the rainy season.
Philodendron findens ranges from Costa Rica to
Panama on the Atlantic slope, from near sea level
to 1400 m elevation in mostly Tropical wet forest
and Premontane rain forest but also in Premontane
wet forest. It probably also occurs in Colombia.
Philodendron findens is a member of P. sect.
Philodendron subsect. Platypodium. This species is
characterized by its generally appressed-climbing
habit and short, thick internodes with sharply two-
ribbed cataphylls, which are finally fibrous and per-
sistent; D-shaped petioles with marginal wings
adaxially; and especially by its usually black-dry-
ing, large, ovate-cordate, leaf blades, which
promptly shred into segments, making the blade
appear almost pinnate (hence the name “findens,”
meaning “tearing or splitting”). Also characteristic
are the paired, green inflorescences with the spathe
tube red within.
Philodendron findens can be confused with P.
pterotum, which has similarly shaped blades and
D-shaped petioles with narrowly winged margins.
That species differs, however, in having blades dry-
ing yellow-green, and major veins drying paler than
the surface below (in contrast to blackened and
darker than the surface in P. findens) and do not
rip into segments, and also by occurring more com-
monly in Tropical moist forest and at elevations of
usually less than 400 m. In addition, the spathe
tube in P. pterotum is purplish on the outside, vs.
solid green (sometimes reddish) for P. findens.
Cuatrecasas 14948, from Valle Department along
Río Digua at Piedra de Moler at 900-1180 m, is
apparently this species. It differs in no significant
way, but there are no field notes and the petiole is
improperly preserved so its cross-sectional shape
ot be confirmed
_ Additional specimens examined. COSTA RICA. v
er Quebrada Volio, 1100-1150 m,
10°08'N, 84°29’ X. Stevens 14115 (МО); San Ramón For-
est Reserve, ca. 10 km of Laguitos, along Río San
Lorencito, 850-1100 m, i 8'N, 84°34’ W, Hammel et al.
(CR); Reserva Biológica вада 950 m, 10^18'N,
84^45' W, Bello & Méndez 2667 (MO); 2668 (CR); 820 m,
10°19’N, 84°43’W, Bello & Eyol 2682 (INB, MO). Car-
tago: Moravia-Quebrada Platanillo, Moravia, 3-5 km
from Finca Racine, 1200-1300 m, сте a (MO); Río
Navarro, El Muñeco, 1400-1500 ley & UN
51355 (US). Heredia: 3 mi. 5 of Cariblan o. 760 m, Croa
35796 (МО); 35813 (MO); 4 mi. N of таки ‘Blanca, 1350
442
Annals of the
Missouri Botanical Garden
m, Croat 35606 (MO); Río Frío de Sarapiquí, Río Sucio—
Finca Zona Ocho, 110 m, 10?18'N, 83°52.5'W, Grayum
o, W of Finca Zona
osé: Braulio Car-
rillo National Park, 600-700 m, -Groas 78751 (СМ, СЕ,
INB, М, МО). PANAMA. Bocas del Toro: Fortuna Dam
area, Chiriquí Grande—Fortuna, 470 m, 8°50'N, 82°15'W,
олет 11646 (МО); ca. O т, 8°55'N, 82709", Croat
1 (МО); енор ае Agua, 3-6 Кт W of Al-
Era iere m, Sees 38218 (BR, CAS, F, K, MO,
PMA, US); 2 of Divide, 850-950 m, 8%45'N,
W, e vr (MO, NY). Chiriqui: Gualaca—
Chiriquí Grande, 1.4 mi. W of Centro de Operaciones,
trail to Río Motta 1010-1130 m, 8°44’N, 82?14'30"W,
Croat 67919 (B, Е, K, M, MO) Croat & Zhu 76293B
(MO); Río Chiriquf, near La Sierpe, ca. 0.5 km N of river,
IRHE Fortuna Hydroelectric Project, 1000-1100 m,
8°46'N, 82°12'W, Knapp 5052 (MO). Coclé: El Valle de
Antón теш, а! А Н Mesa, 860—900 т, Croat 37410 (МО);
775 m, 836 атт W, 74795 (MO); 800-900 m,
8°38'N, 80°09’ W, 67153 (MO); Luteyn & Kennedy 1616
(MO); El Copé region, N of El Copé, 1200-1300 m, Syts-
ma & Andersson 4622 (MO). Colón: 10 mi. SW of Por-
tobelo, 2-4 mi. from coast, 10-200 m, Liesner 1115 (Е,
MO, NY, US); Río Guanche, ca. 2 km E of bridge on main
Puerto Pilón-Portobelo Road, 100 m, ca. 9?30'N
3
n: along headwater of Río Tu uqu SR ca.
2 km air distance due DUE nue Divide, Tyler Kittredge
gold mine, ira 27148 (MO). У : Santa Fe region,
Santa Fe-Río San Luis, beyond Panels Agricola Alto de
Piedra, a ті. N of school, 480 m, 8?33'N, 81%08'W,
е 66937 (МО, РМА, А tret up Cerro Tute, 1050-
Croat 66879A (MO); sees Agrícola Alto Hoda Cal-
ébora, 15.6 km NW of Santa Fe, along trail to Santa Fe,
E of Río Dos Bocas, 450-550 50 m, Croat 27653 (MO); be-
ond Tres Bravos River, 11 km beyond Santa Fe, 650 m,
Goa 25625 (MO).
COLOMBIA. Valle: Cordillera Occidental, Río Digua,
Piedra de Moler, 900-1180 m, Cuatrecasas 14948 8 (F).
Philodendron folsomii Croat, sp. nov. TYPE:
Panama. Coclé: Alto Calvario, 9 km N of El
Copé, N of Continental Divide along path to
са. , 8°40'N, 80737' W, 24 Jan. 1989,
McPherson 13619 (holotype, MO 3693168;
isotypes, B, K, PMA, US). Figures 172, 176.
Planta epiphytica; internodia brevia, 1-1.5 ст longa,
O mm diam.; cataphylla 11-16 cm longa, acute 2-cos-
tata, decidua; petiolus subteres, obtuse sulcatus, 12-17
cm longus; lamina oblongo-lanceolata, 28-33 cm longa,
5-7 ст lata, on ite aut truncata basi; inflorescentia 1–
5 ст и. 2-5 mm diam.; spat
ш Жа extus alba; pistilla 6—7- rare loculi
Ф
l-ovulat
Epiphytic; stem scandent; internodes short, 1—
1.5 ст long, 8— iam., about as long as
broad; cataphylls 11—16 cm long, sharply 2-ribbed,
en deciduous; roots not seen; petioles 12-17
ong, 3-5 mm diam., subterete, obtusely sul-
sed. blades SENE рем narrowly acuminate
at apex, obtuse to almost rounded and decurrent,
sometimes nearly truncate at base, 28—33 cm long,
7 ст wide (4.6-5.2 times longer than wide),
(1.8-2.3 times longer than petiole), upper surface
drying gray-green to brownish green; lower surface
paler, drying yellowish green; midrib sunken above,
convex, drying paler than surface below; basal
veins 1–2, free to base; primary lateral veins 8-10
per side, departing midrib at a 35? angle, arcuate
to the margins, distinct above; minor veins arising
from the midrib only. INFLORESCENCES 1-2 per
axil; peduncle 10-11.5 cm long, 2-5 mm diam.;
spathe 10-12 cm long (about as long as the pe-
duncle), white throughout outside; spathe tube 5.5
cm long, 2.5 mm diam.; spadix short stipitate; 7-
8 cm long; pistillate portion 3—4.1 cm long, 6-
mm diam.; staminate portion 3.8—4 cm long; sterile
staminate portion whitish; pistils 0.8 mm long, 0.6
mm diam.; ovary 6—7-locular, locules 0.6 mm long,
0.3 mm diam., ovule sac 0.6 mm long, with sub-
basal placentation; ovules 1 per locule, contained
within gelatinous transparent matrix (no true en-
velope), 0.2-0.35 mm long, as long as funicle; fu-
nicle 0.2–0.3 mm long (can be pulled free to base),
with glands at base, style similar to style type
style apex flat; stigma subdiscoid, unlobed, + trun-
cate, 0.9 mm diam., 0.3 mm high, covering almost
entire style apex; the androecium truncate, pris-
matic, margins irregularly 5—6-sided, 0.6-0.8 mm
long. by
Flowering phenology in Philodendron folsomii is
uncertain because few collections exist. Flowering
collections are known from January and April, both
within the dry season but in an area that is по!
markedly seasonal.
Philodendron folsomii is endemic to Panama.
known only from the type locality in Coclé Depart-
ment, at 700 to 800 m elevation in Premontane rain
forest.
Philodendron folsomii is a member of P. sect. Ca-
lostigma subsect. Glossophyllum ser. Glossophyllum.
This species is characterized by internodes about as
long as broad; sharply two-ribbed, deciduous cat-
phylls; oblong, narrowly acuminate, oblong-lanceo-
late, green-drying blades with obtuse-attenuate
bases; and solitary, long, slender-pedunculate
inflorescences with white spathes. pe
Philodendron folsomii is most easily confu
with Р. sphalerum Schott from the Guianas аһ
eastern Venezuela, which has similar leaves- је
latter species differs in having smaller, gene у
Sk A —— — ———— ние» |] — — K—
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
more numerous (up to four) inflorescences per axil
with more slender peduncles (drying scarcely 2 mm
diam. vs. 4 mm or more in P. folsomii) and spathes
less than 7 cm long (vs. 10-13.5 cm long in P.
folsomii).
Philodendron folsomii is also similar to P. cor-
reae, which has similarly shaped leaf blades that
also sometimes dry green. The latter species differs
in having generally longer internodes; petioles usu-
ally sheathed to near the apex; blades usually dry-
ing black and lacking conspicuous primary lateral
veins; and spathes not so conspicuously constricted
above the tube.
Additional specimen examined. NAMA. Coclé: El
Copé region, 700 m, Folsom & 1 i 2447 (MO).
Philodendron fortunense Croat, sp. nov. TYPE:
Panama. Chiriquí: Gualaca-Chiriquí Grande
Road, vic. of Fortuna Dam, at junction of road
to IRHE headquarters, 1200 m, 8?45'N,
82°18'W, 23 June 1987, Croat 66714 (holo-
type, MO-3635169; isotypes, AAU, B, CAS,
CM, COL, CR, DUKE, EAP, F, GH, HUA, K,
L, M, MEXU, NY, P, PMA, QCNE, RSA, 5,
SEL, TEX, US, VEN). Figures 179, 181-184.
Planta terrestris aut hemiepiphytica; internodia 3-6 cm
longa, 4-7 ст diam.; cataphylla 18-32 cm longa, acute
2-costata, кен decidas; уут plus minusve D-for-
matus, 49-70 cm longus, 1.2-2.5 ст diam., glaucus, mar-
ginibus manifeste alatus, alis undulatus; lamina ovato-cor-
data 2 Затен, cordata basi, 36-60 ст longa,
; inflorescentia 4; pedunculus 71-8 cm
patha ст longa, omnino vi-
rens, tubo spathae marronino, intus suffuso marronino in
laminam; pistilla 7-8-locularia; loculi 9—10-ovulati.
Terrestrial or hemiepiphytic; stem appressed-
climbing, 2-3 m long, thick, leaf scars conspicu-
ous, 1.2-1.7 cm long, 2-2.5 cm wide; internodes
smooth, to weakly pruinose, 3-6 cm long, 4-7 cm
diam., about as long as broad, green to olive-green,
epidermis drying smooth, semiglossy, light yellow-
brown (B & K yellow-red 9/10), cracking with age,
fissured transversely, sometimes with minute cracks
Perpendicular to axis; roots few per node, elongate,
to 4 mm diam., light brown, smooth, weakly glossy,
the epidermis cracking free upon drying; cataphylls
thick, 18-32 cm long, sharply 2-ribbed (ribs 4 mm
high), pale green, glaucous, dark short-lineate, de-
ciduous. LEAVES erect-spreading, clustered at or
near stem apex; petioles 49-70 cm long, 1.2-2.5
cm diam., + D-shaped, becoming flattened and
much Кал than thick toward apex, medium
green, rounded to flattened abaxially, with adaxial
Margins prominently winged (to 3 mm diam.), un-
dulate to broadly flattened, surface dark short-li-
neate, glaucous; blades ovate-cordate to oblong-
ovate cordate, moderately coriaceous to
subcoriaceous, gradually acuminate to abruptly
acuminate at apex, cordate at base, 36-60 cm long,
—44 cm wide (1.3-1.8 times longer than wide),
(0.85—0.88 times the petiole length), broadest
slightly above point of petiole attachment, margins
moderately undulate, upper surface medium green,
semiglossy, drying greenish yellow, lower surface
medium to pale green, glaucous, matte, paler, dry-
ing greenish yellow; anterior lobe 28-44 cm long,
10-20 cm wide (0.9-1.3 times longer than wide),
(1.8–2.9 times longer than posterior lobes), broad-
est at base; posterior lobes usually overlapping, 10—
20 cm long, 10-20 cm wide, directed downward
and slightly inward, obtuse to rounded; sinus ob-
long-triangular or closed, 10-19 cm deep; midrib
flat to weakly raised, pale green, concolorous
above, weakly raised, paler than surface below; ba-
sal veins 5-10 per side, 1-2 pairs free to base, the
remainder mostly free, sometimes weakly coalesced
14 cm, drying weakly raised; posterior rib usually
not present, naked and straight if present; primary
lateral veins 7-11(15) per side, departing midrib at
a 3045” angle, straight or slightly arcuate to the
margins, flat to sunken and drying darker than sur-
face above, convex and darker than surface below;
interprimary veins prominulous, weakly sunken to
flat, concolorous above, flat and darker than surface
below; lesser veins visible when dried; minor veins
conspicuous, fine, darker than surface below, aris-
ing from both the midrib and primary lateral veins.
INFLORESCENCES 4 per axil; peduncle 7.7–8 cm
long, 8 mm diam., medium green; spathe + erect,
20-21 cm long (2.5-2.7 times longer than pedun-
cle), medium green throughout, glaucous, + acute
at apex; spathe tube maroon, suffused maroon onto
blade inside; spadix sessile; tapered, 20-21 cm
long; pistillate portion tapered toward the apex, 5.2
cm long, 1.3 cm diam. at apex, 1.5 cm diam. at
middle, 1.7 cm wide at base; staminate portion 14.7
cm long; fertile staminate portion tapered, 1.4 cm
diam. at base, 1.3 cm diam. at middle, 1.1 cm
diam. ca. 1 cm from apex, broadest at the base,
narrower than the pistillate portion, narrower than
the sterile portion; sterile staminate portion narrow-
er than the pistillate portion, 1.4 cm diam.; pistils
4.5 mm long, 2.1-2.4 mm diam.; ovary 7-8-locular,
locules 1.9 mm long, with sub-basal placentation;
ovules 9-10 per locule, 0.4—0.5 mm long, slightly
longer than funicle, style 2 mm long, 1.8 mm diam.,
similar to style type B; style apex flat; stigma sub-
discoid, unlobed, 2 mm тры 0.4 mm high, соу-
ering entire style apex; the androecium truncate,
margins 4—6-sided; nina oblong, 0.5 mm wide,
Annals of the
Missouri Botanical Garden
contiguous; a di staminate flowers blunt, 3.2 mm
long, 1.7 mm w
Flowering in Philodendron fortunense is known
only from April, and immature fruits from June.
This corresponds to the late dry season and early
rainy season in most of Panama, though this species
occurs in an area with little seasonality.
Philodendron fortunense is endemic to Panama,
known only from Chiriquí Province in the Fortuna
Dam region (hence the name), from 1100 to 1300
m in elevation in regions of Premontane rain forest.
Philodendron fortunense is a member of P. sect.
Philodendron subsect. Platypodium. This species is
recognized by its thick stems with short, smooth
(when fresh), green internodes; sharply two-ribbed,
glaucous, deciduous cataphylls; ovate-cordate
blades with usually overlapping posterior lobes,
and pale, glaucous lower surfaces; and especially
by the glaucous, broadly flattened petioles with un-
dulate-winged margins.
Philodendron fortunense, one of the showiest,
most distinctive species in the Central American
region, is not easily confused with any other spe-
cies. Из greatest similarity is to P. brenesii, with
which it may occur, but that species differs in hav-
ing proportionately longer, more or less terete pet-
ioles, and one to two inflorescences per ах! (vs. up
to four inflorescences per axil in P. fortunense).
Additional specimens examined. PANAMA. Chiriquí:
Fortuna Dam area, OE e Grande, jct. of roa "d
to IRHE popu жүрү 1200-1300 m, 8^45'N, 82^18'W,
Croat 66529 (CM, ENCB, НАМА, JAUM, eus ње ОСА,
КВ, W); 9.6 mi. beyond Los Planes de Hornito, 1300 m,
Croat 48727 (AAU, COL, F, MO, оа ОСМЕ, 48728А
(MO); 10.1 mi. NW of Los Planes де Hornito, 1300 т
82°17'W, 8'45'N, Croat 49814 (MO); 50000 ny CAS,
CM, L, MO, QCA, SCZ, TEX); 4.5-5 km N of dam over
Fortuna fake: 1100-1135 x 8'43'N, 82%17'W, Croat &
Grayum 60000 (B, CM, K, MO, NY, RSA, US); à mi.
bera: Los Planes de Mcd , 14 mi. W Ш Centro de
Operaciones, trail to Río наан 1010-1130 т, EN
82"14' 30", Croat 67921 (G, M, MO).
Philodendron fragrantissimum (Hook.) G. Don,
in Sweet, Hort. Brit. ed. 3: 632. 1839. Figures
30, 186-191. Caladium fragrantissimum
Hook., Bot. Mag. 61: t. 3314. 1834. TYPE:
Guayana (specimen introduced by C. S. Parker
in 1834 to as Botanical Garden) (ho-
lotype, K
Philodendron latipes K. Koch & Augustin, in A Braun et al.,
Append. Gen. Sp. Hort. Berol. 1854. 6. 1854-1855.
TYPE: origin ил D fotu. B destroyed) Bur-
chell 9452 (neotype, here designated, K
Philodendron ais, Schott, tom Aroid.
TYPE: Braz
ed 84. 1860.
mazonas: (Ega) Tefé, Poeppig s.
(holotype, B). vob dian
а а“ ole Schott, J. Bot. га 1864.
azil. Bahia: Пћеов, Archduke Е Masini
us s.n. n. ole, lost); Schott | ic. 3592 (neotype,
тд designated).
Philodendron sie C. Wright, in Griseb., Cat. Pl.
Cub., 220. 1866. TYPE: Cuba. Near Retiro, Wright
3212 aie td ме designated, К).
Philodendron demerarae Gleason, Bull. bii Bot. а
30. 1I. 1929. "TYPE. Guyana. SE of Lamaha stop-
off, 27 de 1919, Hitchcock 16987 (holotype, Ny)
Philodendron accrescens N. Simmonds, Kew Bull.
1951: 402. 1951. TYPE: енд Long Stretch, 15
Jan. 1949, Simmonds 14256 (holotype, TRIN).
Hemiepiphytic; acaulescent or caulescent, stem
appressed-climbing or rarely scandent with slender,
whip-like branches bearing small leaves, to 1-6 m
long, sap orangish to brownish, sticky, spicy-scent-
ed; internodes short, 1-4 cm diam., usually thicker
than long, dark green, semiglossy, Mc by cat-
aphyll fibers, roots often many per
dark brown, 2-3 mm diam.; pS ot pss
2-ribbed or sharply D-shaped, greenish white,
sometimes drying reddish brown, persisting as fi-
bers, margins weakly upturned below. LEAVES
erect- reni petioles 22—70 cm long, 4-11 mm
diam., = D-shaped to sharply C-shaped, broadly
аи ге ы blades ovate to ovate-triangular,
subcoriaceous, moderately bicolorous, acuminate at
apex (the acumen sometimes inrolled or very short
apiculate, 2-5 mm long), cordate at base, 21.6-59
ст long, 17-37.5 cm wide (1.1-1.7 times longer
than wide), (0.7-1.3 times longer than petiole),
about equal in length to petiole, upper surface se-
miglossy, lower surface semiglossy; anterior lobe
19.5-49 cm long, 17-39 cm wide, (2.1-3.9 times
longer than posterior lobes); posterior lobes 5-16. 5
ст long, 10-18.3 ст wide, obtuse to rounded; mid-
rib broadly sunken, pales than surface above, con-
vex, bluntly angular, r than surface below; ba-
sal veins (3)4(5) per SR pe (0)1(2) free to -—
1-2 coalesced 1—4 cm; posterior rib not nake
naked 1—4 cm along the sinus; primary lateral veins
3-6 per side, etched-sunken above, convex below;
interprimary veins visible and discontinuous above;
minor veins visible and darker than surface below,
arising from both the midrib and primary lateral
veins. INFLORESCENCES erect to semi-erect, 2
per axil; peduncle 3-13.5(17) cm long, 3-15 mm
diam.; spathe 8.5-19 cm long, (1.2-3.6 times lon-
ger than peduncle); spathe blade white to greenish,
rarely reddish outside, white to greenish inside;
spathe tube reddish to dark maroon outside, red to
maroon inside; spadix stipitate 3-4 mm long; Су”
lindrical, 9-16 cm long, + uniform throughout; pis-
tillate portion cylindrical, 2.5-5 cm long, Leve
diam. throughout, 1.6 diam. at apex, 1.8 cm diam.
Volume 84, Number 3
1997
Croat 445
Philodendron Subgenus Philodendron
at middle, 1.5 cm wide at base; staminate portion
6.3-7.8 cm long; fertile staminate portion creamy
white, cylindrical, to 1.3 cm diam. at base, 1.4 cm
iam. at middle, 1 cm diam. ca. 1 cm from apex,
broadest in the middle and as broad as the pistillate
and sterile portions; sterile staminate portion as
broad as or broader than the pistillate portion, 1.2—
1.5 cm diam.; pistils 3.2-3.5 mm long, 2.1-2.9 mm
diam.; ovary 6—10-locular, with axile placentation;
locules 1.6-2.3 mm long; ovules (24)32—36 per loc-
ule, 2-seriate, 0.2-0.3 mm long, + equal in length
to funicle; funicle 0.2—0.3 mm long, style similar
to style type D; stylar canals emerging into con-
spicuous depressions; ira apex slightly concave
to flat; stigma subdiscoid ame like), unlobed,
2.1-2.7 m ^ m high, covering en-
tire 5 apex, inserted on entire style apex; the
androecium 4—6-sided; thecae slightly obovate,
usually bright red to purple-red, sometimes orange,
rarely yellowish white (McPherson 14496) to white
(McPherson 11380)
owering in Philodendron fragrantissimum in
th Panama and Middle America occurs during
the rainy season from May to October (except Sep-
tember). Some post-anthesis collections from Mid-
dle America are scattered in the dry season as well
(January and March). Post-anthesis collections are
also principally from May through December (ex-
collected during May through December (except
July and August).
Philodendron fragrantissimum ranges from Be-
lize to Panama along the Caribbean coast and to
Pacific Colombia (Сћосб), then disjunctly to the
azon basin, where it ranges from Venezuela to
the Guianas, northern Brazil (Roraima and Ama-
zonas), southern Colombia (Meta, Caquetá, Guajira,
and Vaupes), and Peru (Amazonas, Loreto, Ucayali,
and Madre de Dios). It is also known from Cuba
and Trinidad, and it is to be expected in Ecuador
and western Brazil. Ranging from near sea level to
000 m elevation, this species occurs in Tropical
Moist forest, nt wet forest, and Tropical wet
orest life zo
ndron fragrantissimum is a member of P.
Sect. Philodendron subsect. Macrolonchium. This
Species is distinguished by its short internodes; a
tendency to produce slender, whip-like branches
m near the apex; persistent, reddish brown cat-
aphyll fibers; more or less D-shaped petioles with
somewhat elevated, lateral margins; ovate to ovate-
triangular, cordate blades (about equal to the peti-
oles in length); and colorful inflorescences with the
spathes bright red on the tube and white on the
blade.
Philodendron fragrantissimum is not easily con-
fused with any other species in Central America,
since the combination of D-shaped petioles, persis-
tent cataphylls, and red and white inflorescences is
unique, but it has been confused with the Vene-
zuelan P. chimantae G. 5. Bunting, which differs in
having typically more triangular petioles that are
actually winged (not merely ribbed) on the lateral
adaxial margins, brown cataphylls persisting intact
for a time before falling off, and blades with a much
arrower sinus
Philodendron fragrantissimum populations in
South America, particularly in the Guiana region
the type locality), have nearly triangular rather
than D-shaped petioles typical of those in Central
America. In addition, the blade shape is more near-
ly triangular than ovate as in Central America. I
believe, nevertheless, that differences between
Central American and South American populations
do not warrant taxonomic recognition.
~
Additional specimens examined. BELIZE. Cayo:
along Hummingbird Highway at Mile 28, Dwyer 11411
(MO); Mile x bon 9180 (LL). Stann Creek: Middle-
sex, Gentle 2 Ма: = at Toledo: Punta Gor-
da-San Antonio, unction with road, 100 ft.,
Croat 24510 O) COT RICA, Alajuela: Cafias- Upa-
la, near Río Zapote, 1.8-2.7 km S of Río Canalete, 100
m, Croat 36361 (MO). Cartago: Tucurrique, Las Vueltas,
5-700 m, Tonduz 13313 (С, US). Heredia: Zona Pro-
tectora, N slopes of Volcán Barba, along Quebrada Cap-
ta
, Grayum & Hammel 5541 (MO).
le Delirios: Amubri, 100 m, 9°31'N,
pa ay њи et al. 17512 (CR, MO); Barra del Col-
rado, 0-2 m, 10%47'N, 83%55'W, Stevens pee
MO); Río Colorado, 1 1-5 m, 10%47'40'N, 30"W, Dav-
idse & Herrera 30922 , MO); shed de de
Talamanca, са. 5 m, 9°38'N, 82°39'W, Are & Burton
4327 (MO); dur A Uren, Finca de ,
23760 (MO, РМА); il Tap-
8641 (RSA), 8370 (MO, RSA), m et al. 6766 (
; arenas: Piedras Blancas-Rincón, 3.7 mi. У
of Pan-American Highway, 90-105 m , 846'N, 83718",
Croat 67652 (MO); 10 km W of highway, 200 m, 79169
(INB). GUATEMALA. Izabal: ca. 7 mi. S of Puerto Bar-
rios, 50 m, Croat 41810 (GUAT, MO). HONDURAS. At-
Cordillera = de Dios, 5 of San José de Texíguat,
5°30’N, 87726", Davidse et al. 34398
(MO). ÓN Jinotega: Río Bocay, Caño Litutus,
Annals of the
Missouri Botanical Garden
175 m, 13?58'N, 85?21'W, Stevens 16680 (MO). Zelaya:
Comarca del Cabo, Miguel Bikou, Robbins 5874 (F, GH,
MO, NY); еа Neill 2587 (MO); Monkey Point,
Cafio El Pato, 10 11?35'N, 83°42'W, Moreno 12464
(MO); ca. 13 km live ade 200 m, Pipoly 3833 we
ca. 13 km above Kururia (ca. 14?39'N, 84?04'W), on road
to San Jerónimo (14%42'N, 84°11'W), 200 m, Pipoly 3835
(MO); 8 (MO) Colonia Kururia, 50 m, 14°41'N,
84°04'W, Pipoly 3884 (MO); Cerro Вака, .5 km E of
Río C . m, 13%40'N, 84?30' W, Pipoly 4967
ya) , 84°03'W, Stevens 7627 (МО);
fedis Puck: Chee ca. 15.7 km SW of Rio Kukalaya,
100 m, 13%58'N, 84^12'W, Stevens 8526 (MO); Mina Nue-
va América Road, ca. Km 10, Stevens 12663 (MO); El
Empalme—Limbaika, vic. of road to Alamikamba, ca. 25
m, 13°32'N, 84°3 get DN 12738 (MO); ca. 1.5 km SE
of Palmera, 60 m, 13°35'N, 84°20'W, Stevens 12847 (MO);
Bahía de Bue "iid Cay, 0-15 m, 11%56'N
83745", Stevens 20069 (MO); Bluefields, Río Бесс.
Molina 2019 (Е); at junction of road to Alamikamba, с
ње m, 13°32'N, 84730", Stevens 21747 (MO); Río Tkon:
, Waspam-Puerto Cabezas, 200 m, 14?43'N, 84°06' W,
Pipoly 4056 UE ee El Cabo, Pine Savannas, Mo-
lina 15043 (F). P
Grande, near ни Ја, 250 т
Pherson 11148 (M, MISSA,
Wedel 1148 (GH, MO). Canal увид о
5.6, Witherspoon 8606 (MO, SEL); Ty m, arn 6589 (MO
RSA, US); 6591 (MO); Summit Gardens, Croat 10891 (Е
МО, NY, SCZ); Frijoles, Standley 27477 (US); Ft. Ran-
dolph, uU 28728 (US) Cerro Pelado, 1 km N of
Gamboa, 200-220 m, Nee 7769 (MO, US); Cerro Viejo
vicinity, on K16C, Blum 1273 ESU, eN, PMA); Camp
Ba
land, Kenoyer 184 (MO); 185 (US); F airchild 3080 (US);
Croat 6769 Арзы 4529 (МО); 1 г f Mo 11
0911 Mo . SCZ); 9042 (МО); 9.
(MO) Wetmore & Abe i ES (MO); Shattuck
m, 8°47 'N,
, Croat 33539 (MO); Río
Indio-Miguel de la Bonds. As 36922 (MO); re et al.
4546 (MO, SCZ); Santa Rita Ridge Road, 21-16,
зе Кпарр 5851 + мо); Km
У m, 926'N, 79°38’W, Knapp & Schmalzel
1797 (MO); Km 18-20, 1000-1200 ft., 9°24’N, 79°39’ W,
Sytsma 2035 (MO, PMA); Km 16-18 km from highway,
m, 9°26'N, 79%37'W, Knapp 1773 (CM, K, MO).
Darién: ка St. Dorotea, Dwyer 2256 (MO); Alturas de
Nique Region, S of El Real
a, 600 m, Sullivan 669 (MO); Cono
- region, along trail from base camp to Rancho Frio
slope of Cerro Pirre, 200-450 m
0); 58 mi.
abest e Lago а о Azul, 840 m, 9°13'№, 79°22’W, J. Miller
& L. Miller 888 (MO, NY); Cerro Jefe region, 10 mi. from
Tocumen Circle, 800-1000 m, Gentry net O p
mi. from highway, Croat 15188 (MO); K m
9°20'N, 79*W, Cr
9°22'W, Croat 67095 (MO);
erspoon & ا 8551 (МО); 770 m, 9715,
79°29'W, Croat & Zhu 76613 (MO, PMA, US); El Llano-
Cartí Road, Km 5-6, 350-375 m, Croat 34800A (MO); 7
c sy e 21 m, uae La °59'W, Croat 75107
m N off, 300 m, 916'N,
ти 4 Curl 3810 (MO, RSA). 5 km N ef highway,
= m, Nee 7915 (G e baee trail to pum Brewster
m Río e Valley у, , 9°20'N, 79715", Mc-
та 7503 (МО). Ѕап ~ El fais ss Road, 34—
38 km from Pan-American Highway, 100—200 m, 9°25'N,
79W, Knapp & Schmalzel 5467 (MO); km 26.5, 200 m,
9°19'N, 78°55'W, de Nevers et al. 7839 (GH, MO, US);
NW of Nusagandi on Sendero Wedar, 150—200 т, 915'N,
79°W, McPherson 11049 (MO). Veraguas: Boca de Con-
cepción, in Golfo de los Mosquitos, 10 m, 8°50'N, 81°W,
McPherson 11380 (CM, M, MBM, MO)
zy
Philodendron gigas Croat, sp. nov. TYPE: Pana-
ma. San Blas: El Llano—Cartf road, 10.1 mi. N
of COM V usen then 1.9 mi. N of
Nusagandi, 3 9°20'N, 79°W, 20 July
1994, Croat А Zhu 76988 (holotype, MO-
4619563-69; prige B, CAS, CM, COL, CR,
F, K, MEXU, NY, PMA, QCNE, VEN, SCZ,
US). | A 192-196, 205.
е eget ee ig esi ind е cm longa, (3)6–
is cataphylla a, 30-50 ст lon-
=. in sic ui ja y grasso reticulo
(59)98-137 cm longus;
data vel sagittata basi, (44)81- 125 cm longa, (30)37-90
lo
rubriviolacea; tubo spathae extus purpurascentiv
intus atripurpa парале иди pistilla 4—5-locularia; Seal
20-ovulati; baccae viridalbae aut alba.
Hemiepiphytic, growing to 15-20 m; stem ар-
pressed-climbing (or scandent when pre-adult), sap
reddish, viscous, leaf scars conspicuous, to 4 ст
wide; internodes short, 1-2 cm long, (pre-adult
length to 15 cm long), (3)6-10 cm diam., glaucous,
olive-green to dark green, coarsely white striate
near apex, the epidermis with reddish brown patch-
root i 3 cm diam., grayish, geh and conspic-
uously warty; сину 30-50 cm long, to 25-30
cm broad at base (when flattened out), = D-shaped,
unribbed to sharply 2-ribbed (ribs low), greenish
white to reddish, thick, juicy, drying reddish brown,
persisting semi-intact, eventually as a reticulum of
fibers and patches of epidermis; petioles (59)98-
137 ст long, (1.5)2-3 cm diam., subterete, medium
Volume 84, Number 3
1997
oat 447
Philodendron Subgenus Philodendron
to dark green, semiglossy, weakly and obtusely flat-
tened adaxially, especially near apex, surface
densely and finely short-lineate to faintly striate;
blades ovate to narrowly ovate, moderately coria-
ceous, bicolorous, acuminate at apex (the acumen
weakly inrolled), cordate to sagittate at base,
(44)81-125 cm long, (30)37-90 cm wide (1.4—1.7
times longer than wide), (0.9-1.2 times longer than
petiole), subcoriaceous, margins hyaline, upper
surface dark green, velvety, matte, lower surface
slightly paler, semiglossy to matte, sometimes
tinged pale reddish between veins; anterior lobe
33-102 cm long (3.5-6.1 times longer than poste-
rior lobes); posterior lobes 13-26 cm long, 13-24
cm wide, incurved and overlapping with sinus
closed on adult plants, obtuse; midrib flat to weakly
convex, silvery-white to yellowish green, paler than
surface above, narrowly rounded and slightly paler
than surface below; basal veins (3)9-12 per side,
first and second free to base, most of remainder
coalesced to 4 cm, 2 coalesced to 14 cm; posterior
rib naked 1.54 cm; primary lateral veins (4)7-10
per side, departing midrib at a 40-60” angle, ar-
cuate to the margins, narrowly weakly sunken and
slightly paler than surface and marginally discol-
ored above, prominently round-raised, paler than
surface below; minor veins weakly visible but not
distinct below, arising from both the midrib and
primary lateral veins. INFLORESCENCES (post-
anthesis) to 7 per axil; peduncle 13-16.5 cm long,
green, white-striate, especially at apex; prophylls to
22 cm long; spathe 13-16.5 cm long, 5 cm diam.,
moderately constricted above the tube, weakly
glossy; spathe blade 5.8 cm long, white, tinged red-
dish outside, dark red-violet inside; spathe tube ob-
long-ellipsoid, 7.5 cm long, purplish violet, short
white lineate outside, dark violet-purple, short
white lineate inside; spadix sessile; to 12.5 ст
long, ca. 1.5 cm diam., constricted above sterile
portion; pistillate portion pale green, + ovoid, 4.2
cm long in front, 3.4 cm long in back, 2.2 cm diam.;
staminate portion 11 cm long; fertile staminate por-
tion to 2.5 cm diam. midway; sterile staminate por-
tion 2.4 cm diam., slightly broader than pistillate
portion; pistils 4-6 mm long, 1.4 mm diam.; ovary
4—5-locular, with + axile placentation; locules 2.8—
5.2 mm long; ovules ca. 20 per locule, 0.1–0.25
mm long, funicle shorter than to equal in length to
ovules, style similar to style type B; style crown
domed, irregularly lobed; stigma hemispheroid, 1—
1.2 diam., 0.4-0.7 mm high. INFRUCTES-
CENCE with berries greenish white; seeds 3—4 per
locule, 1.6 mm long.
Flowering in Philodendron gigas occurs during
the rainy season beginning in about July and prob-
ably lasting for more than one month, perhaps as
much as two months. Immature fruits have been
collected in March.
Philodendron gigas is endemic to the Canal
Area, where it is known only along the El Llano—
Cartí road in both Panamá Province and in the
Comarca de San Blas at 300 to 375 m in Premon-
tane wet forest and Tropical wet forest.
Philodendron gigas is a member of Р sect.
Philodendron subsect. Philodendron ser. Velutina.
This species is distinguished by its appressed-
climbing habit; short, thick internodes (6-10 cm
diam.); sharply D-shaped, semi-persistent cata-
phylls; subterete petioles; thick, scaly roots; and
especially by its huge, ovate, velvety, dark green
leaf blades. It is found growing high on trees, to
about 20 m.
Philodendron gigas is apparently most closely
related to P. andreanum Devansaye from Colombia.
That species shares with P. gigas dark green, vel-
vety leaf blades, but differs in having more elongate
ovate-triangular blades with the lobes much longer
than broad (vs. rounded and about as long as broad
on P. gigas). Philodendron gigas has blades about
1.6-2.5 times longer than broad, vs. 2.6–3 times
for P. andreanum. In addition, P. andreanum is de-
scribed as having the spathe tube green outside and
becoming whitish within. In contrast, P. gigas has
the post-anthesis spathe tube purplish both inside
and outside.
Philodendron gigas is not easily confused with
any of the other velvety-bladed species of Philo-
dendron from Central America. Philodendron ver-
rucosum L. Mathieu ex Schott and P. squamipetiol-
atum Croat have velvety blades, but both have
scaly petioles. In South America, two other species
with ris ovate leaf blades, P. gloriosum André
and P. mamei André, differ in being terrestrial with
repent stems. gos are probably restricted to the
eastern slopes of the Andes.
Additional specimens examined. PANAMA. Panamá:
(B, F, L, MEXU, MO, PMA, QCA, US);
375 m, nen isi. (F, MO); Mile 10, poe 383714 (F,
MO). San Blas: El Llano-Cartí road, Mile 14, 300 m,
9°15'N, AU Croat 69242 (CM, MO).
Philodendron glanduliferum Matuda, Bol. Soc.
Bot. México 27: 47. 1962. TYPE: Mexico. Oa-
хаса: Sierra de Juárez, along Hwy. 175, 1900
m, 15 Sep. 1961, Matuda 37247 (holotype,
MEXU). Figures 180, 200-202.
Terrestrial; stem to 1 m long; internodes short,
to 8 cm long, to 2 cm diam., about as long as broad
448
Annals of the
Missouri Botanical Garden
or sometimes broader than long, dense; roots drying
2-3 mm diam., dark reddish brown, folded into ir-
regular longitudinal ribs; cataphylls fleshy, sharply
2-ribbed, reddish, drying pale yellowish brown to
brown, persisting semi-intact as a reticulum of fi-
bers which are persistent; petioles 44-51 cm long,
3-6 mm diam., subterete, obtusely flattened near
base adaxially, often purple at apex, surface incon-
spicuously short-lineate to striate, prominently so
toward apex, with moderately dense, hair-like
scales, at least near apex, scales many times longer
than wide; geniculum darker than petiole; blades
ovate-cordate, subcoriaceous, bicolorous, abruptly
acuminate at apex (the acumen sometimes apicu-
late, 2—4 mm long), cordate at base, 34-47 cm long,
1.2-1.4 cm wide (1.2-1.4 times longer than wide),
(0.6–0.8 times the petiole length), upper surface
semiglossy, lower surface glossy, paler; anterior
lobe 24-36 cm long, 11-28(38) cm wide (0.9-1
times longer than wide), (1.9-2.7 times longer than
posterior lobes); posterior lobes 11-14.5 cm long,
broadly rounded to obtuse; sinus spathulate or
V-shaped; midrib flat to convex, paler than surface
above; basal veins 5-7 per side, with 1-2 free to
base, 2-3 coalesced (0.7)1-2.2 cm; posterior rib
not naked, to 1.5 cm; primary lateral veins 2—4 per
side, departing midrib at a 50-65” angle, sunken,
raised below; minor veins arising from both the
midrib and primary lateral veins; tertiary veins
moderately distinct, darker than surface below. IN-
FLORESCENCES 1-2 per axil; peduncle 9 cm
long, 6 mm diam.; spathe smooth, 12-13 cm long
(1.5-2.6 times longer than peduncle); spathe blade
pinkish white outside; spathe tube magenta outside;
spadix to 12 cm long, remaining in the spathe at
anthesis; pistillate portion to 2 cm long, 10 mm
diam.; staminate portion to 10 cm long; fertile sta-
minate portion gradually tapered to apex, 10 mm
diam. at constriction, scarcely constricted above the
sterile staminate portion, narrowly rounded at apex,
11 mm diam. near the middle, 10 mm diam. at
constriction; pistils 2 mm long, 1.9 mm diam.; ova-
ry (5)6(7)-locular, locules 1.3 mm long, 0.6 mm
diam., with axile placentation; ovules ca. 10 per
locule, 2-seriate, 0.5 mm long, longer than funicle;
funicle 0.3 mm long, adnate to lower part of par-
tition; style similar to style type D; style apex some-
what rounded to flat; the androecium truncate, mar-
gins irregularly 4—5-sided, ca. 0.9 mm long.
JUVENILE plants with lower blade surface pur-
plish.
Flowering in Philodendron glanduliferum occurs
during the rainy season, in June and September
(Moore & Bunting 8889). Fruiting collections have
not been seen.
Philodendron glanduliferum occurs in Mexico,
Guatemala, and Venezuela, with P. glanduliferum
subsp. glanduliferum endemic to Mexico and Gua-
temala. In Mexico, it is known only from the Sierra
de Juárez, at 580 to 1900 m elevation in “Bosque
mesófilo.” It has been collected recently in Gua-
temala in the Sierra de las Minas in the province
of Zacapa. Philodendron glanduliferum subsp.
camiloanum Croat is endemic to Venezuela, where
it is known only from the states of Táchira and
Apure at 250 to 1200 m elevation in Tropical wet
forest and Premontane wet forest life zones.
species is characterized by its terrestrial habit;
short internodes; sharply two-ribbed cataphylls per-
sisting semi-intact as a reticulum of pale yellowish
brown fibers; subterete petioles with an area о
moderately dense hair-like glands at least near the
apex; and ovate-cordate blades (about three-fourths
as long as petioles) with a spathulate or V-shaped
sinus (often overlapping in P. glanduliferum subsp.
camiloanum).
Philodendron glanduliferum subsp. glandulifer-
um is not easily confused with any other species in
Central America. Philodendron glanduliferum
subsp. camiloanum, endemic to Venezuela, is dis-
tinguished from the typical subspecies by having
sharply 2-ribbed cataphylls; petioles glandular
more or less throughout; and blades with the sinus
closed or nearly so with the margins of the lobes
overlapping (at least slightly). In contrast, P. glan-
duliferum subsp. glanduliferum has unribbed or
only weakly 2-ribbed cataphylls, petioles glandular
only near the apex, and blades with the sinus
ly larger leaves with the petioles merely warty-ver-
rucose (never glandular) near the apex.
Additional specimens examined. GUATEMALA. d
bal: Sierra de las Minas, 700 m, Forther s.n. (M). =
ICO. Oaxaea: Ixtlán, S side of Río Soyolapan, 1230-12
m, 1734'40"N, 96°20'W, Boyle et al. 2583 (CM, ^
Comaltepec, Highway 185, trail from Puerto Ann
Cuaje, 1280-1725 m, 17?39'50"N, 96^18'05"W, Be f
Coatlan-Mazatlan, Cerro
(MEXU); 409 (MEXU); Tuxtepec-Oaxaca, 55 km S of Tux
A SP |
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
tepec, 630 m, 17%37'N, 96°20'W, Hammel & Merello
15481 (MO); Tenet, Tuxtepec—Oaxaca, 4 km SE of Ме-
tates, Torres et al. 7811 (MO).
Philodendron grandipes K. Krause, in Engl. &
K. Krause, Das Pflanzenr. IV. 23Db (Heft 60):
48. 1913. TYPE: Panama. Colón: Río Fató
ape at Dos Bocas (E of Nombre de Dios),
0-80 m, са. 9?35'N, 79?28'W, Pittier 4228
ва US). Figures 203, 204, 206, 216.
Philodendron pleistoneurum Запа. € L. O. Williams,
eiba 3. 109. 1952. TYPE: Costa Rica. Puntarenas:
Esquinas Forest, 60 m, 27 Mar. 1951, Allen 6036
B Lope, EAP; Ур ва GH, US).
Terrestrial; stem creeping over soil, 20-100 cm
long, leaf scars obscured by cataphylls; internodes
short, 14 cm long, 2.8 cm diam., usually broader
than long, medium green, semiglossy, coarsely
white-streaked at apex; roots + smooth, slender,
elongate, 2-4 mm diam., few per node, descending;
cataphylls moderately coriaceous, 16-22 cm long,
sharply 2-ribbed, green to pale green or reddish or
pinkish, drying brown to tan, persisting semi-intact
or as fibers at lower nodes, acuminate at apex.
LEAVES arching to pendent; petioles 25.5-73 cm
long, 10-12 mm diam., D-shaped, spreading, pale
to medium green to reddish at base, with a medial
rib adaxially, rounded abaxially, with adaxial mar-
gins erect, surface finely and weakly striate; sheath-
ing 3.5-5.5 cm long; geniculum slightly thicker
than petiole when apparent, 1.4—1.7 cm long, paler
than petiole; blades broadly ovate-cordate, subcor-
laceous, moderately bicolorous, acute to abruptly
acuminate at apex (the acumen apiculate and
downturned), cordate at base, 20-50 cm long,
15.5-36 cm wide (0.9-1.7 times longer than wide),
(0.5-0.9 times the petiole length), broadest at or
near the middle, margins broadly undulate, upper
surface dark green, semiglossy to glossy, sometimes
matte or subvelvety, drying brown to greenish
brown, lower surface semiglossy, moderately paler,
drying brown to greenish brown; anterior lobe 16–
37 cm long, 15-38 cm wide (1.8-3.7 times longer
than posterior lobes), broadest slightly above or at
point of petiole attachment; posterior lobes 5.5—
15.5 cm long, 7.4-18.6 cm wide, directed down-
ward, MU held up somewhat at an angle from
the midrib, rounded to obtuse; sinus parabolic to
сетери ко. sometimes closed with lobes over-
lapping before being pressed; midrib concolorous,
at at base, becoming weakly sunken toward apex
above, thicker than broad at base, becoming con-
vexly raised, concolorous below; basal veins 8-10
per side, 1(2) free to base, several remaining coa-
lesced 0. 5—2(4) cm, prominently sunken above,
convexly raised below; posterior rib usually not na-
ked, sometimes briefly so at base; primary lateral
veins (8-10)11-17 per side, departing midrib at a
-30° angle, spreading to 50—70° angle, +
straight to the margins, quilted-sunken and con-
colorous above, prominently convex, matte, and
darker than surface below; interprimary veins about
as conspicuous as primary lateral veins above, flat
and darker than surface below; minor veins arising
from both the midrib and primary lateral veins; ter-
tiary veins distinct, weakly raised above, darker
than surface below. INFLORESCENCES erect, 2—
4 per axil; peduncle 2.5-14 cm long, 5-8 mm
diam., terete, green to reddish, white-lineate;
spathe 6.6-11.8 cm long (0.6-2.5 times longer
than peduncle), constricted + midway; spathe
blade lanceolate, pinkish red, tinged green, pale
green to white or green, short white lineate outside,
4–6 cm long (opening broadly elliptic in face view),
pinkish red to white, pale greenish white or pale
green inside; spathe tube oblong-ellipsoid, dark
reddish maroon to reddish purple to green, weakly
short dark lineate outside, 2.54 cm long, 1.4-3.8
cm diam., pale green to greenish white to pinkish
red inside; spadix sessile or very short stipitate;
tapered, 6.6–11.8 cm long, broadest near the base;
pistillate portion white, yellowish white (post-an-
thesis), slightly tapered toward the apex and base,
(1.6)2–2.7 mm long, 1 mm diam. at apex, 1–
1.5 cm diam. at middle, (6)10—11 mm wide at base;
staminate portion 3.5—7.1 cm long; fertile staminate
portion white, cylindrical to tapered, 7-10 mm
diam. throughout, 1-1.5 cm diam. at base, 7-10
mm diam. at middle, 4—6 mm diam. ca. 1 cm from
apex, narrower than the pistillate portion, as broad
as the sterile portion; sterile staminate portion
about as the pistillate portion, white, 1–1.5 cm
diam.; pistils 1.8-2.3 mm long, 1.1-1.5 mm diam.;
ovary (4)5—6-locular, 1.3-1.5 mm diam., locules 1—
1.5 mm long, 0.4—0.6 mm diam., with axile placen-
tation; ovules (7-10)16-22 per locule, 2-seriate,
0.2–0.4 mm long, longer than funicle, style similar
to style type B; style apex + concave, sometimes
weakly lobed; stigma subdiscoid, brushlike, lobed
or unlobed, 0.9-1.4 mm diam., 0.2-0.5 mm high,
covering center of style apex, depressed medially;
e androecium truncate, margins irregularly 3—6-
sided; м oblong, 0.2-0.3 mm wide; sterile sta-
minate flowers blunt, margins irregularly elongate,
13-18 mm long, 9-13 mm wide. Berries creamy
white, obovoid, apex truncate; seeds 20 per locule,
1.9 mm long, 0.2 mm diam., with prominent stria-
tions running from funicle to apex and slightly spi-
raling, perpendicular to larger veins. JUVENILE
ct
di
Annals of the
Missouri Botanical Garden
blades narrowly elliptic to ovate, acute to weakly
cordate at base
Flowering in Philodendron grandipes probably
occurs throughout most of the year (Febru
through November, except October) but primarily
i Post-anthesis collections have
fruiting collections are known from throughout the
year (except February, March, and June).
Philodendron grandipes ranges from Nicaragua
(Zelaya) to Panama, Colombia, and Ecuador (Es-
meraldas), from near sea level to mostly less than
750 m (sometimes to 1200 m) elevation in Tropical
moist forest, Premontane wet forest, and Tropical wet
forest life zone
Philodendron grandipes is a member of P. sect.
Philodendron od Philodendron ser. Fibrosa.
This species is characterized by its terrestrial habit,
short internodes, persistent cataphyll fibers, broad-
ly ovate-cordate leaf blades, usually green spathes,
and, especially, by its D-shaped petioles with erect
margins and a medial adaxial rib.
Philodendron grandipes is closest to P. jodavi-
sianum, which has a similarly shaped petiole and
other general features in common. The latter spe-
cies differs in being an appressed-climbing hemi-
epiphyte with ovate-triangular (rather than broadly
ovate) blades.
This species frequents stream banks in central
Panama and is one of the few consistently terres-
trial Philodendron species in Central America.
Habitat in Costa Rica, however, is variable. Philo-
dendron grandipes occurs along stream banks on
the Osa Peninsula, but at La Selva (Heredia) this
species is widely scattered in the forest understory
(M. Grayum, pers. comm.).
Spathe tube color in this species is variable to
some extent geographically. For example, on the
Atlantic slope of Costa Rica the spathe tube is usu-
ally greenish, whereas on the Pacific slope it is gen-
erally reddish on both surfaces.
Additional specimens examined. soda RICA. Ne:
juela: 36-37 km NW of San Ram 500-515
10°15'N, 84°34'W, Croat 68198 (MO); "x "eiue Upa
1 mez 18639 (MO, PMA). Hered
R, ر
cienda Tapezco—Hacienda La Suerte, 29 air km W of Tor-
ero, 40 m, 10°30’N, 83°47'W, Davidson & Donahue
8741 (MO, RSA); Turrialba-Limón, along Hwy. 32, ca. 11
mi. S of Siquirres, 650 m, Croat 43330 (MO); Río Chir-
W, Grayum « vs 3525 (MO); Que-
reri Cafiabral-Río Barbilla, 200—400 m, 10?02'N,
6' W, Grayum et al. 8739 b E MO): Hitay Cox
ere da 140 m, 942'N 2'W. Hi
14349 (МО); Parque Natal: a Lomas de Sin
i 00 m, 10?24'N, 83?33' W, Robles et
of Guápiles, Quebrada Danta, 360 m, 10°12'N, 83°49' W,
Croat 68419 (MO); Río Catarata, 50-100 m, 937'N,
82°49'W, Burger & Antonio 10888 (CR, F, MO, PMA, U);
i Colorado, 16 airline km SW of Barra del а
10-120 m, 10%39'N, 83%40'40"W, exei & Herr
31214 (CR, K, MO); Río Sixaola, ca. 3 m
m
rí, 50 m, Croat 43299 (CR, МО);
Росен, Вагта де! донй. Llanura de Tortuguero Sardi-
nas, 15-20 m, 10°38'38"N, 83°44'10"W, Araya 596 (INB).
Puntarenas: Osa Peninsula, Piedras Blancas-Rincón,
3.7 mi. W of Pan-American vs deua ma m, 8
83°18'W, Croat 67651 (K, MO); 2.5 mi. SW of Rin
8°42'N, 83°29’ W, Kennedy 1622 (MO); Ghd 76751 мој
Rincón de Osa, 250-540 m, 8°42'N, 83° Croat &
Grayum 59838 (MO); SW of pinsa a Osa, 40-200 m,
5 (CR, M, MO); Fila
guel, Croat 26519 (MO);
ma а, 1 mi. aa ы Сайаз Gordas, 1150 т, Croat
22262 (МО); АЏеп 6036 (Е, СН); Кови е vr km 183,
ca. 400 m from Santa M NOR Coons
vado National Park, Sirena, 0.200. °36'W,
Liesner 2903 (MO); 5-25 m, 8”29'N, 83°34’ Ху, phan
& Kernan 16662 (CR, MO); 150 m, Kernan & Phillips
512 en MO); Rancho Quemado, 250-3 e m, Marín 39
(CR, INB, МО); vic. Земен а, 50 т, Croat & Hannon
79244 (М 0); Las Cruces Tropical Botanical pese з 1200
m, 8°49'N, 82%58'W, "Oh oat 57234 (CR, MO); Rio Jaba,
Las Cruces, 1200 m, Meerow et al. 2017 (SEL). San José:
San Isidro del General-Dominical, SW of San Isidro, 4. 8
сакай 500 т, Kr
tectora La Cangreja, ca. 2 km N
iscal, m, 942'N, 84°22'W, Grayum 8644 (
Parque Nacional Braulio Carillo, above Río Sucio, 5-600
m, Pennington 11533 (K); 10%24'04'N,
85?03'03"W, Carballo 87 (CR, MO). NICARAGUA. Le-
laya: 150-180 m, 11%43'N, 84°18’ W, Stevens 4965 (MO);
Rio Rama, at Salto ине 15-25 11:576
84°17'W, Stevens 8960 (MO). PANAMA. Bocas de
Toro: Chiriquí Grande-Fo 13.2 mi. W of Chiriquí
of Divide, 700 m, 8°45’N, 8215'W, Croat 60264 >
MO); 450 m, 845 N, 82%15'W, McPherson 7369 (MO);
mi. N of Continental Divide, 650 m, 8247'М, 8271 í
Churchill & Churchill 6209 (MO); 6210 (MO); Chiriqu
Lagoon, von , MO). Canal Area: US):
boa, Standley 28401 (US); «Frijoles; Standley 27471 (
e |
eee e ———— —— ——
Volume 84, Number 3
1997
Croat 451
Philodendron Subgenus Philodendron
Barro Colorado Island, Standley 31361 (US); 40888 (US);
Croat 16574 (MO); 12300 (MO, SCZ); 11886 (MO); 11194
(MO); 11077 (MO); 9526 (MO); 6512 (MO); 5117 (MO);
е 849 (MO); Summit Gardens, Croat 11491 (MO).
qui: Chiriquí Grande—Fortuna, 7.7 mi. W of Chiriquí
угине 80 m, 8°50'N, 82710", Grial & Grayum 60114
(MO); Burica Peninsula, San Bartolo Limite, 12 mi. W of
Puerto Armuelles, 400-500 m, Croat 22188 (MO). Coclé:
La Junta-Limón, 5 hours walk N of Alto Calvario, 800—
1000 m, Folsom 5861 (MO); Coclesito—Llano Grande, 200
m, 847'N, 80°28'W, Churchill et al. 4170 (MO); El Valle
region, 800-900 m, 8*36'N, 80*07'W, Croat & Zhu 76664
(MO, PMA); 67212 (F, MO); 800 m, 25406 (F, MO); ca.
1000 m, Gentry 5662 (F, MO); Croat 14391 (MO); 860 m,
8°37'N, 80°08'W, Croat & Zhu 76710A (MO); 860-900
m, Croat 37398 (MO, RSA); Cerro Pilón, Duke & Dwyer
13977 (MO). Colón: Portobelo-Nombre de Dios, 6-8 km
from Peluca Hydrographic Station, Kennedy & Dressler
3331 (US); о ی Cascajal, vic. of Nuevo Tonosf,
Croat 33648 (K, MO, US); 6 mi. S of Portobelo, Croat
11401 (MO); ы. de Dios, 1.2 mi. beyond
junction of road to Isla Grande, پا и 79°35'W, Croat
o B ms trail, 700
4 (MO); Río
Couche; 0.5-1 km upstream from Pis РА РЈ
s 6 km S of Portobelo, 5-30 m, Nee 7153 (MO, US);
a. 3—5 mi. inland, 10—100 m, Croat 26205 (MO); eni
100 m, 79345 (PMA, МО); 1.5 mi. — ca. 10
Kennedy & Dressler 1513 (SEL) «100 m, 9°27’ N
79°40'W, Croat & Zhu 76245 (MO); d Miguel de la
хе vic. of Guásimo, Croat 9940 (MO). Darién: iem
irre region, vic. Сапа gold mine, 500—600 m, Croa
37637 (MO); 480 m, 37957 (МО); 17 km N of El Real,
trail from base camp along Río Perisenico, 100 m, 801'N,
77447, Croat & Zhu 77177 (MO); Río Cocalito, ше
tefoord & Eddy 132 (BM); Rfo Tuquesa, са. 2 km
from Continental пе vic. of Tyler Kittredge gold ain
Croat 27191 (MO). P а: El Llano-Cartí Road, 9.6
km from Pan Анса T Highway, 410 m, Mori & Kallunki
1835 (MO, PMA); Km 12.4, 300-400
pa
—
ч
Hoover 1313 (MO); Mile 56, 350-375 т, Croat 34802
(МО); vic. of Gorgas Lao Mosquito Control Project Site at
km 12, 26058 (MO); Serranía de Маје, S of Трен, 500-
650 m, Huft et al. 1692 (MO); Уе de Cañazas, Кап-
cho Chorro, above Tortí iue 400—700 m, Folsom 6748
(MO); Cerro Campana, ca. i mi. from Inter-American
Highway, ca. 150 m, old 35969 (MO, RSA). San Blas:
El Llano-Cartí Road, trail along Continental Divide, 400
m, 9°20'N, 78°56'W, McDonagh et al. 323 (BM); SE of
em Obaldía, Croat 16762 (MO); 14 mi. N of Pan-
American Highway, 300 m, 9?15'N, 79°W, Croat 69249
Lis Miria Ubidandup Island, Digole, 0-20 m, 9°26'N,
8°54'W, Herrera 292 (MO); Río Playón Chico, vic. NEBA
поћи 100—450 m, 9°14.5'N, 78715", Herrera et al.
1399 (MO, PMA); 1 mi. S of Nusagandi, Mile 9, 350 m,
9°20'N, 79°W, Croat & Zhu 76997 (MO, PMA); Nusagan-
di, 275-300 m, 920'N, 79°W, Croat 76594 (MEXU, MO).
Veraguas: Santa Fe region, Cerro Tute, 1 1050-1150 m,
әш 48907 (МО); Езсие1а Agricola Alto de
à, Río Dos Bocas Valley, N of Santa Fe,
Croat 27398 (MO); 450 m, 27550 Mo); Escuela vtm
to Piedra-Río Dos Bocas, ca. 10 k m the school,
20 m, Croat 25903 (MO).
Philodendron eS Croat, sp. nov. TYPE:
anama. Darién: Rio Pirre, 14 July 1971,
Croat & 2 rter 15543 (holotype, MO-
2059944; isotype, PMA). Figures 207, 208.
шу: док афа internodia 3.54 cm longa, ca
m diam aphylla acute 2-costata, 15 cm ien: de
um. da 89. cm pe s, 7-8 mm diam., conv
adaxialiter cum 2 costis tco adi bns v Panther
lanceolata, debiliter cordata basi, 19-28 cm longa,
cm lata; nervis primariis lateralibus 5—6 utroque; inflores-
centia 1; RSUN ulus 105 ст m longus; spatha usque 15. 7
m longa, viridis;
extus suffuso intus rubro; pistilla 3-5- locularia; loc-
uli 1-ovulati.
Hemiepiphytic vine; internodes semiglossy, 3.5—
cm long, ca. 1 cm diam., longer than broad, dry-
ing light brown, epidermis vmi deeply
ridged, finely striate on magnification and densely
granular to almost scabrous (but not harsh to the
touch), the vestiture raised to ca. 1.5-2 times lon-
ger than wide; roots drying dark brown, <15 cm
long, 1 mm diam., few per node; cataphylls 15 cm
long, sharply 2-ribbed (ribs low), green, deciduous.
LEAVES with petioles 8-9 cm long, 7-8 mm
diam., convex with acute margins adaxially, drying
dark brown and somewhat sulcate adaxially with
distinct acute marginal rib; sheath short, incon-
spicuous, 1-2 cm long, for «0.25 its petiole length;
blades oblong-oblanceolate, acuminate at apex,
weakly cordate at base, 19-28 cm long, 4—6 cm
wide (3.8—4.7 times longer than wide), upper sur-
face medium green, weakly glossy, drying dark
brown, lower surface slightly paler, drying yellow-
ish brown; midrib flat, concolorous above, convex,
slightly paler than surface below; basal veins 1-2
per side, with 1-2 pairs free to base, indistinct;
primary lateral veins 5-6 per side, weakly sunken
above, convex, slightly paler than surface, drying
paler below; minor veins moderately obscure, bare-
ly visible on drying, the larger weakly undulate,
arising from the midrib only, secretory canals ap-
pearing on magnification as short, pale raphide
cells along the minor veins. INFLORESCENCES
(post-anthesis), 1 per axil; peduncle 10.5 cm long,
cm diam.; spathe to 15.7 cm long, to 7 cm wide
when flat, weakly constricted above the tube, green
throughout; spathe blade green inside with promi-
nent striations from lower 1.5 cm to 9 cm up from
base; spathe tube tinged red outside, red inside;
fertile staminate portion to 5.5 cm long, 9 mm
diam.; spadix with pistils 1.5-3 cm diam., drying
light brown, sparsely granular on the sides; ovary
3-5-locular; ovules with 1 per locule, style similar
to style type E; style funnel 0.5-0.7 mm, distinctly
raised above the surface. INFRUCTESCENCE 10
452
Annals of the
Missouri Botanical Garden
cm long, 2.7 cm diam. in lower У, prominently ta-
pered toward apex to 1.3 cm diam. at apex; berries
whitish; seeds oblong-elliptic, tan, smooth, 1.3-1.9
mm long.
Flowering in Philodendron granulare probably
occurs in the early wet season. Fruiting is recorded
by a single collection made in July.
Philodendron granulare is known only from the
Panamanian type specimen, from Darién Province
near El Real along the Río Pirre at less than 75 m
elevation in Premontane wet forest.
Philodendron granulare is a member of P. sect.
Calostigma subsect. Glossophyllum ser. Glossophyl-
lum. This species is distinguished by its scandent
habit, more or less oblong, short-petiolate blades
with subcordate bases, and especially by its dense-
ly granular dried stem (hence the name) and dis-
tinct funnel-shaped styles.
Philodendron granulare is most easily confused
with P. bakeri, which it resembles in a superficial
way. The latter species differs in having totally
smooth stems, petioles with conspicuous sheaths of-
ten extending to the middle or beyond, leaf blades
with more conspicuous minor veins, and pistils with
a truncate, type B style which lacks a funnel, hav-
ing instead the stylar canals emerging directly onto
the flat truncate apex.
Philodendron grayumii Croat, sp. nov. TYPE:
anama. Bocas del Toro: Fortuna Lake area,
Fortuna—Chiriquí Grande, 0.3 km N of Conti-
пета]! Divide, 970 m, 843'N, 82%17'W, 27
June 1994, Croat & Zhu 76524 (holotype,
MO-4619417-20; isotypes, AAU, B, CAS,
CM, CR, COL, DUKE, F, GH, K, M, MEXU,
NY, PMA, S, SCZ, SEL, TEX, US). Figures
209-215.
Planta hemiepiphytica; internodia 1-10 ст longa,
(2.5)3.5—7 cm diam.; cataphyllum 11-26 ст longa, incos-
tata vel leniter 1-costatata, demum decidua aut acute com-
planta vel 2-costata; petiolus 40-97 cm longus, subteres,
obtuse planatus prope apicem; lamina ovata vel late ovate,
(3150-73 cm longa, 24—44(58) cm lata; cordata basi; ner-
vis basalibus 4-5(6) utroque; nervis lateralibus 1 4-6 ut-
roque; inflorescentia 1-5 per axillam; pedunculus (24—6
cm longus; spatha (11)13-16(18) cm longa, extus pallide
viridis vel virens, albida vel pallide flava vs. apicem, min-
imum interdum maculata purpurea-violacea, tubo pathae
rubro intus; pistilla 7-8-locularia; loculi
(2)3—4(5)-ovulati.
Hemiepiphytic, appressed-climbing; internodes
semiglossy to matte, 1-10 cm long, (2.5)3.5—7 cm
diam., dark green to gray-green or tannish gray,
drying brown to yellow-brown or reddish brown,
epidermis closely ridged or fissured, sometimes
peeling; roots usually 1-2 per node, ca. 3-4 mm
diam., drying dark reddish brown; cataphylls thick,
somewhat spongy, 11-26 cm long, unribbed to
weakly l-ribbed near apex or sharply flattened to
sharply 2-ribbed (ribs close), green, sometimes
weakly maroon-spotted, often persisting intact for a
short time then deciduous, intact; petioles 40—97
cm long, 1–1.5 cm diam., subterete, firm, dark to
medium green, obtusely and often weakly flattened
near apex, sometimes weakly ribbed near apex
adaxially, surface semiglossy, weakly and densely
short pale green lineate, drying yellowish brown to
dark brown, smooth to finely striate, sheath 6-15
cm long; blades ovate to broadly ovate, coriaceous
to subcoriaceous, moderately bicolorous, semiglos-
sy, acute to gradually or abruptly acuminate at
apex, deeply cordate at base, (31)50—73 cm long,
24—44(58) cm wide ((0.7)1—1.9 times longer than
wide), three-fourths as long as to somewhat longer
than the petiole, upper surface dark green, drying
dark brown to yellow-brown or grayish brown, se-
miglossy, lower surface conspicuously paler, drying
yellow-brown, weakly to semiglossy; anterior lobe
(25)31—49(60) cm long, (13)24—32(42) cm wide
(2.3—4.6(8) times longer than posterior lobes), mar-
gins broadly rounded, sometimes sinuate; posterior
lobes rounded to narrowly rounded, sometimes
overlapping, (6)9-13(17) ст long, 12-18(22) cm
wide; sinus hippocrepiform to oblong spathulate,
obovate, or sometimes triangular to parabolic,
rounded to narrowly rounded, (8)10.5-13 cm deep;
midrib flat to broadly raised above, paler than sur-
face above, convex to obtusely acute and sometimes
purplish spotted below; basal veins 4—5(6) per side,
with 1–2 free to base, third and higher order veins
coalesced 3—6(8) cm long; posterior rib usually not
naked, rarely naked for cm; primary lateral
veins 4-6 per side, departing midrib at а (45)55-
70(75°) angle, downturned and splayed ош when
joining midrib, obtusely to narrowly sunken and
paler than surface, sometimes weakly raised, drying
weakly ridged near the midrib above, convex an
paler than surface, sometimes reddish below; mnor
veins moderately indistinct, drying weakly proe
nulous (surface often with minute purplish raised
areas seen on high magnification), arising from both
the midrib and primary lateral veins, secretory
ducts appearing as intermittent lines (less conspic-
uous on highland forms); INFLORESCENCES 1-5
per axil; peduncle (2)4—6 cm long, 1-1.5 cm diam.,
pale green, finely short-lineate; spathe (1 1)13-
16(18) cm long, 2-3 cm diam. (2.1-2.6(3.5) times
longer than peduncle), weakly constricted + above
the middle, pale to medium green, sometimes whit-
ish to pale yellowish toward apex, often tinged or
|
|
Volume 84, Number 3
1997
Croat 453
Philodendron Subgenus Philodendron
spotted violet-purple throughout, especially near
base, sometimes green dorsally with only the front
edges of tube maroon, margins white, outer surface
drying reddish brown throughout inside, violet-pur-
ple to maroon in lower one-half to two-thirds, es-
pecially in lower half; spathe tube 7-8.5 cm long,
2–3.6 cm diam., maroon to red, with раје lineations
extending onto blade area inside; spadix sessile;
cylindrical, 9.5-16 cm long, broadest above the
middle or below the middle; pistillate portion pale
green, cylindrical to weakly tapered toward the
apex, 3.3-3.5 cm long, 4.6 cm long in front, 3.2-
3.5 cm long in back, 9-20 mm diam. throughout;
staminate portion 6.3-12.5 cm long; fertile stami-
nate portion creamy white, cylindrical, 1-1.4 cm
diam. at base, 1-1.4 cm diam. at middle, 8-10 mm
diam. ca. 1 cm from apex, broadest at the base or
at the base broader than or as broad as the pistillate
portion, sterile staminate portion usually not de-
tectable; pistils 1.4-1.9 mm long, 0.8-1.1 mm
diam.; ovary 7—8-locular, 1-1.4 mm long, 0.8-1
mm diam., walls thin and membranous, locules 1-
1.3 mm e 0.3-0.4 mm diam., ovule sac some-
times present, to 0.8 mm long, with sub-basal pla-
centation; cun (2)3-4(5) per Іосше, 2-seriate,
contained within transparent, gelatinous ovule sac,
0.3-0.5 mm long, usually as long as funicle; funicle
0.2-0.3 mm long (can be pulled free to base), style
0.5 mm long, 1.2 mm diam., similar to style type
B; style apex sloping; stigma brushlike, sometimes
cupullate, subdiscoid, unlobed, sometimes truncate
or weakly domed, 0.9-1.3 mm diam., 0.2-0.6 mm
high, covering entire style apex; the androecium
truncate, prismatic, oblong, margins irregularly 4—
6-sided, 0.9 mm long, 1.1 mm diam. at apex; thecae
cylindrical to weakly elliptical, 0.3-0.4 mm wide,
= parallel to one another; sterile staminate flowers
trapezoidal to 4—6-sided, 1.1 mm long, 1.3 mm
wide. INFRUCTESCENCE with peduncle to 11.5
ст long; spathe to 26 cm long, pistillate spadix to
13.5 ст long, 2.5 cm diam.; staminate spadix to
13.5 cm long; berries pale ochraceous, 3-3.3 mm
long; seeds 1.9-2 mm long, 0.7-1 mm diam. PRE-
ADULT petioles sheathing throughout; blades ob-
long, subcordate, 26-31 cm long, 9-13 cm wide.
Flowering in Philodendron grayumii apparently
occurs primarily in the rainy season (March as well
as June and September), but also in the late dry
season. Post-anthesis inflorescences have been col-
lected in March and June through August and No-
vember. Immature fruits are known from June and
November.
Philodendron grayumii ranges from Costa Rica
to central Panama from sea level to 1630 m ele-
vation in Premontane wet forest, Tropical wet forest,
and Premontane rain forest life zones, as well as
Premontane wet forest transition to Tropical moist
St.
Philodendron grayumii is a member of P. sect.
Calostigma subsect. Macrobelium ser. Macrobel-
ium. This species is characterized by из yellow-
brown-drying stems; cataphylls deciduous intact;
more or less ovate blades drying yellow-brown with
the basal veins scarcely or not at all united into
posterior ribs and rarely naked on the sinus; and
primary lateral leaf veins splayed out, somewhat
ridged, and downturned near the midrib. The
spathe is often tinged reddish to purplish outside
and dark red to maroon in the tube within.
This species is polymorphic, especially in the
nature of the resin canals in the leaves and the
number of inflorescences per axil. Collections from
near sea level in Limón Province of Costa Rica
(Grayum 8467, 8469) are unusual in lacking any
clearly visible secretory ducts (normally easily vis-
ible at least on the dried lower leaf surfaces).
Philodendron grayumii may be confused with P.
edenudatum, and the two may prove to be conspe-
cific. Philodendron edenudatum differs in having
proportionately longer leaf blades (1.8-2.1 times
longer than broad) and proportionately shorter pet-
ioles (0.77—0.89 times as long as the blades).
Philodendron grayumii may also be confused
with P. ferrugineum. The latter, occurring princi-
pally in the Canal Area, shares with P. grayumii
brownish-drying blades with the basal veins usually
not at all or only weakly naked. Philodendron fer-
rugineum differs in having generally shorter inter-
nodes; larger cataphylls (26-48 vs. 11-26 cm long);
typically much larger, more coriaceous blades
drying reddish brown (rather than yellowish brown);
and minor veins usually drying markedly longitu-
dinally raised but interrupted, giving the leaf sur-
face a markedly bumpy, irregular, and uneven
appearance.
Also perhaps confused with P grayumii is P.
dodsonii, which has dried leaf blades of a similar
yellowish brown color. The latter species differs in
having conspicuously naked posterior ribs, and
blades drying matte on the upper surface and lack-
ing any obvious secretory ducts on the lower sur-
ace.
Perhaps also confused with P. grayumii is P. aro-
maticum, which has similarly cordate blades that
may dry a similar color and also has deciduous
cataphylls. The latter species is distinguished by
being a shorter-stemmed canopy epiphyte, and by
having spongy petioles and proportionately more
elongated, usually more gray-drying leaf blades
454
Annals of the
Missouri Botanical Garden
(usually more than 1.7 times longer than wide) with
the naked portion of the posterior ribs much longer
(usually 2.5-5 cm long).
A noteworthy collection is Croat 67525, which
is somewhat intermediate between P. grayumii and
P. edenudatum. It has narrowly ovate leaf blades
about 1.6 times longer than wide, mottled petioles
and lower midribs, and relatively small (possibly
immature) inflorescences (to 13 cm long). In these
characters, the specimen is closest to P. edenuda-
tum, but it has prominent secretory ducts visible
on the lower blade surface, and two inflorescences
r axil, features not known in P. edenudatum. In
аса. the blades dry dark brown. In this latter
5 е collection differs from both P. grayumii
› edenudatum. It perhaps represents a new
species, з рени it is being included with P. gra-
yumii for now.
Another noteworthy collection is Croat & Zhu
76613, the only collection from east of the Canal
Area. It is unusual in having the cataphylls semi-
persistent as fibers, rather than deciduous while
still intact. This may prove to represent a different
taxon.
А sterile collection (Croat 66712) is noteworthy
in having stems that dry smoother, darker reddish
brown, and semiglossy; much glossier petioles than
those of P. grayumii; and blades that dry more co-
riaceous, glossier, and with the minor veins prom-
inulous on both surfaces. It might represent another
new species, but is tentatively included with this
species.
The new species is named in honor of Michael
. Grayum, a colleague and aroid specialist, whose
e ca of Araceae from Costa Rica
Panama have played an important role in this
iis He has collected the only Costa Rican ma-
terial of the species.
Additional specimens examined. COSTA RICA. Li-
món: along road between Puerto Viejo de Talamanca and
Manzanillo, vic. of Punta Cocles, ca. m, 9°38'N,
82°43'W, 5 m, Grayum & Sleeper 4302 Pos МО); 1 0-20
m оз, 82°40-44'W, Стауит 3637 (MO); Ріша,
5 km NW of Puerto Limón along маа to Portete,
5: m, шет 83°03'W, Grayum & Hammel 8467 (M 0),
8469 (MO). iria, Bocas del Toro: Fortuna—Chiri-
quí Grande, 1.6 mi. N of Continental Divide, 770—790 m,
8'45'N, 82°17’ W. pas 76456 (AAU, CM, MEXU, MO,
i. N of Divide
, 82°17'W, Croat 60476 (AAU, CR,
MO, PMA), 1170 m, Croat 66653A (MO); 5.8 mi. N of
Fortuna Lake, m, 8°45'N, 82°18’ W, Croat 66794 (B,
CAS, CM, CR, F, K, L, MEXU, MO, NY, PMA, US,
WIS); along Continental Divide (4.5 mi. N of middle of
bridge over Fortuna Lake), 1170 m 81°17'W,
mi. N
13: W, "died
74933 (MO); 4.2 mi. E of Chiriquí asas ca.
8°55'N, 82?09'W, Croat 66814 (MO, TEX); Station Milla
7.5 on Changuin pe oit FOU dde < m, Croat
ag (B, F, K, MO, NY, PMA, US); Cerro Colorado, 6.5
i. W of Chame, 1630 m, SEN 81°50'W, Croat 69164
(MO): Chiriquí pl Palo веб» Guayabo, km 112
Gordon 344 (PMA). Chiriqui: My aliis Grande,
4 mi. N of bridge over Fortuna Lake, 1200 m, 8°46'N,
82°16'W, Croat 68026 (CAS, CR, DUKE, F, С, MO, QCA,
TEX, W); 1170 m, 8°44’N, 81?17'W, Croat 66712 (CAS,
K, L, MEXU, MO, PMA); Río Hornito, ca. 0.5 km S of
Centro de Científicos, 8°45'N, - њива W, Croat 76420
Р s де Hornito, 1130
UK I
M oclé:
67525 E ENCB, MO, P, VDB);
d 8°39'N, 80736'W, 74840 (CM, CR, M0,
USE Sbi Сов 600—800 m, 8738'N,
чулуп Croat & Zhu 77202 (IBE, M, MO, MY, Wy
Llano Grande-Coclecito, 4.3 mi. N of stream in md
Grande, 330 m, Croat 49235 (CM, MO). Veraguas: v
of Santa Fe, 5 mi. Sane Escuela Agricola Alto de Pid,
670 m, 8733'N, 81?08'W, Croat 66922 (BR,
MO), 66969 (AAU, MO, NY, OOM, PMA).
Philodendron hammelii Croat, sp. nov. TYPE:
Panama. Coclé: Alto Calvario, ca. 6 km N of
El te Pacific i on ridge W of saw-
mill, 11 , 8°39'N, 80°36'W, Ham-
mel - фикер MO-2658909). Figure
217.
Planta terrestris; internodia 1-1.5 ст longa, 8-12 mm
e ; cataphylla 11.5 cm longa, 1-costata, peri ii ut
ticulum 31-32 с
We
=
2
lidis prope apicem; lamina ovato-cordata, 21-22 cm
ga, 13-16 ст lata, in en
infra; inflorescentia 1; pedunculus 4.5 c s, 4m
diam.; im atha 11 cm longa; pistilla Gus loculi
14-ovulat
Terrestrial; internodes 1-1.5 cm long, 8-12 mm
diam., drying yellow-brown, semiglossy, conspicu-
ously ridged; roots drying reddish brown to brown,
slender, numerous per node; cataphylls to 11. 5 ст
long, 1-ribbed, drying yellowish brown, thin,
smooth (not scaly), persisting as a thin, pale retic-
ulum of branched tan fibers; petioles 31-32 cm
long, 2-3 mm diam., subterete, drying weakly
glossy, smooth or at most weakly granular, surface
densely covered with broad pale scales near apex,
the scales closed and overlapping, mostly less than
three times longer than broad; blades ovate-cor
ate, subcoriaceous, shortly acuminate at apex, CO
date at base, 21-22 cm long, 13-16 cm wide
(1.4-1.5 times longer than wide), (0.66–0.67 as
long as petiole), upper surface dark green, subvel-
vety and glistening, drying grayish green, lower a
face much paler, semiglossy, drying yellowis
brown and moderately glossy; anterior lobe 15- 6.7
ст long, 15.3 cm wide (2.2-2.6 times longer than
nn EE LN S + — TE
Volume 84, Number 3
1997
Croat 455
Philodendron Subgenus Philodendron
posterior lobes); posterior lobes 6.5—6.7 cm long, 7
cm wide, rounded; sinus obovate to narrow hippo-
crepiform and closed; midrib drying sunken and
concolorous above, convex, matte below; basal
veins 6 per side, with 2 free to base, third and
higher order veins coalesced 1 cm, sunken above,
convex below; posterior rib not naked; primary lat-
eral veins 3 per side, departing midrib at a 40—45°
angle, quilted-sunken above, round-raised or
raised, darker below; minor veins distinct, arising
from both the midrib and primary lateral veins;
"cross-veins" prominently raised upon drying. IN-
FLORESCENCES probably 1 per axil; peduncle
4.5 cm long, 4 mm diam., + terete, glabrous;
spathe 11 cm long (2.4 times longer than pedun-
cle), green, drying dark reddish brown; spadix ses-
sile, 9.5-10.5 cm long; pistillate portion 3 cm long,
1 cm фат.; staminate portion 6.5—7.5 cm long, 1-
1.5 cm diam.; pistils 1.5 mm long, 1.9 mm diam.;
ovary 6-locular, 1.2 mm diam., locules 1 mm long,
0.3 mm diam., оуше sac 0.1–0.2 mm long, with
axile placentation; ovules ca. 14 per locule, 2-se-
riate, contained within small, cloudy, gelatinous
ovule sac, 0.1–0.2 mm long, longer than funicle;
funicle to 0.1 mm long, adnate to lower part of par-
tition, style similar to style type B; style apex flat;
stigma hemispheroid, lobed, 1 mm diam., 0.3-0.4
mm high, covering entire style apex, depressed
shallowly at center; the androecium truncate, +
prismatic, margins irregularly 4—5-sided, ca. 1 mm
long; sterile staminate flowers irregularly 4—5-sid-
ed, 1.6-2.1 mm long.
Flowering in Philodendron hammelii apparently
occurs in the early wet season, based on a single
collection made in June.
Philodendron hammelii is endemic to Panama,
known only from the type in Coclé Province in Pre-
montane rain forest at 850 to 1100 m elevation.
Philodendron hammelii is a member of P. sect.
Philodendron subsect. Achyropodium. This species
is characterized by its short, slender internodes;
persistent, thin, pale network of branched cataphyll
fibers; petioles longer than the blades and densely
covered with broad, pale scales near the apex; and
ovate-cordate, yellow-green-drying blades.
Philodendron hammelii is superficially most sim-
ilar to P. colombianum R. E. Schult. from the Am-
azon basin. These species have similarly shaped
blades drying yellow-green with prominent “cross-
veins” and persistent, pale cataphyll fibers. Philo-
dendron colombianum differs, however, in having
the blade sinus acute at apex and petioles lacking
scales.
In Panama, this species might be confused with
P. squamipetiolatum, which also has scaly petioles.
It differs in having blades with a more arcuate sinus
and petioles conspicuously long-scaly throughout
most of their len
The species is мина in honor of Barry Hammel
of the Missouri Botanical Garden who has collected
many important Araceae in Panama and Costa
Rica, including the type specimen.
Philodendron hebetatum Croat, sp. nov. TYPE:
Panama. Coclé: vicinity el Valle de Antón, at
forested flat area near Finca Macarenita at La
Mesa, 800 m, 8?36'N, 80°07'W, 6 July 1994,
Croat & Zhu 76693 (holotype, MO-4619514—
15; isotypes, B, CAS, COL, CR, F, K, NY,
PMA, US). Figures 33, 218-220, 225, 226.
Planta hemiepiphytica; internodia 14 ст longa, 1.5—
2.5 cm diam.; cataphylla ad 12-35 cm longa, incostata,
persistentia semi-intacta; рене 'subteres, 27-75 cm
lon ngus, 0.8-1.8 cm diam., adaxialiter,
in sicco conspicue flav ibrun nneus; Vua anguste ovato-
ПР cordato-sagittata basi, 24-63 cm longa, 19—
m lata, atriviridi supra, opace Lar hebetata infra;
return 1-5; pedunculus 4—11 cm longus, mm
diam.; x one ll~- 17.5 em longs lamina _враћае extus
viridi у
purpureo vel Was vel rubro aut subroseo, i
tus albido vel rubello vel pallide violaceipurpureo; pistilla
(4)5—8-locularia; loculi (4)20—24-ovulati.
Hemiepiphytic; stem appressed-climbing; inter-
nodes short, semiglossy, 1—4 cm long, 1.5-2.5 cm
diam., longer than broad, green to gray-green, dry-
ing yellow-brown; roots moderately numerous per
node, drying dark brown, ridged; cataphylls 12-35
cm long, unribbed, sometimes sharply l-ribbed
near apex (sharply 2-ribbed in South America), me-
dium green, drying light to medium brown, per-
sisting semi-intact at upper nodes, then deciduous
with large patches of yellow-brown periderm re-
maining intact; petioles 27-75 cm long, 0.8-1.8
cm diam., + terete, obtusely flattened adaxially,
spongy (in South America), dark green to gray-
green, surface semiglossy to matte, transversely fis-
sured near apex, drying pale yellow-brown.
LEAVES erect-spreading; blades ovate-triangular,
subcoriaceous, abruptly acuminate at apex, cor-
date-sagittate at base, 24-63 cm long, 19-44 cm
wide (1.1-2.3 times longer than wide), (0.7—1.2
times longer than petiole), equal to or longer than
petiole, margins concave, broadly undulate, upper
surface semiglossy, dark green, drying conspicu-
ously yellow-brown, semiglossy, lower surface
opaque whitish, matte, paler; anterior lobe 19.5—
49.5 cm long, 14.5-42 cm wide (1.9-3.6 times lon-
ger than posterior lobes); posterior lobes 7.1—20.2
456
Annals of the
Missouri Botanical Garden
ст long, 7.3-20.5 cm wide, obtuse to rounded;
midrib broadly concave, paler than surface above,
thicker than broad, weakly glossy, darker than sur-
face below; basal veins 6 per side, with 1-2 free to
base, second and third veins coalesced ca. 3 cm;
posterior rib naked for 2.5-3.5 cm long; primary
lateral veins 6-11 per side, departing midrib at a
35-50” angle, weakly sunken above, convex and
darker than surface below; minor veins moderately
distinct, fine, intermittent below, arising from both
the midrib and primary lateral veins. INFLORES-
CENCES 1-5 per axil; peduncle 4–11 cm long, 4—
11 mm diam., white-lineate to coarsely white-
streaked toward apex and on spathe base; spathe
11-17.5 cm long, 2-3.5 cm diam. (1.3-3.1 times
longer than peduncle), white-speckled, semiglossy,
constricted above the tube, 9-11 mm diam. at con-
striction; spathe blade green to whitish, sometimes
rose-red outside, green to whitish or sometimes
rose-red and white-speckled inside; spathe tube ob-
long-ellipsoid, 4-7 cm long, usually dark green
or sometimes purple to violet-purple to red or pink-
ish outside, reddish to pale violet-purple (dark ma-
roon in South America) inside; spadix white, cy-
lindrical, 12.9-16.6 cm long, broadest near the
base, constricted near the middle; pistillate portion
whitish, cylindrical to ovoid, 3.8-8.9 cm long, 7—
16 mm diam.; staminate portion 9.3-13.4 cm long;
fertile staminate portion cylindrical, 8-13 mm
diam., broadest at the base, narrower than pistillate
and sterile portions; sterile staminate portion as
broad as or narrower than the pistillate portion, 1—
1.3 cm diam.; pistils glossy, 2.1-4(5.8) mm long,
1.2-2.6 mm diam.; ovary (4)5-8-locular, walls em-
bedded with granular, crystal-like particles, locules
1.3—3.2(4.5) mm long, 0.4—0.6 mm diam., with ax-
ile placentation; ovules (4)20-24 per locule, 2-se-
riate, 0.2 mm long, about equal in length to funicle;
funicle 0.1 mm long, adnate to lower part of axillar
wall, style 0.4-0.7 mm long, 1.2-2.6 mm diam.,
similar to style type B; style apex flat or rounded;
stigma subdiscoid, 1-1.5 mm diam., 0.2-0.5 mm
high, covering center of style apex; the androecium
truncate, margins 4—6-sided, sometimes scalloped;
thecae oblong, 0.3-0.4 mm wide, contiguous, +
parallel to one another; sterile staminate flowers 3—
6-sided, 1.5-2.5 mm long, 1.4-1.8 mm wide. Ber-
ries white, with purple stigmas; seeds 2-3, 6, 10—
12, 18-25 per locule, (1-5)8-11 mm long, 2-3 mm
di
am.
Flowering in Philodendron hebetatum apparently
occurs throughout the dry season and first part of
the rainy season in Panama, from December
through July. Post-anthesis inflorescences have
been collected from January through October (ex-
cept September). Immature fruits have been col-
lected in June and July. South American collections
follow more or less the same pattern as those from
Panama.
Philodendron hebetatum ranges from Panama to
Colombia (Chocó) and Ecuador (Carchi, Esmeral-
das, and Pichincha) along the Pacific coast, at 10
to 1880 m elevation in Premontane wet forest, Trop-
ical wet forest, and Premontane rain forest life
zones. It is to be expected in adjacent Costa Rica.
In Panama, most collections have been made at or
less than 1400 m, rarely to about 1630 m.
Philodendron hebetatum is a member of P. sect.
Philodendron subsect. Philodendron ser. Impolita.
This species is characterized by its generally ap-
pressed-climbing hemiepiphytic habit; short inter-
nodes; usually unribbed, semi-intact persistent cat-
aphylls with large patches of yellow-brown
epidermis remaining intact; petioles equaling or ex-
ceeding the blade in length and drying conspicu-
ously yellow-brown and smooth (thus clearly de-
marcated from the blade); and markedly bicolorous,
narrowly ovate-triangular blades usually concave
along the margin and matte and whitish on the low-
er surface.
The petioles of P. hebetatum are variable, being
typically terete to obtusely D-shaped in Central
America but sharply C-shaped to sharply D-shaped
with acute to bluntly raised lateral margins in South
America. South American populations also have the
major leaf veins drying yellowish to pale red vs.
dark red in Central America. It is possible that the
South American elements may ultimately prove to
be at least subspecifically distinct.
Some South American specimens of P. hebetatum
are particularly noteworthy. Holm-Nielsen et al.
25665, from Ecuador, differs from other collections
in having the petiole epidermis drying paler yellow
and the immature spathes drying a paler reddish
brown. A collection from Narifio Department, Co-
lombia (Croat 72425A), is unusual in having the
margins of the anterior lobe markedly concave, but
perhaps belongs also to this species.
Philodendron hebetatum is easily confused, es-
pecially in live condition, with P. thalassicum,
which may also have triangular-ovate blades much
paler and matte on the lower surface. The latter
differs, however, in having petioles that are sharply
D-shaped and dry blackened rather than conspic-
uously yellow-brown. In addition, P. hebetatum has
spathes reddish within, while in P. thalassicum they
are greenish within.
Philodendron hebetatum is also similar to P.
strictum, which differs by its usually terrestrial hab-
Volume 84, Number 3
457
Philodendron Subgenus Philodendron
it, more or less reclining stems, and broadly ovate
blades. Both species have pale, matte lower blade
surfaces, yellow-brown drying petioles, and cata-
phylls with persistent patches of intact, yellowish
brown epidermis. The lower blade surface in both
species is covered with a wax-like substance, and
dries with a fine reticulate pattern (areolate at 7X
or higher magnification). Philodendron strictum
ranges from Costa Rica to Panama and Venezuela
at 650 to 1900 m elevation. In Panama, where both
species occur together along the Fortuna Dam
Road, P. strictum tends to occur at higher elevations
than P. hebetatum.
Juvenile plants (Croat 56049, 56174, 56213,
69291; Valle and Chocó Departments, Colombia) of
P. hebetatum have oblong-elliptic leaf blades acute
at the base, but the petiolar epidermis already is
colored as in adult plants. Leaves of older plants
become increasingly rounder and broader at the
base and finally subcordate to cordate.
An unusual collection is Croat 49298 from Cho-
có Department, Colombia, with leaves only shallow-
ly cordate, even though they are in the upper end
of the size range for adult plants of this species.
Sánchez et al. 553, a collection from Colombia
(Antioquia), is noteworthy in being from 1880 m
elevation and in being terrestrial. Holm-Nielsen et
al. 25665 from Ecuador (Esmeraldas) differs from
other South American collections in having a paler,
flakier epidermis on the petioles and smaller, more
or less sessile (perhaps immature) inflorescences.
Perhaps it represents another species. South Amer-
ican collections of P. hebetatum differ from those
of Panamanian material in that the petioles dry
dark brown rather than yellowish.
Croat 61396 is unusual in apparently having
only four ovules per loc
Additional specimens examined. PANAMA. Bocas
b Toro: Fortuna Dam area, Chiriquí Grande—Fortuna,
mi. N of Divide, 850—950 m, са. 845'N, 82^15'W,
cid 9663 (MO, NY); 3.2 mi. N of Divide, 700 m,
Pe N, 82*15'W, Croat & Grayum 60263 (CM, yi 13:2
of Chiriquí Grande, 310 m, 8'45'N, 82°10’
60140 (AAU, lee Chiriqui: Cerro e уге
off Pan-Ameri Highway гае тіп 1200-1
road, 1
ч Croat 33141 (MO): 1390-1410 m, Ole 37223 (MO);
9.2 mi. W of Chame, 1450-1480 8°35'N, 81°50'W,
; 8.3 mi. beyond Chame,
630 m, Croat 75042 (MO); 75046 (CM, MO); Fortuna
~
E
=
o
=,
a
£
S.
e
<
o
~
g
a
Е
=
=
$,
—
m, Croat 44686 cle Mor Alto Calvario,
800 m, 8°39'N, 80°36'W, Croat 75075 (CM, МО); 710-
800 m, 8*39'N, 80°36’W, Croat 68712 (ENCB, MO); El
Valle region, vic. of La Mesa, N of El Valle de Antón,
900-1000 m, 8:37'М, 80°08’W, Croat 67279 (MO):
8°37'N, 80907", McPherson 11203 (CM MO);
8°40'N, 80%07'W, Knapp zg (МО); 5753 MERU, ДО
Mesa, 775 m, 8°36'N, 80°07'W, Croat 74783 (MO
QCA); 14372 (MO); те ft, Dwyer et al. 4567
MO, NY); 860-900 m, Croat 37364 (MO); ca. 800 m,
25407 (MO); 800-900 m, 8
(DUKE, MO, US). Darién: Pa oa Nacional Darién, near
gold mine at Men of N branch of Rfo Pucuro, nie
of Cerro Tacarcuna, ca. 6 km N oft Cerro Mali, 1300—1500
m, 8*09'5"N, 77715", Hammel et al. 16541 (COL, MO);
Río Tuquesa, Tyler Kittredge gold mine, ca. 2 air km from
са Divide, = 27240 (МО). Panama: Cerro
Jefe region, vic. of summit, 850 m, 9°14’N, 79°22'W,
perth 67060 (MO, NY): 0.8 mi. beyond turnoff to Altos
de Pacora, 770 m, 9?15'N, 79*29'W, Croat & Zhu 76611
(CM, K, M, MEXU, MO, W); 4.6 km beyond peak on road
to Alto de Pacora, 26.3 km from InterAmerican Highway,
600 m, Croat es (MO); 3-3.5 mi. NE of Altos de Pa-
cora, 7.8-8.2 mi. above highway, 700-750 m, 9?15'N,
79"25' м Croat 68661 (IBE, MO); 5-10 km NE of Altos
~
де Расо i & Kallunki 6031 (МО); El
Мало Can Road, Mile 10, near E о, 330 m, Croat
33773 (MO, US); Cerro Camp 8 40'N,
ana, ca. m,
79°50'W, Thompson 4598 (М0) ‘Groat 17243 (MO), 780-
875 т, 25224 (E, MO, 000 m, Luteyn 3193
DUKE); ca. 1 mi. from aden Ete Highway, ca. 150
m, Croat 35961 (F, MO); above Su Lin Motel, Croat 14726
m, 915'N, 79°W, Croat 69231 (MO, PMA, SAR);
sendero de ciii ] km al este del Campamento
abosques de INRENARE, 800-900 m,
840'N, 79?55'W, Correa et al. 9489 (MO). Veraguas:
Santa Fe region, Santa Fe-Río Calovébora, 0.6 mi. beyond
Hamilton & Krager 3934 (MO); Río Tercero Braso Valley,
beyond Escuela Agrícola Alto Piedra, above Santa Fe,
Yat pares Mo
A. Antioquia: Mpio. Frontino, ene
moe val of the upper r Rí o Cuevas, 1880 m, Sán
си 53 (MEDEL); rud 21 confluence of две:
ae T a and Río rí, ca. 3 km upriver from
Vil ie 28 km ‘SW of Zaragoza, 400-700 m,
Alverson et al. 299 (COL, WIS); Río Anorf valley near
Planta Providencia, 350—600 m, 7°30'N n
herd 924 (COL, WIS). Chocó: Quibdó—Tu
ca. 3 km W к 80 m, Gentr fe ~ al. 30108 (MO):
1 km E of Tutunendo, m, 546'N, 76°35'W, Gentry et
~ 30079 (со; папе E E 136.4, 63 km E
of Tutunendo, 960 m, 5°47’ E 6°22'W, Croat 56348
(COL, JAUM. MO»); Km 2 m W of Tutunedo,
<100 m, 5?39'N, 76740", Croat Sis (MO); Río Atra-
to, 39 km W of Bolívar, 1600 m, Croat 49261 (MO); 78
km W of Bolívar, 466 m, Croat 49298 (MO); 27 mi. W of
Bolívar, 1190 m, 5?50'N, 76716'W, Croat & Cogollo
52089 (MO); e Km 137-138, 79-80 km E
of Quibdó, 910-920 т, 5*45'N, 76'21'30"W, Croat 57345
bre е Ro Ae ó-Las Animas, ca. 1 km N of
m, 5?14'N, 76°40'W, Croat 55958
pet MO): Sis As en Palmar-Cartago, Vereda La Bella
tween San José del Palmar and turnoff to El Cairo, 1430
m, 4%53'N, 76°13'W, Croat 56715 (COL, MO, QCA);
Pueblo Rico (Risaralda)-Istmina (Chocó), Quebrada An-
tón, 15 km W of Santa Cecilia, 6 km W of Chocó-Ris-
aralda border, 240—350 m, 5°20'30"N, 76^13'45"W, Croat
70961 (MO); Quibdó-Istmina, Km 4, <100 m, 6728'N,
458
Annals of the
Missouri Botanical Garden
76°36'W, Croat & Cogollo 52234 (MO); Serranía de Bau-
do, Las Animas-Pato on Río Pato, са. 150 m, 5?30'N,
76°46 'W. Croat 56131 (COL, JAUM, MO, PMA); 16 km
NW of junction with Quibdó-Istmina road near Las Ani-
mas, 100 m, 5?20'N, 76742", Croat 56174 (COL, JAUM,
MO); 5?16'N, 76°41'W, Croat 56049 (COL, JAUM, МО);
Río San - eic 0 m, LIN 77722'W, Fi огего а
Chirrincha, Rfo Aguita, 950 m, Ea: et al. 3522 (MO);
corregimientos de Geguadas—Puerto de Oro, 1550 m,
Alonso et al. 9819 (МО); 9742 (MO). Valle: Bajo Calima
area, Croat 62761 (CM, MO); ca. 15 km N of Buenaven-
ura, Самбп de Colombia concession, Juanchaco region,
m, 3^56'N, 77°08'W, Gentry et al. 53711 (MO); pn
Gasolina road junction, SSW of San Isidro, 50 m
et e а (MO); 11 km N of main tap
, ca. 90 m, 3°56'30"N, 77?01'W, Croat & Mon
AE 61396 (MO, QCA); Buenaventura-Río digi dd 3
km N of Cali-Buenaventura Highway, ca. 50 m, 4%02'N,
77707" М, Croat 61279 (COL, MO, NY, TEX); Km m 50-
80 m, 3° 77°01' W, Croat 69291 (CAS, F, L, MO,
US); ca. 4 km fi a, near Km 14 marker, <50
m, 3?56'N, 59'W, Croat 57530 (AAU, DUKE, K,
MEXU, uenaventura-Mála d, 5
Croat &
Bay 75629 (МО); Km 18-20. on Lo Me a High-
way, Finca Zingara, 1500-2000 m, Cabrera & van der
Werff 15779 (MO); Rio Cajambre, 5-80 m, Cuatrecasas
17054 (US); Río Naya, «pitos from Puerto Мена, са.
10 m, 3^15'N, 7725", Gentry & Juncosa 40673 (MO).
ECUADOR. Carchi: Tulcán reris аны нений
лив sector Sabalera, parroquia Тођаг Don
A he 78°24'W, Tipaz et al. 1298 (MO): pi ee
eraldas: Lita-San Lorenzo, 15.5 km W of Lita, 705
m, 0°55" N, 78°28'W, Croat 72372 (MO); San José de Cay-
apas, 80 m, 0752'N, 7857'W, Holm-Nielsen et al. 25665
(AAU, MO); Eloy PCs Reserva ees амо
Cayapas, Charco Vicente fo San Miguel, afluente del
Río Cayapas, 150 m, O ^43'N, 78:53! е Palacios & Tirado
11276 (MO, QCNE). Pichincha: Reserva ENDESA, Qui-
to-Puerto Quito, Km 113, 0°05'N, 79°02’ W, 750 m, Croat
& Rodríguez 61454 (CM, K, M, MO, QCA); 61455 (K,
MO, QCA); 710 m, 0%03'N, 7 79°07'W, Croat 73155 (AAU,
B, COL, F, K, MO, NY, ОСА, US).
|
Philodendron pp ton ite ae Schott, Wie-
ner Z. Kunst. 1829: 780. 1829. Arum heder-
aceum Jacq., Pih. de Pl. 31. 1760. TYPE:
t. 51, fig. D in Plum, Pl. Amer, 1756
(holotype). Figures 221—223.
Hemiepiphytic vine; growing to often high in
trees, stem appressed-climbing, eventually scan-
dent, often pendent, sap clear, turning honey-col-
ored, leaf scars 7-15 mm long; internodes weakly
flattened on one side, sometimes with 2 sharply
raised on the side above the petioles, usually weak-
ly glossy, sometimes matte, pale to medium green,
minutely speckled to striate, usually smooth when
fresh, but drying minutely ridged, sometimes prom-
inently ribbed throughout its circumference (the
ribs smooth to prominently warty), (2)10-28 cm
long, 1-2.5(3.5) cm diam., dark green, usually dry-
ing green, sometimes reddish; roots brown, to 10
cm long, many at nodes; cataphylls 6-10 cm long,
unribbed, weakly l-ribbed, or bluntly to sharply
2-ribbed, pale green, deciduous; petioles (6)9.7—
27(33) cm long, 6-10 mm diam., terete to subter-
ete, pale green, firm, flattened adaxially, pale green,
surface smooth, weakly glossy to matte; blades
broadly ovate, subcoriaceous to coriaceous, semi-
glossy to matte, acuminate to long acuminate,
sometimes cuspidate at apex (the acumen inrolled,
0.5-0.9 mm long), 11—40(50) cm long, 8-24(34) cm
wide (1.2—1.9 times longer than wide), (ca. 1.3(1.5)
times longer than petiole), margins hyaline to pale
yellowish, upper surface dark green, sometimes
subvelvety, lower surface slightly paler, often pur-
plish violet, drying gray-green to yellow-green; an-
terior lobe 9-30(41) cm long, 9-24(29) cm wide
((2)2.4—3.7(6.3) times longer than posterior lobes);
posterior lobes 3—10(14) cm long, 4.3-15.7 cm
wide, directed inward and sometimes overlapped,
obtuse to rounded; sinus usually deeper than broad,
mostly spathulate, rarely hippocrepiform, 3-7 cm
deep; midrib convex to flat or sunken, concolorous
or slightly paler than surface above, convex, con-
colorous below; basal veins (3)4—5(6) per side, with
(0)1-2 free to base, part of the remainder coalesced
to 1.5-2 cm; posterior rib not naked; primary lat-
eral veins 2-6 per side, departing midrib at a 35-
55° angle, + straight to the margins, sunken to
weakly raised, slightly paler than surface above,
convex and paler than surface below; minor veins
obscured to moderately distinct, arising from both
the midrib and primary lateral veins. INFLORES-
CENCES erect or pendent, 1 per axil; peduncle (2–
34—15.7 ст long, 8-12 mm diam., pale green,
sometimes tinged purple, matte; spathe subcoria-
ceous to coriaceous, 9-16.6(24) cm long, (0. 9-
2.6(3.3) times longer than peduncle), weakly con-
stricted above the tube, 1.3-3.6(5.7) cm diam. at
constriction, usually green, sometimes yellowish
white, yellowish green, or cream to creamy-white
throughout; spathe blade sometimes purple tinged
outside, 1.2-3.6 cm diam. when furled, pale green,
greenish yellow, sometimes tinged red inside;
spathe tube dark green, sometimes tinged reddish
maroon outside, 5-6 cm long, 1.5—4.9(6.9) cm
iam., maroon, dark red, crimson, or purple at base
inside; spadix stipitate to 5-10 mm long, dark ma-
roon; 12-20 cm long; pistillate portion pale green-
ish white to green, 3.5-6 ст long, 1.5 cm diam. а!
base, 1.7 cm diam. at middle and near apex; sta-
minate portion 7-11 cm long; fertile staminate por-
tion creamy white to pinkish, broadest at base,
Volume 84, Number 3
1997
Croat 459
Philodendron Subgenus Philodendron
weakly constricted ca. 1 cm above vmi: portion
then + uniform to near apex, 1.4-2. iam. at
middle, 9 mm diam. ca. 1 cm bud apex; sterile
staminate portion 1.6 cm diam.; pistils 4—9.2 mm
long, 1.8-3.1 mm diam.; ovary 4—6(7)-locular, 8
mm long, locules 8 mm long, 1.4 mm diam., with
axile placentation; ovules 20-25 per locule, 0.1
mm long, 2-seriate; funicle 0.1–0.3 mm long, ad-
nate to lower part of partition, style 1.1 mm long,
29 mm diam., similar to style type B; style apex
flat or somewhat rounded, drying concave with a
pale margin and 4—6 paler, flat to weakly sunken
circular areas associated with the stylar pores; stig-
ma 3—4-sided, light brown to reddish, drying light
brown, 1.8-3.3 mm diam., 0.35 mm high, margins
thin; sterile staminate flowers blunt, irregularly 4—
5-sided, 2.2 mm long, 1.3 mm wide. INFRUC-
TESCENCE pendent, often on leafless stems;
spathe dark green, weakly glossy outside; pistillate
spadix 5-8 cm long, 3.5—4 cm diam.; berries green-
ish white; seeds 1-2 per locule, somewhat orange,
many per berry, + ovoid to oblong ellipsoid, (1.5)3—
5 mm long, 2.5—4 mm diam., with weak constriction
(nipple) and densely covered with raphide cells.
JUVENILE plants with upper blade surface dark
green, sometimes reddish green, with glistening
minute close papillations, lower surface somewhat
maroon; veins less conspicuous.
Philodendron hederaceum ranges throughout the
West Indies and from Mexico throughout Central
America and much of South America, at sea level
to 1200(1500) m elevation. In South America it ex-
tends from as far south as Los Rfos Province in
coastal Ecuador to Trinidad, Venezuela, the Guian-
as, Brazil, Ecuador, Peru, and Bolivia on the At-
lantic drainage of the continent.
Philodendron hederaceum is a member of P. sect.
Philodendron subsect. Solenosterigma. This species
is distinguished by its scandent habit, long inter-
nodes, deciduous isti ovate-cordate, long-
petiolate leaves, and s inflorescence with
usually green spathes ow m tube reddish to pur-
plish within.
Philodendron hederaceum is most easily con-
fused with P. purpureoviride, which is a vine with
similar leaves. See that species for differences.
Philodendron hederaceum is also somewhat sim-
ilar to P. jacquinii, but the latter differs in its gen-
erally pubescent stems, petioles and major veins of
the lower blade surface, its thinner leaf blades,
swollen spathe tube, and broad pistillate portion of
the spadix with elongate styles.
The taxa here treated as Philodendron hedera-
ceum and P. jacquinii have long been confused no-
menclaturally. The former name is based on Arum
hederaceum, first validly published by N. J. Jacquin
(1760) in his Enumeratio Systematica Plantarum.
Jacquin cited only a Plumier (1756) plate (t. 51,
fig. d), which thus must be accepted as the holotype
of the name (Greuter et al., 1994; Art. 9.1, Note
1). Three years later, € (1763), in his Selec-
tarum Stirpium Americanarum Historia, published
his own plate (t. 152), identified as A. hederaceum
but actually depicting a different species from Car-
tagena, Colombia, namely, that treated in this re-
vision as P. jacquinii Schott. These two species are
fortunately sufficiently different in appearance that
even drawings of sterile plants, such as that of Plu-
mier, are unmistakable. This mistake by Jacquin
has caused considerable confusion, since several
authors, including Kunth (1841), Engler (1899),
Krause (1913), Dugand (1945), and Bunting
(1963b, 1995), have misapplied the name P. hed-
eraceum based on Jacquin's (1763) publication.
Schott, however, understood the problem. He had
already transferred Arum hederaceum to Philoden-
dron (Schott, 1829), and in his treatment for Syn-
opsis Aroidearum (Schott, 1856) he described P.
јасашти, explicitly basing it on Jacquin's (1763)
plate. Schott (1856) also placed P. hederaceum sen-
su Kunth into synonymy under his newly described
P. jacquinii. However, he included P. hederaceum
in his grex Macrobelium, while treating other syn-
onyms of P hederaceum in grex Solenosterigma.
Names so treated were: P. scandens, P. POP NN
P. oxycardium, P. cuspidatum, and P. m
Engler (1899), followed by Krause (1913), treat-
ed the species herein called P. hederaceum as four
distinct species: P. prieurianum, P. scandens, P. ox-
ycardium, and P. micans. Both Engler and Krause
erred in treating Arum hederaceum Jacq. as a ques-
tionable synonym of P. hoffmannii (= P. jacquinii),
citing Jacquin's (1763) t. 152 as the type.
Despite the confusion by Engler and Krause,
Standley and Steyermark (1958b) correctly dealt
with the taxonomy of P. hederaceum, citing P. scan-
dens, P. oxycardium, and P. miduhoi in synonymy.
Their treatment of P. jacquinii was incorrect, since
they cited that name under the later synonym P.
hoffmannii Schott (1858). In this regard they fol-
lowed Krause (1913). Thus, despite the confusion
by Engler and by Krause, the nomenclature of these
species was essentially rectified as early as 1958
to the species as P. oxycardium or P. cordatum hort.
(non Vell.).
Bunting (1963b), apparently following the lead
of Dugand (1945), was aware of the Plumier illus-
tration cited by Jacquin but seemed to believe that
it had no bearing on the application of the name.
460
Annals of the
Missouri Botanical Garden
He also referred to Jacquin (1763) as the “initial”
publication of Arum hederaceum, presumably in the
erroneous belief that the name had not been validly
published in Jacquin (1760). Bunting (1963a) ac-
cordingly applied the name P. hederaceum to the
species here called P. jacquinii and the obscure P.
scandens K. Koch (1853) for the species herein
called P. hederaceum. Certainly the epithet P. scan-
dens was not in general use up until that time be-
cause most horticultural works (Birdsey, 1951) still
referred to the species as P. oxycardium Schott.
Other, more practical matters substantiate that
Plumier’s (1756) rather crude drawing depicts the
species here treated as P. hederaceum, rather than
the one called P. jacquinii (accurately illustrated by
o 1763). Plumier's text associated with t. 51,
. D states that the plant grew in Martinique,
^e wn to have been visited by Plumier (Urban,
1898: D а hederaceum is а wide-
spread species in the West Indies and occurs on
Martinique, while P jacquinii, though also wide-
spread, is not known from the Lesser Antilles. The
epithet hederaceum connotes an ivy-like growth
habit and aspect, apt for the species to which the
epithet is here applied, but not for P. jacquinii. The
application of the name Р. hederaceum, as ex-
plained above, is unambiguous, whereas that of P.
scandens, the name used during the last 30 years
for this plant, is highly dubious, as it is based on
a sterile Koch specimen of unknown origin, lacking
an extant type specimen or even illustrations.
Article 57.1 of the Tokyo Code (Greuter et al.,
1994) states that “А name that has been widely and
persistently used for a taxon or taxa not including
its type is not to be used in a sense that conflicts
with current usage unless and until a proposal to
deal with it under Art. 14.1 or 56.1 has been sub-
mitted and rejected." I consider that this Article
does not apply in the present case, since the name
> hederaceum was used in the sense, including its
type, as recently as 1958, in a higher regional flora
(Flora of Guatemala), which is the most recently
published treatment of Araceae for any Central
American country and still reigns as the standard
work throughout the region.
In Central America, P. hederaceum can be divid-
ed into three varieties. Philodendron hederaceum
var. hederaceum an raceum var. oxycardium
(Schott) Croat are only distinguishable on the basis
of their juvenile leaf blades. In P. hederaceum var.
hederaceum, the juvenile blades are velvety with a
silky sheen on the upper surface, whereas the j ju-
venile blades of P. hederaceum var. oxycardium are
glossy on the upper surface. These juvenile forms
were formally treated by Bunting (1968) as P. scan-
dens forma micans
The third newly recognized variety, P. hedera-
ceum var. kirkbridei Croat, is distinguishable by its
adult stems that dry brown and are deeply sulcate
with prominent ridges. This taxon also differs in
occurring at higher elevations.
The following key separates the three varieties
of P. hederaceum in Central America. For anatom-
ical differences see Bunting (1968).
KEY TO THE VARIETIES OF P. HEDERACEUM
la. Adult stems weakly sulcate on living plants,
moderately to conspicuously sulcate and ce
brown upon drying, usually densely
spathe tube dark red to red-purple inside; Ciis
veu Panama, € and pease mostly
var. kirkbridei
e, never eo a he tube green
in Mexico, Central America, and to the West
fend E e. America E Ama-
r. hederaceum
N
Е"
E
=
<
c
=
= =
[87]
Е Б
ББ
as
Ф
Ф
T
Ф
Ф
ча
©
=
S-
o
=
"9
T
O
~
un
т
w
о
p
green on the lower surface; known only from
the Gulf slope
ern Oaxaca, and Tabasco, but pn also
found in Jamaica ar. oxycardium
Philodendron hederaceum (Jacq.) Schott var.
ederaceum
Philodendron scandens K. Koch & Sello, in A. Braun et
al., Append. sp. Hort. berol. 1853: 14. 1853-1854.
TYPE: A cultivated plant grown in Berlin, origin not
stated (holotype, B lost
Philodendron harlowii Y. M. Johns. Sargentia У 9], t.
14, fig. 1. 1949. TYPE: Panama. Panamá: San José
ine area 11B, Johnston 1 030 (holotype, GH; iso-
Hist.
Philodendron nubis burn Revista Soc. Mex.
95, ай fiet. 0. TYPE: Mexico. Chia-
a Espe ru m, 23 Dec. 1949, Mat
m et MEXU: UR U
Hann cuspidatum & ВоВе in A. Braun
Append. Gen. sp: Hort: berol. 1854: 7. 1854-
1899. TY cultivated at Berlin Botan
(holotype, B? destro yed). Schott ic. 2619 (ose
here d designate p
Philodendron micans K. Koch, in + ume et al., Appen
1854-1855. hilo
unknown (hol olotype, B? kotoi: Schott ic. 2709
W).
(neotype, here designated,
Volume 84, Number 3
1997
Croat 461
Philodendron Subgenus Philodendron
Philodendron sit ghar K. Koch, in A. Braun et al.,
ppend. Gen. s rt. berol. 1854: 7. 1854-1855.
ie cultivated атат at Berlin (holotype, В?
w lost).
Philodendron oxyprorum Schott, Syn. Aroid. 82.
E: Venezuela. [Synonymized by Engler rox
bot specimen or illustration now exists.
Philodendron bi Schott, Oesterr. Bot. Z. 8: 1
E: Guatemala, Wendland (holotype, че
dese Schott ic. 2498 (neotype, here designat-
Philodendron жеө Engl., Вог. Jahrb. Syst. 26: ore
: Cost : Rica. Puntarenas: Ujarrás de
Dici nos Hid 912'N, 83"17' e Feb. 1897, Pis
11132 пита B; isotype,
Hemiepiphyte; internodes 10-25 cm long, 1–
2.5(3.5) cm diam., weakly flattened on one side,
medium green, minutely speckled to striate or
smooth when fresh but drying minutely ridged,
greenish; cataphylls 6-10 cm long, unribbed,
weakly 1-ribbed or bluntly to sharply 2-ribbed, de-
ciduous intact; petioles (6)10-27(33) cm long, 6–
10 mm diam.; blades (11)16–40(50) ‹ cm long, 8-
greenish brown, lower surface medium green,
glossy, drying gray-green to yellow-green; primary
lateral veins 2-6 per side, departing midrib at a
35—55° angle. INFLORESCENCES with peduncle
(24—16 cm long; spathe 9-17(20) cm long, spathe
usually green, sometimes yellowish white, spathe
blade sometimes tinged purple outside; spathe tube
dark, sometimes tinged reddish maroon outside;
spadix stipitate to 5 mm, 12-18 cm long; ovary 4—
6-locular, 20-25 ovules per locule. INFRUCTES-
CENCES with many seeds per locule.
Flowering in Philodendron hederaceum var. hed-
eraceum is rare (known only in October and No-
vember), although the species has been found in
pre-anthesis condition in every month of the year.
Post-anthesis collections predominate in the dry
season and earliest part of the rainy season from
December through May, but there are a surprisingly
large number of post-anthesis collections made in
November, a month when very little flowering gen-
erally takes place (Croat, 1975, 1978). Mature
fruiting collections have been made from April and
May, also indicating that the species may flower
predominantly in the dry season.
The range of P. hederaceum var. hederaceum is
essentially that of the species. It is the most wide-
spread taxon of Philodendron and, indeed, perhaps
of all neotropical Araceae, ranging from San Luis
Potosí State in Mexico to the Greater and Lesser
Antilles, and down both slopes of the Andes, east
о the Guianas and south to Brazil and Bolivia. In
Central America, the variety occurs on both slopes
of the Continental Divide. In Mexico, it ranges from
sea level to 1200(1500) m elevation, whereas in
Middle America and Panama it ranges mostly to
450 (rarely to 900) m. It occurs principally in Trop-
ical moist forest but ranges into Premontane wet
forest and even Tropical wet forest. A single sterile
pre-adult collection from Tropical wet forest transi-
tion to Premontane wet forest in Bajo Calima (Bay
237) is apparently also this species.
Additional specimens examined for P. hederaceum var.
hederaceum. BELIZE. Gentle 617A (MICH). Cayo: S of
Guacamallo ee m ney 2837 (BM); DE
Grano de Oro, 1700 at 23697 (MO
on Hummingbird Highs “жїл DU. 12718 (МО);
Rio Frio, 1.5 mi. W of Augustine, 450 т, Sutton et al.
205 (BM); 0.9 km before Caracol, 16°46'N, 89°07'W, Bal-
ick et al. 3149 (MO). Stann Creek: Cockscomb Basin,
Jaguar Preserve, 10 km W of
Highway, 400 m, 16°
(MO). Toledo: along highway to Punta Gorda,
road to San Antonio, ca. 100 ft., Croat 24511 (MO); Union
Camp, 750 m, Whitefoord 1773 (BM, MO); Columbia Riv-
er Forest Reserve, Gloria Camp, Holst 4422 (MO);
Southern Maya рама poss PAPA ¿rio
16°29'31"N, 8854'37"W,
боце COSTA RICA. Мей) s.n. quos пи тај or dd
m, Rojas 178 (MO). Alajuela: Сапаз—Џраја, 4 km
NNE of icm f ca. 400 m, Croat 36320 (MO); Escuela
Centroameriana de Ganadería, (near Atenas) 425 m,
9?58'N, 1
Guanacaste: Tilarán, 500—6 tandley Valerio
44563 (US): ine SK июн mee 46563 (US; 7 km
N of Cañas, Janzen T-24 (MO); Quebrada Desprendimien-
to- Quebrada XS RR along Río Las Flores, ca. 450
m, 10%40'N, 85*04'5"W, Grayum et al. 4912 (B, CAS, CR,
K, MO, US); Nandayure, Península de Nicoya, 120 m, A.
Rodriguez & Estrada 141 (CR, INB). Heredia: La Seb
Field Station, ca. 100 m, McDowell 810 (MO); 100 m
ind (MO); Grayum 2565 (MO). Limón: 20 mi. SE of Li-
road to Punta Cahiuta, near sea level, Croat 43174
(МО); ca. 1 km N of Shiroles, са. 50 т, Croat 43278 (МО);
Siquirres-Río Pacuare, 5 of Río Pacuare, 50-100 m
10%5'N, 8329'W, Burger & Liesner 6953 (MO, NY); la
Bomba-Cahuita, 20 m, Gómez & Hampshire 20129 (МО);
r rte 0-50 m, Taylor & Skotak 4443 (DUKE);
um tion on Indiana Branch, 30 m, Standley
& Valerio 48392 (US); Parque Nacional Bahía de Portete,
Río Jiménez, S of Guápiles, 10^17'N, 83
2393 (CR, MO). утуу Сапібп de Buenos Aires,
along Río Ceibo, m, 914'N, 83718", Gra-
yum 10229 (CR, IBN, ri MO, US); Golfito, 20-200 m,
8°38'N, 83^11' 15"W, H. Schmidt 603 (CR, MO); San Vito,
Ан CR.CB.7C. 109 (CR); твата, Cordillera de
Tilarán, 1500-1620 m, Pounds 94 (MO); Cantón de Tur-
rubares, along Río om pes between Río Carara and Que-
brada veg m, 9?46'30"N, 84?32' W, Gra-
yum 10420 (CR, Moi EL SALVADOR. Ahuachapan:
San Francisco Menéndez-Tacuba, 1–3 mi. above road to
Río Clara Sucia, 1000-1250 m, Croat 42135 (MO). San
Miguel: ca. 50 mi. NW of San Miguel, along highway
CA-1, Croat 32799 (MO). San Salvador: vic. of Tona-
catepeque, Standley 19533 (CH. NY, US). Sonsonate:
462
Annals of the
Missouri Botanical Garden
Родові de San Isidro, са. 3 ті. 5 of Lake Coatepeque,
850 m, 2 42241 a GUATEMALA. Los
Amate, sree n. (US); Watson 427 (GH). Alta Ver-
o El ine (Lago Izabal), 7 mi. E of Highway
0 (MO). Escuintla:
Santa Lucia, 1045 ft., Kellerman 4547 (US); 5285 (US).
Izabal: Montaña del Mico, between Milla 49.5 and ridge
6 mi. from Izabal, 6 m, Steyermark 38486 (Е); ca.
7 mi. S of Puerto Barrios, 50 m, Croat 41811 (MO). Que-
Rafael, 600 m, Grat 40760 (MO). Suddiephque ет: 1 т
Е of Mazatenango, <500 т, Croat 43757 (МО). HON.
DURAS. Atlantida: Lancetilla Valley, Tonacatepeque,
Pfeifer 2130 (BH, US); San José de Texíguat-El Chorizo,
00 m, Nelson 10565 (TEFH); 4 km S of Tela, 0-100 m,
Trias 190 (ТЕЕН, ОМАН); ca. 10 mi. SE of Tela, along
Río Lancetilla, 10-150 m, Croat 42639 (MO); bito
gore a. 10 km SW als Ceiba, 80–140 m, 15?42'N,
^51'W, Liesner 26335 (MO). Colón: Río Sle, 3 km E
of a «os Howler Site, Saunders 192 (MO). Comayagua:
junction Río Yure-Río Humuya, 200 m, NU et al. 6182
(MO). Copán: 13 mi. E of Copán, road to La Entrada,
750 m, Croat 42529 (МО); Sta. Rita village, 650 m, Mo-
lina 33668 (MO). Cortés: Puerto Cortés-Guatemalan bor-
of Omoa, sea level, Croat 42555 (MO);
N of Lago de Yojoa, 2-6 mi. from highway, 600 m, Croat
42739 (MO). Gracias a Dios: Ahuas Bila, 200 m SW
of Puerto Lempira, 100 m, Nelson & Cruz 9316 (UNAH);
9292 (TEFH, UNAH). Olancho: Mpio. San Estebán, near
Santa María del Carbón, 21 mi. NE of San Estebán, along
road to Bonito Oriental, 440 m, 15%25'25"N, 85?34'45"W,
Davidse et al. 35571 (MO; Río Guyape, San Pedro de
Catac gE чы Blackmore & Heath 1984 (BM); Río
Wampú, 8 km Pisijire, 500-700 m, 15?15'N,
85°25'W, Nelson & ‘Clewell 594 VT MO). MEXICO.
: i E of Escuintla, 300 m
Croat 43859 (MO); 2 mi. SW of Cu atemalan border, CAE
way 200 to Tapachula, 300 m, Croat 43771 (MO); Bonam-
pak, near ruins, 500 m, Matuda 38715 (MO); Mpio. Ocos-
ingo, 5 km SW of Santo Domingo, 600 m, Davidse et al.
20425 "T Esperanza, Bows 150 m, Matuda 17789
(NY); 6 mi. N of Ocoz ps 1000 m, Croat 40584
р Ра зар . SE of Palenque, са.
400 m, Croat 40167 (MO); els MR ад Highway
199, 27 mi. SW of Palenque, 210 m, Croat 40302 (MO);
Cerro Vernal, NW side, 25-30 km SE of Tonalá, 400—600
m, Breedlove 25617 (DS). Guerrero: Tierra Colorada—
харор Tierra Colorada, Río Comitlán, 900-1000
m, Croat 457 55 (MO); Pinotepa Nacional-Tlaxiaco, High-
way 125, ca. 8.4 mi. S of Putla de Guerrero, ca. 1000 m,
Croat (MO); Tierra SRR Кони | 366-
a. 380 m, Moore & Bunting 8840 (BH). Jalisco:
اا a 100 m, Mexta 1314 (ОС); 24.1 mi. from
Autlán, ca
m, re & Bunting 8737 (BH); Quimix-
to, hair 1201 UN. CAS, Ма G, GH, МО, NY, UC a
t: Miram W of Jalcocotán, Dres
a. 1
Wirth 2708 (US) Sing pe s.n. (BH); Philbrick A
(BH); San Blas, Lewis s.
Chayuco, 220 m, Miller & Tenorio L. 524 (MO); y subici
Rincón del Tigre, Mpio. Acatlán, 2 km from Acatlán on
road to Capilla, ca. 100 m, ca. 18°31'N, 96°36'%, Gereau
et al. 2190 (CAS, MO, RSA); Tux xtepec-Oaxaca, 10 mi. S
of Valle Nacional, 700 m, Croat 39802 (MO); 0.5 mi. S of
Valle Nacional, 120 m, Croat 39694 (MO); Esmeralda-
m, 17°10°№, 94745", Croat & Hannon 63233 (МО).
Puebla: Teziutlán-Nautla, Rancho Las Margaritas, Huey-
tamalco, near border with Veracruz, 19%57'N, 97?16'W,
Conradi 218 (MEXU). San Luis Potosí: 6
Tamazunchale, 250 m, Croat 39271 (MO); Tamar e
tezuma, 1000-1150 m, "АВА edt (MO); Mpio. Nautla,
Cuatro Caminos, near Cerro Chico, 30 m, Ventura 3600
(DS); Mpio. San Andrés Er Estación de Biología Trop-
ical Los Tuxtlas, LOTE 71, 400 m, Љатта & Colin 3126
MO), 400 m, 18°34-36’ N, 95°04—09'W, Manríquez :
Colin 3126 (MO). NICARAGUA. Boaco: along Hwy.
from Río DE ca. 300—310 m, 12°35! №, 85°32’ W E.
vens 9335 a.
12?24—25'N, 85 132-33 W,
(MO); Quebrada Río Grande, NE
600—700 -29 N, Ва 32: W, Moreno 354 (MO). Chin-
andega: Río poes El Viejo, 0-100 m, Atwood 2635
(MO). Chontales: Río Bizcocho-Río El Jordán, 350-550
m, са. 12%12-16'N, 85%15-17'W, Stevens & Montiel
22589 (MO); ca. 2.8 km N of Cuapa, 400-500 m, ca.
12%17'N, 85°23'W, Stevens 3696 (MO); З km N of Santo
Tomás, 280-300 m, 12%05'N, 85°07’W, Moreno 16066
sep Juigalpa, La Libertad, Río se Bizcocho, © 17.4
of Rio Mess 35 m, са. 12°12'N,
М, МО); 4 aa NW of villa San-
MO); Highway 8, ca. 2, 4 ien SW of intersection with Hwy.
2, km 2 . 700 m, 11%57'N, 86"20'W, Stevens 3990
a 4 km pee Highway 8 to Highway 2 intersection,
11*58'N, 86°18-19'W, Stevens 4539 (MO);
IRA Nacional de Agricultura and Ganadería, d h
12 km E of Managua, Atwood 2930 (MO). Ma
M ی ігу, km 140, ca. Mr Ceu"
al. 229 ; Cerro Musün, .
85°16'W, jin 12032 (MO); jean Malacal, Haci-
nda La Bonanza, ca. 20 km from Matagalpa dae
13°01'N, 85%47'W, Castro 2391 (MO). Nueva Segovia: 7
m SE de Santa Clara, 600-700 m, 13%40'N, 86°14'W,
Araquistain A Moreno 2191 (MN kr n El i
4 kms NE of El Jícaro, N, 86°07’
Stevens & ines 2215 (MO). Rio p jud. 'Río Indio,
San Juan del Norte, 2 m, thes ae d (K, M. xis
MEXU, MO). Rivas: Volcán Con La Esperan
ПА 11°31'N, 85°37'W, “Robleto 1618 үт
MO); Isla Onietepe, 140-350 m, 11°33-34’N, m i СР
Robleto 997 (MO); SE of “La Flor,” 300-800 m
Rio Copalar, ca. 29 km E of Rio Blanco, 200-400 a? :
12550—55'N, 85%0-05'W, Stevens 12179 (MO); “Las evi
cedes," 160-180 m, Vincelli 331A (MO); Matagalpa- Was-
lala, near Río Las Carpas and Río Babasca, 540-580 m.
Volume 84, Number 3
1997
463
Philodendron Subgenus Philodendron
13%15'N, 85°32'W, Sandino 2426 (MEXU, MO); La Luz,
ca. 200 m, 13°44’N, 84%47'W, Ortiz 1606 (F, MO); Cerro
Waylawás, E side of Central Range, ca. m,
13°38-39'N, 84?48—49' W, А еа 4172 (MO); 6 km SE of
Waslala, 520-560 m. 13 16" 5°24'W, Moreno 1 ae
M n Experimental fo ecreo,” ca
W, Rios 176 (MO); ca. 80 m, 13°30'N,
84°48’ W, Pipoly 4450 (MO); he M cota Es
: 13°42'N, 84%50'W, Pi-
poly 4593 Мын ca. 0—
Moore 897 ie rk Caño Monte Cristo, “La Grupera,” ca.
А , Moreno & Sandino 14709 (MO);
E of campo Germán Райа, ca. 60-90 m, 11?36'N,
°52'W, Moreno 15137В (МО); са. 10 т, hohe
83°51'W, и (MO); Саћо Monte Cristo, 1 Кт before
the camp Germán Pomares, са. 10 m, 12%35'N, 83°51'W,
Moreno 14839 (Mo) 14843B (MO); “Kurinwacito,” 80—
к 84°55'W, Moreno 23699 (Е, МО); Cerro
El Ocote, 700 m, 13°38'43’N, 85707'06"W, Ortiz 1093
(MO): E ira El Hormiguero. ы of Rio Uli, Age 1921
(MO) Mpio. Rama, Loma Buena Vista m,
12%08'N, 84°12'W, сене 620 (MO); Mpio siii Siun-
awás, ca. 200 m, 13%43'N, 84%45"W, Ortiz 1481 (MO);
Santa Rosa, Ortiz 102 (MO); Negro Wás Sector, El Em-
palme-Rosit ta, «200 m, ca. 13°45'N, 84?25'W, Ortiz не
(МО); Río E ca. 7 km W of Ко Iyas, са. 200 m
13°29'N, 85° Stevens 19278 (МО); Ro
Matis, она а. 13°43'N, 84°49
Pipoly 4685 (MO); ca. iD m, gri ds (MO. PAN-
AMA. Bocas del Toro: S end, Cayo García, 0-5 m
terson & Annable 7287 (MO); Chiriquí Lagoon, von Wedel
1253 (GH, MO); 2668 (GH, MO); Gualaca-Chiriquí Gran-
= 4.2 mi. S of Chiriquí Grande, near sea level, 8°55'N,
2°09’ W, Croat 66813 m TEX); Changuinola—Almiran-
+ Station Milla 7 DU 100 m, чы 38130 (MO); Río
Oeste, S of Al 5 uh n & Anna 2
anal Асе um of Pi dic Road, 19 km NW of
Gamboa, 25-50 m Ans & J. D. Smith 11070 (MO, US);
pe San Lorenzo n & Valentine 7001 re A 40 m,
o Island,
S); Wetmore & Woodwo
(GH); Welch 19833 (МО, NY); Croat 6066 (MO); 11758
(MO); 9223 (MO); 5878 (MO); 10357 (MO); 12802 (MO);
10744 (MO); Starry 24 (MO); Netting 60 (MO); Hutchison
& Wright 2920 (NY, US); Croat 11744 (MO, SCZ); 10383
(MO, SCZ); 7129 (MO, SCZ); Madden Forest Preserve,
vic. George W. Green Park, Welch 19853 (MO, NY, RSA);
Rio Pequení, near Salamanca Hydrographic Station, 70—
8°58'N,
3
E
T
кә
y
A
=
m
=
3
PON
a
Ы
~
+
near Palmas Bellas, Thoms 47 (МО); vic. of Miguel de la
Borda, Croat 9895 (МО); ens (MO); 10 mi. SW of Por-
tobelo, 2-4 mi. from coast, 10-200 m, Liesner 1103 (MO,
NY, US); Portobello, 5-100 m, Pittier 2424 (US); ушы
Field—Catival, Standley 30439 (US); Río Guanche, D'Arc
9724 (MO); Cour 6313 (MO); 100 m, 930'N, 79^39' Y.
3 (MO). Darién: 3 mi. N of Santa Fe, Tyson
al. 4616 (MO, SCZ); Canglón-Yaviza-Río Chucunaque,
TIm mi. E of Canglón, 50 m, 3°20'N, 77°50'W, Knapp &
Mallet 3944 (MO, US). Los ыштен —
80 m, Croat 34454 (MO). Panamá: Corozal Road, near
Panamá, Standley 26861 (US); "26841 (US); Río Tapia,
28112 (US); 4 mi. E of Panama City, road to Tocumen,
Tyson 3483 (FSU, MO, SCZ); Cerro Jefe region, D'Arcy et
al. 15520 (M
919 (GH): 92] (СН, US). San
Cartí road, Mile 10.3, 300 m, 9°20'N, 79°W, Croat & Zhu
76576 (MO). Veraguas: 5 of Santa Fe, ca. 450 m, Nee
8041 (МО); Bahfa Honda, Elmore H30 (US); са. 1 km
above Cañasas on road to Los Valles, 230 m, Croat 37063
(MO).
Philodendron hederaceum var. kirkbridei
Croat, var. nov. TYPE: Costa Rica. Alajuela:
eastern rim of Laguna Hule, ca. 1.5 km N of
Angeles, row road through pasturelands,
840—860 m, 8 July 1972, Luteyn 3350 (holo-
type, M0-2173874; isotype, DUKE). Figure
224
Planta hemiepiphytica; owe giam 2-28 cm longa, 1—
25 ст diam.; cataphylla 9 cm longa, incostata aut
niter 2- a ag decidua intacta; petiolus
(6.5)8-22 га cm longus, 3-6 mm diam.; lamina (11)16—
29.5(42) ст longa, e 3-23(35) cm а: inflorescentia
1; pedunculus 5.5-7.5 cm longus; spatha 14-24 ст longa,
lamina spathae ‘ios dde extus cremascenti vs. apicem,
intus pallidius viridi; pistilla 5- acai loculi 20-25-
ovulati; loculi cum ca. 24-seminibus
Hemiepiphyte; internodes 2-28 cm long, 1-2.5
cm diam., weakly flattened on one side with two
sharply raised ribs on the side above petiole, very
weakly sulcate on the opposite rounded side, prom-
inently ribbed throughout its circumference, the ribs
smooth or prominently warty, 2-28 cm long, 1-2.5
cm diam., medium to dark green, matte, drying red-
dish brown; cataphylls 6-19 cm long, unribbed or
sometimes rr зА 2-ribbed, deciduous intact; peti-
oles (6.5)8-22.5 cm long, B
(11)16-29.5(42) cm long, (8)10.3-23(35) cm wide;
upper surface medium to dark green, drying brown
to greenish brown, lower surface medium green,
glossy, drying brown to greenish brown; primary lat-
eral veins 3—4 per side, departing mdirib at a 45—
50? angle. INFLORESCENCES 1 per axil; peduncle
5.5-7.5 ст long; spathe 14—24 cm long, spathe
blades yellow-green becoming cream-colored toward
apex outside, lighter green inside; spadix stipitate
to 1 cm; 13-20 cm long; ovary 5-locular, 20-25
ovules per locule. INFRUCTESCENCE with about
24 seeds per locule.
Flowering in Philodendron hederaceum var. kirk-
bridei occurs during the late dry season and early
rainy season (probably as early as March and as
late as September) based on post-anthesis and early
fruiting collections. No collections have been made
464
Annals of the
Missouri Botanical Garden
at anthesis. Immature fruits have been collected in
April, July, and August.
Philodendron hederaceum var. kirkbridei ranges
from Costa Rica to Ecuador at (250)300 to 900 m
in Premontane wet forest and Tropical wet forest.
The variety has been collected only once in both
Suriname and Ecuador. It was collected in Ecuador
at Reserva ENDESA (0%5'N, 79°02’W), an area of
Premontane rain forest. It is to be expected in Co-
lombia.
s variety is recognized by its scandent habit,
conspicuously ribbed. Specimens from Panama
have stems also conspicuously and densely warty
while specimens in Costa Rica have stems reddish
brown and conspicuously ribbed but are smooth
rather than warty.
The variety differs from the autonymic variet
primarily in occurring at generally higher eleva-
tions in wetter forests and by its reddish brown,
conspicuously ribbed stems. In parts of Central
America and in South America where both species
occur, P. hederaceum var. kirkbridei occurs usually
above 500 m in Premontane wet and Tropical wet
forests. In contrast, P. hederaceum var. hederaceum
has stems that dry smooth and green and occur
primarily in tropical moist forest. Though there is
overlap in the elevation range and even life zone
th may sometimes be found in Premontane wet
forest), the two varieties do not occur in the same
sites and can be readily distinguished by their
stems.
Philodendron hederaceum var. kirkbridei is
named in honor of Joseph Kirkbride (BARC), who
first collected the taxon in 1968 on Cerro Caracoral
in Coclé Province in Panama while a graduate stu-
dent at the Missouri Botanical Garden.
Additional specimens examined. COSTA RICA. Ala-
juela: Cafias-Upala, 8 km N of Bijagua, 300 m, Croat
36498 (MO); Cariblanco, 900 m, 10%15'N, 84?11"W, Lent
3591 (F). Heredia: 3 mi. S of Vener ES m, Croat
35837 (MO). PANAMA. Coclé: , 9'39'N,
80°36'W, Croat 68758 (MO, PMA, m Cora] Caracoral,
Pons 1098 (MO); Alto Calvario, 9.4 km a
opé, 750—900 m, Croat 44741 (MO); La Mesa, a. El
Valle de Antón, 900-930 m, Croat 37471 (MO); Cerro
Gaital, 900-1000 m, 8°37'N, 80°08’W, Croat 67236
Т Colón: Río Piedras, Río Piedras Lumber R оад, 6.7
i. E of Sabanitas, 250 m, 9*22'30"N, 79*41'30"W, Croat
75158 (MO, PMA). Panamá: El Llano-Cartí Road, p
above Pan-American Highway, т 500 т, Croa
22909 (MO); Mile 7, near El Llano, 460 m, 919'N,
79°59'W, Croat 75108 (MO, PMA, US): Km 12-16, Ken-
nedy et al. 3146 (MO, US). San Blas: 1 mi. S of Nusa-
gandi, 9 mi. N of Interamerican Highway, 350 m, 9%20'N,
79°W, Croat & Zhu 77001 (CM, MO).
ECUADOR. Pichincha: онов Quito and Puerto
Quito, Km 113, 800 m, 0%5'N, 79°2'W, Rodríguez 282
(MO).
Philodendron hederaceum var. oxyeardium
Schott) Croat, stat. et. comb. nov. Basionym
mr Gs: Bunting, Gentes E ud
TYPE: Mexico: exact o bs own
EE ype, W? lost). Schott ic. 2714 (neotype,
here designated, W). [See fig. 17 in Bunting,
1968.]
Internodes smooth, sometimes with 2 ridges or
weakly angular on one side, semiglossy, 5-26 cm
long, to 1 cm diam., usually longer than broad, dry-
ing greenish to pale brownish green; cataphylls 19
cm long, unribbed, green; petioles 8-23 cm long;
blades broadly ovate, semiglossy, long-acuminate
to abruptly acuminate at apex, cordate at base, 26-
32 cm long, 18-22 cm wide (ca. 1.5 times longer
than wide), 2.5-3 times longer than petiole, broad-
est near the middle, upper surface bright green,
lower surface yellow-green; sinus narrowly trian-
gular to closed, to 5 cm deep; basal veins 2-3 per
side, with 1-2 free to base, and with the first and
or second free to base, 2-3 coalesced to 1 cm; pri-
mary lateral veins 2-3 per side, departing midrib
at a 45” angle. INFLORESCENCES (based on
dried specimen) with peduncle 4 cm long, 7 mm
diam.; spathe 14 cm long; spathe blade 6.5 cm
long; spathe tube 7.5 cm long; spadix sessile, су-
lindrical, 12 cm long; pistillate portion 4 cm long;
staminate portion 8 cm long; fertile staminate por-
tion drying reddish brown.
Flowering in Philodendron hederaceum var. 0%-
ycardium is poorly known. A single fertile collec-
tion with immature fruits in January was seen.
Philodendron hederaceum var. oxycardium 15 ap-
parently endemic to Mexico, known only from the
Gulf slope in the states of Jalisco, Nayarit, Oaxaca,
San Luis Potosf, and Veracruz at 120 to 580 m,
principally in areas of Tropical moist forest. Material
of P. hederaceum [as P. scandens] collected in Ja-
maica (Bunting, 1968) is sterile.
ron hederaceum var. oxycardium is
similar in appearance to variety hederaceum, слона
that leaves of the former are firmer in texture ап
glossy in both the juvenile and adult forms. In out-
line, the leaf blades of P. hederaceum var. охуса"
dium are often somewhat broader and more abrupt-
ly acuminate than for variety hederaceum (Bunting,
1968). The epidermal cells have a more OF ess
Volume 84, Number 3
1997
Croat 465
Philodendron Subgenus Philodendron
flattened epidermis on the upper surface (Bunting,
1968), which causes it to be glossy. In contrast, the
epidermal cells of P. hederaceum var. hederaceum
are mounded and sometimes conical, giving leaves
of that variety their characteristic velvety sheen.
Though Bunting (1968) treated this taxon as a
subspecies, it appears to overlap geographically
with the typical material so it will be treated here
as a variety.
Additional specimens examined. EXICO. Nayarit:
uis Potosi: N of Tamazun-
chale, Clark 7408 (MO, dem Evene 120 m,
лайм & Reko 334 silla Veracruz: 5.7–6 mi. from Ca-
on road to Sontecomapan, ca. 380 m m, Moore *
coni 8937 (BH); just а дн on road fro
ecolutla, Moore & Bunting 53 (BH); NW of она.
Puente Nacional, slopes of barranca de Santa Мапа : across
sy from Hacienda El Mirador, Moore & Bunting
69 (BH): Dist. Papantla, Kelly 16 (BH).
Philodendron heleniae Croat, sp. nov. TYPE:
anama. San Blas: El Llano—Cartf road, 14 km
N of Pan-American Hwy., 300 m, 915'N,
79°W, 12 July 1988, Cris 69244 (holotype,
MO-3599872; isotypes, B, CAS, COL, CR, F,
K, MEXU, NY, PMA, SCZ, US, VEN). Figures
227-230.
Planta ренти Bag возите raro terrestris; inter-
zn ia 4-12 ст m diam.; cataphylla 15-36
m longa, 1-2-costata, qe petiolus teres aut D-for-
зш 14–41.5 ст longus, 6–10 тт diam.; lamina an-
guste ovata, 25—52 cm longa, 6–26 cm lata, in sicco fla-
vibrunnea; es 2-10; pedunculus 2–6 cm
ongus, 4-6 mm diam.; spatha 5-10 cm longa, rubrimar-
ronina; i рейин extus pallidiori vs. apicem, intus
alba, dilute suffusa marronina; tubo spathae intus a
marronino; pistilla (5)6-9-locularia; Јосић (1)3-4-ovulati.
Usually hemiepiphytic, rarely terrestrial; vining
or appressed-climbing, stem appressed-climbing to
somewhat scandent, sap tannish; internodes scurfy,
4-12 ст long, 2-3 cm diam., much longer than
broad, medium green, drying reddish brown, epi-
dermis smooth or irregularly folded and ridged,
sometimes fissured with minute cracks perpendic-
ar to axis, drying with folds longitudinally and
usually with fissures transversely; roots dark brown,
to 50 cm long, 1-2 mm diam.; cataphylls thin,
15-36 cm long, l-ribbed to sharply l-ribbed, or
sharply 2-ribbed, green, tinged red, speckled vio-
let-purple, deciduous, rounded, apiculate at aen
petioles 14—41.5 cm long, 6-10 mm diam., + te-
rele or to slightly thicker than broad, or bluntly
D-shaped, firm, medium green, tinged violet-purple
at base and apex, bluntly flattened adaxially; gen-
iculum 9 mm long, thicker and paler than petiole;
blades narrowly ovate, subcoriaceous to coria-
ceous, strongly bicolorous, acuminate at apex (the
acumen sometimes apiculate, inrolled, 5 mm long),
usually weakly subcordate, sometimes obtuse, trun-
cate or rounded at base, 25-52 cm long, 6-26 cm
wide (2-3.37(4.2) times longer than wide), (1.1-
1.8(2.30) times longer than petiole), upper surface
dark green, semiglossy, drying yellow-brown, lower
surface much paler and with dark secretory canals,
weakly glossy to matte, drying yellowish to reddish
brown; sinus arcuate, 1-1.5 cm deep when present;
midrib broadly convex or weakly raised to flat, pale
green above, bluntly acute and thicker than broad,
reddish slightly paler than surface below; basal
veins lacking; primary lateral veins (6—7)8—10(11—
13) per side, departing midrib at a 30—40? angle,
weakly and narrowly sunken above, raised and pal-
er than surface below; minor veins weakly visible,
arising from the midrib only, often interspersed with
intermittent secretory ducts, the surface often mi-
nutely granular below. INFLORESCENCES 2-10
per axil; peduncle 2-6 cm long, 4—6 mm diam.,
green, tinged red to reddish; spathe recurved
(curved downward), semiglossy, 5-10 cm long (1.2—
2.5 times longer than peduncle), reddish maroon
throughout; spathe blade paler toward apex outside,
white, weakly tinged maroon inside; spathe tube 3—
4 cm long, dark maroon inside; spadix sessile; cy-
lindrical to clavate, protruding forward at anthesis,
5.8-7.5 ст long, + uniform throughout; pistillate
portion na green to yellowish green, cylindrical
to slightly ovoid, 2-3.5 cm long, 5-7 mm diam.
throughout; staminate portion cream to white, cy-
lindrical or clavate, 3.4—5 cm long, 6-9 mm diam.
at base, 9—10 mm diam. at middle, 5-9 mm diam.
са. 1 cm from apex, broadest at or above the 9
dle, or equally broad throughout, as broad as
slightly broader than the pistillate portion; кь
staminate portion not detectable; pistils 1.4—1.
mm long, 0.5-1.4 mm diam.; ovary (5)6-9-locular,
0.7 mm long, 0.7 mm diam., locules 0.9-1.2 mm
long, 0.3-0.4 mm diam., ovule sac 0.4-1 mm long,
with sub-basal placentation; ovules (1)3—4 per loc-
ule, 1-seriate, contained within translucent оуше
sac, 0.2-0.4 mm long, longer than funicle; funicle
0.1–0.2 mm long (can be pulled free to base), style
0.2-0.4 mm long, 0.4-0.6 mm fg sh similar to
style type B; stylar canals tiny, difficult to see
emerging; style apex flat; stigma ‘discoid or subdis-
coid, unlobed, 0.4—0.5 mm фат., 0.1 mm
high, covering entire style apex pu for the cen-
ter; the an m truncate, margins 4—6-sided,
sometimes кайр» рей thecae Med 0.1-
466
Annals of the
Missouri Botanical Garden
0.3 mm wide, + parallel to one another. INFRUC-
TESCENCE with berries white, 3.3 mm long, 2.4
mm diam.; seeds ca. 6 per jota reddish brown,
1-1.1 mm long, 0.3-0.5 mm
Flowering in Philodendron unio occurs in
Panama during the late dry season and much of the
rainy season, March through October (except April,
May, and September). South American collections
broaden that range to include the whole year (Jan-
uary through December, except May and Septem-
ber). Post-anthesis collections have been made
from June to October.
Philodendron heleniae ranges from Panama (to
be expected in the Cordillera de Talamanca in east-
етп Costa Rica) to Ecuador, from 20 to about
1040(–1450) m elevation in Tropical wet forest and
Premontane rain forest in Panama, Colombia, and
Ecuador. In addition, it occurs in pluvial wet forest
in Colombia
Philoden. i heleniae is a member of P. sect.
Calostigma subsect. Oligocarpidium. This species
is characterized by its scandent habit; internodes
much longer than broad drying with longitudinal
folds and usually transverse fissures; petioles terete
to bluntly D-shaped; and narrowly ovate, subcor-
date, yellow-brown-drying blades with dark secre-
tory canals visible on the lower surface. Especially
characteristic are the clusters of 2-10 small inflo-
rescences with externally red spathes.
Philodendron heleniae can be confused with P.
lentii, which also comprises more or less scandent
plants with subcordate leaf blades. That species
differs, however, in having one to two large, mostly
white inflorescences per axil and more broadly
ovate blades drying with many, pale, sunken veins
on the upper surface and no dark secretory vessels
on the lower surface.
Specimens from the Pacific slope of South Amer-
ica (e.g., Croat 58424, Tipaz et al. 1318) are on
average larger, with leaves ranging up to 60 cm
long and 30 cm wide and with spathes occasionally
to 12 cm long. Still, these probably do not differ
sufficiently to represent even another subspecies.
Lawrance 817 from Boyacá Department at El
Humbo (at 914 m in the Río Magdalena drainage)
in Colombia closely matches the material from the
Pacific slope. If it proves to be P. heleniae it would
be the first collection from the Eastern Cordillera
of the Andes.
Some South American collections from the Am-
azon basin may belong to this species. These col-
lections are from both Colombia (Pipoly et al.
15892, 16027 in Amazonas) and Ecuador (Gudiño
1160 and Thomas & Ríos 6708 in Pastaza Province
and Cerón 3360 and Bennett et al. 4526 in Napo
Province). However, most of the collections from
the Amazon basin do not mention spathe color, al-
though Gudifio 1160 indicates the spathes to be
greenish red.
Particularly interesting is Thomas & Ríos 6708,
with spathes described as becoming creamy white,
something that never happens with Central Amer-
ican material of Philodendron heleniae.
The species was first collected in 1970 by Jim
Luteyn and Helen Kennedy in the vicinity of El
Valle in Coclé Province, Panama. It is named in
honor of one of the collectors, Helen Kennedy, who
at the time worked for the Missouri Botanical Gar-
den as the Curator of Summit Herbarium.
Additional specimens examined. PANAMA. Восав
6.6 mi. N of bridge over Fortuna La
82°18' "W, Croat 66780 (MO). Coclé: El Valle region, La
Mesa, 800-900 m, 8°38'N, 80°09'W, Croat 67154 d
CM, KYO, L, = МО, PMA, TEX, US); El Valle, 5
mi. N of town, 2200 ft., Hammel & Kress 13427 (DUKE);
(DUKE); 880 m, Croat 37576 (MO); 860 m,
80°08'W, Croat & Zhu 76738 (AAU, CM, DUKE, 2 мо,
5, SEL, TEX); slopes of Cerro Gaital, 800-900 m, 8°37'N,
80°07'W, McPherson 11209 (L, MO, NY, P). mb Río
Tuquesa, 2 km f nental Divide, vic. of
US). Pan
above desc ideam Mia. 325—350 m, Croat 67351
(MO, NY, PMA, US, WIS); Mile 6.8, 350 m, 49106 (MO);
Mis 10, 330 m, 33778 (F, MO), 33817 (MO); Mile 7, 460
19'N, 79°59'W, 75114 (CAS, мау 550 т, ша
Ten W, 60502 (MO); Mile 10.6 400 m, 9°17'N,
78°58'W, Miller et al. 869 (HUA, мо, PMA): El Valle de
Madroño-La Saena, 2.5 mi. N of El Valle de Madroño,
180 m, 9°14’ 25'N. 79°05'W, Croat & Zhu 77048A (MO);
Cerro етен е сънна н n Blas”],
т, 9°17'N, 79°17'%, Hi & de Nevers 13553 (MO).
San Blas ат is C1 Ro. —5 mi. N of Nusagandi,
250-300 m, 9°16'N, 79°W, Thompson "4660 (СМ, MO);
Nusagandi-Cartí Road, 400 m, 9°18'N, 78 8°58" W, T.
Donagh et al. 93 (BM, мох са 79
10. lm e N of Intesemerican Highway, then ca.
aseo Mariska near road, 300 m, 920'N, a јон &
Zhu 77020 (MO); El Pia Road, 23-29 km from
ап-Атепсап Highwa , 922'N, 7869
Knapp 1830 (MO); 1840 мө, PMA); Km 15, 350 т,
9°20'N, 78%58'W, Galdames et al. 1288 (PMA); km 17-
19, 350 m, 9°19'N, 78°55'W, de Nevers & Herrera qa
(CM, MO, US); Cerro Obu, 400-500 m evers el а
8068 (MO, NY). Veraguas: Santa Fe region, Alto dient
Calovébora, Río Dos Bocas Valley, 350-400 m Croa
27370 (F, K, MO, PMA); 27497 M 0).
COLOMBIA. Antioquia: Medellín-Bogotá, e "y.
from entrance to San yes 800 = 5°50'N, 74 7 74
0); 4
m, 6°18'N, (АД he & Roldán 1486 (HUA); Quebrada
Honda, Río Gua Finca
, 1250 m, = et al. 955 (COL); San Carlos, Fine
Volume 84, Number 3
1997
Croat 467
Philodendron Subgenus Philodendron
“E] Desespero,” Alto de Затапа, near Miraflores, 4 hrs.
SW of Alto de Samaná, 710—820 m, 6%5'N, 74°56’ W, Cal-
lejas et al. 8628 (NY); San с Ba ii узан» ap La Te-
baida, 1010-1060 m, 6*8'N, 75^10'W, Callejas et al. 4015
(MO); Sector Río Samaná—Río Айы ‚ 750 m, Hernández et
al. 551A (HUA); San Luis—Puerto ‘asks, SE of ced
S of San Carlos, ca. 3 km SE of San Luis, ca. 800 m
5°57'№, 74^57'W, Croat 52038 (COL, MO, NY); Gómez
Plata—Yolombo border, Sector La Cancana, along Río Por-
ce, km 14 via Amalfi, 1030 m, Callejas et al. 2338 (MO);
Sonson, Rioverde region, Hacienda “La Soledad,” on the
pes of Quebrada “Curubital,” Gutiérrez 35552 (UC);
Amalfi-F layers NE of Salazar, 23-26.5 km from center
of Amalfi, 00 m, 6°58'N,
et al. 4042 ИО). Сћосб: Bahia Solano, 5 of a
100 m, 5°13'30"N, 76°21'30’W, Croat 57458 но,
COL, JAUM, MO, US); са. 2 km Е of Playa де Oro, са.
200 m, 5°20'N, 76°23'W, Croat 57424 (CHOCO, COL,
yi Mecana (N of Bahía Solano), Quebrada Resaquita,
0 m, berita 2536 (MO); cee i es 6.6 km 5 of
Quibd <100 т one 6°37'W, Croat & ee
tpm (MO); Rio Pat 8 cum imas-Pató, Serranía de
Baudó, 10 km SE of des 5°17'N, 76^45' W, Cr = и
MO), Croat 56107 (MO); Rio Sucio, Cerro del Cuchillo,
zona de Urabá, 520 m, Cárdenas 4
và de Utría, Río San Pichf, oie m, ын
—
—
sera del Taparal, 5-20 m
y TIO
MO); 22.3 km beyond Pulpapel Headquarters,
d Y 7707", Croat 61275 (СОУС, MO). ECUADOR.
Car Tulcán, Reserva Indígena Awá, 650-100 m, 1°N,
T8 24^ W, Tipaz et al. 1318 (MO, мав, Esmeraldas:
Lita, 550-650 m, Madison et al. 4990 (F, K, SEL); Lita—
San Lorenzo, Km 18, re 78°28W, dud 72389 (MO);
San Lorenzo, Mun. aurte, Reserva га lee a Awá, 3
m, VION, 78232", Траг et al. 2 MO); Río Jordán,
NE of Las Golondrinas, near San Is Yel. 300 m
79°12'W, Palacios 11485 (MO, QCNE). на TE
600 m, Cobb 29 (MO). Pichincha: Río Verde, 2 km SE
of Santo Domingo de Los Colorados in Coop
Marta #2, 530 m, Dodson 7403 (MO, ОСМЕ); Santo De.
mingo de p Colorados, Río Baba, 28 km S, 350 m, Dod-
son e Thien 1187 (MO).
У
Philodendron immixtum Croat, sp. nov. ТҮРЕ:
anama. Panamá: Comarca de San Blas, Río
Playón Chico, 50-100 m, H. Herrera et al.
1167 (holotype, MO-4256423; isotypes, AAU,
K, MEXU, PMA, US). Figures 233-236.
5
mm) 1-1.5 cm dia ber actue D-form
ad debiliter io Pee an ovata vel ad
Я M» 16-36 cm lon a, 5-12 cm lata; i inflorescentia sol-
a; pedunculus 5-13. cm a 9-19 cm longa,
aca albo-viridis vel alba, tubo. сан extus, rubo ve
De "eir lat internodia (2)10-15 cm M: б
; petiolus
purpureo intus; spad мы ст longus; ovarium (3)4—
6(8)-loculare; loculi 12) ovulat
Hemiepiphytic; stem scandent, sap reddish, wa-
tery; internodes long, slender, + glaucous, (2)10—
15 cm long, (0.7)1-1.5 cm diam., much longer than
broad, dark olive-green matte to weakly glossy, be-
coming gray-green, drying light brown, epidermis
sometimes cracking with loose сан roots са. 5
per node, to 15 ст long, 2-3 mm ., tannish
gray, drying reddish brown, + uio) du cata-
phylls somewhat spongy, to 17 cm long, unribbed,
weakly 2-ribbed or sharply (1—)2-ribbed (ribs 2 mm
high), pale green, dark short-lineate, drying pale
brown to yellowish brown, usually some deciduous,
sometimes persisting somewhat intact, eventually
fibrous, obtuse at apex, margins paler, yellowish;
petioles 7.5—16(21) cm long, 2-10 mm diam., sub-
terete, sharply D-shaped to slightly flattened adax-
ially, rounded abaxially, surface dark green-striate,
matte to semiglossy, shortly dark lineate, with dark
green ring around apex; blades narrowly ovate to
ovate-elliptic, or rarely oblong-elliptic, subcoria-
ceous, semiglossy, strongly bicolorous, abruptl
apiculate, 2-4 mm long), SEU Uu rarely round-
ed at base, 16–36 cm long, 5-12 cm wide (1.9—3.7
times longer than wide), (133.188) times longer
than petiole), about twice as long as petiole, broad-
est below middle, margins narrowly hyaline, upper
surface drying green to grayish to brownish green;
posterior lobes short, about as broad as long and
held close to petiole; sinus shallow, 0-2.5(3) cm
deep; midrib flat to weakly convex, concolorous or
slightly paler than surface above, convex to bluntly
acute, paler than surface below; basal veins lack-
ing; primary lateral veins 4-8 per side, departing
midrib at a 50-60” angle, + straight to weakly ar-
cuate to the margins, sunken and paler than surface
above, weakly raised and darker than surface be-
low; interprimary veins weakly raised and darker
than surface; minor veins distinct to weakly visible,
darker than surface below, arising from both the
midrib and primary lateral veins. INFLORES-
CENCES 1 per axil; peduncle 5-13 cm long, 3—
7(11) mm diam., dark green; spathe 9-19 cm long,
(1.1-2.1 times longer than peduncle), green, dense-
ly and obscurely short-lineate, narrowly acuminate
at apex; spathe blade pale greenish white to white
outside, pale greenish white, dark lineate inside;
resin canals appearing as continuous lines, drying
reddish brown, resin droplets forming on spathe
surface within; spathe tube dark green in back, pal-
er in front on open edges outside, red or violet-
purple inside; spadix gradually tapered to apex,
468
Annals of the
Missouri Botanical Garden
protruding out of the end of spathe, bluntly rounded
to obtuse at apex, 10-11 cm long, broadest above
the middle, constricted slightly above sterile sta-
minate portion; pistillate portion pale green, cylin-
drical, 3.4-4.2 cm long, 3.7 cm long in front, 2.7
cm long in back, 9-11 mm diam. at apex, 10-11
mm diam. at middle, 9-11 mm wide at base; sta-
minate portion 6.8-7.5 cm long; fertile staminate
portion white, cylindrical to clavate, 9-12 mm
diam. at base, 1.1-1.3 cm diam. at middle, 7-12
mm diam. ca. 1 cm from apex, broadest at the mid-
dle, broader than the pistillate portion, broader
than the sterile portion; sterile staminate portion as
broad as, narrower than, or broader than the pistil-
late portion, + 12 mm diam.; pistils 1.9-2.3 mm
long, 1.3-1.9 mm diam.; ovary (3)4—6(8)-locular,
walls sometimes embedded with granular, crystal-
like particles, locules 0.7-1.3 mm long, 0.4—0.7
mm diam., ovule sac 0.6-1.3 mm long, with sub-
basal placentation; ovules 1(2) per locule, con-
tained within transparent ovule sac, 0. m
long, longer than funicle; funicle 0.2-0.6 mm long
(can be pulled free to base), style 0.3 mm long,
0.9-1.4 mm diam., similar to style type B; drum
subdiscoid, Баг 0.7-0.9 mm diam., 0.2
high; the androecium truncate, margins 4—6- side:
thecae oblong to slightly elliptical, 0.4 mm wide,
+ parallel to one another; sterile staminate flowers
4—6-sided, 1-1.8 mm long, 1.1-1.7 mm wide. IN-
FRUCTESCENCE with spadix yellowish orange, 8
cm long, 3 cm wide; berries concave at apex; seeds
1 per locule, 1.5-1.8 mm long, 0.6-0.7 mm diam.,
yellow-brown.
Flowering in Philodendron immixtum occurs
during the last half of the dry season and the first
part of the rainy season, March through May, with
post-anthesis inflorescences seen from March
through August, and fruits seen in July.
Philodendron immixtum ranges from Panama to
Colombia (Golfo de Џтађа), occurring at or near sea
level (to ca. 140 m elevation), primarily along the
Atlantic slope in Colón and Panamá Provinces but
also in Panamá Province on the Pacific slope along
the Río Маје. It occurs in Tropical wet forest, Pre-
montane wet forest, and in wetter parts of Tropical
moist forest life zones.
Philodendron immixtum is a member of P. sect.
Calostigma subsect. Glossophyllum ser. Glossophyl-
lum. The species is characterized by its scandent
habit; long, slender, mostly light-brown-drying in-
ternodes; slightly flattened petioles (about one-half
or less as long as the blade); narrowly ovate to
ovate-elliptic, green-drying blades, which are usu-
ally cordulate at the base; as well as by the solitary
inflorescence with a green spathe, the base white
on the blade at anthesis and red within on the tube.
The species has been confused with P. inconcin-
num Schott (1856) (hence the name “immixtum,”
meaning “intermixed or mingled with”) from Ven-
ezuela (Standley, 1944; Croat, 1978). The Schott
paintings of P. inconcinnum show a single unat-
tached leaf and unattached inflorescences and
probably represent the same species as P. sphal-
erum Schott (1860). Schott paintings of P. sphaler-
um show some leaves that are virtually identical to
the above-mentioned drawings. Philodendron
sphalerum differs from P. immixtum in having sev-
eral small inflorescences per axil (rather than one,
somewhat larger inflorescence for P. inconcinnum).
Schott's original description of P. sphalerum did not
mention an inflorescence, so his drawing of that
must have come later than his description. Schott's
original description of P. inconcinnum cited Vene-
zuela as the type locality, not Panama, and the solid
greenish inner spathe surface depicted in his paint-
ings corresponds well to P. sphalerum but not to the
material in Panama, which has a reddish interior
spathe surface.
Philodendron immixtum has also been confused
with P. ligulatum. The latter species differs in hav-
ing blades that are typically more nearly oblong-
elliptic to narrowly obovate-elliptic, frequently
broadest above the middle, and drying usually
much blackened. In addition, it has petioles that
bear a conspicuous purple ring around the petiole
at the apex. In contrast, the blades of P. immixtum
typically dry grayish to brownish green and are
broadest frequently below the middle. The species
also differs in having petioles that lack the purple
ring at the apex.
A number of sterile or immature collections from
Nicaragua may represent this species, but if so it
is curious that the species has not been collected
in Costa Rica (or for that matter, not further west
in Panama than Miguel de La Borda in Colón Prov-
ince).
Additional specimens examined. PANAMA. ps
Area: Balboa, Standley 28542 je Ft. ^ Rasa 2861 :
ге road between Сашп Locks and Ft. Sherman, са.
E of Ft. Ек <25 m, 971 ON. 79%57'30"W, Croat
TEX, US); Ft. Sher-
0);
Aviles 89 (F); Fi irchil 3079 (US); Parque eee Sob-
erania, 140 m, 9°06'20"N, 79°37'20"W, Croat & 2%
76987 (MO). Colón: 0.25 mi. N of María deseri on E
to Portobelo, 71360 (F, MO); хаа Indio—Miguel де la a
a, along ocean trail, O m, 36898 (MO); vic. of San enn
ie la Borda, 9896 (MICH, MO, NY, PMA, SCZ); 2 m.
of Portobelo, 33574 (BR, DUKE, MO); Ades Bellas-Sal-
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
ud, near sea level, 36865 (MO); 4.5 km SW of Раћа, 0-5
m, Nee 11713 (MO); 3 km S of Pifia, 50 m, Sullivan 137
(MO); Portobelo, Croat & Porter 15613 (MO); Nuevo To-
0 i i
, along Rio Perisenico,
100 m, 8°O1'N, 77744", Croat & Zhu 77093 (CM, MO);
77162 (CM, MEXU, MO, PMA, US). Panamá: Río Maje,
W of Bayano Lake, ca. 2 mi. upstream, 30—60 m, Croat
34592 (MO).
COLOMBIA. Chocó. Mpio. Acandí, Golfo de Urabá,
Bahía Zardí, 0-5 m, 8°23'N, 77%07'W, Betancur et al.
1238 (HUA, MO).
Philodendron jacquinii Schott, Syn. Aroid. 90.
1856. TYPE: Plate 152 in Jacq., Select Stirp.
Amer. Hist. 1763 (holotype). Figures 231—232,
231, 241-242.
oe i uius Schott, ee Bot. Z. 8
1858 (as “Р. hoffmanit”). : Costa Hg - $4
mann s.n. (hol otype, B epica Schott ic. 2507
(neotype, here designated, W).
Philodendron deviatum anes Bonplandia 7: 29. 1859.
PE: Venezuela, Fendler 1329 (lectotype, here
designated, K).
Philo emen erlansoni I. M. Johnston, Pig acia 8: 89,
. TYPE: Panama. Panamá: Perlas Archi-
pelago, ca José Island, along — bod one-half
mi. E of Red Hill, 31 Dec. 1945, Johnston 967 (ho-
type, GH).
london apocarpum Matuda, Madrofio 10: 171. 1950.
TX exico. Chiapas: along Río lene at Gil-
guero, us km E of Escuintla, Chia ca. 200 m,
10 Aug. 1948, Matuda 18313 oe MEXU; iso-
types, F, MEXU).
Hemiepiphytic vine; stem scandent, green to
gray-brown, glossy, setose-pubescent; leaf scars
conspicuous, 1 cm long, 1 cm wide; internodes
elongate, 6-25 cm long, 0.8-1.5 cm diam., longer
than broad, brownish gray, semiglossy, weakly as-
perous, epidermis paper-thin, sometimes peeling
with age, drying gray-green to yellowish green,
matte, sometimes almost blackened, larger stems
drying yellow-brown and finely ridged; roots few
Per node, short, ca. 1 mm diam., light brown,
sparsely scaly; cataphylls unribbed, itia: ари
10 cm long, light green, translucent, drying cream-
colored, deciduous intact; petioles 7.5-44.5 cm
long, 5-10 mm diam. ., subterete, medium green,
weakly glossy, narrowly and obtusely sulcate adax-
lally, surface setose-pubescent, often puberulous;
blades broadly ovate-cordate, thin, chartaceous,
acute to shortly acuminate, sometimes mucronate
at apex (the acumen sometimes apiculate, 1-3 mm
long), cordate at base, (10.5)11.5-39 cm long, 7.5-
28 cm wid e (0.99-1.8 times longer than wide),
(0.6-1.85 times longer than petiole), usually about
equal in length to petiole, broadest near point of
petiole attachment, upper surface dark green, matte
to subvelvety to weakly glossy, lower surface mod-
erately paler, weakly glossy to almost matte, mod-
erately paler; anterior lobe 8.3-27 cm long, 9-28
cm wide (1.6-3.4 times longer than posterior
lobes); posterior lobes + rounded, 3-13.5 cm long,
3.4-13.6 cm wide, directed inward at maturity,
rounded or obtuse; sinus + narrowly parabolic;
midrib broadly convex, concolorous above, broadly
convex to round-raised, often setose-pubescent
near base, slightly paler below; basal veins 3-6 per
side, with 0—1 free to base, 1-2 coalesced 0.5-2.5
cm; posterior rib not naked or briefly naked for 0.5
cm (rarely to 1.5 cm); primary lateral veins 2-
3(4—5) per side, departing midrib at a 45-55” an-
gle, + straight to the margins, prominently im-
pressed to weakly convex above, convex to round-
raised below; minor veins arising from both the
midrib and primary lateral veins; tertiary veins ob-
scurely visible to raised and conspicuous, darker
than surface below. INFLORESCENCES erect, 1
per axil; peduncle (2)4-15 cm long, (2)3-6(7-13)
mm diam., green, dark-striate, glabrous; spathe
10-16.5 ст long, (0.96-2.3(2.7-3) times longer
than peduncle); spathe blade green outside, red in-
side; spathe tube inflated bulbous, green outside, 4
cm long, white or green inside; spadix sessile; 8—
13 cm long, narrowly rounded at apex; pistillate
portion 3-3.5 cm long, 3.5 cm long in front, 3 cm
long in back, 3.5 cm diam. throughout; staminate
portion 9.2 cm long; Tue staminate portion 1.
1.6 cm diam. at base, 1.1 cm diam. at middle, 1
cm diam. са. 1 cm from apex; sterile staminate por-
tion narrower than the pistillate portion, 1.2-1.5 cm
diam.; pistils 6.5—7.2 mm long, 2 mm diam.; ovary
4-locular, locules ca. 1.3 mm long, ca. 0.6 mm
diam., with sub-basal placentation; ovules 2 per
locule, contained within transparent, gelatinous
matrix (no true envelope), 0.6-1 mm long, as long
as or longer than funicle; funicle 0.4 mm long (can
be pulled free to base), style similar to style type
B; style apex quite rounded; styles usually elon-
gated to 5 mm long, sometimes essentially sessile;
stigma hemispheroid to sometimes somewhat glob-
ular, lobed, 3.3 mm diam., 1.5 mm high, covering
entire style apex; the androecium truncate, pris-
matic, some oblong, margins irregularly 4—6-sided,
mm long; thecae oblong, 0.5 mm wide, +
contiguous; sterile staminate flowers irregularly 4—
5-sided, 2.2 mm wide. INFRUCTESCENCE usu-
ally pendent, often from leafless stems; peduncle
7-15 cm long, recurved; spathe falling free; spadix
(2.5)5-9(14) cm long, (1.5)2.5—7 cm wide; berries
greenish when immature, pale orange to red to red-
dish orange when mature, irregular, 1 cm long, 5
mm diam.; seeds r berry, white, ovoid, 4 mm
long, 3—4 mm diam., moderately sticky. JUVENILE
pand
|
470
Annals of the
Missouri Botanical Garden
plants with petiole margins vaginate-winged almost
to one-half their length; blades narrowly ovate-cor-
date.
Flowering in Philodendron jacquinii occurs in
August but most post-anthesis collections are from
the rainy season, mostly June through September
(but also November, January, and March). Immature
fruits are known from virtually throughout the year,
and mature fruits from March, June, and July.
There seems to be little phenological variation in
this species.
Philodendron jacquinii ranges from Mexico to
Panama on both slopes of the Continental Divide
to northern Colombia, Venezuela, and the Guianas,
as well as western Cuba and the Cayman Islands,
from sea level to 1500(2500)m elevation (mostly
below 300 m), mostly in Tropical moist forest but
also in Tropical dry forest, Premontane moist forest,
and Premontane wet forest life zones.
This species, the only member of P. sect. Macro-
gynium, is distinguished by occurring in moderate-
ly low, dry habitats, and by its scandent habit, elon-
gate internodes, setose-pubescent stems, petioles
are the generally swollen spathe tubes and
elongated styles, making the pistillate portion of the
spadix much broader than the staminate portion.
ilodendron jacquinii may be confused with P.
brevispathum, another species with scaly petioles
and thin blades with prominent posterior lobes. The
latter species differs in having ovate-triangular
blades with more slender, frequently flaring poste-
rior lobes and a reddish brown stem with simple or
branched scales lacking the elongated setae typical
of P. jacquinii, which has typically greenish stems.
In addition, the berries of P. jacquinii are pale or-
ange to reddish, and the spathe is reddish within
(vs. berries whitish and the spathe green within for
P. brevispathum).
For a discussion of the long-standing nomencla-
tural confusion involving P. hederaceum, see under
the latter name. Although Standley and Steyermark
(1958b), in their treatment of Araceae for the Flora
of Guatemala, had the taxonomy correct in regards
to this confusion, their illustration labeled P. hed-
eraceum (fig. 58) appears to be a mixture of the two
species, with the leaves of P. hederaceum and an
inflorescence of P. jacquinii. The former species
differs in lacking the setose-pubescence and scales
characteristic of P. jacquinii, and in having more
coriaceous blades as well as normal, short, closely
compacted pistils. The spathe tube of the latter is
also not markedly infla
A collection from El Salvador (Croat 42075) is
unusual in lacking the usual trichomes on the
stems.
Philodendron lundellii Bartlett ex Lundell (Bart-
lett, 1937) also corresponds to this species, but the
name was invalidly published (without a descrip-
tion or Latin diagnosis).
Additional specimens examined. BELIZE. Belize:
Mile 35, Croat 24784 (F, MO); Bel-
Est pe
Min MO). Co 98, Gentle 530 (CM, MICH); Cerro Maya
ns, Lowry’s Bight, Crane 510 (LL), 418 (LL, MO); Gen-
de 345 му. Orange Walk: Honey Camp, Lundel
N of herr ad Walk, Sutton et al. 122 (BM).
& Brant 31955 (B, MO). COSTA RICA. Alajuela. Oro-
tina, Valle del Tárcoles, 100 m, Ham быа & Стауит 1 9932
R, INB). Guanacaste: El Mira
b 1343 (INB, MO). Heredia: La Selva Field Station, ca.
m, Hammel 10500 (MO); 1.4 km NW of Puerto lic
m, А
FO
MO); j diet = е, 1500 10%
84748", Croat 61192 (CM, МО); 1300 1 ii pu & nd
7983 (CR, MO); Cantón de Buenos Aires, along Río Cei-
bo, Ujarrás, 500 m, 9714"М, 83 ea W, Grayum 10228 (CR,
INB, MO, US). San José: Mora, Co lón-Hacienda El Ro-
84°16" W, Hammel 18579 (INB, MO).
i n-
dez-Tacuba, 0-2 mi. NE an
00—450 m, Croat 42075 (MEXU, MO). GUAT ATEMALA.
Without locality: Heyde 463 (US); Aguilar 309 (F). Chi-
quimula: Río Chiquimula, Santa Bárbara- Petapilla, 4-6
mi. N of Chiquimula, 350—420 m, Standley 30245 (F);
Las Mamacas, 16 mi. S of Guatemala City on CA- 9
ft., Dziekanowski et a 3472 (UMO). Jalapa: Jal apa-San
Раб Pinula, 1400-1800 m, Standley 77051 (F). Jutia-
pa: Standley 75385 (F); 75695 (P mesi Ui We
a y 757 паре
Las Tunas, NW of Jutiapa, 850-900 m, Standley 76294
én: 31 k res, 175 m, Harmo mon & Daye
2797 (MO); La Libertad, Lundell 2551 (MICH); 1k
on old trail to Mahaquila, Contreras 2912 (LL); Tikal [^
tional Park, Lundell 15292 (LL); 16792 (y Contreras
343 (LL); Dolores, Contreras 3763 (LL). Retalhuleu:
Nueva Linda-Chaperico, 120 m, Standley am 4
Santa Rosa: SE of siam 1100- wer
77740 (F); Сиара, 900 m, ¡ras 78087 (P: a
pultura region, W of i € m, Sta
de
79331 (F); Chiapas, 3500 pp. Hey 3867 ab
аннга, ћоодрјат оѓ i Santa Cruz МА зар p ne
ry of Rio Usui), 560 se Pe s" w, pr "it
ling 93 (CM). Zacapa: trail кон п Rfo Hon
terfall, 250-400 m, Standley 720396 (F); pleri
ño APP A AAA E
Volume 84, Number 3
1997
Croat 471
Philodendron Subgenus Philodendron
Alejandria, summit of Sierra de Las Minas, Finca Alejan-
dria, 2500 m, Steyermark 30856 (F); Mpio. Ocozocoautla,
E edge of Selva del Ocote, 900 m, Breedlove 70760 (CAS,
MO). HONDURAS. Colón: Trujillo, Capuchin Monkey
әче m Saunders 523 (MO); Bonito Oriental-Limón
50 m of Río Piedra Blanca, 40 m,
15°46" 30'N, 85°41’ б Evans 1086 (МО). Сорап: 10 mi.
А о Га Еп 700 m, Croat 42517
Williams 15908 (F); Rfo Yeguare, E of
750 m, Standley 15448 (F); Rio de la Orilla region, SE of
El Zamorano, 900-950 m, Standley 22404 (F). Olancho:
Juticalpa, 380-480 m, Standley 1 7524 (F); Río Olancho,
Gualaco—San Bonito Oriental, 7.4 mi. NE of San Esteban,
m, 15°20'N, 85
ieba
Sula, SW of Tela, near border of Yoro and Atlántida, 50
m, 15°30'N, 87743'W, Croat & Hannon 64666 (MO).
MEXICO. Hahn s.n. (P). Campeche: Yucatán Peninsula,
Champoton, Steere 1742 (CM); Maskall, Gentle 1186 (CM,
Та — PE 1074 (F). Chiapas: Arria-
160 m, Moore & Bunting 8923
47'N, 93"18'W, Thompson
et al. 432 (CM, MO); 5 mi. N of Ocozocoautla, along road
i 544 fo Cintalapa,
Aguas Calientes, Escuintla, Matuda 18312 (NY); Esper-
anza, Escuintla, Matuda 18314 (MEXU, MO); Mpio. Ber-
riozábal, 5 km E
5 . Eo
Kimnach 527 (US); Río Grijalva, 10 km S of Hwy. 190 on
i, 1600 ft., en he 2811 (F).
Guerrero: Acapulco, Palmer 462 (US). Oaxaca: Tuxte-
pec—Matias Romero, <100 m, Croat 78724 (CHIP, MO);
7.5 mi. SW of Tuxtepec on road to Valle Nacional, Moore
& ne 8908 (HB); 14 mi. N of Puerto Escondido on
Rte. 131 1613-A7'N, 97°5-8'W,
Temazcal, Temazcal, 150 m, 18725'М, 96°2
& Merello 15454 (MO). Quintana Roo: Mpio. Chetumal,
6.5-7 km N of Tomás Garrido on the road joining Hwy.
186 W of Nicolas Bravo, 150 m, 18°6'N, 89°3’W, Saun-
ders et al. 9950 (MO ); 1 km E of Chanca Veracruz, É.
Cabrera £ H. Cabrera 4981 (MEXU). San Luis Potosí:
Tamazunchale, Edwards 549 (F, MO). Veracruz: Córdo-
cren, Ejido San José de Gracia below Penuela,
oore & Bunting 8877 (HB); San Lorenzo МЕ
22-75 m, si 50 (GH). Yucatán: Schott 4
пда, Gaume
. Boaco: Las Pitas, Сатоара, 400 т
12°28'N, 85°35'W, Moreno 10643 (MO); Rfo Las Cafias,
along Hwy. 33, ca. 3.1 km N of wy. 35 intersection, ca.
275 m, 12°38’N, 85°33'W, Stevens 5838 (MO). Chonta-
les: viu 1, ~ km SE of Juigalpa, Moore 1622 (MO).
Gri na de Apoyo, 110-180 m, 11%53'N,
86 OW. страница 11178 (MO). Masaya: Parque n
Volcán Masaya, N slope of Volcán epo ca. 375 m
" 107, Stevens 5267 (BM, MO); summit of
olcán Masaya, 500 m, Stevens 2950 (MO); Piedra Po
тада, 2 km E of Volcán Masaya, 250 m, Stevens
(BM, MO). Matagalpa: Ranchería, 11 km NE de Muy
Muy, ca. 280 т, 12746" М, 85731", Moreno 24483 (AAU,
MO); Río Yasica, Tuma, ca. 28 km NE Matagalpa, El Dia-
mante, са. 350—400 m, 13°04’N, 8546", Guzmán т x
915 (MO). Río San Psi: Boca de Sábalos, *La Tobo
70-90 m, 11?03-04'N, 84?28-29' W, Robles 1833 mois
Rivas: Tola—Las Salinas, El Coyol, с 3 km beyond
entrance of Hda. Miramar, ca. 11?23'N,
85°58' W, Stevens 9749 (BM, МО). Rivas-Carnso-Gra-
nada: Rio
. Zelaya: SW
Саров 0-2 m, 14%01'N, 8372 4'W, Sevens 7880 (MO);
Mpio. Siuna, Wany, Ortiz 59 (MO). PANAMA. Canal
Area: Balboa, Standley 29256 (US); ‘toad oe N of Es-
cobal, Croat 12458 (MO, NY, U); Rodman Marine Base,
Rodman Naval pe Supply Depot, W of Balboa,
0-80 m а 6'25"W, Howard 147 (MO); road
to Cerro
(MO); Gamboa, Croat 74755 (F, MO); Geologic Test Site
N of Paraíso, Croat 12977 (MO); near Summit Hills val
Course, Croat 10956 01 rds Hwy., vic. Summ
Pore ка € 69835 (AAU,
pod ач E of Summit Gar-
dens, Croat 9080 (MO); Barro teks Island, Shattuck
115 (Е, GH); Zee 4675 (MO, US); Wheeler 5, Shattuck
215 (MO); laboratory clearing, Croat 10261 (MO); 9259
(MO); Colorado Point, Croat 6138 (MO); Vista Alegre, Ze-
; gres River, vic. of Juan Mina, Flat
vic. of end of C-16 road, Blum 2243 (MO, SCZ); Ft. Sher-
man, Standley Sane (US); Madden Dam Area, Boy Scout
Road, Porter et al. 4062 (MO); Madden Forest, Las Cruces
Trail, Croat 11878 (MO); 140 m, 9*06'20"N, е 20"W,
Croat & Zhu 77072 (MO); Pipeline Road, 2 . N of
Gamboa, ca. 100 po се 65 2 (МО); Río кит road
K- ^ ДУР у al. 348 (GH, MO, US); Parafso, ~~
194 a Coclé: Penonomé and vic., 50-1000 ft.,
Georgia Cuipo Forest Site #2, 15 m, Duke 14258 (F, МО).
па Ocú, Ebinger 1090 Ў . Los Santos: са.
m S of Las Tablas, Burch et al. 1236 (MO, UC); Pocrí,
Doyer 1189A (MO); Las Tablas, Dwyer 1189 (MO). Pan-
ong road to Bique, 5 km SW of Arraiján, 20—40
m, a Na 7699 (MO, US); Panamá жы Козе 18505 (МУ,
US); road K-15 near Huile, vic. Gatún Lake, Smith et al.
3277 (F, US); El oer Gentry & Tyson 1727 (MO,
SCZ); Tocumen, Dwyer 4220 (MO); vic. Macambo, Croat
14911 (MO); ie Campana, Croat 12018 (MO, к
Penonomé ап ., 50-1000 ft., Williams 381 (NY); San
José Island pe " Archipelà ie. "Erlanson 234 (US); 402
beri US); Johnston 974 (GH, MO), 918 (GH), 12 (GH),
60 (GH).
Philodendron jefense Croat, sp. nov. TYPE: Pan-
ama. t Cerro Jefe, along road short of
50-800 m, 9"15'04"М, 79°30'04W,
Fac usen 10038 Пишите, MO-3475849;
isotypes, K, PMA, US). Figures 238, 239.
Planta hemiepiphytica; internodia brevia, in sicco
usque 3 ст Са cataphylla 20-24 cm longa, acute
2-costata, m па ара ЛИ et persistentia ut fibrae pal-
lide мери petiolus teres, 38-76 cm longus, 4 mm
472 Annals of th
Missouri Botanical Garden
diam.; lamina late ovata, 39-54 cm longa, (22.5)34—49 Philodendron jefense is a member of P. sect. Ca-
cm lata, cordata basi, in sicco canoviridis; inflorescentia
1; pedunculus 10-13 cm longus, 4—5 mm diam.; spatha
9.5-14 cm longa, omnino viridis, in sicco cum magnis
maculis albis; pan (6)7—8-locularia; locules cum 6
seminibus; bacc e
Hemiepiphytic; stem appressed-climbing; inter-
nodes short, to 3 cm diam.; roots moderately few
per node, drying to га mm diam., sharply ridged,
brown; cataphylls 20–24 cm long, sharply 2-ribbed,
persisting, promptly weathering to light brown,
semi-organized fibers with small, thin fragments of
epidermis persisting; petioles 38-76 cm long, 4
mm diam., terete, drying black; blades broadly
ovate, short acuminate at base, cordate at base, 39—
54 cm long, (22.5)34—49 cm wide (0.8-1.4 times
longer than wide), (0.5—0.6 times the petiole
length), about one-half as long as the petiole, mar-
gins sinuate, upper surface glossy, drying gray-
green and semiglossy, lower surface glossy, drying
minutely granular; anterior lobe 22-25 cm long,
24.5-29.7 cm wide (1.8-2 times longer than pos-
terior lobes); posterior lobes 12-14 cm long, 12.2—
13.2 ст wide, obtuse; midrib drying darker than
surface above; basal veins 5 pairs per side, first
and second free to base, the remainder coalesced
3—6 cm; posterior rib naked to 3-3.5 cm long; pri-
mary lateral veins 5-6 per side, departing midrib
at a 35—45" angle; minor veins frequently
branched, arising from both the midrib and primary
lateral veins, minute pustules visible between veins
on both surfaces; *cross-veins" many. INFLORES-
CENCES 1 per axil; peduncle 10-13 cm long, 4—
5 mm diam., terete, black-drying; spathe 9.5-14
cm long (ca. 1 time longer than peduncle), green
throughout, drying with large white flecks; spathe
blade 6 cm long; spathe tube 6 cm long; pistillate
portion 5 cm long; spadix to 11 cm long; staminate
portion 5.5—6 cm long, 7-10 mm diam.; fertile sta-
minate portion to 1 cm diam.; sterile staminate por-
tion 7 mm diam.; pistils 3.9 mm long; ovary (6)7—
8-locular, bidi 3.2 mm long, 0.4—0.5 mm diam.,
with axile placentation; ovules 2-seriate; funicle
0.3-0.4 mm long, adnate to lower part of partition,
style similar to style type D; style apex flat; style
boss small. INFRUCTESCENCE with berries white
(immature); seeds 6 per locule, 1-1.1 mm long,
0.2-0.3 mm diam., translucent.
Flowering in Philodendron jefense is documented
by a single collection with immature fruits collected
in December. Flowering apparently occurs during
the wet season.
Philodendron jefense is endemic to Panama on
Cerro Jefe in Premontane rain forest at about 800
m elevation.
lostigma subsect. Macrobelium ser. Reticulata. This
species is distinguished by its appressed-climbing
habit, short, thick internodes, sharply 2-ribbed cat-
aphylls promptly weathering to light brown, semi-
organized fibers with small, thin fragments of epi-
dermis persisting, terete petioles (about twice as
long as the blades), broadly ovate, gray-green-dry-
ing blades, and solitary long-pedunculate inflores-
cences with the spathes green outside.
Philodendron jefense is probably closest to P. la-
zorii Croat, which shares broadly ovate, gray-green-
drying leaf blades and semi-organized, brownish
masses of persistent cataphyll fibers. The latter spe-
cies differs in having matte-drying blades with the
lower surface smooth and minutely B or
whitish-speckled, with the minor veins
darker than the surface and weakly puckered «i
out any sign of branching or cross-veins. In con-
trast, P. jefense has blades semiglossy on drying
with the lower surface sparsely granular with fre-
quently branched minor veins and with cross-veins
not drying darker than the surface. The upper sur-
face lacks any signs of raphide cells.
The species name refers to the type locality, Cer-
ro Jefe, which has proven to be one of the most
endemic-rich areas for its size of any site in Pan-
ama.
Philodendron jodavisianum G. S. Bunting, Gen-
tes Herb. 9: 337. 1965. TYPE: Mexico. Chia-
pas: Cerro de Madrugal, along Teapa—Tacotal-
pa (Tabasco) Highway, at km 4, W of hwy., on
rock face of steep slopes, Bunting 1526 (ho-
lotype, US). Figures 243, 245-248, 253.
Usually hemiepiphytic, terrestrial; stem appressed-
climbing, gray-green to brown or green, sap sweet-
pol СЕ blackened, leaf scars conspicuous, 2
cm long, 1.7 cm wide; internodes weakly flattened on
one side, 2-ribbed, weakly glossy, to 8 cm long, but
surely longer than wide and 2.5-3 cm diam. at apex
to medium green, soon gray, epidermis brown,
flaking, fissured sometimes; roots whitish, few per
Be iy ездөрү» 10—20 cm long, bluntly ог sharply
2-ribbed or unribbed, sharply D-shaped, somewhat
spongy, light to medium green or reddish to b brownish,
sometimes streaked pinkish, semiglossy, drying
rown, persisting semi-intact as pale fibers at upper
es, disorganized below; petioles (18)29-91 ст
bing (2)3-7(8-12) mm diam., sometimes pr usu-
ally D- or U-shaped, spongy or firm, dark gre n, flat-
tened or somewhat sulcate to convex, often que me-
dial rib, margins raised adaxially, rounded abaxially,
surface semiglossy, minutely and short-lineate; blades
Volume 84, Number 3
1997
Croat 473
Philodendron Subgenus Philodendron
triangular-ovate, + subcoriaceous, moderately bico-
lorous, long acuminate at apex, deeply cordate at
base, 25-74 cm long, 18-34 cm wide (1.1–2.8 times
longer than wide), (0.5-2 times longer than petiole),
broadest + near the middle, upper surface dark
green, drying blackish, semiglossy, lower surface pale
green to yellow-greenish, drying blackish, glossy to
semiglossy or matte; anterior lobe 18-58 cm long,
10.1–40.4 cm wide (1.7—4.3 times longer than pos-
terior lobes); posterior lobes 8-17 cm long, (3.7)5-17
cm wide, obtuse to rounded; midrib broadly sunken
to broadly convex or flat, dark i
narrowly convex, thicker than broad, concolorous or
darker than surface below; basal veins 5—10 per side,
in part coalesced to 4 cm, barely naked or naked to
2 cm; primary lateral veins 7-11 per side, departing
midrib at a 50—60° angle, + straight to the margins,
obtusely to weakly or narrowly sunken and sometimes
weakly quilted above, convex and darker than surface
below; interprimary veins almost as prominent as pri-
mary lateral veins; minor veins in part discontinuous
and darker than surface below, arising from both the
midrib and primary lateral veins. INFLORES-
CENCES erect, 2-6 per axil; peduncle 3-13 cm long
(to 20 cm long in South America), 2-6 mm diam.,
pale green to whitish, with raised, prominent white
striations; spathe semiglossy, 6-14.5 cm long (0.9-3
times longer than peduncle), sometimes greenish with
short white lineations throughout; spathe blade white
to pale green (at anthesis), reddish (post-anthesis) (B
& K red-purple 7.5/7), short-lineate outside (opening
4.6 cm long, 3.6 cm wide), very pale green to white,
matte, few, sparse, dark-lineate inside; spathe tube
semiglossy and greenish to qae brown or reddish
(maroon or reddish at anth , green to white
inside; spadix weakly exserted from the spathe,
1.6–9.2 cm long, constricted near base of fertile sta-
minate portion; pistillate portion pale green or whitish,
cylindrical, 1.4-2.4 cm long, 1 cm diam. at apex, 1.1
cm diam. at middle; staminate portion 5.3—7.6 cm
long; fertile staminate portion creamy white, cylindri-
cal to weakly tapered, 0.85-1.5 cm diam. at base,
0.7-1.3 cm diam. at middle, 6-10 mm diam. ca. 1
cm from apex, broadest at the base, narrower than the
pistillate portion, narrower than the sterile portion;
sterile staminate portion broader than the apical area
of the pistillate portion, gray-white, 1.1—1.5 cm diam.;
pistils 1.8-2.5 mm long, 1-1.4 mm diam.; ovary 4-
6(5—7) ocular; locules 1.3-1.7 mm long, 0.5-0.7 mm
., With axile placentation; ovules 18-26(23-28)
Per locule, 2-seriate, 0.2-0.4 mm long, longer than
funicle; funicle 0.1-0.2 mm long, ois to lower part
of partition, style 0.2-0.3 mm long, 1-1.4 mm diam.,
similar to style type B; style apex flat to slightly
rounded; stigma subdiscoid, brushlike, unlobed, 1–
1.3 mm diam., 0.2-0.6 mm high, covering entire style
apex; the androecium truncate, margins 4—6-sided;
thecae oblong to obovate, 0.3-0.5 mm wide, + par-
allel to one another, contiguous; sterile staminate flow-
ers blunt, 1.6-1.8 mm long, 1.4-1.5 mm wide. IN-
FRUCTESCENCE with berries white (mature), 6 mm
long, 2.6 mm diam.; seeds 14—16(24—28) per locule,
1.3-1.7 mm long, 0.2-0.3 mm diam. JUVENILE
plants with internodes 8 cm long, 5 mm diam.; peti-
oles sharply U-shaped in cross section, usually lack-
ing medial rib adaxially, margins erect.
Flowering in Philodendron jodavisianum occurs in
the late dry season and throughout much of the rainy
season (April through October) in Panama. In Costa
Rica it has been collected in flower in February, and
post-anthesis collections have been made in January.
ts have been collected August through
November.
Philodendron jodavisianum ranges from southern
Mexico to Colombia, Ecuador, and Venezuela (Mérida)
from near sea level to 1500 m elevation. It occurs in
Tropical wet forest and Premontane wet forest in Pan-
ama and Costa Rica but also in Tropical moist forest
in some parts of Central America.
Philodendron jodavisianum is a member of P. sect.
Philodendron subsect. Philodendron ser. Fibrosa. This
species is characterized by its short internodes, sharp-
ly 2-ribbed cataphylls persisting as pale fibers, peti-
oles flattened or somewhat sulcate above and usually
with a medial rib, usually triangular-ovate, blackish-
drying blades with many impressed primary lateral
veins, and spathes usually greenish on the tube and
white on the blade.
Philodendron jodavisianum 15 quite variable
throughout its range, and the type (being one of the
most northerly collections) differs from most in having
more elliptic blades.
Philodendron jodavisianum is most easily confused
with P. grandipes, which differs in comprising terres-
trial plants with broadly ovate leaf blades. It has also
been confused with P. panamense К. Krause, which
differs in having subterete petioles, promptly decid-
uous cataphylls, and longer peduncles often bent be-
neath the spathe (see P. panamense for additional de-
tails)
Similar collections from South America, especially
Croat 57441 from Bahía Solano, Chocó Department,
Colombia, differ in sometimes having distinct “cross-
veins." Collections from Pichincha Province, Ecuador
(Croat 55737 and Grayum & Zamora 9431), differ in
having peduncles 16-20 cm long.
Two Darién collections (Hammel et al. 16252 and
McPherson 11517) are unusual in having leaf blades
with somewhat flaring posterior lobes and markedly
474
Annals of the
Missouri Botanical Garden
concave margins. However, the leaves on the second
sheet of the McPherson collection are typically
shaped with a convex blade margin
Additio: BELIZE. Toledo: Blue
na е 3234 (BM). а RICA. Alajuela: Los
Ensayos, Buena Vista de San Carlos, Barquero 10 (UC); Fin-
= nsayos, са. 11 mi. NW of Zarcero, са. 850 m, Croat
43583 (MO); Reserva Biológica Monteverde, along Río Peñas
Blancas, 820 m, 10°21'N, 84740", Bello & Haber 2881
(INB, MO); Río Zapote, 5 km S of Canalete, along new road
to Upala, 100-200 m, 10°48’N, 85°02’W, Burger & Baker
:M
km from Finca Racine, 1200-1300 m, Croat 36648
(MO); along Camino Raiz de Hule, SE of Platanillo, 1200—
1400 m, Croat 36756 (MO); Jicotea Valley, Jicotea-Río Pac-
uare, Croat 36539 (MO). G —
above Bijagua, ca. 1500 m, Gómez 19048 (MO). Heredia:
11 km E of Cariblanco, 1060 m, Loiselle 255 (MO); Paracito—
6
illo, 1000 m, 10°16’38N,
84°04’ 57 W. jon 1285 (MO): Zona Protectora “La Selva,”
along trail from main road across Quebrada Cantarrana to
Río Guácimo m, Grayum « 6 3216 (DUKE); La
Selva Field вена = са. 100 т 44300 (МО); 100–150
m, 44320 (МО); Стаушт 1915 (Moy. Jiménez V (MO); Murrell
nie T ie Frío, W of Finca Zona Nueve, ca. 110
, Grayum 3562 ا = Sucio, near
e oes Tx m, Croat 35703 (MO). Limón: Manzanillo
de Talamanca, ca. 5 m, 938'N, 82°39' W, Grayum & Burton
4340 (MO); ca. 11 mi. S of Siquirres, 650 m, Croat 43329
(MO); Cerro Coronel, 10—40 m, 10°40'N, 83:40" W, Stevens
24346 (MO); Rio Sixaola, ca. 0.5 mi. SW of Bambú, ca. 3
0 m, 8°34! N, 83°25’ W,
Croat & Grayum 59807 (CM, CR, K, MO): Osa Península,
i. W of Pan-
с y, 46'N, 83° 18' W, Croat
67696 (CR, MO); 67693 са E мо); "pus G. Herrera &
ава она R). San José: above Río Sucio, 5-6000 m,
Pennington et al. 11529 (K); San Isidro del Dui ла
ical, SW of San Isidro, 4.8 mi. from Río Pacuare, 1000 m,
Croat 35253 (CR, F, MO); 9 mi. SW of Rio Pacuar, 680 m,
Croat 35344 (MO); C
500-800 m, Croat 41607
(MO); 41698 (K, MO, NY). Izabal: 12 km NW of El Estor,
650 m, Martinez & Stevens 23305 (MO). HONDURAS. At-
lantida: Lancetilla Botanical
MO); Lancetilla, 100 ft., Yuncker 5008 (F, MICH, MO, NY).
MEXICO. genie Mpio. Teapa, 7 km SE of Teapa on road
to Tacotalpa, Ran neas, 70 m, 17735'N, 9250",
Hammel & Merello 15516 (MO). NICARAGUA. Chontales:
4 km NNW of Cuapa, Cerro Oluma, 700-775 m, 12°18’N,
85723'30"W, Nee 28413 (MO, NY). Río San
Chontalefio, 20 km NE of El Castillo, 200 m, Neill T Илге
Norte, 11%07'N, 83 :
6 km upriver from Barra nta Gorda, 8-10 m, 11%30'N,
83°49'W, Stevens 20756-a (MO); Mpio. Siuna, Santa Rosa,
Load 67 вина PANAMA. Bocas del Toro: Escudo de Ver-
coast, 5 m, 9°05’N, 81°35'W. McPherson
11412 (MO. ы Fortuna Dam area, road along Continental
Divide, 2.8 mi. S of Continental Divide, 850-950 m, 8°45'N,
82°15'W, McPherson 9676 (MO, US); Chiriquí те
tuna, 1.2 mi. N of Divide, 5.3 mi. N of bridge over Fortuna
Dam, 910 m, 844'N, 82°17'W, Croat 60450 (MO, PMA,
RSA); along Divide, 1.1 mi. W of highway, 1200 m, 8'44'N,
82°17'W, Croat 60368 (MO); E of camp Corriente Grande
(IRHE), 200 m, Correa et al. 3697 (PMA). Chiriquí: Río
cm Valley, vic. of Fortuna Dam, 11
66587 (F, IBE, M, MEXU, MO, US); 1000-
о
82437, 82177, Croat & Grayum 60080 (СМ, МО); Gual-
aca—Fortuna, 10 mi. NW of Los Planes de Hornito, 1260 m,
845'N, 82717", Croat 50074 (MO); Gualaca—Chiriquí
Grande, Rio Hornito, ca. 0.5 km S of Cem de Científicos,
1010-1130 m, 8°44’N, 82713'30"W, C 67922 (MO);
8°45'N, 82718", Croat & Zhu 76376 (о, SEL); 1100-
1200 m, 8°45'N, 82*15'W, Thompson 5026 (CM); Río Chi
riqui, beyond Gualaca, 10.8 mi. beyond Los Plantes de Hor-
nito, ca. 1400 m, Croat 48721 (MO). Coclé: Santa Rita
Ridge Road, 4–6 km from Transisthmi 1 00
34280 (MO); Mile 6.5, 370 m, 9°21'15’N, 7944",
Croat & Zhu 76960 (MO); El Copé — near Continental
Divide, ca. Mile 1.5, ca. 900 m, Croat 44572 (МО); ca. Mile
5.6, 800 m, Croat 75068 (МО); La caes ~ Copé,
ft, Hammel 2633 (MO); El Valle region, N of El Valle de
Anión, La Mesa, 900-1000 m, 840, € 10'W, Hamilton
et al. 4111 (INB, MO); 800-900 m, 8°38 N, 80°00", Croat
67122 (CAS, CM, COL, HNMN, L, MEXU, jn aie 860-
900 чуо (МО); a (AAU, DUKE, MO, N Y, PMA,
TEX); 67213 (AAU, CM, EAP, MO, PMA, TEX, 5с;
67121 UT А МО); 67129 ee igs СМ, COL, CR,
F, HUA, , NY, P, PMA, SEL, ТЕХ); 775
m, 8°36 cane W, 74793 (KUN кој 860 m, 8°37'N,
80°08'W, Croat & Zhu 76742 (MO, SEL); 76710 (MO). Co-
lon: Río Guanche, ca. 3-5 mi. inland, 10— m, Ur
26175 (MO). Darién: Cero Pire age Alturas de Nique
region, S of El Real, near Cana min m, 7°45'N,
— McPherson 2 (MO); 11517 (B. MO); Ciel d
e, 480 m, Croat 38003 (MO); 38039 (MO); 600-1
nM 37743 (MO); Parque Nacional Darién, N of Tacarcuna,
m, 805'N, 77716", Ham-
MO); ca. 17 km E of Pucuro, 850 т,
8°03.5'N, 77°17’ a 16252 (MO). Panamá: El Llano-Cartí
road, 10 m from highway, near El Llano, 330 m, are 33779
, NY); Mile 5-6, m, 34771 (F, МОЈ М
68, 350 m, 49128 (AAU, CAS, CM, MEXU, MO); Mile 10.
u Lin Hote otel, Croat
O, NY); at end of road МО);
E M beyon
14227 (MO); 800 m, 8'41'N, 79°56'W, ur ON (СМ,
summit and upper tail Witherspoon &
(MO); ca. 1 a a sa З 450 т, Croat 35950 (MO,
PMA); Cerro Jefe region, Campo Tres, ca. 700 er
27064 (F, MO); 700-150 | m, т N, 7925'W, Croat ders
(IBE, MO, TEX). San Blas: El Llano-Cartí, 23-29 ме po
Pan-American Highway, км 9°22'N, 78'
Volume 84, Number 3
1997
Croat 475
Philodendron Subgenus Philodendron
Нив. 1839 (МО, PAX B rte Road, vicinity Nu-
sagandi, 300-350 m 797, Croat 69279 (DUKE,
MO); 300 m, 92207, a m. & Zhu 76542 (MO); 350
m, 920'N, 797, Croat & Zhu 77011 (МО); 450 m, 918'N,
7959'W, Gon 75121 (MO). Veraguas: Río Concepción—
Río Barrera, 300-600 ft., Hammel 5252 (MO); Santa Fe re-
m is Г уе Сайты», 0.5 mi. N of Escuela Alto
Nacional Cerro Tute, 800-1030 m, Croat &
йш Зе 14 (MO): 5-8 5-8 km from achiodl; 730-770 m, Croat
967 (MO); Río Primero Brazo, 5 mi. NW о
e,
450—550 m, Croat 27639 (MO); 0.6 mi. beyond Escuela
Agrícola Alto Piedra, 730 m, Croat & Folsom 33997A (MO);
34062 (MO); ca. 1200 m, 8732'N, 81707", Hamilton et al.
1280 (MO); Escuela Vh чэр Alto Piedra, Mori & Kallunki
2529 (MO); Santa Fe-Río San Luis, past Escuela Agrícola
Alto Piedra, Río ай Brazo, 480 m, 8°33’N, 81708",
Стоа 66901 (СМ, МО).
Philodendron knappiae Croat, sp. nov. TYPE: Рап-
a. Chiriquí: Cerro Hornito, in elfin forest on
ridges and summit, approached from Los Planes
de Hornito, oe. m, 842'N, 82°06'W, 14 Mar.
1982, Knapp, Kress & Hammel 4219 (holotype,
NO ЊЕ Figures 240, 244, 249, 254.
Planta terrestris aut hemiepiphytica; internodia 0.5-7 cm
longa, 1-1.7
— mm diam.; lamina ovato-triangularis, cordata
23.5 ст longa, 7.5-11.5 cm lata; inflorescentia 1; siad
culus 2-3 em longus, 5-6 mm diam.; spatha 7-8 cm longa,
> viridis, intus subrosea; pistilla 4-locularia; Јосић 4-ovu-
ап
Terrestrial ог hemiepiphytic; stems usually erect;
internodes short, somewhat flattened on one side with
marginal ribs, moderately glossy, 0.5-7 cm long, 1-
1.7 cm diam., longer than broad below, short near
apex, medium green, epidermis smooth, light brown;
cataphylls 10-18 cm long, sharply 2-ribbed, green,
sulcate adaxially, margins weakly raised abaxially, de-
ciduous; petioles 14-19 cm long, 2-7 mm diam.,
bluntly D-shaped, green, tinged reddish; blades
ovate-triangular, subcoriaceous, Montem long-apic-
ulate at apex, cordate at base, 16-23.5 cm long,
rowly to е ibündeds sinus Мрака: 3-
5.5 ст deep; midrib broadly convex, paler than sur-
face above, convex, reddish or paler than surface be-
low; basal veins 3(5) per side, with 0-1 free to base,
2-3 coalesced 1—1.5 cm; posterior rib naked; primary
lateral veins 5-7 per side, departing midrib at a
(40)55—70" (lowermost to 80°) angle, arcuate-ascend-
ing to the margins, weakly visible above, we
raised, often reddish below; minor veins distinct be-
low, arising from the midrib only; secretory ducts
present but inconspicuous. INFLORESCENCES 1
per axil; peduncle 2-3 cm long, 5-6 mm diam.;
spathe 7-8 cm long (2.3-3.6 times longer than pe-
duncle), weakly constricted above the tube, 1.5 cm
iam. at constriction, green throughout, e with-
in; spathe tube 3—4 cm long, 1.9 cm diam.; spadix
5.5-7 cm long, white throughout; pistillate dcin 12
cm long, 1 cm diam.; staminate portion 4.3-5.8 cm
long, 8-12 mm diam.; pistils 1-1. dn mm long, 0.7 mm
diam.; ovary 4-locular; locules 0.8 mm long, 0.3-0.4
mm diam., with axile placentation; ovules 4 per loc-
ule, 2-seriate, 0.2 mm long, longer than funicle; fu-
nicle 0.1 mm long, adnate to lower part of partition,
style similar to style type B; style apex flat to concave;
stigma = hemispheroid, unlobed, 0.8 mm diam., 0.2
mm high, covering entire style apex; the androecium
truncate, ^ prismatic, margins irregularly 4—6-sided,
mostly 4—5-sided, 0.8-1.1 mm long; thecae oblong,
0.4 mm wide, = parallel to one another, not contig-
uous; sterile staminate flowers irregularly 4—6-sided,
0 mm long.
Flowering in Philodendron knappiae apparently oc-
curs in the dry season and is documented by only two
fertile collections, one flowering and one post-anthe-
sis, both made in March
Philodendron knappiae is endemic to western Pan-
ama, known only from the type locality in Chiriquí
Province on Cerro Hornito in Tropical Lower Montane
rain forest at 2100 m elevation.
Philodendron knappiae is a member of P. sect. Ca-
lostigma subsect. Macrobelium ser. Macrobelium. This
species is recognized by its relatively small stature;
short internodes (distally) with smooth, brown epi-
dermis; adaxially sulcate, sharply two-ribbed, green
but densely short-red-lineate, deciduous cataphylls;
bluntly D-shaped petioles (about as long as the
blades); small, ovate-triangular dark brown-drying
blades with a hippocrepiform sinus; and solitary in-
florescences with the spathe green outside and pink
inside.
The species is perhaps most easily confused with
P. wilburii, which has leaf blades of similar size and
shape. The latter species differs in having longer in-
ternodes drying with a tan, glossy epidermis, thicker
blades drying yellowish green to dark olive brown
above, 2-3 inflorescences per axil, and 2 ovules per
locule bs 4 per locule for P. knappiae).
specimens examined. PANAMA.
ен йе. above Los Planes de Hornito, 1750 m, 8°41 N.
82°10'W, Croat 67982 (CM, MO); 2100 m, Kress et а!
82-1363 (DUKE)
476
Annals of the
Missouri Botanical Garden
Philodendron lazorii Croat, sp. nov. TYPE: Pan-
ama. Canal Area: vic. Madden Lake, along
both sides of stream SSE of pumping station
(SE of dam), 140 m, 9?13'N, 7937'W, 18 Jan.
1990, Croat 69833 (holotype, МО–З789003—
5; isotypes, AAU, B, CAS, CM, COL, CR, F,
GH, K, MEXU, NY, PMA, QCNE, RSA, SCZ,
TEX, US, VEN). Figures 13, 14, 250-252,
255-258.
Planta hemiepiphytica; internodia 1—4 cm longa, 5 cm
diam.;
32-54 cm longa, (22)34—49 cm lata, cordata basi, in sicco
canoviridis; sinus hippocrepiformis vel obovatus; inflores-
centia 2; pedunculus 8.6-17 cm longus; spatha 12-13.5
em longa; pistilla (4)5—6-locularia; loculi 10—14(18)-ovu-
lati.
N
Hemiepiphytic; stem appressed-climbing (scan-
dent as juvenile); internodes weakly striate, 1—4 ст
long, 5 cm diam., + broader than long, light olive-
green to dark yellow-green, glossy, roots 1-3
per node, dark brown to reddish brown, matte,
smooth to densely scaly, to 30 cm or longer, 3-5
mm diam., feeder roots to 8 mm diam., densely
scaly; cataphylls 17-19 cm long, pale yellow-green,
sharply D-shaped, margins acutely raised,
persisting semi-intact at upper nodes with a dense
reticulum of coarse fibers, often overlain with a
thin, fragmented epidermis, becoming dilacerated,
eventually deciduous; petioles 37-63(76) cm long,
5—6 mm diam., terete, dark green, firm, flexible,
surface matte, faintly dark green striate; blades
broadly ovate, subcoriaceous, long-acuminate at
apex, cordate at base, 32-54 cm long, (22)34—49
ст wide (0.97—1.4 times longer than wide, aver-
aging 1.13 times), (0.65-0.9 times the petiole
length), slightly shorter in length than petiole, up-
per surface dark green, drying gray-green, weakly
to semiglossy, lower surface drying gray-green,
matte, much paler; anterior lobe 27-33 cm long, 36
cm wide (1.6-2.2 times longer than posterior
lobes); posterior lobes 13-17 cm long, 16–19.5 cm
wide, obtuse; sinus hippocrepiform to obovate, 9.5—
11.5 cm deep; midrib flat to broadly raised, con-
colorous to slightly paler than surface above, nar-
row-rounded to bluntly acute, darker than surface
below; basal veins (5)6—7 per side, with 0—1 free
to base, 3rd and higher order veins coalesced 2—
4.5 ст, obscure; posterior rib 2-3 cm long along
the sinus; primary lateral veins 3-4 per side, de-
parting midrib at a 40-50” then to 70° angle,
straight to weakly arcuate to the margins, weakly
and obtusely sunken and concolorous or paler
above, convex and darker than surface below; mi-
nor veins moderately distinct, arising from both the
midrib and primary lateral veins, drying weakly
puckered and darker than surface below, upper sur-
face usually drying with whitish cells visible, lower
surface drying smooth and minutely brownish to
whitish speckled. INFLORESCENCES 2 per axil;
peduncle 8.6-17 cm long, to 1.1 cm diam., medium
green, lightly white-striate, heavily white-striate
nearest apex; spathe 12-13.5 cm long (spathe
equal in length to or slightly shorter than pedun-
cle), constricted above the tube, 2.9 cm diam. at
constriction, margins pale to creamy; spathe blade
yellowish green outside, 8 cm long (opening 4 cm
wide), creamy on outer margins, glossy inside;
spathe tube medium green, finely white-striate with
pale margins outside, 5 cm long, 3.3 cm diam., pale
yellowish green, glossy inside; spadix bluntly
pointed at apex, 10.9-11.3 cm long, constricted 1.5
cm above base of fertile staminate portion; pistillate
portion pale yellowish green, 3.34 cm long, 1 cm
diam. at base, 1.5 cm diam. midway, 1.4 cm diam.
at apex; staminate portion 7.8-9.3 cm long; fertile
staminate portion creamy white, 1.7 cm diam. at
middle, 11 mm diam. ca. 1 cm from apex; sterile
staminate portion barely detectable, creamy white,
drying darker than fertile staminate flowers, 1.7 cm
diam.; pistils 1.9-2.5 mm long; ovary (4)5—6-locu-
lar, 1.1 mm diam., locules 1.1-1.6 mm long, 0.3-
0.4 mm diam., with axile placentation; ovules 10-
14(18) per locule, 2-seriate, 0.3 mm long, longer
than funicle; funicle 0.1–0.2 mm long, adnate to
lower part of partition, style 0.9 mm diam., similar
to style type D; style apex + rounded to bluntly
pointed; style boss broad and pronounced; stigma
subdiscoid, unlobed, + truncate, 1.5 mm diam., 3
mm high, covering entire style apex, depressed
shallowly and medially; the androecium truncate,
prismatic, oblong, margins irregularly 4-6-sided,
0.7-1 mm long; thecae oblong, 0.4-0.5 mm wide,
not contiguous, + parallel to one another; sterile
staminate flowers irregularly 4—6-sided, slightly
clavate to prismatic, 1.2-1.9 mm long. J UV
plants with internodes matte, gray-green,
long, 7 mm diam.; petioles terete; blades weakly
velvety; upper surface + glistening-glossy, los
surface matte, much paler, with flecks of brilliance;
minor veins very distinct, darker than surface.
Flowering in Philodendron lazorii occurs п pei
late dry season and early rainy season (Marc
through June), with mature fruits in August. Et
mature fruits have been collected in March -
June, which indicates that there must be flowers r
the late wet season as well (or perhaps it indicates
bimodal flowering).
6 cm
Eu A E EO ed
—— ——— ——— ——— — — —
MM NM M MM аса савана каса EE a NN ННН
Volume 84, Number 3 Стоа! 477
1997 Philodendron Subgenus Philodendron
Philodendron lazorii is endemic to Panama, basi; inflorescentia 1-2; pedunculus 3-10 cm longus; spa-
known from the type locality, a region of limestone
outcrops near Madden Lake, at about m ele-
vation in Tropical moist forest, and in Darién Prov-
ince at 250 to 1050 m in Tropical moist forest and
Premontane wet forest.
Philodendron lazorii is a member of P. sect.
Philodendron subsect. Philodendron ser. Fibrosa.
This species is distinguished by its thick, short in-
ternodes, cataphylls persisting semi-intact with an
organized network of yellow-brown fibers, terete
petioles averaging 1.25 times longer than the blade,
and broadly ovate, gray-green-drying blades matte
on the lower surface.
Philodendron lazorii is probably most closely re-
lated to P. panamense, which has similar inflores-
cences. The latter species differs in having semi-
glossy, ovate-triangular blades usually 1.3 or more
times longer than wide (averaging 1.5 times longer
than wide), and petioles commonly shorter than the
blade. In addition, it has generally shorter pedun-
cles (usually shorter than the spathe).
This species is also similar to Р. jefense. See that
species for a discussion of the differences.
Philodendron lazorii is named in honor of one of
its original collectors, Robert Lazor (Army Corps of
Engineers, Vicksburg), who collected in Panama
while a student at Florida State University.
Additional specimens examined. PANAMA. Сапа!
ea: Madden Lake Cave a а. 913'N, pue W, Ty-
son & Lazor 6266 (FSU, IBE), Salvador Hill, near Juan
Mina, Bartlett & Lasser 16785 (MICH, MO). Darién:
along trail from base camp to Rancho Frio on slopes of
Cerro Pirre, 2 m, 7°58'N, 77*43'W, Croat & Zhu
77126 (CAS, CM, CR, "MO ); W side of Cerro Pirre, 800-
1050 m, 7°56'N, 77°45'W, Croat 68953 (DUKE, "HUA,
M, MO).
Philodendron lentii Croat & Grayum, sp. nov.
TYPE: Panama. Coclé: El Valle region, be-
ca. 1 km off road, vic. La Mesa, N of El Valle
de Antén, 800-900 m , 838'N, 80?09'W, 11
i 1987, Croat 671 63 (holotype, MO-
, B, BR, CAS, CM,
QCA, RSA, S, SCZ, TEX, US, VEN). Figures
259-265
Planta gy caulis scandens; internodia 4—
12 ст lon nga, 1.54.5 cm diam.; cataphylla 20-23(40) ст
longa, plerumque casio enda m 1-2 costata, decid-
ua; petiolus teres vel late D-formatus, interdum
6-5 a, 9-25 cm lata, truncata vel
ordata basi; nervis basalibus 1—3(4) utroque, liberis
tha 12-21 ст longa, lamina spathae extus viridalba vel
marronino vel violaceipurpureo; pistilla RE locularia;
loculi (1-2)4—6-ovulati; baccae aurantiaca
Hemiepiphytic; stem scandent, creeping, assur-
gent, sap soapy-scented; internodes elongate, se-
miglossy to matte, 4–12 cm long, 1.5—4.5 cm diam.,
usually slightly longer than broad, medium to dark
green, conspicuously pale striate at upper edge of
each node, drying brownish, often narrowly ribbed
ribs irregularly ridged, sometimes warty), epi-
dermis weakly fissured transversely; roots moder-
ately few, to ca. 30 cm long, drying to 2 mm diam.,
reddish brown; cataphylls 20—30(40) cm long, usu-
ally unribbed, sometimes 1-ribbed, sharply
l-ribbed, or weakly to sharply 2-ribbed, greenish,
tinged reddish, deciduous, intact; petioles 22-44
cm long, 4-12 mm diam., terete to broadly D-sha-
ped, firm, dark green, sometimes purplish below,
broadly sulcate, flattened or broadly convex adax-
ially, convex abaxially, with adaxial margins obtuse
to rounded, surface semiglossy, densely and mi-
nutely white or dark striate-lineate; blades ovate
to narrowly ovate or ovate-elliptic, coriaceous to
subcoriaceous, conspicuously bicolorous, acumi-
nate to narrowly acuminate at apex, rounded to
truncate or subcordate at base, 21.6-51 cm long,
9-25 cm wide (1.4-2.7(3.3) times longer than
wide), (0.7-1.4(2.25) times longer than petiole),
usually about equal in length to petiole, upper sur-
face dark green, semiglossy, drying dark grin
brown to yellowish brown, lower su
to matte, paler; anterior lobe 28—45.5 cm hue 9-
25 cm wide (4.9-10.2 times longer than than pos-
terior lobes); posterior lobes 3-7 cm long, broader
than long, broadly rounded to obtuse; sinus +
V-shaped to arcuate, to 4 cm deep; midrib flat to
broadly convex, paler than surface above, convex,
tinged reddish or darker than surface, drying mi-
nutely granular below; basal veins 1–3(4) per side,
with all free to base; primary lateral veins 8-14 per
side, departing midrib at a 60—70^ angle, weakly
arcuate, ascending to the margins, weakly to nar-
rowly sunken or weakly quilted, drying paler than
surface, raised along the margins with the center
collapsed (forming a channel) above, convex to
weakly raised or weakly pleated, darker than sur-
face, drying minutely granular, paler than surface
below; interprimary veins weakly sunken or ob-
scure above, visible below; minor veins visible, few,
darker than surface, arising from both the midrib
and primary lateral veins, minutely etched in upper
surface of fresh leaves, drying raised. INFLORES-
—
478
Annals of the
Missouri Botanical Garden
CENCES 1-2 per axil; peduncle 3-10 cm long, 3—
7(14) mm diam., pale green; spathe 12-21 cm long
(1.5-3.98(4.5-5.1) times longer than peduncle),
margins reddish; spathe blade greenish white to
white outside, 4 cm diam., dark maroon to cream
inside; spathe tube white to greenish white, some-
times tinged pinkish or red, densely white-lineate
outside, 6-9 cm long, dark maroon to violet-purple,
densely white-lineate inside; spadix sessile; pale
greenish to white throughout, cylindrical, tapered,
11-14(18) cm long, broadest near the base or
slightly above the middle, constricted below the
middle; pistillate portion green, cylindrical to
ovoid-tapered, 6.6 cm long in front, 4.5 cm long in
back, 1.1 cm diam. at apex, 1.3-1.4 cm diam. at
middle, 8-11 mm wide at base; fertile staminate
portion greenish white, tapered to clavate or cylin-
drical, 6.8-9.4 cm long, 9-16 mm diam. at base,
9-15 mm diam. at middle, 8-10 mm diam. ca.
cm from apex, broadest at or near the base, broader
than the pistillate portion, sterile staminate portion
not detectable; pistils 1.6-2.4 mm long, 1-1.8 mm
diam.; ovary (5)6-8-locular; locules 1.2-2 mm long,
0.3-0.6 mm diam., ovule sac 0.8-1.5 mm long,
with basal or sub-basal placentation; ovules (1—
2)4—6 per locule, 1- or 2-seriate, 0.3-0.5 mm long,
longer than or equal in length to funicle; funicle
0.1—0.4 mm long (can be pulled free to base); style
0.1—0.5 mm long, 1–1.6 mm diam., similar to style
type B; style apex flat or weakly rounded; stigma
discoid, brushlike, unlobed, 1-1.3 mm diam., 0.1—
0.2(0.6) mm high, covering almost entire style apex
or just the center of style apex; the androecium
truncate, margins 4—6-sided (4—5-sided); thecae
oblong, 0.3-0.5 mm wide, contiguous or + parallel
to one another; sterile staminate flowers 4—6-sided,
1.1-1.5 mm long, 0.8-1.3 mm wide. INFRUC-
TESCENCE with berries orange; seeds 3—4 per loc-
ule, 1-1.1 mm long, 0.5-0.6 mm diam.
Flowering in Philodendron lentii appears to oc-
cur during the dry season and early rainy season
(February, March, June, and July). Post-anthesis
collections exist from January, March through July,
and November, indicating а somewhat broader
range of flowering. Most post-anthesis inflores-
cences were collected between March through July.
Immature fruits are known from January, April, and
July while mature fruits are known from April,
June, and July.
m
Philodendron lentii ranges from Costa Rica (Car-
tago) to Panama (Chiriquí to Coclé), from (210)670
to 1800 m elevation in Premontane rain forest and
Tropical Lower Montane rain forest life zones. In
Costa Rica, this species is apparently rare, having
been collected only near Tapantí and Moravia in
artago Province and near Cariblanco in Alajuela
Province. In Panama, it has been collected mostly
in the Fortuna Dam region, at Cerro Colorado (both
Chiriquí), at Santa Fe in Veraguas and at E] Valle
and El Copé in Coclé Province.
Philodendron lentii is a member of P. sect. Ca-
lostigma subsect. Macrobelium ser. Ecordata. This
species is distinguished by its somewhat scandent
habit; elongate internodes; deciduous cataphylls;
D-shaped to broadly sulcate petioles (about as long
as the blades); ovate, subcordate blades with the
few basal veins free; and one to two large inflores-
cences with the spathe white externally and pur-
plish internally on the tube. Also characteristic are
the orange berries. Especially useful for recognition
is the upper dried blade surface, which usually
dries dark with the lateral primary and minor veins
weakly raised.
Philodendron lentii may be confused with P. he-
leniae, which has blades of similar size and shape
with few pairs of free basal veins and petioles about
as long as the blades. That species differs in having
more slender stems, subterete petioles, and several
to many small, red inflorescences. Although a few
collections of P. lentii (Knapp 4970, Thompson
22) report the spathe to be red, these cannot be
confused with P. heleniae, owing to the much larger
size of the spathes (mostly more than 12 cm long
vs. mostly less than 10 cm long for P. heleniae).
Possibly also belonging to this species is a sterile
collection from Ecuador (Esmeraldas), Croat
72298, which differs in having internodes to 30 cm
long, more numerous primary lateral veins (to ca.
20), and interprimary veins also sunken on live
plants giving the blade the appearance of having
very close primary lateral veins. This collection
also differs in having the cross-veins prominulous
on the lower surface toward the margin. Despite
these differences, this Ecuadorian collection surely
represents either P. Јети or another closely related
new species.
Philodendron lentii is named in honor of Roy
Lent, who first collected the species in 1967. Lent,
a resident of Costa Rica, has been an important
collector over many years and has collected many
new species of Агасеае.
Additional specimens examined. COSTA RICA. Ala-
juela: 3 mi. S of Cariblanco, 760 m, Croat 35777 (MO)
Cartago: Moravia-Quebrada Platanillo, Moravia. 3-5 km
from Finca Racine, 1200-1300 т, Croa t 36592 (E.
MEXU, MO, US); Río Grande de Orosi, 0.5 km W у
Tapantí, 1200 m, Lent 909 (CR, F). PANAMA. Bocas de
Toro: Fortuna Dam area, A equo ри RTT NOR near
Continental Divide, 1170 8°44'N, 17'W, d
7 (B. CAS, CR, К, MO, PMA, US); 1200 m, 603
Volume 84, Number 3
1997
Croat 479
Philodendron Subgenus Philodendron
(CAS, MO); 1.2 mi. N of Divide, 910 m, 8'44'N, 82°17'W,
hiriquí: Cerro Colorado, vic.
m
e
„В.
e
Boquete, 1800-2200 m m,
Grayum et al. 6358 (MO); vic. of Boquete, 1630-1780 m,
8°46'N, 82°25'W, Croat 66395A (MO); 66391 (MO);
66355 e MO); Fortuna Dam area, 1100-1200 m,
82?05'W, Тарі 5022 (СМ, MO); 8 km N of
es de Hornito, 1250-1300 m, 8?45'N, 82°12’W,
Knapp 4970 (МО, NY); 11.8 mi. N of Los Planes де Hor-
nito, 1400 m, Croat 48682 (CM, MO); 10.8 mi. beyond
Los Planes de Hornito, ca. 1400 m, Croat 48712 (M
10.1 mi. NW of Los Planes de Hornito, 1250 m, 8°45’ М,
82°17'W, Croat 50041 (СМ, MO); 10 ті. NW of Los
Planes de Hornito, 1260 m, Croat 501 M (MO, NY, PMA);
1 N of dam over For-
YT
yum 60007 (K, MO, US); 6006
Chiriquí Grande, 8 mi. N of Los
nito, 1010-1130 m, 8”44'N, 82^14'30"W, Croat 67923
, MO); near Fortuna Lake, 8°45'N, 82*18'W, Croat 4
Zhu. 76388 (MEXU, MO); N edge of lake, ca. 1100 m, c
8°45'N, 82%15'W, McPherson 9078 (MO, NY); tm
Dam, 1200 m, gone 2196 (МО); yupi аи Arena,
n m, 8'45'N, 82°16'W, Hammel et al. 14706 (CR,
MO); 26 km past Gualaca, 670 m
ver 1329 (CAS, MO). Cosik Penonomé-Coclecito, above
А °40'N,
80°26'W, Croat 67538 (MO, PMA); El Copé region, Alto
Calvario, Continental Divide, 5.2 mi. above El Copé, 930
m, Croat 49204 (MO); Alto Calvario, 710-800 m, 8°39'N,
80°36’ E Croat 68724 (MO); El Valle region, Grea 14367
(MO); 2700 ft. puse et he 4369 (CR, MO); ca. 800 m,
Croat Pip (F, M ry & ded 3683 (MO): 900-
930 m, Croat in ЧЕ, МО); 860 m, 8°37'N, 80708",
Croat & Zhu 76740 (MO, US); Cerro Gaital, 800-900 т,
8°37'N, 80°07'W, McPherson 11211 (L, MO); 800-909 m,
of Cerro Ga s 800 m, 8°38'N,
e Croat & Zhu 76897 (MO); 1000-1250 m, Croat 48931
0).
Philodendron ligulatum Schott, Prod. Syst. Ar-
oid. 224. 1860. TYPE: Cultivated from Central
America, Wendland s.n. (holotype, W? de-
stroyed). Costa Rica. Limón: Ref. Nac. Barra
del Colorado, between Río Chirripocito and
Río Sardina (Sardinal), 10°38'N, 83°45'W, 12
m, 22 Apr. 1990, Grayum 9823 gets MO;
isoneotype, CR). Figures 269-271
Hemiepiphytic, usually scandent or appressed-
climbing or sometimes epiphytic, sometimes occur-
ring high into canopy, rarely terrestrial; stem ap-
pressed-climbing or scandent (flowering stems often
ose, semi-erect or spreading), green, sometimes
with white, waxy coating; leaf scars conspicuous,
1.5-2 cm long, 1-1.8(3) cm wide; internodes some-
times weakly flattened on one side, (1)3-9(20) cm
long, 0.5—3 cm diam., usually longer than broad,
medium to dark green or gray-green to brownish,
weakly glossy, drying gray-green to pale yellow-
brown, sometimes irregularly ridged or cracked,
sometimes closely tranverse-fissured; epidermis
peeling, bubbling or with loose flakes; roots several
per node, 15-45 cm long, drying 1-2 mm diam.;
cataphylls usually sharply 2-ribbed, sometimes
weakly and bluntly 1-2-ribbed, sometimes sharply
D-shaped with adaxial margins winged to 6 mm
high, 14—19 cm long, sharply flattened, with obtuse
medial rib, green to whitish, sometimes tinged red-
dish, densely dark-lineate, sometimes densely
diam., subterete to obtusely flattened, rarely D-
shaped, sometimes with a thin, medial rib toward
apex, sometimes with adaxial margins winged to 6
mm high, spongy but brittle (fresh), medium to dark
green, obtusely to sharply flattened to sulcate adax-
ially, the margins at least sometimes acute, broadly
rounded abaxially, surface matte to semiglossy,
densely short dark green-lineate or speckled, with
a deep maroon to purple ring around apex; sheath
flattened, unopened, to 3—10 cm long, for up to half
the length of the petiole; blades oblong-elliptic to
arrowly ovate or narrowly oblanceolate-elliptic,
subcoriaceous, weakly to moderately bicolorous,
acuminate to long-acuminate at apex (the acumen
inrolled or apiculate and sometimes downturned,
14 mm long), subcordate, obtuse to narrowly
И or weakly cordate at base, (14)18—61 cm
ong, 8-19 cm wide (1.57-5.4 times longer than
mel (1.1-3.89(6.3) times longer than путне
margins weakly undulate; upper surface dar
green, semiglossy to moderately glossy, ne
blackish brown to dark grayish or grayish brown,
lower surface moderately paler, matte to weakly
glossy, mottled violet-purple or maroon, drying dark
olive-green to yellow-brown, dark olive-brown or
dark brownish black; posterior lobes, when present,
rounded to narrowly rounded, about as broad as
long, 4-5 cm long, 5.5-6.6 cm wide, held close to
petiole; sinus somewhat V-shaped, 0.5-3 cm (most-
ly 2.5 cm) deep; midrib flat to broadly convex, con-
colorous to paler than surface, sometimes sparsely
purple-speckled near base above, narrowly rounded
to convex to broadly convex, sometimes round-
raised, paler or darker than surface, or sometimes
concolorous below, sometimes sparsely purple-
speckled near base below; basal veins not evident
or 1-3 per side; primary lateral veins 4—8(12) per
Annals of the
Missouri Botanical Garden
side, departing midrib at a 40-70(75)” angle, +
straight or slightly curved to the margins, dark
green, weakly sunken to etched, concolorous above,
weakly raised to convex and paler than surface be-
low (sometimes with purple spots on older leaves),
drying brownish; interprimary veins obscure to im-
pressed, paler than surface, weakly sunken above,
weakly raised, darker than surface below; minor
veins weakly visible above, very close, obscurely
and weakly raised, darker than surface below, aris-
ing from both the midrib and primary lateral veins.
INFLORESCENCES erect, 1(2) per axil; peduncle
(5)7.5-17.5(28) ст long, 7-10 mm diam., subter-
ete, obtusely angular to 3-sided, pale to medium
green; spathe 10-19.5 cm long, 1.5-2 cm diam.
(to 6.5 cm wide when flattened), ((0.8—1.3)1.4—
1.6(1.8) times shorter than peduncle), semiglossy,
acuminate at apex, barely or not at all constricted,
ribbed on back, margins revolute; spathe blade
white or cream, sometimes sparsely red-spotted
outside, sometimes tinged purple-violet on outer
margin, 7-8.5 cm long, 1.7 cm diam. (opening 7.5—
7.8 cm long, 3.24 cm wide), whitish inside; resin
canals bright orange, appearing throughout lower %
of spathe and to near base of tube inside; spathe
tube oblong, pale to medium green with pale violet
nearest base, sparsely red-spotted outside, 6.5—7
cm long, 1.2-1.5 cm diam., glossy, whitish or pale
green, sometimes red to violet-purple (B & K pur-
ple 5/10) near base or throughout tube inside; spa-
dix + tapered with slight bend, held + erect, pro-
truding forward from spathe, rounded at apex,
(8.3)11–15.5 cm long; pistillate portion light green,
(1.4)3.7–6.5 cm long, 9-13(16) mm diam. at apex,
10–14(17) mm diam. at middle, 9-12(15) mm
diam. at base; staminate portion (6.8-7.2)10.5-11.2
cm long; fertile staminate portion (9)11-16 mm
diam. at base, (7)11-16 mm diam. at middle,
(5)9-11 mm diam. ca. 1 ст from apex, broadest
slightly above the middle, as broad as or broader
than the pistillate portion, as broad as or slightly
narrower than the sterile portion; sterile staminate
portion generally broader than pistillate portion,
whitish or tinged faintly violet, 1-1.2 ст diam.;
pistils 1.2-2.7(3.5) mm long, (0.8)1.1-1.9(2.6) mm
diam.; ovary (5)6-8-locular, 0.8-1.8(2.4) mm long,
(0.8)1.1—1.9(2.6) mm diam., walls embedded with
granular, crystal-like particles; locules (0.8)1.1—
1.8(2.4) mm long, (0.1-0.2)0.4-0.7 mm diam.,
ovule sac 0.5—0.7(1—1.3) mm long, with sub-basal
placentation; ovules 1 per locule, contained within
translucent, gelatinous envelope, 0.2-0.6 mm long,
usually longer than funicle; funicle 0.1-0.2(0.4)
mm long, style 1-6 mm long, (0.8—10)14—19(26)
mm diam., similar to style type B or type D (rarely
to type C); central style dome sometimes present;
style apex flat to rounded or sloping; style boss
sometimes present; funnel shallow when present;
stigma somewhat cupulate, truncate, unlobed to
subdiscoid, (0.4)0.9-12 mm diam., 0.1-0.3 mm
high, covering center of or entire style apex, in-
serted on center of style apex or style boss or funnel
if present, sessile, papillate, semiglossy; the an-
droecium truncate, margins irregularly 4—6-sided,
1-3 mm long; thecae oblong to ovate, sometimes
elliptical, 0.3-0.6 mm wide, = parallel to one an-
other or contiguous; pollen subspheroidal to oblong
or obovoid; sterile staminate flowers usually 4—6-
sided, 1.2-2.2 mm long, 1.3-2 mm wide. INFRUC-
TESCENCE with berries oblong-elliptic, 3 mm
long; seeds oblong, light yellow-brown, 1.4 mm
long, 0.4 mm diam., narrowly ribbed longitudinally.
JUVENILE blades with lower surface sometimes
tinged maroon.
Philodendron ligulatum ranges from Nicaragua
(Chontales and Zelaya) along the Atlantic slope of
Costa Rica and along the Atlantic slope and Con-
tinental Divide of Panama to Colombia (Antioquia
and Chocó). It occurs in Tropical wet forest and Pre-
montane rain forest life zones, from sea level to
1200 m elevation (to 600 m in Nicaragua and 650
m in Costa Rica).
Philodendron ligulatum is a member of P. sect.
Calostigma subsect. Glossophyllum ser. Glossophyl-
lum. This species is characterized by its somewhat
scandent to appressed-climbing habit; generally
elongate internodes; sharply 2-ribbed to obtusely
l-ribbed cataphylls; more or less spongy, obtusely
flattened to somewhat D-shaped petioles clearly de-
marcated from the blade by a purple ring; usually
more or less oblong blades narrowly cordulate at
the base (or with the blades ovate and cordate) with
usually 1–2 free basal veins; and usually 1-2 in-
florescences per axil.
This species comprises three varieties. Philoden-
dron ligulatum var. ligulatum exists throughout the
range of the species, whereas varieties ovatum and
heraclioanum are endemic to Panama. See under
those varieties for the differences separating them
from variety ligulatum. Philodendron ligulatum Var.
ovatum is endemic to central Panama in Coclé and
Veraguas Provinces, while P. ligulatum var. hera-
clioanum is known from far eastern Panama in San
Blas and Darién.
Philodendron ligulatum is probably close 10 Р
correae from upland Chiriquí in western Panero
The latter species differs in lacking distinct pe
mary lateral veins. It also differs in occurring "
generally higher elevations (780-1400 m).
Volume 84, Number 3
1997
Croat 481
Philodendron Subgenus Philodendron
Philodendron ligulatum may also be confused
with P. immixtum, also a vine but with smaller
leaves. See under the latter species for a discussion
of the differences. Philodendron ligulatum may also
be confused with P. wendlandii. The latter has sim-
ilar blades and spongy petioles, but differs in being
a true epiphyte with short internodes and petioles
sharply flattened adaxially and usually much
broader than thic
The protologue of P. ligulatum describes a sterile
plant, whereas Schott's color plates (Icones) depict
fertile material. None can therefore be used as type
material. In addition, Engler’s Araceae No. 180 pre-
pared in 1883 and said to have been collected from
the living type plant was not studied by Schott. This
species is represented by excellent illustrations of
fertile material by Schott, one of which could have
served as a neotype, but it was deemed best to
select a new fertile specimen for the neotype.
The names Philodendron ligulatum and P. lin-
gulatum (L.) K. Koch (P. subg. Pteromischum) have
been confused in the past. Pursuant to an unpub-
lished request for a recommendation by the Com-
mittee for Spermatophyta, the two names have of-
ficially been ruled not confusable, hence not
homonymous (see Nicolson, 1994: 280).
KEY TO THE VARIETIES OF P. LIGULATUM
la. Petioles sharply D-shaped with undulate lateral
axially; lower leaf blade surface mot-
ded purple-violet or maroon; ovaries with 2
ovules per кен (lacking an obvious ovule sac);
eastern Pana m raclioanum Croat
. Petioles кана пе at most ob ha
flattened adaxially; lower
colored; ovaries with 1
tained within an obvious а sac; Nicaragua to
~
c
anama.
2a. Leaf blades 1.5-1.7 times longer than vids
peduncles longer than spathe; Pana
1200 m var. ais
. Leaf blades 2-4.5 times longer than wide;
peduncles shorter than or about as long as
spathe; usually occurring below 800 m (rare-
ly to 1200 m) var. ligulatum
N
[-
Philodendron ligulatum Schott var. ligulatum
nternodes sometimes weakly flattened on one
side, (1)3-9(2) cm long, 0.5-3 cm diam.; cataphylls
usually sharply 2-ribbed, sometimes weakly
2-ribbed; petioles 7-38 cm long, 5-15 cm diam.,
subterete, usually at most obtusely somewhat flat-
tened adaxially, sometimes sulcate, sometimes
sharply flattened on one margin adaxially, rarely
D-shaped; lower leaf blade surface not nass
blades рер, i 24—60 cm long, 8—19 с
wide (2.1-4.4 times longer than wide), (1. ы
3.9(5.1) times longer than petiole), upper surface
drying dark olive-green to brownish; sinus 3
cm (mostly 2.5 cm) deep; basal veins not evident
or 1-3 per side; primary lateral veins 5-8(9) per
side, departing midrib at 60—70° angle. INFLO-
RESCENCES 1 per axil; peduncle 7.5—17(28) ст
long; spathe 10–19.5 cm long, spathe blade white
or cream, sometimes sparsely red-dotted outside,
spathe blade whitish or sometimes violet-purple
near base of tube or throughout inside; spathe tube
pale green thoughout, pale violet nearest base,
sometimes sparsely red-violet outside; spadix
(8.3)11-15.5 cm long; pistillate portion (1.4)3.7—
6.5 cm long; ovules 1 per locule, contained within
an obvious ovule sac; style similar to style type B
or D, central dome sometimes present; stigma
somewhat cupulate, truncate, unlobed to subdis-
coid.
Flowering in Philodendron ligulatum var. ligu-
latum occurs in the early rainy season, between
May and August, with a few flowering collections
from October through March at the end of the rainy
season and early dry season. Post-anthesis inflores-
cences have been collected from February through
August (as well as November). It is possible that
this species flowers bimodally, once near the be-
ginning of the rainy season and again near the end
of the rainy season or the early dry season. Im-
mature fruits are known from July, August, and De-
cember, but mature fruits only from December.
Philodendron ligulatum is typically somewhat
scandent in Costa Rica but is often more nearly an
appressed hemiepiphyte in Panama. In Costa Rica,
the number of inflorescences per axil is never more
than two but in some areas of Panama up to three
inflorescences per axil may be encountered, and in
the Santa Rita Ridge area of Panama, a particularly
unusual specimen (Croat 76954) has up to five in-
florescences per axil. That collection also has pro-
portionately somewhat longer petioles than other
collections. This unusual specimen is otherwise
identical to other plants of the species, even those
from the same region. It is perhaps of hybrid origin.
A few Panamanian specimens of P. ligulatum
var. ligulatum from the vicinity of Santa Fe in Ver-
aguas Province (Croat 25692, 48906A, 66914) and
in Coclé (Croat 49195) differ in drying yellow-
green and having somewhat more coarse venation.
These are somewhat intermediate with P. ligulatum
var. ovatum.
Croat 33306, from 1200 m elevation on Cerro
Colorado in eastern Chiriquí Province, has black-
drying blades averaging slightly more than two
Annals of the
Missouri Botanical Garden
times longer than broad, and is also somewhat in-
termediate with P. ligulatum var. ovatum.
A collection from the southern slopes of Panamá
Province in the basin of the Río Мадгоћо (Croat &
Zhu 77041) is — between P. ligulatum
var. heraclioanum in having
sharply D-shaped аси lacking undulate-winged
margins.
A noteworthy collection from Antioquia Depart-
ment in Colombia (Fonnegra et al. 1957) perhaps
ts P. ligulatum var. ligulatum, but dif-
fers in having the primary lateral veins drying paler
than the surface below rather than darker as is the
general case.
Additional specimens examined for P. ligulatum var. lig-
ulatum. COSTA RICA. Alajuela: 3 mi. N of San Miguel,
380 m, Croat 35643 (MO); Cañas—Upala, 4 km NNE « of
Bijagua, ca. 400 m, Croat 36262 (MO); near Rfo Zapote,
1.8-2.7 las 5 of te Сани ca. 100 m, Croat 36360
(MO); Naranjo-Aguas Zarcas, 8.5 km NE of Villa Que-
say, Schatz & Grayum 707 (DUKE); Finca La Selva, at
confluence of Río Sarapiquí and Río Puerto Viejo, Atlantic
slope, 50-80 m, 10°26'N, 84*01'W, Grayum 7673 (CR,
MO); Folsom 9973 (MO); 100 m, Grayum 2800 (MO);
Hammel 8827 (MO); Croat 61220 (MO); between Río Su-
. MO); near Mee Viejo along road
30 (MO). Limón: Hitoy
W of Valle La Estrella, 150-550 т,
na,
12 m, 10738'N, 83°45'W, Grayum 9823 (CR, MO). NIC-
ARAGUA. Chontales: Cerro Las Nubes, El Tamagás-
Gregorio, ca. 2 km N of Santo Domingo, 600
m, Grijalva & Ríos 3455 (MO). : NW of Cerro
Musún, 500-800 а et raquistain & Moreno 2570 Loe
Zelaya: Colonia К 50 m, са. 14°41'N, 84
Pipoly 3974 (MO); Bante of Neptune Mining cto
Bonanza, 2 , са. ME 84°35'W, Stevens
13029 (МО); Río lyas, ribet
rante—Ojo
Croat 38212 (F, K, diis vic. of Chiriquí Lagoon, Wedel
1479 (F, GH, MO). Canal a: along Río Indio de Ga-
tún, near sea level, Pittier 2794 (US). : Cerro
Colorado, along Continental Divide
Félix, 1200-1500 m, Croat 333 06 (MO). Coclé: La Mesa,
N of El Valle de Antia: 785 т, 8°37'N, 80*08' W,
MO, PMA, TEX, WIS): vaai m, 44734 (MO); 2700
ft., Sytsma et al. 4343 (MO); Quebrada Mollejón, ca. 5
пи. N of El yat 700 m, Croat 75050 (MO); road to
Coclesito, 12 mi. from Llano Grande, 200 m, 8°47'N,
80°28'W, Churchill et al 4018 (F, MO). Colón: Santa Rita
MO, NY, PMA); 26 km from highway,
500 m, 9°26'N, 79°57'W, Knapp et al. 1737 (MO); Mile
6.5, 370 m, 921'15"N, 79744", Croat & Zhu 76954 (В
CAS, COL, CR, F, K, MEXU, MO, NY, PMA, SCZ, US,
VEN); Sabanitas-Portobelo, Río Piedras Lumber R
922'30'N, 79^41'30"W, A НИ
M 75171 (MO); penu de Di АЈ mi.
4..4 (AAU, CAS, CM, COL CR, DUKE, Е С, MR
К, KYO, L, LE. MBM, MO, NY, Р, PMA, QCA, RSA,
E.
Croat 26100 (МО); Río Boquerón, past Salamanca,
400 m, 9°20'N á « Vict e мо;
Río San Augustin n flowing into Rfo Guanche, с
¥30'N, 79740", Churchill et al. pice (CAS, мо. NY NY
(МО); v vic. of Guásimo on Río Miguel de la Borda,
9983 (MO, SCZ); Río Fató, 10-100 m, Pittier 3867 (NY,
US). Panamá: El Llano-Cartí Road, 6.8 mi. from high-
way, 350 m, Croat 49102 (MO); Cerro Jefe, 21 mi. from
Panamerican Hi 9092 (МО);
Serranía de Маје, hi int of ridges тіне.
730 ши al. € (MO); El Valle de
ыл N of El Valle де Мадгоћо,
"у, Croat & Zhu 77041 (MO).
Agricola Alto de Piedra, at Rio Segundo Brazo, 480 m,
833'N, 81°08’ W, Croat 66914 (MO, US); са. 5-8 km be-
agricultural school, 730-770 m, Croat 25962 (F.
MO); Santa Fe-Calovébora, 1.7 mi. past Alto Piedra
School, 1.5 mi. beyond Quebrada Cosilla, 570 m, 8°33'N,
81°08'W, Croat & Zhu 76862 (MO); Río Concepción,
Lewis 2793 (MO, NY).
Philodendron ligulatum var. heraclioanum
Croat, var. nov. TYPE: Panama. Darién:
Río Perisenico,
77°44'W, 26 July 1994, Croat & Zhu 77
(holotype, MO- 04619347-9; isotypes, AAU,
B, CAS, CM, COL, CR, DUKE, F, GB, GH,
HUA, IBE, K, L, M, MEXU, NY, oom, 5
PMA, QCA, SEL, TEX, U, US, VEN,
Figures 274-276.
B diam.; cataphylla
Internodia 1 jc eed 1.1 на decidua: petiolus jac
: bus alatis: |
ina (2946-61 cm longa, (8.5)12-15 cn ай oblongo
liptica vel anguste oblanceolato-elliptica, obtusa basi,
Volume 84, Number 3
1997
Philodendron Subgenus Philodendron
sicco nigrescens; inflorescentia 1; ся nes
ст longus, 1 cm diam.; spatha 10-12 c
Internodes short, to 1 cm long, 1.1-2.5 cm diam.,
usually longer than broad, medium green, semi-
glossy, drying pale yellow-brown, conspicuously
and irregularly ridged and cracked; roots moder-
ately few, to 30 cm long, 4 mm diam., reddish
brown, weakly glossy, finely scaly; cataphylls 15—
20 cm long, bluntly 1-2-ribbed, sharply 2-ribbed
near apex; petioles (8.5)20-28 cm өгү; sharply
D-shaped, with adaxial margins winged (to 6 mm
high); blades oblong-elliptic to narrowly oblance-
olate-elliptic, obtuse to narrowly rounded at base,
29)46—61 cm long, (8.5)12-15 cm wide ((2.9)4.3—
5.4 times longer than wide), (2-2.6(6.3) times lon-
ger than petiole), upper drying blackish, lower sur-
face matte, heavily tinged or mottled violet-purple
or maroon, drying dark brownish black; midrib nar-
rowly rounded and paler than surface below with
maroon spots on older veins; basal veins lacking;
primary lateral veins 5-12 | side, departing mid-
rib at a 40—50° angle to th i i
maroon spots on older Pa INF LORESCENCES
l per axil; peduncle (5)13-15 cm long, 1 cm diam.;
spathe pale green, 10-12 cm long, to 6.5 cm wide
when flattened, tinged violet-purple on outer mar-
gin and at base outside and at base inside; spathe
blade greenish inside; spathe tube pale green,
glossy inside; spadix 9.5-12 cm long; pistillate
portion 4.3 cm long in front, 3.6 cm long in back;
ovules 2 per locule, contained within transparent
matrix, style similar to style type C; style funnel
shallow to moderately deep. INFRUCTESCENCE
with berries oblong-elliptic, 3 mm long; seeds ob-
long, light yellow-brown, 1.4 mm long, 0.4 mm
diam., narrowly ribbed longitudinally. JUVENILE
Мад» with lower surface heavily tinged maroon.
Flowering in Philodendron ligulatum var. hera-
clioanum is poorly known. Pre-anthesis flowering
collections were seen from March and in July. The
inflorescence on the July collection was nearly fully
emerged so that it would certainly have opened in
less than one month. Immature fruits were seen in
October.
Philodendron ligulatum var. heraclioanum is en-
demic to Panama, known definitely only from the
type locality at the base of Cerro Pirre in Darién
Province from 50 to 200 m elevation in a Premon-
tane wet forest life zone.
is variety is characterized by its bluntly two-
ribbed cataphylls; sharply D-shaped, marginally
Winged petioles; and the oblong-elliptic to narrowly
ьан. blackish drying blades heavi-
ly tinged with maroon on the lower surface when
young.
Philodendron ligulatum var. heraclioanum dif-
fers from both varieties ligulatum and ovatum in
its sharply D-shaped petioles with undulate-winged
margins; in having 2 ovules per locule without an
obvious ovule sac (vs. 1 ovule per locule contained
in an ovule sac for the other two varieties); and in
the maroon coloration of the lower surface of the
young blades. In contrast, the other two varieties
have petioles that are typically terete or subterete,
merely obtusely flattened adaxially.
Philodendron ligulatum var. heraclioanum may
be confused with P. pseudauriculatum, but the lat-
ter differs in having the petioles unwinged and, at
most, obtusely flattened adaxially. In addition, Р
udauriculatum has never been reported to have
the blades purplish on the lower surfaces.
A collection from San Blas in far eastern Panama
is probably also this variety; it differs in having
much longer internodes (to 10 cm long and drying
ca. 5 mm diam.) and proportionately shorter peti-
oles.
Philodendron ligulatum var. heraclioanum is
named in honor of Panamanian collector Heraclio
Herrera, who was one of the first to collect the tax-
on.
Additional specimens examined. PANAMA. Darién:
€ cad National Park, W side of Cerro Pirre, base
p. 77°48'W, Croat 68963 (MO); Parque
май — Базен Rancho Frio, М base of Cerro
Рите, ca. 9 km S of El Real, Quebrada Perisenico, 70—
270 m, 801'N, 77744", Hammel et a 16145 (MO). vs
Blas: Quebrada Masargandí, Isla de Tubuala, 2
8*54'N, 71*46' W, Herrera 1304 (CAS, CM, MO, PMA).
Philodendron ligulatum var. ovatum Croat, var.
nov. TYPE: Panama. Veraguas: vicinity of
Santa Fe, along road between Alto Piedra and
Calovébora, 0.5 mi. N of Alto Piedra, on slopes
COL, CR, F, K, NY, PMA, US, VEN). Figure
273
Internodia 1-6(8) cm longa, 1-1.7(3) cm diam.; cata-
ylla 13-16 cm longa, rig pl vel acute saga pe-
р
tiolus (10)15—19 ст longus, т diam.; lamina anguste
ovata, subcordata basi, Айлы ст longa,
(8. 5)12. 5—16 ст "andi in sicco viridibrunnea supra, эё
brunnea infra; inflorescentia А pedunculus 12-16.5 с
longus, 7-10 mm et pararlo 11.5-18.5 cm longa, aram
gus,
omnino viridalba; tubo spathae intus rubro basi.
Internodes 1-6(8) cm long, 1-1.7(3) cm diam.,
sometimes to 3 m or more long, sometimes covered
with a thin layer of translucent white wax; cata-
484
Annals of the
Missouri Botanical Garden
phylls 13-16 cm long, unribbed to bluntly 1-ribbed
or sharply 2-ribbed (ribs to ca. 4 mm high, in-
curled); petioles (10)15—19 cm long, 7 mm diam.,
subterete and obtusely flattened toward apex;
blades narrowly ovate, weakly cordate at base,
(14)18-26(39) cm long, (8.5)12.5-16 cm wide
(1.57-1.65 times longer than wide), upper surface
drying greenish brown to blackened, lower surface
drying yellowish brown to dark olive-brown; pos-
terior lobes rounded to narrowly rounded, 4-5 cm
long, 5.5—6.6 cm wide, narrowly rounded to obtuse;
sinus somewhat V-shaped, 2.5-3 cm deep; midrib
convex to round-raised below; basal veins 2-3 per
side, free to base; primary lateral veins 4—6 per
side, departing midrib at a 55-65(75”) angle. IN-
FLORESCENCES 1 per axil; peduncle 12-16.5 cm
long, 7-10 mm diam.; spathe 11.5-18.5 cm long,
1.5-2 cm diam. (0.8-0.95 times as long as pedun-
cle), greenish white throughout; spathe tube some-
times pale to medium green outside, pale to me-
dium green, sometimes red to purplish violet at
base inside; spadix 10-12.5 cm long; pistillate
contained within transparent, gelatinous ovule sac,
style similar to style type B; stigma subdiscoid, un-
lobed, often truncate.
Flowering in Philodendron ligulatum var. ovatum
occurs in the rainy season. Collections at or near
anthesis have been made in July, October, and No-
vember. Observations on a single plant in July
showed a series of inflorescences with one at an-
thesis and one other on the same stem in post-
anthesis condition. No fruiting collections are
known.
Philodendron ligulatum var. ovatum is endemic
to Panama, known only from Santa Fe in Veraguas
and at El Copé in Coclé Province in Premontane
rain forest at 770 to 1200 m elevation.
This taxon is characterized by its smooth, brown-
drying stems, elongate internodes, sharply 2-ribbed
to bluntly 1-ribbed deciduous cataphylls, subterete
to D-shaped petioles about as long as the blades,
ovate, subcordate blades with two to three pairs of
free basal veins, and solitary inflorescences with
the peduncle longer than the spathe.
Philodendron ligulatum var. ovatum differs from
variety ligulatum in having blades 1.5-1.7 times
longer than wide (vs. 2—4.5 times longer than wide
in var. ligulatum); peduncles longer than the spathe
(vs. shorter than or about as long as the spathe for
var. ligulatum); and two ovules (rather than one)
per locule. In addition, P. ligulatum var. ovatum
usually occurs at higher elevations (770 to 1200 m)
vs. usually below 800 m for P. ligulatum var. lig-
ulatum. Curiously, leaves of a few specimens of P.
ligulatum var. ovatum dry yellowish green or
brown, as opposed to the somewhat blackened color
typically associated with the species.
Philodendron ligulatum var. ovatum may also be
confused with P. lentii Croat & Grayum with which
it also occurs in the El Copé region. That species
differs in having usually more than ten pairs of pri-
mary lateral veins with several pairs of interprimary
veins visible between them on the dried upper
blade surfaces (vs. only about five pairs of primary
lateral veins and no interprimaries visible on the
upper dried blade surface). In addition, the spathes
of P. lentii are typically short-pedunculate.
Additional specimens examined. PANAMA. Сосје:
Alto Calvario, ca. 6 ті. N of El Copé, 770 m, 8738'N,
80°35'W, Croat & Zhu 76754 (MO); Croat 67572 (МО);
68767A (CM, MO); 1200-1300 m, 8?38'N, 80736" W, Syts-
ma 1903 (MO); 650—850 m, Folsom 6221 (MO, РМА); ca.
5 mi. N of El Copé, 900-1000 m, Croat & Zhu 75055
(K, MO, SCZ). Veraguas: vic. Escuela Agrícola Alto Pied-
ra near Santa Fe, 1050-1150 m, Croat 48906A (MO).
Philodendron llanense Croat, sp. nov. TYPE:
anama. Panamá: vic. of Cerro Jefe, along road
to summit which leads S off main road to La
Eneida, 750—800 m, 9?14'N, 79?22'W, Croat
67092 (holotype, M0—3582669—71; isotypes,
B, CAS, COL, CR, F, G, HUA, K, M, MEXU,
NY, P, PMA, SCZ, SEL, TEX, US, VEN). Fig-
ures 268, 277, 278.
inter-
Planta plerumque hemiepiphytica, raro terrestris;
nodia 5-10 cm longa, cm diam.; cataphylla 30-41
ст longa, plerumque incostata, persistentia semi-intacta
aut ut fibrae nodis supremis; petiolus subteres, 49-76 cm
longus, 1-2 cm diam.; lamina 51-77 cm longa, 39-604
cm lata, late ovato-cordata; nerviis lateralibus I (4)5—6(7)
utroque; inflorescentia (1)2—3; pedunculus 3-10.5 ст lon-
gus; spatha 12.5-24 cm longa, lamina spathae extus atri-
viridi, intus alba; tubo spathae atrirubroviolaceo basi; pis-
tilla (5)6-8(9)-locularia; loculi (6)12(20)-ovulati.
Usually hemiepiphytic, rarely terrestrial; stem
coming brown, 3—4 mm diam.; cataphylls 30-41 cm
long, usually unribbed, sometimes weakly 2-ribbed,
light to medium green, sharply and broadly D-
shaped, persisting semi-intact or as fibers at UP”
permost nodes, fibrous below, the fibers thin, pale,
disorganized, light gray. LEAVES erect-spreading.
clustered at or near stem apex; petioles 49-76 cm
эы AAA
Volume 84, Number 3
1997
Croat 485
Philodendron Subgenus Philodendron
long, 1-2 cm diam., subterete, somewhat spongy,
pale to medium green, rarely with medial rib adax-
ially, with adaxial margins sometimes acute, surface
semiglossy to almost matte, densely and in con-
spicuously short-lineate, at least sometimes with a
dark green ring at apex; blades broadly ovate-cor-
date, moderately coriaceous, abruptly acuminate,
sometimes acute at apex, conspicuously cordate at
base, 51-77 cm long, 39—60.4 cm wide (1.2-1.6
times longer than wide), (0.9—1.2 times longer than
petiole), about equal in length to petiole, margins
= hyaline, upper surface dark green, semiglossy to
moderately glossy, drying dark brown, lower surface
semiglossy to almost matte, paler, drying yellow-
brown; anterior lobe 41-60 cm long, 39-60.4 cm
wide (0.9-1.4 times longer than wide), (2.2-3 times
longer than posterior lobes); posterior lobes 18.5—
24 cm long, 16-31 cm wide, obtuse; midrib almost
flat to broadly convex to broadly flattened, paler
than surface above, convex, paler than surface be-
low; basal veins 4—6 per side, with 0—1 free to base,
the remaining coalesced 6-9 cm; posterior rib usu-
ally naked for 2-3 cm; primary lateral veins (4)5—
6(7) per side, flat to weakly sunken, paler than sur-
face above, convex, paler than surface below; minor
veins moderately obscure above, darker than sur-
face, arising from both the midrib and primary lat-
eral veins, drying prominulous above and below;
secretory canals distinct. INFLORESCENCES
(1)2-3 per axil; peduncle 3-10.5 cm long, 1.1-1.2
cm diam., coarsely white striate near apex; spathe
12.5-24 ст long (1.7-5 times longer than pedun-
cle), oblong, dark to medium green throughout out-
side, markedly white-lineate near the base, faintly
so above the base, moderately constricted above the
tube; spathe blade weakly and densely lineate out-
side (opening broadly elliptic in face view), white
inside; spathe tube cylindrical, markedly white-li-
neate near the base outside, 6–8.5 cm long, 3—5.5
em diam., dark reddish violet, suffused onto base
of blade inside; spadix sessile, greenish white
throughout, + cylindrical, protruding prominently
forward from and out of the end of spathe (at an-
thesis), 10-17 cm long; pistillate portion creamy
white (anthesis), reddish (pre-anthesis), weakly ta-
pered toward apex, 4.1-5 cm long, 2.5-5 cm long
in front, 1 ст diam. throughout, 1.1-1.5 cm diam.
at apex, 1.4-1.6 cm diam. at middle, 1-1.5 cm
wide at base; staminate portion 6-14.4 cm long;
fertile staminate portion white, + cylindrical 1.1-
1.7 cm diam. at base, 1.1-1.9 cm diam. at middle,
7-10 mm diam. са. 1 cm from apex, broadest + at
the middle, slightly broader than the pistillate por-
tion, narrower than the sterile portion; sterile sta-
minate portion 1.1-1.9 mm diam.; pistils 1.2-
4.8(7.2) mm long, 1.1–2.4(4.3) mm diam.; ovary
(5)6-8(9)-locular, 1.5-3.5(6.8) mm long, 1.1-
2.4(4.3) mm diam., walls sometimes embedded with
granular, crystal-like particles, locules 1.5-3.5(6.8)
mm long, (0.2)0.6-1.5 mm diam., thin and mem-
branous, ovule sac 1.9-2 mm long, with axile pla-
centation; ovules (6)12(20) per locule, 1-2-seriate,
contained within gelatinous ovule sac, 0.2—0.45
mm long, slightly longer than funicle; funicle 0.1—
0.3 mm long, style 0.3-0.7 mm long, 1.3-1.6(2.5,
4.5) mm diam., similar to style type B; style crown
truncate at apex with lobed depression; style apex
rounded; stigma button-like, unlobed or weakly
lobed, truncate, 1.1-2.2 mm diam., 0.5-1.0 mm
high, covering entire style apex, sometimes de-
pressed medially; the androecium truncate, margins
irregularly 4—6-sided; thecae oblong to cylindrical,
0.4 mm wide, + parallel to one another, + contig-
uous; sterile staminate flowers acutely to bluntly 4—
6-sided, 2.1-2.4 mm long, 1.1-2 mm wide. IN-
FRUCTESCENCE with spathe green outside, or-
ange-brown inside; berries white, 7.2-10 mm long,
3.6 mm diam.; seeds more than 10 per locule, pale
yellow. JUVENILE plants terrestrial or epiphytic;
internodes short, brown, weakly glossy; petioles
29.2 cm long, + terete, moderately spongy; blades
28.6–37.7 cm long, 17-28 cm wide; midrib broadly
convex above; minor veins distinct below.
Flowering in Philodendron llanense occurs at the
beginning of the rainy season, in July. Post-anthesis
collections exist from January through August, with
immature fruits and mature fruits found only in the
dry season and early rainy season, mostly January
and March but also in June.
Philodendron llanense is endemic to Panama
(though to be expected in adjacent Colombia), rang-
ing from 250 to 800 (mostly below 500) m elevation
in Tropical wet forest and Premontane rain forest life
zones. This species appears restricted to the region
of the Cerro Jefe and the El Llano-Cartí Road
(hence the name “llanense”).
Philodendron llanense is a member of P. sect.
Philodendron subsect. Philodendron ser. Fibrosa.
This species is characterized by its short, thick in-
ternodes; mostly unribbed, semi-intact, or fibrous
cataphylls; obtusely flattened to D-shaped petioles
(about as long as the blades); thick, broadly
ovate-cordate blades with a deeply spathulate to
closed sinus and with prominently raised minor
veins upon drying; and two to three green inflores-
cences with the spathe tube dark reddish violet in-
side.
Philodendron llanense is perhaps most easily
confused with Р. schottianum, which occurs at
486
Annals of the
Missouri Botanical Garden
mostly higher elevations (generally 1000 to 1600
m) in Costa Rica, rarely to as low as 490 m or as
high as 2200 m elevation. It differs from P. Џапепзе
in having sharply two-ribbed cataphylls persisting
more intact and with a prominent yellow epidermis,
and blades usually drying with the lower surface
more yellowish with the secretory ducts more out-
standing and usually with partial *cross-veins" be-
tween the minor veins. In addition, P. schottianum
also has blades proportionately somewhat longer
(averaging 1.65 times longer than broad vs. 1.4
times longer than broad for P. llanense) and spathes
shorter and nearly elliptic with almost no constric-
tion (vs. oblong and noticeably constricted above
the tube for P. llanense).
Philodendron llanense looks superficially much
like P. ferrugineum in live condition, and since they
occur together, they can be confused. However, P.
ferrugineum differs in having promptly deciduous
cataphylls and somewhat more elongate blades,
which dry conspicuously reddish brown.
A single sterile collection (Croat & Grayum
60209) from the Atlantic slope near the Continental
Divide at 590 m elevation in Bocas del Toro Ргоу-
ince may also belong to this species. In addition to
being out of range, this collection has the major
veins much darker than the surface below.
Additional specimens examined. PANAMA. Bocas
del Toro: Fortuna—Chiriquí Grande, 4.3 km N of the Con-
tinental Divide, 590 m, 8°46’N, 82°14’ W, Croat & Gra-
yum 60209 (MO). Panamá: El Llano—Cartf Road, 10-12
km from El Llano, 400 m, Maas & Dressler 1709 (0); Km
9.5 of Pan-American Highway, 200-300 m, 9°15’N, 79°,
McPherson 10822 (B, MO); Km 12, 350 m, Mori et al.
4628 (MO); Mile 7, 460 m, 9°19’N, 79*59"W, Croat 75104
(CAS, MEXU, MO, NY); Km 8-12, ca. 400-450 m, Мее
et al. 8775 (MO, NY, SCZ); Mile 5-9, 200-250 m,
9?15-16'N, 78°59'W, Thompson 4621 (CM); ca. Km 16–
18, 400 m, Tyson & Nee 7361 (MO, PMA, US); Mile 10.1,
325—350 m, Croat 67367 (DUKE, K, M, MEXU, MO, NY,
US); Mile 12, 200-500 m, Croat 22910 (MO, PMA); Km
7-12, 360-400 m, Croat 25118 (MO), 25172 (MO); Mile
10, 330 m, Croat 33774 (MO), 33819 (MO); Mile 6.8, 350
m, Croat 49127 (MO); Km 12, Croat 26031 (MO); Cerro
Jefe, near summit, <2900 ft., Gentry et al. 3501 (MO,
NY); 12 km E of Lago Azul, 800-1000 m, Gentry & Mori
13450 (MO). San Blas: El Llano-Cartí Road, 300—500 m,
Liesner 1317 (F, MO, NY, US); Mile 14, 300 m, 915'N,
79°W, Croat 69251 (F, MO); Mile 7, 550 m, 9?43'N,
78'68'W, 60505 (CM, MO, PMA); Km 22, 350 m, 9?19'N,
78°55'W, de Nevers & Herrera 7859 (MO, PMA); Nusa-
gandi, Sendero Wedar, 300-400 m, 918'N, 78°58'W,
McDonagh et al. 216 (BM, MO); 1-2 mi. N of Nusagandi
on road to Cartí, 250-275 m, 9°20’N, 79*W, Croat & Zhu
76577A (CM, MO); Mile 10.1, 300 m, 9?20'N, 79°W,
76541 (MO, SEL); 1.9 mi. N of Nusagandi, 310 m, 76993
(AAU, CR, GB, MO, NY).
Philodendron madronense Croat, sp. nov.
anama. Panamá: Valle de Madroíio,
ca. 10 road mi. N of La Margarita (near Che-
po), just S of Continental Divide along main
trail to Cangandí, 350-500 m, 9?19'N,
7908'W, 21 Feb. 1986, Hammel & McPher-
son 14526 (holotype, MO-3490432; isotypes,
CM, M, PMA). Figures 279, 280.
Planta hemiepiphytica; internodia usque 2.5 cm diam.,
longiora quam lata; cataphylla decidua; petiolus subteres,
que 68 cm longus, 1.5 cm diam., aliquantum spongio-
sus; lamina trisectus, 39.5 i i
mentis medianis ellipticis; segmentis lateralibus valde
i i 8 cm longis, 12.7-13 ст latis; in-
squ
cm longa, ca. 4 cm diam.; spadix pistillata 11.8 cm longa,
i i 7.5 cm longa, ca. 1
cm diam.; l semen in quoque loculo.
Hemiepiphytic vine; internodes to 2.5 cm diam.,
longer than broad, epidermis drying light brown
and semiglossy but conspicuously and irregularly
folded; roots several per node, drying dark brown;
cataphylls deciduous; petioles to 68 cm long, 1.5
cm diam., subterete, somewhat spongy, drying
blackened, surface closely and finely ridged;
sheathing to 8.5 cm long; blades trisect, subcor-
iaceous, moderately bicolorous, drying weakly
glossy, upper surface dark green, drying dark
brown, semiglossy, lower surface drying yellow-
green, matte; median segments elliptic, somewhat
inequilateral, 39.5 cm long, 15 cm wide (ca. 2.5
times longer than wide), (1.5 times longer than lat-
eral lobes), somewhat acuminate at apex, base at-
tenuate and somewhat inequilateral with leafy tis-
sue extending to base on one side, ending 2.5 cm
above the base of the other side; lateral segments
markedly inequilateral, 32–32.8 cm long, 12.7-13
cm wide, acuminate; the outermost lobes with the
inner margins attenuate, tissue ending 1-2.5 cm
from base of petioles; outer margins rounded, bare-
ly or not at all naked along the sinus; midrib weakly
raised, concolorous above, + round-raised, drying
blackened and irregularly ridged below; basal veins
lacking; primary lateral veins ca. 25 per side, ca.
4-5 mm apart, departing midrib at a 65—70° angle
toward apex, 85—100° angle toward base, gradually
curved to the margins, weakly sunken, scarcely
more conspicuous than interprimaries above, prom
inently raised below; interprimary veins numerous,
1-3 between each pair of primary lateral veins; mr
nor veins in part undulate upon drying, close, mod-
erately visible, arising from both the midrib and
primary lateral veins. INFLORESCENCES (post
anthesis) 1 per axil; peduncle to 22 cm long, lem
diam., drying blackened, somewhat spongy; spathe
Volume 84, Number 3
1997
Croat 487
Philodendron Subgenus Philodendron
to 20 cm long, ca. 4 cm diam., semiglossy, green
outside, red at base inside, drying blackened; pis-
tillate spadix 11.8 ст long, са. 3 cm diam.; sta-
minate spadix 7.5 cm long, ca. 1 cm diam., mod-
erately tapered to a bluntly acute apex; pistils +
cylindrical, 3-4 mm long, 2.5 mm diam.; o
7-locular, with sub-basal placentation; locules 2.3
mm long, 0.5 mm diam.; ovules 1 per locule, con-
tained within a transparent evelope; funicle ca. 0.4
mm long (can be pulled free to base), style similar
to style type D; style apex flat; style boss pro-
nounced but narrow; seeds 1 per locule, 1.4 mm
long.
Flowering in Philodendron madronense appar-
ently occurs in the rainy season, since immature
fruits have been collected in January.
Philodendron madronense is endemic to central
Panama along the border of Panamá and San Blas
at 350 to 450 m elevation in a Tropical wet forest
life zone.
Philodendron madronense is a member of P. sect.
Tritomophyllum. This species is recognized by its
scandent habit; elongate internodes; subterete,
more or less spongy petioles; and especially by its
trisect leaf blades with the medial segments ellip-
tic, inequilaterally attenuate at the base, and with
many close prominent primary lateral veins only 4–
5 mm apart and departing the midrib at about a
angle.
Philodendron madronense is superficially most
similar to P. cotobrusense, which has deeply 3-lobed
blades with more or less elliptic, closely veined di-
visions. The latter species differs in having the
lobes united and confluent at the base and up to
five inflorescences per axil. It is also similar to
some broad-leaved forms of Р. tripartitum (e.g.,
Whitefoord & Eddy 223), which occur in eastern
anama, but those differ in having no more than
12 pairs of primary si А; veins, which depart the
midrib at a 50—60? a
Philodendron malesevichiae Croat, sp. nov.
TYPE: Panama. Coclé: vicinity El Valle de
Antón, La Mesa, 4 mi. E of El Valle at base
of Cerro Gaital, along trail which goes to the
S edge and leading to the summit, 830-900
m, 8°36'N, 80%07'W, 25 Mar. 1993, Croat
74818 (holotype, MO-4342656-57; isotypes,
B, COL, CR, F, K, MEXU, NY, PMA, QCNE,
US, VEN). Figures 3, 21, 25, 27, 28, 281-284.
Planta terrestris; caulis repens; internodia 2—4 cm lon-
ва, 1.5-5 cm diam.; cataphylla 21-29 cm longa, leniter
Mc. tata, in s sicco ганнан, persistentia semi-in-
tacta; petiolus teres, 50-56 cm longus, 8-19 mm diam.,
anguste sulcatus, sparsim squamus in triente superiore;
lamina ovato-cordata, 33-65 cm longa, 23-36 ст lata, in
sicco atribrunnea supra, flavibrunnea infra, nervis basa-
libus 6-9 paribus, superioribus 2-3 liberis ad aliquot bas-
im; reliquiarum pom 24.5 cm; inflorescentia 1-3; pe-
dunculus 4. cm longus; spatha 8-20 ст longa; pistilla
5—6-locularia; loculi ca. 19-25-ovulati.
Terrestrial, usually less than 1 m tall, sometimes
to 1.5 m, reclining against trees; stem creeping over
the ground but well rooted in the soil; internodes
24 cm long, 1.5-5 cm diam., pale green to whit-
ish, moderately glossy, usually totally hidden by
cataphylls, coarsely but faintly white-short-lineate
at apex, drying pale brown, epidermis closely and
mostly bluntly ridged with ridges close and cracked
transversely; roots moderately few per node, drying
-3 mm diam., dark brown, closely ridged and
weakly scaly; cataphylls 21-29 cm long, unribbed,
weakly 1-ribbed or weakly 2-ribbed, whitish to ma-
roon or russet, heavily tinged pink toward lower
half, drying dark brown to reddish brown, persisting
semi-intact at least toward apex with an underlying
network of pale, anastomosing fibers, the outer sur-
face becoming fibrous toward the base. LEAVES
erect to erect-spreading; petioles 50-56 cm long,
diam., terete or subterete, dark green,
tinged purple-violet, weakly glossy, drying dark
brown, weakly, obtusely and narrowly sulcate to ob-
tusely flattened adaxially, weakly flattened and of-
ten tinged purplish toward apex, faintly striate on
surface, sparsely scaly in upper one-third to one-
half, scales fine, + terete, 1-2 mm long, 0.1 mm
diam., green; blades ovate-cordate, subcoriaceous,
moderately bicolorous, acuminate to long-acumi-
nate at apex, prominently cordate at base, 33-65
cm long, 23-36 cm wide, upper surface subvelvety-
matte, dark green, drying dark brown, lower surface
semiglossy, moderately paler, drying medium yel-
low-brown; anterior lobe 23—48 cm long, 23-36 cm
wide; posterior lobes narrowly rounded, 8-25 cm
ong, 12-14 cm wide; sinus hippocrepiform, 8-14
cm deep; midrib concave, concolorous or paler
above, convex or thicker than broad, much darker
olive-green and matte, sometimes tinged maroon
near base below, drying darker than surface; basal
veins 6-9 per side, with upper 2-3 free to base,
part of the remainder coalesced 2-4.5 cm; posterior
rib naked for 0.5-3 cm, moderately straight; pri-
mary lateral veins (3)5-10 per side, 1.7-2.2 cm
apart, departing midrib at a 45-60” angle, down-
turned acutely at the midrib, weakly sunken to
weaky quilted above, convex and darker below, of-
ten branching toward the margins, drying black-
ened; interprimary veins persistent in lower half of
blades; minor veins moderately visible but not dis-
488
Annals of the
Missouri Botanical Garden
tinct when fresh, moderately distinct on drying,
weakly undulate, arising from both the midrib and
primary lateral veins. INFLORESCENCES 1-3 per
axil; peduncle 4.5-8 cm long, to 4 mm diam., pink-
ish red, coarsely white-lineate toward apex, drying
blackened; spathe 8-20 cm long, 1-2 cm diam.
when furled, to 5.5 cm wide when open (7-11 cm
wide when fully flattened), ((2.5)3—3.7 times longer
than peduncles), acuminate at apex (acumen 2.5
cm long), semiglossy, in Central America pale yel-
low-green, darker green toward middle, white along
margins, sometimes tinged with pink, the open mar-
gin sometimes violet-purple outside, greenish white
and matte throughout within, in South America
spathe tube sometimes reddish outside, dark red to
red-violet, suffused onto lower one-half of blade in-
side; spathe blade to 15 cm long, white within;
spathe tube 7.5—8.5 cm long, to 4.3 cm diam. at
anthesis; spadix weakly stipitate; 14—15.5 cm long;
pistillate spadix (3.4)4—4.3 cm long in front, 2.84
cm long in back, 11-16 mm diam. at apex, 1.2-
1.8 cm diam. at middle and base; staminate portion
10-13 cm long, 1.2-1.5 cm diam. at base, 1.5-1.8
mm diam. midway, 8-12 mm diam. 1 cm from apex,
constricted to 1.5 cm diam. above the sterile por-
tion; sterile staminate portion to 3.4 cm long, 20
cm diam. at base, 10-18 mm diam. at apex; pistils
1.1-2.4(3.5) mm long, 1.2-5 mm diam.; ovary 5—
6-locular, with axile placentation; locules 1.3-1.6
mm long; ovules 19-25 per locule, 2—3-seriate, ca.
(0.1)0.25—0.3 mm long; funicle as long as or shorter
than ovules, 424 as long as the ovule; stigma 1—
1.5 mm diam., depressed medially; style similar to
style type B, 1 mm diam.; style apex flat to weakly
concave; stylar canals arising at base of apical de-
pressions; stigma discoid, 0.8 mm thick, 1.5-18.
mm diam.; the androecium truncate to + prismatic,
margins irregularly 4—6-sided, many 6-sided, ca.
0.6 mm long; thecae oblong, + parallel to one an-
other; sterile staminate flowers clavate, irregularly
4—5-sided to ovoid, 0.9-1.3 mm long
Flowering phenology in Philodendron malesevi-
chiae in Panama is uncertain. All flowering collec-
tions are from South America. Flowering collections
have been made in February and March and post-
anthesis collections in March, July, and December.
Cultivated collections from Panama grown at the
Missouri Botanical Garden flowered in May, June,
July, November, and December.
Philodendron malesevichiae ranges from Panama
to Colombia. In Panama it is only known from the
type locality in Tropical wet forest at 830 to 860 m
in Coclé Province. In Colombia it has been col-
lected only along the Pacific Coast in Chocé and
Valle at 50 to 150 m elevation.
Philodendron malesevichiae is a member of P.
sect. Philodendron subsect. Achyropodium. This
species is characterized by its terrestrial, creeping
habit; short internodes; weakly 1-2-ribbed cata-
phylls drying reddish brown and persisting semi-
intact; subterete, narrowly sulcate petioles, which
are conspicuously scaly in the upper one-third; and
ovate-cordate brown-drying blades with up to eight
basal veins, largely coalesced and naked along the
hippocrepiform sinus.
This species is most easily confused with P.
glanduliferum. The latter species differs in having
fewer primary lateral veins (2—4 vs. 6-10 for P.
malesevichiae); posterior ribs that are not at all na-
ked; and a narrow closed or spathulate sinus (vs.
hippocrepiform in P. malesevichiae).
In Panama there are three other species that
have petiolar glands of some form and thus might
be confused with Р. malesevichiae. These are: P.
hammelii, P. verrucosum, and P. squamipetiolatum.
Philodendron hammelii differs in its smaller, green-
drying blades and petiolar scales mostly less than
three times longer than broad. Philodendron ver-
rucosum differs in its mostly appressed-climbing
habit; scaly cataphylls, inflorescences, and even
parts of the lower blade surface; and velvety and
matte (rather than semiglossy) upper blade surface.
Philodendron squamipetiolatum differs in its ap-
pressed-climbing habit, long internodes, deciduous
cataphylls, green-drying blades, and scaly inflores-
cences.
Zarucchi & Escheverry 4776, from 2000 m ele-
vation in Antioquia Department, Colombia, may
also belong to this species. It is described as having
a deep wine-red (rather than green) spathe.
This species is named in honor of Petra S. Ma-
lesevich, who has loyally worked with me on
aspects of the Philodendron revision. This species
is in cultivation at the Missouri Botanical Garden
and is deemed a beautiful addition to horticulture.
dditional specimens examined. PANAMA. Coclé: En
Mesa, Lgs El Valle de Antón, 860—900 m, Croat Mb
(MO); b of Cerro Gatital, 860 m, 837'N, 80^ ,
Croat & Zhu 76707 (CAS, MO, PMA).
COLOMBIA. Chocó: hills above junction of Río J hes
and Río Mumbá, up river from Lloró, 80-120 m ‚9 ae :
76°25" W, jose Pia: (MO). Meis Buenaventura, aj
4°4'N, 77°09'W, 70162 (CAS, COL, L, MEXU, Bor
PMA); Km 49, 150 m, 4%02'N, 77%04'W, Croat & Bay
75810 (B, CM, COL, F, GB, M, MEXU,
Duenbventura- Rio muni 33.3 km
Buenaventura Hi
Volume 84, Number 3
1997
Croat 489
Philodendron Subgenus Philodendron
marker, «50 m, 3°56'N, 76°59’W, Croat 57547 (CM,
COL, G, JAUM, MO).
Philodendron mexicanum Engl., in Mart., Fl.
Bras. 3(2): 143. 1878. TYPE: Mexico. Vera-
cruz: vic. of Córdoba, Bourgeau 2176 (holo-
type, P; isotype, G). Figures 285-288.
onini m Matuda, Anales төр Biol.
Univ. Nac. Méx : STL: Fig 2. 1951. TYPE:
Morin: Sik Pent Siltepec, iei mixed
humid forest, over humid rocks, 1200 m, 4 Маг
1951, Nakamura 31 (holotype, MEXU).
Usually hemiepiphytic, sometimes terrestrial or
epilithic; stem scandent, leaf scars conspicuous, 1—
1.5 ст long, 1.3-1.6 cm wide; internodes scurfy,
glaucous to semiglossy, 18-21 cm long, 1-2 cm
diam., longer than broad, medium green to gray-
green, epidermis blistering, fissured + transverse-
ly; roots branched at tips, sometimes with swollen
nodes along length; cataphylls fleshy, 10-23 cm
long, unribbed or bluntly 1-ribbed, pale green,
glossy, drying yellowish tan to yellowish green, de-
ciduous intact; petioles 22-66.5 cm long, 2-13
mm diam., terete, moderately spongy, medium
green, somewhat flattened adaxially, surface semig-
lossy; blades narrowly triangular-sagittate to tri-
angular-hastate, subcoriaceous, acuminate to long-
longer than wide), (0.6—1.5 times longer than pet-
iole), about equal in length to petiole, margins
weakly undulate, upper surface dark green, drying
dark brownish green, semiglossy, lower surface dry-
ing yellowish green, weakly glossy, paler; medial
lobe 19-38 cm long, 8-20 cm wide (1.7-2.7 times
longer than posterior lobes), usually 3-3.5 times
longer than wide (rarely to 1.7 times longer than
wide); posterior lobes 7-19.5 cm long, 4-12.6 cm
wide, directed somewhat toward the base, rounded
to rarely rounded; sinus parabolic to hippocrepi-
form or spathulate; midrib broadly sunken, concol-
orous above, convex, sparsely orange-spotted,
slightly paler than surface below; basal veins 0—
1(2-5) per side, with 0—1(2—5) free to base, or 1
coalesced, the third and fourth coalesced 3.6-4(13)
Cm; posterior rib not naked or naked for 0.5-2 cm,
directed straight toward the tip of the blade and
remaining 1.5-3.5 cm distant from blade margin;
primary vem = (2)4—5(6) per side, departing
midrib at ? angle, + straight to the margins,
weakly ae tios raised below; minor veins
moderately distinct below, arising from both the
midrib and primary lateral veins. INFLORES-
CENCES 1 per axil; peduncle 4-15 cm long, 4-12
mm diam.; врате glossy, 8-16.5 cm long, 1.9-2.3
cm diam. (0.8—1.4(1.5-2.4) times longer than pe-
duncle), acute at apex, the margins paler to clear
within; spathe blade greenish to whitish outside;
resin canals pale range and appearing as continu-
ous lines inside; spathe tube greenish, sometimes
pale reddish tinged outside; red-violet to maroon (B
& K red-purple 5/7.5) inside; spadix sessile;
bluntly rounded to somewhat acute at apex, 10.5—
14.7 cm long, broadest toward the apex, constricted
below the middle of fertile staminate portion; pis-
tillate portion pale green to green to pale yellow,
cylindrical to obovoid, 2-6 cm long, 1 cm diam. at
apex, 1.1 cm diam. at middle, 1.3 cm wide at base;
staminate portion (6.4)10.3-12.7 cm long; fertile
staminate portion creamy white, = cylindrical, 6—
17 mm diam. at base, 7-15 mm diam. at middle,
1.1 cm diam. ca. 1 cm from apex, about as broad
as the pistillate and the sterile portions; sterile sta-
minate portion usually broader than the pistillate
portion, white to light gray, 1-1.2 cm diam.; pistils
1.6-2(3.5) mm long, 1.1-1.3(2.6) mm diam., trans-
parent white; ovary 5—7-locular, 0.9-2.7 mm long,
1.2-1.4(2.3-2.6) mm diam., with sub-basal placen-
tation; locules 0.9—1.1(2.5) mm long, 0.2—0.4(0.6—
diam.; ovule sac 0.6—0.7 mm long; ovules
1-2(3) per locule, digitate, contained within trans-
parent ovule sac, 0.3—0.6(1.1) mm long, longer than
funicle; funicle 0.3 mm long, style 0.7(1) mm long,
1.2-1.5(2.4) mm diam., similar to style type B; style
apex domed; stigma discoid, at least sometimes
lobed both laterally and vertically, sometimes = cy-
lindrical, 0.7-1.0 mm diam., 0.1-0.3 mm high,
covering center of style apex, at least sometimes
drying with radial arms sunken between the central
peak and the peaks on the end of the arms (Croat
& Hannon 64520); the androecium truncate, mar-
gins irregularly 4—6-sided; thecae = cylindrical,
.4 mm wide, * parallel to one another; sterile
staminate flowers bluntly or acutely 4—6-sided,
3-1.9 mm long, 1.3-1.8 mm wide. INFRUC-
TESCENCE with seeds 1 per locule, yellowish or-
ange, 1.5-1.9 mm long, 0.6-0.8 mm diam.
Flowering in Philodendron mexicanum occurs in
the mid-dry season and early rainy season (Feb-
ruary through May), with post-anthesis collections
known February through June (except May) and
also in November. Mature fruits are not known. A
cultivated collection at Missouri Botanical Garden
(Croat 59933) flowered perhaps twice per year, in
March and in October.
т
Philodendron mexicanum ranges from Mexico to
Colombia, from near sea level to 1900 m elevation,
ranging from dry habitats in west-central Mexico
(both “Selva Baja Caducifolia” and “Bosque Pino-
490
Annals of the
Missouri Botanical Garden
Encino”) to more humid sites on the Atlantic slope
in Vera Cruz to as far south as Honduras in Tropical
moist forest and to Costa Rica in Premontane wet
forest. Though widespread, this species is appar-
ently rare and has been collected from relatively
few localities, often restricted to mesic enclaves in
otherwise arid regions as in the state of Morelos,
Mexico. It is one of the most ecologically versatile
species in the genus
ilodendron mexicanum is a member of P. sect.
mug subsect. Macrobelium ser. Macrobel-
um. This species is recognized by its scandent
n internodes longer than broad; unribbed, de-
ciduous cataphylls; moderately spongy, somewhat
flattened petioles (about as long as the blades); and
i by its narrowly triangular-sagittate to tri-
gular-hastate blades, which dry dark brownish
een above and yellowish к кар Sterile col-
lections from Los Ríos and as Provinces of
Ecuador (Dodson & Valverde 6959) at Jauneche and
in Guayaquil Cantón (Rubio et al. 2008) may also
represent this species. Dodson et al. (1985) erro-
neously reported the former collection as P. barro-
soanum G. S. Bunting, a species restricted to the
eastern side of the Andes
Philodendron mexicanum is most similar to P.
angustilobum, a species ranging from Honduras to
Panama. The latter species differs in having leaves
that dry more blackened (rather than green to yel-
low-green as in P. mexicanum), and which are more
decidedly three-lobed with the medial lobe propor-
tionately narrower and broadest at the middle or
even above the middle.
Although the type specimen of P. mexicanum was
collected in Mexico near Córdoba, most Mexican col-
lections are from the Pacific slope. These have some-
what less prominently narrowed posterior lobes than
the type. Moore & Bunting 8873, from near the type
locality, has the posterior lobes even more conspicu-
ously narrowed than those of the type specimen. In
this regard, its blades approach those of Р. angusti-
lobum in overall shape, but differ in drying greenish
to yellowish brown rather than blackened
A collection from Olancho Department, Hondu-
ras (Croat & Hannon 64520), is unusual in having
a style that dries with radiating arms from a central
peak.
The sole Costa Rican collection (Grayum 5418)
is unusual in having prominently hastate blades.
This collection has only a juvenile inflorescence.
Further collections may prove it represents a dis-
tinct species.
е = examined. COSTA RICA. Pun-
taren cimal downstream from tedio
Cordillera de’ Tilarán. 1300 т, 10^18'N, 84°49'W, Grayu
5418 (MO). LEA Quezaltenango: Finc
reneos-Patzulín, 1200-1400 m, Standley 86917. ©
87007 (F); 87022 (F). San Marcos: Volcán Tajumulco,
00 m, Steyermark 37968 (F). HONDURAS. At-
lántida: Quebrada Grande, ca. 10 km SW of La Ceiba,
80-180 m, 15?42'N, 86?51'W, Liesner 26138 (MO).
Olancho: San Esteban-Bonito Oriental, Río Grande,
350—400 m, 15?31'N, 85°42’W, Croat & Hannon 64520
(B, CAS, CM, CR, HNMN, K, L, MEXU, MO, NY, USCC).
Yoro: Puente Grande, Río Puente Grande (tributary of the
Río Agua), Blac n Chorley 4077 (BM, MO). MEX-
ell Tie dime Ithaca, New York,
—
ac аа above Fila de Caballo, El Parafso in Parque
Nacional de Guerrero, Croat 67442 (MO). Morelos: Cul
pas 95 (U); 5000 ft., Pringle 8093 (BH, B
G, GH, H, HBG, ISC, K, LL, MASS, XR it
POM, RSA, UC, US); Rose & Hough 4439 (US); Río Pollo,
below Salto San Antonio, E 1
W of Colonia Carolina, NW of С
m, 18?57'N, 99°
G EXU,
25982 (Е MEXU); (e & Bunting 8820 (BH); Barranca
Santa Clara, N de Acatlipa, 1450-1550 m, Vázquez 3
MEXU). Veracruz: Córdoba- Veracruz, Ejido San José 2
Gracia, below Peñuelo, ca. 730? m, Moore & Buntin,
8873 (BH, MO).
“~
01.1.4 1 з Kee
tandl. & L. O. Wil-
liams, Ceiba 3: 108-109. НЕ TYPE: Costa
Rica. Puntarenas: Esquinas Forest Reserve,
sea level, 10 Jan. 1951, Allen 5755 (holotype,
EAP; isotypes, F, GH, US). Figures 289, 290.
Hemiepiphytic vine, stem scandent, green to
gray-green, drying pale yellowish brown, unscent-
ed, leaf scars inconspicuous, 1.2 cm long, 8 mm
wide, obscured by cataphylls; ees smooth,
semiglossy, 6-12 cm long, 1-2 cm diam., longer
than broad, moderately green, nes khaki-col-
ored, epidermis fissured transversely; roots drying
light reddish brown, smooth, weakly glossy. 20-30
cm long, 2-3 mm diam., 4—6, at the nodes; cata-
phylls ts bluntly 2-ribbed or unribbed,
pale green to cream-colored, promptly deciduous;
petioles 12-24 cm long, 2-4 mm diam., subterete
to broader than thick to broadly D-shaped, weakly
spongy, bluntly flattened to broadly sulcate adaxi-
ally, rounded abaxially, with adaxial margins blunt,
surface pale or dark green streaked and demarcated
than wide), (0.65-2 times longer than petiole),
about equal in length to petiole, broadest + ne
the middle, margins straight, upper surface d
green, semiglossy, somewhat pruinose, lower SU
Volume 84, Number 3
1997
Croat 491
Philodendron Subgenus Philodendron
face glossy to weakly glossy, paler; anterior lobe
12-20.5 cm long, 10.7-22 cm wide (1.85-3 times
longer than posterior lobes); posterior lobes 4–8 cm
long, 3.5-9.5 cm wide, broadly rounded to obtuse;
sinus arcuate, sometimes parabolic, 3.4—6 cm deep;
midrib broadly convex to weakly raised, concolor-
ous to paler than surface above, weakly to broadly
convex, paler than surface below; basal veins about
3 per side, obscurely and scarcely more conspicu-
ous than primary laterals; posterior rib lacking; pri-
mary lateral veins (3)4—5 per side, obscure vites
minor veins about as conspicuous as the prim
laterals, arising from the midrib only. INFLORES-
CENCES spreading, as long as leaves, 1 per axi
peduncle 14—25 cm long, 2-6 mm diam., de a REN
green inside; spathe tube ca. 5 cm
side, red or violet-purple inside; spadix sessile, 8
cm long, broadest below the middle; pistillate por-
tion medium green to pale greenish yellow, = cy-
lindrical, 1.9-2 cm long, 6.5-7 mm diam. at apex,
7-8 mm diam. at middle, 6.5—7 mm wide at base,
with 13 (per cm) flowers per spiral; staminate por-
tion 5.9-8.3 cm long; fertile staminate portion
white, tapered toward apex, 5-9 mm diam. at base,
mm diam. at middle, 0.9 mm diam. ca. 1 cm
from apex, broadest at the base, mostly narrower
than the pistillate and sterile portions; sterile sta-
minate portion broader than the pistillate portion,
9 mm diam.; pistils 1.2-1.4 mm long, 0.
mm diam.; ovary 6—7-locular, 0.8-1 mm long, 0.8—
1.1 mm diam., with sub-basal placentation, walls
drying weakly warty; locules 0.8 mm long, 0.3 mm
diam., ovule sac 0.7—0.8 mm long; ovules 1 per
locule, contained within transparent ovule sac, 0.4
mm long, as long as funicle; funicle 0.4 mm long;
style 0.2 mm long, 1 mm diam., similar to style
type B; style apex flat; stigma unlobed, subdiscoid
to somewhat cylindrical, 0.8-0.9 mm diam., 0.2
mm high, covering almost entire style apex, cen-
tered on stylar canal pores; the androecium trun-
cate, margins irregularly to bluntly 4—6-sided, 1.2
mm long, 0.8 mm diam. at apex; thecae + cylin-
drical, 0.3 mm wide, + parallel to one another,
contiguous; sterile staminate flowers acutely or
bluntly and irregularly 4—6-sided, 1.2-2 mm long,
0.7-1.1 mm wide. INFRUCTESCENCE with fruits
whitish. Seeds (dried) ca. 20 per locule, tan, 1-1.2
mm long, 0.5-0.6 mm diam., finely ridged with
much finer cross-etching.
Flowering in Philodendron microstictum occurs
during the dry season and early rainy season, with
uin collections known from January, April,
and May, and post-anthesis collections from Feb-
ruary, March, May, and July. Immature fruits have
been collected in May.
Philodendron microstictum is endemic to Costa
Rica (though to be expected on the Burica Penín-
sula in adjacent Panama), where it is restricted to
the Pacific slope, at 50 to 450 m elevation in wetter
parts of Tropical moist forest, Tropical wet forest, and
Tropical wet forest basal belt transition life zones.
Philodendron microstictum is a member of P.
sect. Calostigma subsect. Glossophyllum ser. Ovata.
This species is characterized by its scandent habit;
long, moderately slender stems drying pale yellow-
ish brown; subterete petioles, which are about as
long as the blades; and broadly ovate, weakly sub-
cordate blades (which may be as wide or wider than
long) with an arcuate sinus and obscure primary
lateral and basal veins (sometimes with only the
basal veins visible). Also characteristic is the ex-
ternally green spathe (red within the tube) with the
peduncle as long as or longer than the petiole.
Philodendron microstictum is not easily confused
with any other species. It has been confused by
some with P. scandens, which differs in having more
narrowly ovate blades with prominent major veins
and a spathulate to parabolic sinus, and much more
short-pedunculate inflorescences (with the pedun-
cles much shorter than the petioles).
This species is perhaps related to P. chirripoense,
which is also a vine with inflorescences longer than
leaves and has blades of similar color and texture.
That species differs in having longer, more slender
internodes that dry dark brown and by its narrowly
ovate, subcordate blades, which are more than
twice as long as broad and have more prominent
primary lateral veins.
Additional specimens examined. COSTA RICA. Pun-
tarenas: Palmar Norte, 100-200 m, Croat 35107 (CR,
MO); Palmar Norte, along trail to Jalisco, 50-700 m, Croat
35189 (F, MO); 110 m, Croat & Hannon 79210 (CR, INB,
MO); Villa Neily-San Vito de Coto Brus, Cuesta Fila de
Cal, 300 m, 8°41'N, 82°57'%, Grayum ^ > 7580 (CR,
MO); El General — Río sim ed Río n, 450—500
m, Williams et al. 24214 (CR jd sa о. 20–300
m, Liesner 1836 (B, MO. ca. 5 km W of Rincón de Osa,
m, 8'42'N, 83731 W, ass & Liesner 7300 (CR,
Е MO, PMA): Fila Huacas, ca. 4 km NE of Las Huacas
Pisani along road to Sinaf, 4
2177, Grayum & Hammel све (СЕ, a
vici пее M La
0’ , Herrera 4066 (СВ, L о, NY): аде 0-
150 m, 8?27-30'N, 83°33-38' W, vis 463 (CR); in
Monkey Woods (just W of airstrip), 5 m, 8°29'N, 8335'W,
ammel et al. 16643 (CR, MO); Claro Ridge, 1-10 m
8°28'№, 83°35'W, Kernan & و 1028 (CR, МО); Sip
ena Woods, 1-50 m, 8?28'N, 83°35'W, Kernan & Phillips
1089 (CR, MO); € Sorpresa ca. 1 km NE of Golfito, E
of microwave tow a. 400 m, 839N, 83°10'W, Croat
& Grayum 59941 (CR. MO): Golfito, Cerro Las Torres, 500
492
Annals of the
Missouri Botanical Garden
m, G. Herrera 5045 (CR, INB, MO); Reserva Forestal Gol-
fo Dulce, W of Rancho Quemado, 1-300 m, 8°44’N,
83°36'W, Saborío et al. 127 (INB, MO); Río Volcán, 48
km SE of San Isidro General, 300 m, Molina et al. 18162
(F, GH, NY, US). San José: R gro, Cangreja,
са. 1. E of Santa Rosa de Puriscal, 320 m, 9?42'N,
84°23'30'W, Grayum et al. 8340 (MO); Parque Nacional,
sector Esquinas, vic. Fila Gamba, 200-300 m, Croat &
Hannon 79288 (MO).
Philodendron morii Croat, sp. nov. TYPE: Pan-
ama. Panamá: Valle de Madroño, ca. 10 mi. N
of La Margarita (near Chepo), in forest S of
and on Continental Divide, near border of
Comarca de San Blas, along trial to Cangandí,
350—450 m, 9?19'N, 79?08'W, 21 Feb. 1986,
Hammel & McPherson 14530 (holotype, MO-
3398570). Figures 291, 292.
Planta epiphytica; internodia 1–1.5 ст longa, 1.5-2 ст
diam.; cataphylla 13 ст longa, obtuse 2-costata, decidua;
petiolus subteres, 23-37 cm longus, (24—6 mm diam.,
subspongiosus; lamina ovato-triangularis, leniter cordata
basi, 25—27.5 cm longa, 11.5-16 cm lata, in sicco cana-
viridis; inflorescentia 2; pedunculus 9.5-14 cm longus, 3—
6 mm diam.; spatha 8.7-12 cm longa, viridis vel flavivir-
idis omnino; pistilla (3)4—5(6)-locularia; loculi cum 3-6
seminibus; baccae albae.
Epiphytic; stem appressed-climbing; internodes
semiglossy, 1-1.5 cm long, 1.5-2 cm diam., about
as long as broad, dark green, drying light brown;
roots drying reddish brown; cataphylls 13 cm long,
bluntly 2-ribbed, green, deciduous, intact; petioles
23-37 cm long, (2)4-6 mm diam., subterete, some-
what spongy, dark green, obscurely flattened adax-
ially, surface unmarked, often dries with loose,
puffy epidermis; blades ovate-triangular, narrowly
acuminate at apex, weakly cordate at base, 25-27.5
cm long, 11.5-16 cm wide (1.7–2.4 times longer
than wide), (0.7-1.2 times longer than petiole),
about equal in length to petiole, upper surface se-
miglossy, lower surface drying green, weakly glossy,
moderately paler; anterior lobe 24—28 cm long,
12.6-16 cm wide; posterior lobes broadly rounded,
5-7 mm long, 3—7.5 cm wide, broadly rounded to
obtuse; sinus arcuate with blade decurrent on pet-
iole; midrib prominently raised above, slightly paler
than surface below; basal veins 2-3 per side, with
0-1 free to base, 0—1 coalesced less than 1 cm;
posterior rib weak, to 1.3 ст long, naked through-
out its length; primary lateral veins (2)5-8 per side,
departing midrib at а 40-50” angle, straight to the
margins, sunken and concolorous above, convex
and darker than surface below; minor veins fine,
numerous, and distinct below, arising from both the
midrib and primary lateral veins. INFLORES-
CENCES (post-anthesis) 2 per axil; peduncle 9.5-
14 cm long, 3-6 mm diam.; spathe 8.7-12 cm
long, (0.8-1.1 times longer than peduncle), green
to yellowish green throughout; spathe tube 3.5-5
cm long; spadix 8-11 cm long; pistillate portion
(post anthesis) 5.7 cm long in front, 4.8 cm long in
back, 1.5 cm diam. midway, 1.2 cm diam. near
apex, 8 mm diam. near base; fertile staminate por-
tion 5 cm long, narrowly tapered to apex, the nar-
rowest portion to 5 mm diam., ca. 1.3 cm above
base; sterile staminate portion broader than
constricted area, to 5.5 mm diam.; pistils 1.4-1.6
mm long; ovary (3)4—5(6)-locular, with sub-basal
placentation; locules 1.3 mm long, 0.5 mm diam.;
ovules 1 per locule, 0.3 mm long; funicle 0.2-0.3
mm long (can be pulled free to base), style similar
to style type D; style apex flat to weakly rounded,
0.6–0.7 mm long, style boss small; stigma covering
entire style apex and inserted on style boss; the
androecium truncate, margins irregularly 4—6-sid-
ed, 0.7 mm long. INFRUCTESCENCE with pistil-
late spadix 3-5 cm long; berries white, + oblong-
ellipsoid, 1.1-2 mm long, 0.4–0.8 mm diam.; seeds
3-6 per locule, tan.
Flowering phenology in Philodendron morii is
poorly known, but it is perhaps bimodal with flow-
ering collections known in March and November
and immature fruits in December, February, and
June.
Philodendron morii is endemic to Panama,
known only from highlands east of the Canal Area
in Panamá Province and Comarca de San Blas and
from Cerro Pirre in Darién Province, at 450 to 850
m elevation in Tropical wet forest and Premontane
rain forest life zones.
Philodendron morii is a member of P. sect. Са-
lostigma subsect. Glossophyllum ser. Glossophyl-
lum. This species is characterized by its slender
internodes (about as long as broad); bluntly two-
ribbed, deciduous cataphylls; terete petioles (about
as long as the blades), which often dry with loose,
puffy epidermis; ovate-triangular, scarcely cordate,
green-drying blades; and peduncles equaling or ex-
ceeding the green spathes.
Philodendron morii is most similar to P. sphal-
erum Schott from the Guianas, which has similar
green-drying, long-petiolate leaves. The latter spe
cies differs in having the leaf blades acute 10
rounded or truncate at the base and proportionately
longer petioles (fully as long as or much longer than
the blades). In addition, P. sphalerum has up to four
much smaller inflorescences with spathes 5.5-1 cm
long, whereas P. morii has one to two much larger
inflorescences per ах! (spathes 9-12.5 em long).
Philodendron morii is also similar to P. —
especially in terms of size and color of its le
uri م co а AD کار وای cli шл ГА E
Volume 84, Number 3
1997
Croat 493
Philodendron Subgenus Philodendron
blades. The latter species differs, however, in com-
prising more or less scandent plants with usually
long internodes.
Philodendron morii is not easily confused with
any other Central American species. It is named in
honor of one of its earliest collectors, Scott Mori
(NY), who collected for the Missouri Botanical Gar-
den during 1974-1975.
Additional specimens examined. PANAMA. Darién:
Parque Nacional Darién, W side of Cerro Pirre, 800-1050
m, 7°56'N, 77%45'W, Croat 68700B (MO). Panamá: Cerro
Jefe region, Altos de Azul-Río Chagres, 700-850 m
M N, 79°30'W, Г аад 11899 (МО, РМА, 05); a
(CM, MO); Campos Tres, 3 mi. NE of Altos
500-800 m, Liesner 567 (MO); Gorgas Memorial Labs
“Campamento Quatro,” 5-10 km NE of Altos de Pacora,
78"34'W, Herrera et al. 1489 (MO, PMA, US).
Philodendron niqueanum Croat, sp. nov. TYPE:
Panama. Darién: Serranía de Pirre, along
headwaters of Río Escucha Ruido, ca. 16 km
due N of Alto de Nique, ca. 7°47'N, 77%45'W,
27 July 1976, Croat 37942 (holotype, МО–
2416709). Figures 293-296.
Planta hemiepiphytica; internodia brevia, usque 5 cm
diam.; cataphylla incostata vel leniter 1-costata, rubra,
pers istentia plus minusve intacta; petiolus teres, (32)46—
59 с
m longus, in sicco mm diam.; lamina ovata vel
late ae 2842.55 5 cm longa, (14)2 528.5 cm lata,
onga quam petioli; sinus plus minusve V-for-
matus; indices: immatura; pedunculus 4 cm longus;
spatħaé vi virid
Minds stem scandent when young, ap-
pressed-climbing when adult; internodes short, to 5
cm diam. (younger stems with internodes to 8 cm
long, 1.5 cm diam.), epidermis moderately smooth,
dark reddish brown; cataphylls unribbed to weakly
l-ribbed, red, persisting + intact; petioles
(32)46-59 cm long, 6-8 mm diam., terete; blades
ovate to broadl ovate, acuminate at apex, truncate
to subcordate at base, (28)42-55.5 cm long,
(14)25-28.5 ст wide (1.7-2 times longer than
wide), (ca. 0.9 times the petiole length), about equal
in length to petiole, upper surface weakly glossy,
drying dark brown, lower surface paler, drying dark
yellowish brown; anterior lobe (27)37-49.5 cm
long, 25-28.5 cm wide (4.7-5.4(13.5) times longer
than posterior lobes); posterior lobes (2)7-9 cm
long, 9-12 cm wide, broadly rounded to obtuse;
sinus * V-shaped, to 4 cm deep; midrib promi-
nently raised, paler than surface above, raised be-
low; basal veins 3 per side, with О free to base,
some of the lowermost coalesced to ca. 1 cm; pos-
terior rib never naked; primary lateral veins 5-9
per side, departing midrib at а 45-65°(70°) angle,
weakly arcuate to the margins, sunken above,
raised below; minor veins obscurely visible, slightly
raised on drying below, arising from the midrib
only. INFLORESCENCES immature; peduncle 4
cm long; spathe green, 7 cm long; spadix imma-
re.
Flowering in Philodendron niqueanum is poorly
known owing to too few collections overall. The
species was collected with flower buds in late July
and probably both flowers and fruits within the
rainy season (although, since it flowers so late, the
fruits may mature in the dry season).
Philodendron niqueanum is endemic to Panama,
known only from the type locality on the Serranía
de Pirre, at 1530 to 1550 m elevation in Tropical
Lower Montane wet forest.
Philodendron niqueanum is a member of P. sect.
weakly one-ribbed cataphylls per-
sisting mostly intact; terete petioles; and narrowly
ovate, dark brown-drying blades about as long as
the petioles.
Philodendron niqueanum is apparently close to
P. lentii, which ranges from Costa Rica to central
Panama, but no further east than the province of
Coclé. Both species are similar in having truncate-
to subcordate-based leaf blades with more or less
V-shaped sinuses. Philodendron lentii differs in
having the primary lateral veins sunken, paler, and
much more conspicuous on the upper dried blade
surface. In contrast, the primary lateral veins of P.
niqueanum are scarcely or not at all paler than the
surface on dried leaves and are raised rather than
sunken. In addition, the epidermal pattern is al-
veolate and moderately smooth at 10X magnifica-
tion on the upper blade surface of P. niqueanum,
whereas P. lentii lacks an alveolate pattern and the
adaxial surface is densely covered with round, pale
inclusions.
Philodendron niqueanum is named for the type
locality near the Alto de Nique, hence the name.
Additional specimen examined. PANAMA. Darién:
Cerro Pirre region, ca. 9 km from Alto de Nique, 1480—
1520 m, Croat 37886 (MO, PMA, US).
Philodendron panamense K. Krause, in Engl. &
K. Krause, Pflanzenr. IV. 23Db (Heft 60): 65.
1913. TYPE: Panama. Canal Area: at Frijoles,
25-30 m, 9°10'N, 79487, Pittier 3753 (ho-
lotype, US). Figures 297-2
Usually hemiepiphytic; stem appressed-climb-
494
Annals of the
Missouri Botanical Garden
ing, to 1.3 m long, sap reddish, sticky; internodes
short, semiglossy, 2.5—4 cm di ometimes lon-
ger than broad, dark green; roots several per node,
drying 2—4 mm diam., dark brown, semiglossy,
sparsely scaly; cataphylls 18-20 cm long, sharply
l-ribbed to sharply 2-ribbed, green to whitish, dry-
ing light brown, persisting briefly + intact, even-
tually fibrous, sometimes persisting for a time,
eventually deciduous; petioles erect-spreading,
(23)34—70(79) cm long, 4-13 mm diam., terete to
subterete, sometimes weakly flattened or with nar-
row flattened rib adaxially, sometimes weakly and
narrowly sulcate at base, dark green, sometimes
pink at base, surface sparsely to densely pale
greenish striate or striate-lineate, minutely grooved
upon drying, geniculum to 6.5 cm long, sheath 1—
4 cm long, usually inconspicuous, to 10 cm long
when subtending an inflorescence; blades broadly
triangular-ovate or more infrequently ovate, sub-
coriaceous, semiglossy, moderately bicolorous,
abruptly acuminate, sometimes acute at apex (the
acumen sometimes inrolled, to 4 mm long), deeply
cordate at base, 32-72 cm long, 24-38 cm wide
(1.2-2.3 times longer than wide, averaging 1.5),
((0.3)0.7—1.4 times longer than petiole, averaging
1.07), broadest near point of petiole attachment;
upper surface dark green and glossy, drying semi-
glossy, dark gray-brown to olive-green, often some-
what blackened, lower surface slightly paler, se-
miglossy; margins sometimes broadly undulate;
anterior lobe 25—41(58) cm long, (15)20—36(41) cm
wide (1.94 times longer than posterior lobes); pos-
terior lobes 7.5-18 cm long, 4.5-17.7 cm wide,
rounded to broadly rounded to broadly obtuse; si-
nus hippocrepiform to parabolic (arcuate on youn-
ger blades), 4-12 cm deep; midrib broadly sunken,
concolorous or paler than surface above, weakly as-
perous, thicker than broad, matte, sometimes short-
white-striate, darker than surface below; basal
veins 5-8 per side, with 1 free to base or nearly
so, third and higher order veins coalesced 4—7 cm
long; posterior rib naked for 2-3 cm long; primary
lateral veins 4—7 per side, departing midrib at a
55—65° angle, spreading to a 65-75” angle, usually
curved down gradually before merging with the
midrib, narrowly sunken, concolorous or paler than
surface above, convex, matte, slightly darker than
surface below; interprimary veins narrowly sunken
above; minor veins distinct, darker than surface be-
low, arising from both the midrib and primary lat-
eral veins; secretory ducts moderately visible on
lower dried surface, alternating with minor veins.
INFLORESCENCES = erect, 4(6) per axil; pedun-
cle (4.5)6.5-20 (most more than 15) cm long, 4—12
mm diam., pale green, strongly white-lineate,
slightly to moderately bent just below the spathe;
spathe 10.5-18.5 cm long, (0.6—1.7(2.7) times lon-
ger than peduncle), acute at apex; spathe blade
white outside, (opening 4—7.5 cm wide), pale green,
moderately glossy to pale-punctate inside; spathe
tube ellipsoid, medium green, densely pale-speck-
led outside, 6–9 cm long, to 4.5 cm diam., pale
green, moderately glossy to pale-punctate inside,
spadix sessile; protruding forward at anthesis, 12-
16 cm long, broadest at upper two-thirds constrict-
ed to ca. 1.5 cm diam. between sterile staminate
portion and fertile staminate portion; pistillate por-
tion pale green, cylindrical, 3-6.5 cm long in front,
2.7-3.6 ст long in back, 1.3-1.7 mm diam. at
apex, 1.3-1.9 mm diam. at middle, 1.1-1.5 mm
wide at base; staminate portion 10.5-14.7 cm long;
fertile staminate portion creamy white, + clavate,
1.5-1.8 cm diam. at base, 1.3-2 cm diam. at mid-
dle, 9-12 mm diam. ca. 1 cm from apex, about as
broad as the pistillate portion; sterile staminate por-
tion 1-1.9 cm diam.; pistils 2.1-3.4 mm long, 1-
1.6 mm diam.; ovary 6(7)-locular, 1.5-3.1 mm long,
1-1.6 mm diam., with axile placentation, walls
sometimes embedded with granular, crystal-like
particles; locules 1.5-3 mm long, 0.5-0.7 mm
diam.; ovules 20-31 per locule, 2-seriate, 0.3 mm
long; funicle 0.2 mm long, adnate to lower part of
partition, style 0.5-0.6 mm long, 1.3-1.6 mm
diam., similar to style type B; style apex rounded
or domed; stigma truncate, hemispheroid, 1-13
mm diam., 0.3-0.5 mm high, covering entire style
apex; the androecium truncate, prismatic, oblong,
margins acutely and regularly 4—6-sided, 0.9-1 mm
long, 0.7-2.2 mm diam. at apex; thecae oblong to
cylindrical, 0.3-0.5 mm wide, + parallel to one
another, contiguous; sterile staminate flowers blunt-
ly, irregularly 4—6-sided, 1.9-4.1 mm long, Lbs
mm wide. INFRUCTESCENCE with seeds many
per berry, white, narrowly cylindrical, 1.3 mm long.
sticky. JUVENILE petioles terete, sheathing broad-
ly, for % to 34 its petiole length, acute to rounded,
eventually weakly to strongly cordate; blades
broadest at the middle.
Flowering in Philodendron panamense occurs
during the dry season and early rainy season
(March through May), with post-anthesis inflores-
cences collected from May through August and im-
mature fruits from July through November.
Philodendron panamense is endemic to Panama,
but it is likely to occur also in adjacent ah
In Panama, it occurs in Tropical moist forest on bo
slopes of the Canal Zone, and Premontane wet forest
and Tropical wet forest in Panamá, Colón, and Da-
Volume 84, Number 3
1997
Croat 495
Philodendron Subgenus Philodendron
rién Provinces, at sea level to 800 m elevation (but
mostly below 300 m).
Philodendron panamense is a member of P. sect.
Philodendron subsect. Philodendron ser. Fibrosa.
Philodendron panamense is characterized by its ap-
КО ae’ hemiepiphytic habit; short inter-
nodes; weakly one-ribbed, mostly deciduous cata-
phylls vet ibn persisting intact or as fibers);
terete, pale-striate petioles; usually broadly trian-
gular-ovate, moderately thin blades with parabolic
to hippocrepiform sinus; and by the long-pedun-
white blade (greenish white throughout within).
Also characteristic is the peduncle, which is often
bent abruptly just beneath the spathe.
his species is most easily confused with P. la-
zorii, which has similar greenish inflorescences.
The latter species differs in having more broadly
ovate blades (0.97—1.4 times longer than wide and
averaging 1.1 times) matte on the lower surface. In
contrast, the blades of Р. panamense are usually
ovate-triangular, 1.3 or more times longer than
broad (averaging 1.5 times longer) with the lower
surface semiglossy. In addition, the peduncles of Р.
lazorii are proportionately shorter and usually
shorter than the spathe (vs. often longer than the
spathe in P. panamense).
Philodendron panamense has been confused with
P. jodavisianum, which has leaves drying a similar,
somewhat blackened color. The latter species dif-
fers, however, in having D- to U-shaped petioles,
typically with a medial rib adaxially, as well as
more typically persistent cataphyll fibers and much
shorter peduncles (typically less than 10 cm long
in P. jodavisianum, vs. typically more than 10 cm
long in P. panamense). In addition, the peduncles
of P. jodavisianum are usually straight, not bent.
А collection from the Serranía del Pirre at Cana,
Croat 37600, is unusual in having several persis-
tent cataphylls and shorter-than-usual peduncles.
Collections from Cerro Sapo and Cerro Pirre
(Croat 55184 and 68951 respectively) differ from
typical material of P. panamense in having ovate
(rather than triangular ovate) leaves and more
short-pedunculate inflorescences. In addition, they
have sharply two-ribbed cataphylls that persist in-
tact (in the case of Croat 55184, on plants in the
living collection. at MO) ог semi-intact (Croat
951, collected in the wild on Cerro Pirre). Most
ied collections of P. panamense have по cata-
Phylls and longer peduncles.
Earlier (Croat, 1978), the species was reported
to be much more widely distributed (to Honduras
and Ecuador), but collections from outside of Pan-
ama have since proven to be misidentified (most
now P. jodavisianum).
e examined. PANAMA. Canal ر
: Ват Island, Croat 5101 (MO); 5530
(МО); 5840 (E. MO, 5; 6188 (МО); ад = РМА);
8993 (МО); 9292 (МО, PMA, SCZ); 3 (MO, SCZ);
10264 (MO, SCZ, US); brie (MO); ence (MO); 11016
(MO); 14876 (MO); Bailey & Bailey 199 (BH); Parque
Nacional Soberania, Pipeline Road, N of Gamboa, Luteyn
& ои 1556 (MO); Río Macho ae 10 km NW of
°4 Wy, roat & Zhu
(МО); cibis Cais: Croat 10867 (F. MO). edem т,
tobelo-Nombre de Dios, 0.5 mi. beyond junction of road
to Isla Grande, 9°40'N, 79°35'W, Croat 49804 (МО); near
Nuevo Tonosf, <100 m, Croat 33517 (MO, RSA). Darién:
Cerro Sapo, Croat 55184 (AAU, K, MEXU, MO, PMA,
US); ca. 5 km 5 of Garachiné, Rfo San Antonio, 130 m
TSO'N. 78?25'W, Hammel et al. 1481 (MO); Cerro Piers
region, NW of Cana, 600 m, Sullivan 672 (MO); vic. of
gold mine at Cana, 500-600 m, Croat 37600 (MO); Ser-
ranfa Sapo above Casa Vieja along boundary trail of Da-
rién National a 150-300 m
Pherson et met 5359 (COL,
5378 (MO, US); Parque Nac
Pirre AE. camp, trail E side of Rio Paracida, 0-80 m,
8°№, 77°48'W, Croat 68991 (СМ, L, MEXU, MO, PMA,
US); Estación Rancho Frío, at N base of Cerro Pirre, ca.
9 km S of pe Real, 70-270 m, 801'N, e Hammel
al. 16131 (MO); Cerro Pirre, ВО
°45'W, Croat ie (CAS, COL, Mb. Т. PMA): Río
Thane за, са air distance from Continental Divide,
vic. of Tyler ттан гуй mine, Croat 27193 (МО);
Parque Nacional Cerro Rio Perisenico, 110 m,
8°O1'N, TEMA W, Croat "s Zhu 77100 (CAS, CM, F, MO).
erro Azul,
Cerro Nene er Dwyer et al. 4848 (MO), Croat 12074
(МО, SCZ).
Philodendron pirrense Croat, sp. nov. TYPE:
anama. Darién: Cerro Pirre, middle slopes on
western approach, 800-1050 m, 7%56'N,
77°45'W, 29 June 1988, Croat 68952 (holo-
type, MO-3610823-24; isotypes, B, COL, F,
K, NY, PMA, US). Figures 301—305.
Planta hemiepiphytica aut terrestris; internodia 1-2 ст
longa, diam.; cataphylla 25-30 cm diam., acute
2-costata, persistentia ut fibrae grossae pallidae, cum frus-
tris rubrobrunneis epidermidis; petiolus 56.5—100 ст lon-
gus, 5-8 mm diam., aliquantum comp lanatus adaxialiter
cum costa inconspicua; lamina late ovato-cordaa, 3
ст longa, 27—46 с m lata, i in sicco brunnea; costa айын
uda 1—5 ст; ЗЕ изаар 4; pedunculus. manifeste al-
iy 8.5-12 cm longus, 6-8 mm diam.; spatha 12—
7 cm longa; lamina spathae extus viridis, suffusa mar-
ronina, intus a usa marronina; tu bo 8 s ae extus
p
loculi атда j albae. я
Hemiepiphytic or sometimes terrestrial; stem to
ст long; internodes short, semiglossy, closely
ribbed, completely enclosed in cataphyll fibers,
л
496
Annals of the
Missouri Botanical Garden
1-2 ст long, 4-5 cm diam.; roots to ca. 30 cm
long, drying reddish brown, smooth, semiglossy, ca.
2 mm diam., closely ridged; cataphylls 25-30 cm
long, sharply 2-ribbed (ribs prominently raised),
reddish, drying reddish brown, persisting semi-in-
tact, ultimately as coarse pale fibers with fragments
of reddish brown epidermis; petioles 56.5-100 cm
long, 5-8 mm diam., subterete, somewhat flattened
near base, weakly flattened toward apex, with faint
medial rib adaxially, surface densely short-lineate;
blades broadly ovate-cordate, acuminate to narrow-
ly acuminate at apex, cordate at base, 35-58 cm
long, 27-46 cm wide (1.2-1.5 times longer than
wide), (0.6-0.9 times longer than petiole), upper
surface drying reddish brown, semiglossy, lower
surface much paler; anterior lobe 24.5-48.5 cm
long, 27-50 cm wide (1.7-2.3 times longer than
posterior lobes); posterior lobes + rounded, 12.5—
19 cm long, 12.5-23.5 cm wide, broadly obtuse;
sinus hippocrepiform, 14 cm deep; midrib flat, pal-
er than surface above, drying reddish brown below;
basal veins 7-11 per side, with 0—1 free to base,
1-2 coalesced (4)6-8(10) cm, posterior rib well
developed, naked for 1-5 cm; primary lateral veins
6-10 per side, departing midrib at a 55—65° angle,
+ straight to the margins, prominently sunken
above, prominently raised below; interprimary
veins distinct, fine, mostly continuous, drying
darker than surface below; minor veins arising from
both the midrib and primary lateral veins. INFLO-
RESCENCES 4 per axil; peduncle 8.5-12 cm long,
diam., prominently white-striate, clearly
demarcated from spathe; spathe 12-16.7 cm long
(1.3-1.5 times longer than peduncle), moderately
constricted above the tube; spathe blade green,
tinged maroon, conspicuously and densely pale li-
neate-striate outside, white, tinged maroon inside;
spathe tube red-maroon, inconspicuously short-li-
neate outside, 4-6 cm long, dark maroon inside;
spadix sessile; + ovate, 11-13.4 cm long, broadest
below the middle; pistillate portion greenish white,
2.7 cm long, 1.4 cm diam. at apex, 1.3 cm wide at
base; staminate portion 10.8 cm long; staminate
portion 5.7-8.3 cm long; fertile staminate portion
white, tapered toward apex, 1.4 cm diam. at base,
1.3 cm diam. at middle, 8 mm diam. ca. 1 cm from
apex, broadest at base, about as broad as pistillate
portion; sterile staminate portion broader than the
pistillate portion, 1.4–1.5 ст diam., pistils 2.5 mm
long, 1.2-1.4 mm diam.; ovary 5-6-locular, 1.7 mm
long, 1.3 mm diam., with axile placentation, walls
embedded with granular, crystal-like particles; loc-
ules 1.7 mm long, 0.4 mm diam.; ovules 20 per
locule, 2-seriate, contained within gelatinous ma-
trix (no true envelope), 0.2-0.3 mm long, longer
than funicle; funicle 0.1-0.2 mm long, style 0.3 mm
long, 1.3 mm diam., similar to style type B; style
apex flat; stigma subdiscoid to slightly hemispher-
oid, weakly lobed, 1.2 mm diam., 0.1-0.3 mm high,
covering entire style apex; the androecium trun-
cate, prismatic, oblong, margins irregularly 4—6-
sided; thecae oblong, 0.5 mm wide, + parallel to
one another; sterile staminate flowers blunt, irreg-
ular, 4—6-sided, 2.1—3 mm long, 1.7 mm wide. Ber-
ries white.
Flowering in Philodendron pirrense occurs during
the early rainy season, judging by post-anthesis col-
lections from June and July.
Philodendron pirrense is endemic to Panama,
where it is known for certain only from the Serranfa
de Pirre, at 1000 to 1560 m elevation in Premon-
tane rain forest.
Philodendron pirrense is a member of P. sect.
Philodendron subsect. Philodendron ser. Fibrosa.
This species is distinguished by its short inter-
nodes; sharply two-high-ribbed cataphylls persist-
ing as pale fibers with tiny fragments of thin, red-
dish brown epidermis; petioles somewhat flattened
adaxially with a faint medial rib; broadly ovate-cor-
date blades with a well-developed posterior rib na-
ked to the sinus up to half its length; inflorescences
up to four per axil; and whitish peduncles clearly
demarcated from the red spathe tube.
Philodendron pirrense is similar to P. copense,
with which it shares dried leaves of similar color
and persistent, reddish brown cataphylls. The latter
species differs in having longer blades with the an-
terior lobes frequently concave along their margins
and petioles with a yellowish or reddish brown flak-
ing periderm.
Additional specimens examined. PANAMA. Darién:
Serranía de Pirre, ca. 12 km N of Alto de Nique, 1520-
1560 m, Croat 37916 (MO); ca. 9 km from Alto de Nique,
1480-1520 m, Croat 37887 (MO); Río Escucho Ruido,
ca. 16 km N of Alto de Nique, 1530-1550 m, Croat 37944
Philodendron platypetiolatum Madison, Selby-
: 22. 1977. TYPE: Ecuador. Los Ríos:
Río Palenque Science Center, km 56 on road
to Santo Domingo and Quevedo, 150-220 m,
ca. 0°35'S, 79°22'W, Dodson 6638 (holotype,
SEL; isotypes, MO, QCA, US). Figures 306-
308.
Hemiepiphytic; stem + scandent, loosely ap-
pressed-climbing, to 3 m long, semiglossy, sap
clear, watery, sticky, leaf scars to 2 cm wide; inter”
nodes slender, 14-20 cm long, 1-2 cm төнө <
ger than broad, sometimes somewhat flatten
Volume 84, Number 3
1997
Croat 497
Philodendron Subgenus Philodendron
side above, semiglossy, green to grayish green, epi-
dermis drying tannish brown, cracking, loosening
and flaking; roots pale to brownish, less than 20 cm
long, thin, 2 mm diam., smooth; cataphylls (6)10—
17 cm long, sharply 2-ribbed, sharply and deeply
sulcate with margins flared, pale to medium green,
drying yellowish to olive-green, deciduous; peti-
oles 15.5-56 cm long, 3-9(12) mm diam., broadly
flattened to markedly flattened and turned slightly
upward adaxially, broadly convex abaxially, firm,
medium to dark green, surface + unmarked;
blades ovate-triangular to broadly ovate, subcor-
iaceous, semiglossy to glossy, weakly to moderately
bicolorous, acuminate, sometimes long, narrowl
acuminate at apex (the acumen tightly inrolled, 2—
3 mm long), weakly cordate at base, 17-39 cm
long, 12.7-29 cm wide (1-1.5 times longer than
wide), (0.6-1.4 times longer than petiole), usually
about equal in length to petiole; anterior lobe 15—
34 cm long, 13-29 cm wide (2.4—4.5(5.2—5.4) times
longer than posterior lobes); posterior lobes 4-10
ст long, 5-12 cm wide, broadly rounded to obtuse;
sinus hippocrepiform, rarely arcuate with blade de-
current on petiole; midrib sunken or flat to broadly
convex above, slightly paler than surface to con-
colorous above, bluntly low-triangular to convex be-
low, paler than surface below; basal veins (2)3—4(5)
per side, with 0-1 free to base, second and third
coalesced 1-2 cm; posterior rib naked for 0-1 cm;
primary lateral veins 3-5 per side, departing mid-
rib at a 50-60” angle, straight to the margins, sunk-
en to weakly sunken above, convex below; minor
veins moderately indistinct, arising from both the
midrib and primary lateral veins. INFLORES-
CENCES 1(2) per axil; peduncle 5-20 cm long,
2-11 mm diam., subterete ог bluntly 1-2-angled,
pale green, semiglossy, unmarked; spathe semi-
glossy, (8)11-18 cm long (0.7-2(2.8) times longer
than peduncle); spathe blade green or red outside,
pale yellow-green (dark red in South America), in-
side; spathe tube green to olive-green, usually
tinged red or maroon outside, 4.5-7.5 cm long,
tinged red or maroon inside; spadix sessile; weakly
tapered, + acute at apex, 8-9.8 cm long, broadest
at the base or + uniform throughout, constricted
weakly between sterile and fertile staminate por-
tions; pistillate portion pale greenish, cylindrical or
y tapered toward the apex, 2.5(3.5) cm long,
11 mm diam. at apex, 10-13 mm diam. at mid-
dle, 10-13 mm wide at base; staminate portion 5.7—
7.3 cm long; fertile staminate portion white, weakly
tapered or cylindrical, 5-7 mm diam. at base, 4-5
mm diam. ca. 1 cm from apex, broadest at the base
or + uniform, narrower than the pistillate portion,
broader than or as broad as the sterile portion; ster-
ile staminate portion as broad as or slightly narrow-
er than the pistillate portion, 5-10 mm diam.; pis-
tils (0.8)1.7-2.5 mm long, (0.6)0.9-1.4 mm diam.,
white; ovary (6)7—8-locular, (0.7)1.1—1.8 mm long,
0.8-1.2 mm diam., with sub-basal placentation;
locules 0.7—1.5 mm long, 0.3 mm diam., ovule sac
0.8 mm long; ovules 3 per locule, l-seriate, con-
tained within translucent to transparent, gelatinous
envelope, (0.2—0.3)0.4—0.5 mm long, longer than
funicle; funicle 0.2-0.3 mm long (can be pulled
free to base), style (0.1)0.4—0.7 mm long, 0.6–0.8
mm diam., similar to style type B; style apex steep-
ly sloping, sometimes with small medial depression;
stigma subdiscoid to hemispheroid, somewhat cu-
pullate, 0.7-0.9(1.3) mm diam., 0.2-0.3 mm high,
covering entire style apex; the androecium pris-
matic, truncate, oblong, margins irregularly 4—6-
sided, sometimes weakly scalloped; thecae cylin-
drical to oblong, 0.3—0.4 mm wide, = parallel to
one another, sometimes + contiguous; sterile sta-
minate flowers irregularly 4—6-sided or rounded,
0.9-1.5 mm long, 0.7-1.2 mm wide. INFRUC-
TESCENCE turning red outside; berries green
(immature).
Flowering in Philodendron platypetiolatum ос-
curs in the early rainy season, June and September,
in Central America based on the few available flow-
ering collections. Post-anthesis collections from
Costa Rica and Panama are mainly from August
through November, but also from March. Immature
fruits have been collected only in January, Febru-
ary, and November. Ecuadorian populations have a
similar phenology, but flowering collections have
been made earlier, in March and April, with post-
anthesis collections from March through August.
Philodendron platypetiolatum ranges from Nica-
ragua to Ecuador, from sea level to 1500 (though
most collections are from below 400) m elevation,
in Premontane wet forest, Tropical wet forest tran-
sition to Premontane wet forest, and Tropical wet
forest life zones. This species has been collected at
several localities in Chocó and Valle Departments;
in Ecuador, it has only been collected at and near
the type locality.
Philodendron platypetiolatum is a member of P.
sect. Calostigma subsect. Macrobelium ser. Macrobel-
ium. This species is characterized by its scandent
habit moderately long, slender internodes with tan-
nish brown, frequently flaking epidermis; markedly
flattened petioles about as long as the blade; and
ovate-tri to broadly ovate, weakly cordate
blades. Though the spathes may be entirely green out-
side, they usually are heavily tinged with red or ma-
roon outside and the tube within is likewise colored.
498
Annals of the
Missouri Botanical Garden
There are no other species in Cental America
with which P. platypetiolatum might be confused.
Madison compared this species to P. lechlerianum
Schott, a species from Peru and Bolivia that has
terete petioles.
Panama, this species ranges along the Atlantic
slope but may also occur on the Pacific slope near
the Continental Divide. In Ecuador, it is restricted
to the Pacific slope.
Ecuadorian specimens have blades that are more
broadly ovate than those from Panama, but no other
differences have been detected.
Additional specimens examined. COSTA RICA. Ala-
juela: Upala Road, 3 km NNE of Bijagua, 450 m,
10°45'N, 85°3’W, Burger € Baker 9890 (CR, F, MO, NY,
SEL); 5 km 5 of Canalete, near Río Zapote, 100-200 m
10%48'N, m W, Burger & Baker 9972 (F, MO); Dos
Rios, 5 de Brasilia, Río Pizote, 500 m, 10%55'N,
85°20’ W, poda 1001 (CR, MO, SAR); 17 in NW of
San op ор 10%14'14”N, 84733", Croat 68136
| МО); В Fires Tigra, vic. of La Tigra, 330
10°22’ N. 84 84°38'W, Croat 68205 (B, CM, G, M, MO,
SAR, US); Monteverde Cloud Forest Nature male Río
Peñas Blancas, 1250-1350 m, 9°17'N, 84 . Burger
et al. 10745 (F, MO); Vara pic аи oa 3 mi. N
of San Miguel, 380 m, Croat 35666 (MO); Cafias—Upala,
13.8 km N of Bijagua, 100-150 m, Croat 36433 (MO); 4
NNE of Bijagua, ca. ак m, Croat 36269 (MO); slopes
of Miravalles, above Bija Mit et al. 19066 (MO);
as Zarcas, A ا pm 8.5 km NE of Villa
da, 600 m, Croat 46973 (СЕ, P Heredia: “Star-
m, 10°26'N, 83°58’W, Stevens 13489 (MO
Station, "ast 2228 (MO); 100 m, — 329 (MO)
100 m, 647 (CAS); 50-80 m, 10?26'N, 84?01'W, Grayum
7665 (CR, Mox 6 km from Río н pei 5 km SSE
of pd pe 340 Ad 10*21'N, 84%04'W, Schatz & Grayum
, MO); Puerto Viejo 6 de Sarapiquí, 100 m, Croat
pes pon 10°26'N, 84701", 61213 (MO); Parque Na-
cional Braulio Carrillo, Río Peje-Río Sardinalito, Volcán
B та, 10°
(МО); 8874 (МО, RSA); Barra del Colorado, 0-2 m,
10%47'N, 83%35'W, Stevens 24214 (CR, :
„
6 (MO, NY); Braulio Carrillo-Guáp-
iles, 250-270 m, Croat 78744 (CR, INB, M MO); Río Сој-
orado, 14 km by air SW of Barra del tr 10-120
m, 10740'N, 83°40'W, Davidse & Herrera 31060 (CR,
MO); 16 km by air SW of Barra del Colorado, 10°39'N,
83°40'40"W, 31211 (MO). Puntarenas: Osa Península,
Piedras Blancas, 3.7 mi. W of Pido Aarón Highway,
m, 8°46'N, 83*18'W, Croat 67687 (CR, К, MO);
Rincón e Ом башти 100 m, Pennington et al. 1 a
(K); Fila Gamba, ca. 6 km from Golfito airport, <1
8°41'30"N, 83*12'W, Croat 59926 (СМ, K, MO); 200-300
m, Croat & Hannon 79290 (MO). San José: El General
Valley, Finca Volcán Angel, Schubert & Rogerson 769 (A,
GH); Braulio Carrillo National park, 600—700 m, Croat
78778 (CR, INB, MO). NICARAGUA. Río San Juan: Río
Santa Cruz-Cafio Santa Crucita, La Palma, 40–60 m
11°2-4'N, 84724—26' W, Stevens 23496 (MO). Zelaya: Río
Sucio, E of Bonanza, ca. 140 m, 14?01'N, 84?34' W, Ste-
vens 12347 (MO); Mpio. Siuna, Coiharoi Danlí, 100—130
m, Ortiz 218 (MO). PANAMA. Bocas del Toro: Fortuna
Dam area, b gba cie] pus 9.4 mi. N of Con-
tinental Divide, gs 8°46'N, d W, Croat 66822
AAU, MO). Col : Ко Guan *s m above
bridge on road to abel J: a io & F. Wither-
spoon 8662 (CAS, MO); Sabanitas—Portobello, Río Piedras
Lumber Road, 6.7 mi. E of Sabanitas, 250 m, 9"22'30"М,
19*41'30"W, Croat 75166 (MO, PMA). Darién: Parque
Nacional Cerro Pirre region, Cana, 500-600 m, Croat
37661 (MO); near station along Río Perisenico, 110 m,
8°O1'N, 77°44'W, Croat & Zhu 77116 (MO); Parque Na-
cional Darién, Ко Topalisa-Río Pucuro, ca. 17 km E о
~
13.8 km N of Pan-American Highway, Folsom et al. 5788
(MO); Mile 6.8, 350 m, Croat 49125 (MO, PMA); Cerro
Jefe region, 0.8 mi. beyond turnoff to Altos de Pacora,
770 m, 9°15'N, 79°29'W, Croat & Zhu 76645 (MO); 4.6
79°22'W, Croat 67083 (MO); 3-3.5
Pacora, 700-750 m, 9°15'N, 79 ai Ww. His 68680 (CM,
MO). San Blas: El Llano-Ca , vic. Nusagandi,
300—350 m, 9?15'N, 79W, Geant 69278 (CM, MO); 450
is ем oe W, Croat 75118 (CAS, MO, NY, PMA);
1. No n highway, 300 m, 920'N, 79°W, Croat
Fete he (CM. M, КМО); Mile 9, 350 т, pede 79W,
Croat 76999 (MO); Río Playón Chico, 8 80-200 m
9°13.5'N, 78°15'W, Herrera & Arosemena 1778 (AAU,
COL, CR, K, MEXU, MO, NY, P, PMA, STRI, US).
Philodendron pseudauriculatum Croat, sp. nov.
TYPE: Panama. Panamá: El Re. road,
4 mi. from Inter-American H El Llano,
ca. 300 m, 27 Mar. 1976, Са 33730 (holo-
type, MO-2381528; isotypes, PMA, RSA,
SEL). Figures 35, 300, 309, 310, 313, 314.
Planta hemiepiphytica; internodia 1.4(9) cm longa, 2-
4 cm diam.; cataphylla 18-27 cm longa, acute 2-costata,
decidua; patindi subspongiosus, 11—42 с m longus, 0.7
2. pa m diam; T (S ARS deg epis adaxialiter,
um annuluo viridi apice:
od оаа E ‘oblanceolato-elliptica, уй
мама“ асша уе p+ pared rotundata, peng subcordata
cata m longa, 7.5-25 с
airieaniviridis: eg 2-3; reee 5.
m longus, 8-12
omnino alba vel subrosea, marginibus cremels;
—
3
argines, vivide rubriviolaceo vel a 1-2(4)-
Ah ueste рде ш ا 5-8(9)- e adi loculi 1 e
ovulati; baccae aurantiacae.
Hemiepiphytic; stem appressed-climbing, 10 1m
long; internodes gray-green, glossy to semiglossy.
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
1—4(9) cm long, 2—4 cm diam., usually somewhat
flattened on one side, frequently with a short series
of transverse ridges just below the nodes on both
sides of the rounded portion of internodes, usually
about as broad as long, or broader than long on
flowering plants, sometimes slightly longer than
broad; roots dark brown, slender, few per node; cat-
aphylls moderately spongy, 18-27 cm long, sharply
2-ribbed (ribs to ca. 1 cm high), medium green,
densely dark green short-lineate, deciduous, apic-
ulate (apiculum >1 cm long) at apex. LEAVES
erect, + rosulate, somewhat clustered at or near
stem apex; petioles 11-42 cm long, 0.7-2.6 cm
iam., subterete, somewhat spongy, slightly flat-
tened, bluntly and broadly sulcate midway, less so
toward apex, more so toward base adaxially, surface
medium green, semiglossy, with dark green ring
around apex; blades oblong-elliptic to oblanceo-
late-elliptic, subcoriaceous to coriaceous, somewhat
to markedly bicolorous, semiglossy, acuminate,
sometimes long-acuminate at apex, mostly acute to
rounded, sometimes broadly subcordate, or some-
times truncate at base, 27-80 cm long, 7.5-25 cm
wide (2.2-4(5) times longer than wide), (0.97-3.6
times longer than petiole), upper surface dark
green, drying dark gray-green, lower surface much
paler, yellow-green, drying yellow-green to yellow-
brown; sinus (when present) to 2.5 cm deep; midrib
flattened at base, slightly sulcate midway, broadly
convex at apex and concolorous above, convex,
short-green-lineate, paler than surface below; basal
veins (0)1-3(4), all free to base; posterior rib lack-
ing; primary lateral veins 8-14 per side, departing
midrib at a 65-75” angle (45-55” angle at apex),
= straight to the margins, sunken above, convex
and slightly paler than surface or darker than sur-
ace below; interprimary veins flat, darker than sur-
face below; minor veins moderately distinct and
fine below, arising from both the midrib and pri-
mary lateral veins. INFLORESCENCES 2-3 per
axil; peduncle 5.5-21(25) cm long, 8-12 mm
diam., pale to medium green, finely white- or dark-
striate; spathe (10.6)12-23 cm long, 1.7 cm diam.
0.7-2.6 times longer than peduncle), constricted
midway above the tube, white to pinkish through-
out, margins cream; spathe blade creamy white to
yellowish green with faint green tinge along center
on backside (B & K yellow-red 7.5/9), yellow-or-
ange striate outside, 8.5-9 cm long (opening ellip-
tic и — view, 7.5-9 cm long, 2-5.5 cm wide),
n to white inside; resin canals orange;
ње “a oblong-ellipsoid, medium to dark
green, sometimes weakly tinged purplish along
margins outside, densely short white-lineate,
Semi-glossy outside, 5.5-10.5 cm long, 1.84 ст
diam., bright red-violet to dark maroon (weakly so
toward apex) to yellowish orange, sometimes broad,
white-lineate or orange striate inside; spadix weak-
ly stipitate to 3-5 mm long, cylindrical to weakly
tapered, 7.5-15.3 cm long, broadest below the mid-
dle; pistillate portion cylindrical to clavate, pale
lime green to pale yellow to medium or dark green,
3–5.5 cm long in front, 1.7— cm long in back, 1—
1.2 cm diam. at apex, 1.5 cm diam. at middle, 8—
ll mm wide at base; staminate portion 4.5—
10.7(12) cm long; fertile staminate portion broadest
in middle, slightly tapered toward both ends and
broadened before the sterile portion, 9-16 mm
diam. at base, 9-10 mm diam. at middle, 5-10 mm
diam. ca. 1 cm from apex, broadest at the base, as
broad as the pistillate portion, narrower than the
sterile portion; sterile staminate portion broader
than the pistillate portion, white, 8-15 mm diam.;
pistils 1.7-3.2 mm long, 1.3-2 mm diam.; ovary 5—
8(9)-locular, 1.4-2.3 mm long, 1.5-2 mm diam.,
with sub-basal placentation, walls embedded with
granular, crystal-like particles; locules 1.4-2.1 mm
long, 0.3-0.5 mm diam.; ovule sac 1-1.2 mm long;
ovules 1-2(4) per locule, contained within trans-
parent, gelatinous ovule sac, 0.4-0.5 mm long, lon-
ger than funicle; funicle 0.1-0.5 mm long (can be
pulled free to base), style 0.4-1.4 mm long, 0.9—
2.1 mm diam., similar to style type B; style apex
sloping to rounded, with small medial depression;
stigma brush-like, cupulate, subdiscoid, 0.9-1.2
mm diam., 0.2-0.6 mm high, covering entire style
apex; the androecium truncate, prismatic, oblong,
margins irregularly 4—6-sided, 1.3-1.4 mm long,
6-1.8 mm diam. at apex; thecae oblong to cylin-
drical, 0.4-0.5 mm wide, nearly contiguous and +
parallel to one another; pollen ellipsoidal to sphe-
roidal, <1 mm long, <1 mm diam.; sterile stami-
nate flowers irregularly 4—5-sided, margins bluntly
rounded, 1.2-1.9 mm long, 1.8 mm wide. Berries
orange (mature) or white; seeds 5 per berry, 1.3—
1.9 mm long, 0.5-0.7 mm diam.
Flowering in Philodendron pseudauriculatum oc-
curs during the dry season and the first half of the
rainy season (January through September). Imma-
ture fruits have been collected during March, June,
and July, with mature fruits known only from Sep-
pal
–
@
Philodendron pseudauriculatum is definitely
known only from Panama and adjacent Colombia
(N Antioquia), ranging from 20 to 1400 m elevation
in Premontane wet forest and Tropical wet forest life
zones. Most collections have been made at La Mesa
(Coclé), Cerro Campana (Panamá), and along the
El Llano-Cartí Road (Panamá), but the species is
Annals of the
Missouri Botanical Garden
also known from a single collection in Bocas del
Toro сеже in the Serranfa де Cafiasas along the Pa-
cific Oce
Philodendron Sors ripe is a member of
' sect. Calos sect. Glossophyllum ser.
Glossophyllum. This fea is recognized by its ap-
pressed-climbing habit; short internodes; more or
less rosulate habit; somewhat spongy, subterete pet-
ioles; oblong-elliptic to oblanceolate-elliptic, dark
gray-green-drying blades with mostly narrowly
rounded bases, and two to three inflorescences per
axil; and white to pinkish spathes clearly demar-
cated from the peduncles.
Philodendron pseudauriculatum is most easily
confused with P. ligulatum, especially P. ligulatum
vars. heraclioanum and ligulatum, which have sim-
ilarly shaped blades. Philodendron ligulatum var.
heraclioanum differs in having sharply D-shaped
petioles with undulate-margined wings. While the
aforementioned varieties of P. ligulatum differ in
their usually vining habit and typically elongate in-
ternodes, these features are particularly apparent
in P. ligulatum var. ligulatum, which has inter-
nodes much longer than wide (vs. about as long as
broad or scarcely longer than broad as in P. pseu-
dauriculatum). In addition, the leaves of P. ligu-
latum frequently dry much darker, mostly some-
what blackened, rather than the typical yellowish
or brownish green of P. pseudauriculatum. Another
feature separating live material of the species is the
line of demarcation at the apex of the petiole, which
is purple in P. ligulatum rather than green as in P.
"e
ndron pseudauriculatum may also be
idus with both P. auriculatum (hence the epi-
thet *pseudauriculatum"), from the Pacific slope of
southwestern Costa Rica and P. wendlandii Schott
from the Atlantic slope of Costa Rica and Panama.
Philodendron auriculatum is distinguished by hav-
ing leaf blades drying a paler yellow-green color
and more narrowed toward the base with minute
narrow auriculate posterior lobes. It also has pro-
portionately somewhat longer petioles. Philoden-
dron wendlandii differs in having a much shorter
stem and petioles typically broader than thick with
sharp margins
Two collections (Croat 16908 and Knapp & Mal-
let 4658) from San Blas at Puerto Obaldía are prob-
ably also this species. They differ in having broader
blades and drying somewhat blacker. If these prove
to represent P. pseudauriculatum, the species is
most assuredly present in adjacent Chocó. A col-
lection from the Department of Santander in Colom-
bia at 1460 to 1700 m (García-Barriga & Jaramillo
19671) may also be this species.
Additional specimens examined. PANAMA. Bocas
del Toro: Quebrada Huron, Duwebdulup Peak, N of Río
Terebe, 300—900 ft., Kirkbride & Duke 558 (MO). Canal
Area: Summit Gardens, — que (F, MO, SCZ). Co-
0—710 m, Mori et
D
a
—
0
m
ES
g EZ
®
3
dà
m.
©
=
=
mae 37420 (MO); 900
(MO); 14362 ue , Croat & Zhu 76692 (M0);
Bartlett & Lasser 16694 E MICH, MO); Kennedy et al.
3189 (МО, PMA); 785 m, 8°37'N, 8008", Croat 67112
(MO); summit of Cerro Caracoral, near La Mesa, N of El
Valle de Antón, 1100 m, Knapp 1 i - MO); Río Cas-
cajal, Penonomé-Coclecito, 5.6 m of Llano Grande,
150 m, 8°46'N, 80°27'%, Croat zd (AAU, CAS, COL,
F, 1, MEXU, МО, NY, PMA, QCA, TEX).
Peluca, on road to е is Dios, Kennedy 2
MO); Portobelo-Nombre de
Croat 33526 (CM, COL, K, : мо, NY, PMA, QCA, RSA,
US); eng асанд а belo, Río Piedras drainage, 250 m,
922'30"N, 79°41'30"W, Croat 75160 (MO); Portobelo,
Croat & Porter duds (MO); Río Boquerón, Wo
Portobelo, ca. 1.5 ge.
79?4'W, Croat & Zhu 76251 (CR, MO, PMA); Río Guan-
che, 30-100 m, Croat 79322 (PMA); ca. 5 km ups
from road to Portobelo, 50 m, 9?30'N, 79°40'W, pee
& Trainer 14765 (MO); ca. 3-5 km above bridge, 50-200
m, Dt 37002 ee 10–100 m, 26147 (BR, F, MO); ca.
ca. 9°30'N 40'W, McPherson 8507 (MO); Río
panita, near bodie
Croat & Zhu 77094 (CM, МО); Cerro Pirre region, trail
1
of Tres Bocas, Kirkbride & Duke 1203 ( Ri
Cocalito, Whitefoord & Eddy 224 (BM) 162 а ~ MO);
Río Jaqué Valley, Quebrada Luka, 100-200 m TN,
7805 17, Knapp & Mallet 3199 (MO, NY); Río Tes
a. 2 km by air from Continental Divide, Croat 27160 (F.
МО); Clezio 168 (MO). P anamá: El Llano-C $
mi. from Pan-American Highway, 350 m, Croat 6734
(CAS, MO); Km 8-12, ca. 400—450 m,
(МО); ca. Mile 8, 225-275 т, 9°15’ oic
McPherson 10492 (ААЏ, MEXU, МО); Km 19.1, 350 ds
9*19'N, 78°55'W, de Nevers et al. {я мо; Ro Te
9°16'N, 79W, Knapp &
around Pilota de Toro, Folsom et al. ( :
Tortí-Pilota del Toro, above Tortí Arriba, Folsom e
» Ji
f highway, ca. 150 m ‚ 3599
74762 (F, МО); ca. ] mi. from E pe (M o. 5
PMA, US); ca. 850 m, 8/42'N, 79*56' W, Miller et al. ja
(MO); upper slopes, 207 Р deae 2595 (MO); above of
Lin Motel, Porter et al. 4250 (MO); Rio Torti, mere
Serranía de Cañazas, ca. S kw SW of Caaza, 159 ™
8°52’N, 78°22’ ^W. | Stein 1342 (МО); Sendero de Interpre-
Volume 84, Number 3
1997
Croat 501
Philodendron Subgenus Philodendron
tación, 1 km al este del Sonpanele. de los guardab-
osques de INRENARE, 800—900 m, 8?40'N, 79*55'W,
Correa & Montenegro 10681b (STRI). T Blas: El Llano-
Cartí Road, Km 19, 350 m, 9?19'N, 78°55'W, de Nevers
et al. 5598 (MO); Puerto Obaldía, 0—50 m, Pittier 4398
(US); beach E of Puerto Obaldía, Croat 16908 (MO); Puer-
to Obaldía-La Bonga, ca. 2 hours walk from Puerto Ob-
00 m, 9°20'N, 79°W, zu s e 76554
(CM, MO); ко Playón Chico, 80-200 m, 9?13'05"N,
78°15’ W, Herrera & Arosemena 1784 (MO, PMA, STRI).
COLOMBIA. Antioquia: Mpio. Turbo, carretera tapón
del Darién, sector Rio Le León-lomas aisladas, km 37, 20
m, Brand 1070 (COL, MO). Chocó: Mecana, N of Bahía
So Ла ano, 1-100 m, 6?16'N, 77°21'W, Juncosa 1609 (MO).
aralda: Mistrató, Jeguadas rr Serilis От
5*24'N, 76701", Betancur et al. 9 (MO); „соте.
miento de Santa Cecilia, 500-550 т, geet N, 76°13'W,
Betancur et al. 2930 (MO); Pueblo Rico, Santa nth
ueblo Rico, Km 13, Quebrada Pionda, 700-900 m,
5°17'N, 76°13'%, наба et al. 3052 (МО).
—
Philodendron pterotum K. Koch & Augustin, in
A. Braun et al., Append. gen. sp. Hort. berol.
1854: 6. 1854-1855. TYPE: Cultivated at
Berlin [received from Warszewicz in Venezue-
la] (holotype, B? lost). Panama. Canal Area:
vic. Fort Sherman, along road uw Gatün
Locks and Fort Sherman, ca. 3 mi. W of Сашп
Locks, «50 m, 919'N, 79°57'30°W. 17 July
1994, Croat & Zhu 76982 (neotype, MO-
4619421-26, here designated; isoneotypes,
AAU, B, CAS, CM, COL, CR, DUKE, F, GB,
GH, K, MEXU, P, PMA, QCA, RSA, SEL,
VEN, W). Figures 311, 312, 315, 316.
(drame espe Standl. & L. O. T Ceiba
PE: Costa Rica. Punt
apa i. de Osa, 75 m, 24 Mar. 1951, “Allen 6031
(holotype, EAP; isotype, F).
Usually hemiepiphytic or epiphytic; stem ap-
pressed-climbing, semiglossy, sap watery, weakly
turpentine-scented, leaf scars conspicuous, 2 cm
long, 1 cm wide; internodes sometimes obscured by
cataphylls, sparsely short-striate, about as long as
broad or sometimes longer than broad, 2-8 cm
diam., dark green to gray-green, eventually brown,
epidermis sometimes cracking, fissured longitudi-
nally; roots short, few per node; cataphylls 20-30
cm long, sharply 2-ribbed, C-shaped, semiglossy,
ark green, densely pale lineate, persisting as red-
dish brown, semi-intact fibers, eventually decidu-
ous; petioles (37)47-111 cm long, (3-5)6-13(16-
2) mm diam., erect-spreading, D-shaped, margin-
ally winged, broadly convex adaxially, with adaxial
Margins slender, erect, undulate, medium green,
weakly glossy, conspicuously pale striate; sheath
with margins involute; blades ovate, subcoriaceous,
semiglossy, moderately bicolorous, abruptly acu-
minate at apex (the acumen inrolled, 2-4(6) mm
long), broadly cordate at base, 36-93 cm long, 22—
84 cm wide (0.7-1.5(2.9) times longer than wide),
(0.6–0.9 times the petiole length), margins weakly
undulate, upper surface dark green, lower surface
much paler, glossy, drying yellow-green; anterior
lobe 23.5-65(84) cm long, 22.1-75(86.4) cm wide
(1.1-2.8 times longer than posterior lobes); poste-
rior lobes 12-33 cm long, (11)15-34 cm wide, di-
rected inward, broadly rounded to broadly obtuse;
sinus spathulate to rhombic; basal veins 7-10 per
side, first free to base, part of remainder coalesced
1-6 cm, loosely so distally; posterior rib naked to
6 cm; midrib flat to broadly sunken, paler than sur-
face above, convex to narrowly rounded, paler than
surface below; primary lateral veins 3-6 per side,
departing midrib at a 45-55” angle, + straight to
weakly arcuate to the margins, deeply sunken, pal-
er than surface above, raised to convex, paler than
surface below; interprimary veins weakly raised,
darker than surface below; the minor veins mod-
erately distinct, arising from both the midrib and
primary lateral veins. INFLORESCENCES erect,
2—4 per axil; peduncle (2)4–12 cm long, 4-13 mm
diam., medium green, coarsely white streaked to-
ward apex; spathe 11-29 cm long (1.4-3.8(5)
times longer than peduncle), + cuspidate at apex;
spathe blade light green outside, greenish white in-
side; spathe tube abruptly delineated from tube,
reddish to purplish to dark purple-violet (B & K
purple 2/10) or dark green and raised-white-striate
at base outside, red to magenta inside; spadix
weakly stipitate, exserted from the spathe, con-
stricted above sterile staminate portion; pistillate
portion pale green, 4.3 cm long in front, 3.8 cm
long in back, 2 cm diam. at apex, 1.7 cm wide at
base; staminate portion 16-18 cm long; fertile sta-
minate portion white, 1.7 cm diam. at middle; ster-
ile staminate portion 1.4-2.2 cm diam.; pistils
2.2-2.8 mm long, 1.1-1.4 mm diam.; ovary
6-locular, with axile placentation; locules 1.5-1.9
mm long, 0.4 mm diam.; ovules ca. 20 per locule,
2-seriate, somewhat translucent, 0.2-0.3 mm long,
longer than funicle; funicle 0.1-0.2 mm long, ad-
nate to lower part of partition, style similar to style
type B; style apex flat to weakly rounded; stigma
subdiscoid to weakly hemispheroid, sometimes
weakly lobed, 1.5 mm diam., 0.5 mm high, covering
entire style apex; the androecium truncate, pris-
matic, margins irregularly 4—6-sided, 0.7-1.5 mm
long; thecae oblong, 0.4 mm wide, not contiguous,
+ parallel to one another; sterile staminate flowers
irregularly 4—5-sided, 1.5-2.5 mm long. INFRUC-
TESCENCE with spathe green at base and tip, pur-
502
Annals of the
Missouri Botanical Garden
ple around fruiting area when ripe, berries pale
brown to white. JUVENILE and PRE-ADULT
plants with petioles flattened adaxially; blades
broadly ovate. PRE-ADULT blades broadly ovate,
28.5 cm long, 22.5 cm wide
Flowering in Philodendron pterotum occurs in
the early rainy season from May through August
based on both specimens and field observations; it
certainly must flower during part of the dry season
as well, since post-anthesis collections have been
made as early as March. Other post-anthesis col-
lections have been made in April and May but es-
pecially in June. Immature fruits have been col-
lected in February, March, May, July, October, and
November, especially October. Mature fruits have
been collected only in November.
Philodendron pterotum ranges from Nicaragua to
Central Panama, from sea level to 1900 (mostly be-
low 700) m elevation in Tropical moist forest and
Tropical wet forest life zones. In Nicaragua it occurs
only on the Atlantic slope, but in Costa Rica and
Panama it occurs on both slopes.
Philodendron pterotum is a member of P. sect.
Philodendron subsect. Platypodium. This species is
characterized by its broadly ovate juvenile leaves
with flattened petioles; appressed-climbing adult
habit with short internodes; persistent cataphyll fi-
bers; D-shaped petioles with slender, erect, undu-
late, marginal wings; and large, ovate, yellow-
green-drying blades with large inflorescences with
the spathe tube reddish to purplish on the outside
and much darker magenta within.
No type material has survived for P. pterotum.
The species was described from cultivated material
of a juvenile plant purportedly received from the
“little garden of Cl. Augustin,” which the tireless
traveler (Mr. Augustin) collected from Warszewicz
in Venezuela. It is important to note that it was not
said to have been collected in Venezuela (where it
does not occur), but only that it was obtained from
Warszewicz in Venezuela. There is much uncertain-
ty involved, since the extrapolation of juvenile
forms to adult plants is at best risky, even at the
type locality. Given the frequency of mishaps in-
volving the dispersal of living material among hor-
ticulturists and botanical gardens, there is always
the chance of a mislabeled plant.
Still, while it is not possible to confirm the true
nature of Koch and Augustin's plant, it is certain
that the plant which Schott illustrated in detail (Ico-
nes #2478, #2480, #725, and #726) really does
represent the plant currently being called P. pter-
otum. Although this Central American species does
not occur in Venezuela, Koch and Augustin seemed
not to state that the plant received from Venezuela
had been collected there, only that it had been re-
ceived from Warscewicz in Venezuela.
Krause (1913), in his revision “of Philodendron,
cited only a Wendland collection from Costa Rica.
That collection is neither at Göttingen (GOET) nor
Berlin (B) and must be lost. Since Engler reported
the Wendland collection to be alive at the Berlin
Botanical Garden and since Wendland made much
of his material available to Schott, it might have
been that it was the Wendland material from Costa
Rica that Schott illustrated, rather than the Koch
material received from Venezuela.
Whether these two elements corresponded to the
same species is conjectural, but Schott would have
conceivably had the opportunity to compare both,
and it is reasonable to assume that he made the
correct interpretation and that his use of P. pterotum
for the Central American species is correct. In any
event, there is a need for a neotype since no spec-
imen actually seen by Koch, Schott, or Engler and
Krause still exists. Therefore, a modern collection
has been chosen here.
Philodendron pterotum may be confused with P.
findens, but that species differs in having blades
that dry usually blackened and promptly split pin-
nately into segments.
guas Zarcas, 60 m, r & Stol-
ze 5186 (CR, F, US). поља La Selva Field _
. Puntar
(MO); Palmar Norte-Panamanian border.
79197 (CR, INB, MO); San Yi: de Coto iN Fila
de Cal and Cuesta Fila de 00-600 m, 8°41'N,
82^56.5"W, Hammel 14161 (MO): Carara Reserve, Que-
brada Bonita, ca. 35-80 m, 9%47'N, 84°36’ W, Grayum el
al. 5/21 (CR, MO). Isla del се 40 km NW of Corco-
vado National – Gó 63 (MO); Corcovado Na-
P 1546
(MO); 11600 (MO) 0-200 m, 8”29'N, 83 ~
2850 (CR, MO); 1-10 m, Kernan & Phillips 1026 (С
MO); Osa Península, са. 5 km W of Rincón de Osa, д
200 m, 842'N, 83?31'W, Burger & Е А 8867 (CR, >
МО, МУ uebrada Aguabuena-Quebrada Banegas. '
5 km W irem de Osa, 300-400 m, 8°42'N, 83733 А,
Grayum 4066 (CR, MO); vic. Жонси; C
79245 (INB, MO); Río Claro, along Inter-American High
way, 30 m, Croat 32945 (MO). San José:
Gen siib а he 9 mi. SW а
Croat 35372 (М О); Paris riscal, Z.P. La Can, и
Mora 2031 (CR). NICARAGUA. Zelaya: Biana T
pa, m before Cafio Piedra del Balsamo, is
Stevens 8812 (MO); near Bil Tingnia, 6 km N y
Bonanza, 150 m, Neill 3995 (MO); с Вака, ci "Y
km E of Río Coperna, 200-300 m 3°40'N, 84° =
Pipoly 4927 (MO); 4844 (МО); ae Waylawas,
Volume 84, Number 3
Croat 503
Philodendron Subgenus Philodendron
00-200 m, са. 1338-39'N, 84?48—49'W, бан vei
MO) пар e Caño Calcamo, ca. 5
Siuna, ca. 13%40'N, 84^45'W, Grijalva & Burgos 1536
(MO); El ин Emile са. 65 m 1339'N,
- ded W, Stevens 12902 (MO); Cerro Duas E km NE of
na, 500 m, Neill 3633 (MO). PANAMA. Bahía Soldado,
Cowell 224 (NY). Canal Area: Gatún—Piña, ca. 3 km 5
of Pifia, са. 50 m, Croat 36929 (MO); Сашп Lake Hutch-
ison & Wright 2885 (BH, UC, US); Barro Colorado Island,
Imore X20 (F, № Нела 3081 (US); Croat 10903
E
(MO); 10265 (MO, SCZ); 7143 (MO); 6640 (MO, PMA);
6581 (MO, SCZ); 5136 aioe Bailey & Bailey 328 (BH);
along road between Gattin Locks and Fort Sherman, ca. 3
mi. У of Gatün Locks, 1.4 mi. E of Ft. Sherman, 9?18'N,
79738", Croat 69860 (CM, MO); Summit Gardens, Croat
10792 (MO, SCZ). Chiriqui: Puerto Armuelles—San Bar-
.6 m j
m, Croat 21933 (MO). Colón: 4 km E of Buena Vista,
Quebrada Ancha, 80 m, Nee 7781 (MO, US); Portobelo—
Nombre de Dios, 1.2 mi. beyond the junction of the road
to Isla Grande, 79?35'W, 9°40'N, Croat 49810 (MO);
Santa Rita Ridge Road, 6.5 mi. E of Boyd-Roosevelt
Highway, 370 m, 9°21'15’W, 7944", Croat & Zhu
76965 (MO). Panama: 26.8 km E of Bayano ab. Fol-
som 3529 (MO).
Philodendron purpureoviride Engl., Bot. Jahrb.
Syst. 26: 526. 1899. TYPE: Ecuador. Guayas:
Balao, Eggers 14710 (holotype, B). Figures
317-320
Hemiepiphytic; stem appressed-climbing, scan-
dent, often pendent, green becoming brownish to
gray-green; internodes glossy, to 25 cm long,
(0.8)1-2 cm diam., longer than broad, gray-green,
semiglossy, + terete, epidermis drying light yellow-
brown, conspicuously fissured or ridged but
smooth, frequently flaking free; roots thin, + twist-
ing or sinuous, few per node; cataphylls 10-29 cm
long, unribbed to obtusely 1-ribbed or bluntly to
sharply 2-ribbed, cream to medium green, magenta
speckled, quickly deciduous, fragile; petioles 9—
24 cm long, 8-10 mm diam., + terete, somewhat
spongy, somewhat flattened adaxially, surface se-
m
ovate-cordate, subcoriaceous, concolorous or weak-
ly bicolorous, acuminate to long-acuminate at apex
(the acumen e sese at base, 12-23(27)
ст long, 7.4-19 c de (1.3-1.8 times longer
than wide), (0.8-2. rine times longer than petiole),
broadest just below point of petiole attachment, up-
Per surface drying yellow-green, semiglossy, lower
surface glossy; anterior lobe 10–21 cm long, 7.4—
19 cm wide (2.5—4.2 times == than posterior
lobes) posterior lobes (2.8)3.6–7(8.2) cm long,
(3.14.3—7.7 cm wide; sinus hippocrepiform; midrib
Convex to broadly convex and slightly paler above,
convex to broadly convex and paler below; basal
veins ca. 3 per side, with 0-1 free to base, 2-3
coalesced to 5 mm long, flattened to raised; pos-
terior rib 0.5-1 cm long, never naked; primary lat-
eral veins about 3(4) per side, departing midrib at
angle, + straight to the margins, convex
to weakly raised above, convex below; interprimary
veins drying darker than surface below; tertiary
veins + obscure to visible and darker than surface
below; minor veins fine below, arising from both the
midrib and primary lateral veins; “cross-veins”
conspicuous (in Central America). INFLORES-
CENCES erect, 1 per axil; peduncle (3.5)5-7(11.5)
cm long, 5—7 mm diam., subterete, purplish tinged,
whitish streaked; spathe 11-15 cm long (1.2—
3(4.3—4.7) times longer than peduncle), constricted
only slightly midway above the tube; spathe blade
green to greenish white, with reddish speckling out-
ong, dark violet-purple inside; spadix
sessile; ca. 13 cm long; pistillate portion white to
pale greenish white, 4.5—5 cm long, 1.5 cm diam.
throughout; staminate portion 8.5-9.2 cm long; fer-
tile staminate portion white, drying reddish brown,
1.2 cm diam. throughout; sterile staminate portion
1.5 ст diam.; pistils 7.5 mm long, 1.6 mm diam.;
ovary 4—5-locular, 6.2 mm long, 1.6 mm diam., with
axile placentation; locules ca. 6.2 mm long; ovules
15-25 per locule, 0.2-0.25 mm long, 2-3-seriate,
style similar to style type D; style apex with low
style boss. INFRUCTESCENCE wi pistillate spa-
dix 6.5—7.5 ст long, 2.5-3.5 cm wide; berries 5.9
cm long, 2.5 cm diam.; seeds 24-25 per locule,
yellow-orange, 1.2 mm long, 0.5 mm diam., thin
and faintly striate.
Flowering in Philodendron purpureoviride is ap-
parently aseasonal with post-anthesis material col-
lected virtually MES Post-anthesis or early
fruiting collections have been made in every month
except September, but mature fruits have been col-
lected only in January and August.
Philodendron purpureoviride ranges from Costa
Rica and Panama to the Pacific slope of Colombia
and Ecuador (to Los Ríos and Guayas Provinces)
from sea level to 1600 m elevation in Premontane
rain forest, Tropical wet forest, Tropical wet forest
transition to Premontane wet forest, and Premontane
wet forest. In Central America, it is known primarily
from the Pacific slope of Costa Rica and adjacent
Panama, but also from the Atlantic slope in both
ountries.
Philodendron purpureoviride is a member of P.
t. Philodendron subsect. Solenosterigma. This
species is reportedly the dominant climber in west-
о
Annals of the
Missouri Botanical Garden
ern Ecuador in forests around San Sebastian, south
of Jipijapa in the province of Manabf (A. Gentry,
pers. comm.).
Philodendron purpureoviride is recognized by its
scandent habit; stems with the epidermis drying
yellow-brown, conspicuously exfoliating and fre-
quently fissured; terete to somewhat flattened pet-
ioles about four-fifths as long as the blades;
narrowly ovate-cordate blades drying yellow-green;
solitary inflorescences, with the spathe tube vio-
let-purple on both surfaces.
Philodendron purpureoviride is closest to and
perhaps inseparable from P. lechlerianum Schott
from Peru. The latter species is known from the
type collected at “San Gavan” (San Gabon) in the
Department of Puno, Carabaya Province, as well as
a more recent collection (D. N. Smith 6386) from
Pasco Department. Philodendron lechlerianum dif-
fers in having more conspicuous and prominent
cross-veins. If these names prove to be synony-
mous, P. lechlerianum is older and would extend
the range of the species into the Amazon basin.
Among sympatric species, Philodendron purpu-
reoviride is most easily confused with P. hedera-
ceum, also a vine with ovate-cordate, greenish-dry-
ing blades. The latter species is distinguished by
having stems usually drying green or dark brown
and without a peeling epidermis. In addition, the
blades are more coriaceous, typically more broadly
ovate with more (four to six) pairs of basal veins
(vs. two to three pairs for P. purpureoviride), and
generally have more prominent primary lateral
veins. Philodendron purpureoviride may also be
confused with some sterile specimens of P. wilburii.
The latter species has much longer peduncles and
stems drying darker brown and more closely fis-
sured with the epidermis seldom exfoliating.
Central American material of P. purpureoviride
differs from the Ecuadorian type by having con-
spicuous *
blade. The Ecuadorian material, by contrast, has
less conspicuous minor veins which lack “cross-
veins” except at or near the margins. In addition,
Engler's description of the inflorescence was based
solely on Eggers’s field label. Since the holotype is
sterile and no fertile material associated with Eg-
gers's original collection is apparently extant, fur-
ther studies should investigate whether these spec-
imens all represent the same species.
Additional specimens examined. COSTA RICA. Ala-
juela: 3.5 km W of Fortuna, 2.5 km NW of New Volcán
Arenal, 1500 m, 10%28'N, 84°41’ W, Taylor & Taylor
d (MO, NY, US); Cafias—Upala, 4 km NNE of Ала
. 400 m, Croat 36267 (МО); Upala, m, Rivera
1559 (INB, MO). Cartago: 12 km S of Turrialba by air,
throughout most of the,
4 km SE of Pejibaye along Río Gato, 700 m, 9°48’N,
83"42'W, Liesner 14355 (CR, MO); Río Reventazón, Tur-
, 500-600 m, 9°53.5'N, 83°38.5'W, Grayum &
Schatz 5242 (CR, MO); Tucurrique, Las Vueltas, 635-700
m, desi Il Yes Limón: Turrialba-Limón, along
Hig of Siquirres, 650 m, Croat
43332 yo Río Tlie Bajo Te Telire, 400-600 m, Gémez
24119 (MO). Pun : Zona Protectors Las “Tables,
Parque International ys yu istad, Finca Cafrosa, 1600-
1800 m, 8?53'20"N, 82*50'30"W, Mora 139 (CR, МО);
1680 m, Alfaro & Navarro 29 (INB); Cerro Ааай
950-1150 т, 8”49'18"N, 83°11'15"W, Grayum 1
(СВ, МО); Рајтаг Note to Jalisco, 780—960 m, 8°59.5'N,
83°28'W, Grayum 9141 (СК, F, К, MO, US); 50-700 m,
Croat 35203 (MO); Las Cruces Neil, Fila de Cal,
400 m, Gómez 19635 (MO d Xen Golfito,
100—500 m, Morales et al. 1903 (CR NB); 9 km W of
Rincón, Grant & rs ndell - 2.02203
Croat & Grayum 59857 (CAS, м CR, MO, NY); Villa
— Fila Gamba, ca. 6 km from Golfito air-
, «100 m, 841'30'N, 83°12'W, Croat 59902 (CR,
мо); Parque Nacional Corcovado, Dos Brazos de Rio
Madrigal, 600 m, 8”29'50'N,
со
. O2
o
N
eo
сл
e
=
i
a
£
~
со
bo
о
a
t NE
45
un
= 3
©
ES
un
@
=
Isidro del General—Dominical, 9 m
680 m, Croat Aes (MO); Weis КӨЕН, W Montañas
m^ са, т МЕ of Bijagual де Turrubares, 500-
600 m earn. 84°33’ w. iting et al. 5851 (CR, F.
MO). Ae AMA. Bocas del Toro: Fortuna Dam area,
near road to Chiriquí Grande, 650 m, 8°45’N, 82°15' i
губна 9925 (МО). Chiriqui: Burica Peninsula, 1
mi. W of Puerto Armuelles, vic. of San Bartolo pu
100—500 m, Liesner 84 (F, MO, US); 450 m, Busey 595
(F, MO); “Ojo de Agua,” Finca Hartmann, vicinity of Santa
850'N, 8
km from 3s Sereno, McPherson
K, MEXU, МО, PMA, US). Coclé: El Valle region, La
Mesa, above El Valle Be Antón, 860-900 m, Croat 37421
(MO); Finca А ч 800 m, 8°36'N, 80°%07' W, Croat
7 (MO
Pirre region, Cana,
senico, 110 m, 801'N, 77°44' W, Croat & Zhu Zl
(MO). ое Santa Fe area, between Santa Fe and
Calovébora, 1.7 mi. past Alto Piedra School, 570 m,
8°33’N, 81°08’ W, Croat & Zhu 76865 (MO).
Philodendron purulhense Croat, sp. nov. TYPE
Guatemala. Alta Verapaz: El uae >
on Hwy. CA-14, 2-3 mi Purulhá,
1500-1720 m, 15°13’S, 90°12'W, 21 July
1977, Croat 41752 (holotype, MO-2582045;
isotype, GUAT). Figures 321-323, 325, 326.
dne hemiepiphytica aut raro terrestris; internodia
, cataphylla 20-30 cm longa.
"itenim acute D-formata, persistentia semi-intacta; P"
tiolus subteres, aliquantum spongeosus, obtuse complan
atus adaxialiter, 32-52 ст longus, 1.5-2 ст
ina ovata vel он 25-48 ст longa, 18. а
lata, in sicco atribrunnea vel e nervis basali
liberis ad basim; fra a 1; pedunculus 2. 51 "i
longus; spatha 10-17 cm een lamina alae extus V
PPP AÑ kk ee -— — РО ——— > Ao x» c]
Volume 84, Number 3
1997
Croat 505
Philodendron Subgenus Philodendron
idi, breve pene intus viridi; pistilla 6—7-locularia; loculi
13-20-ovulat
Hemiepiphytic or rarely terrestrial, growing 3—6
m high in trees; stem appressed-climbing or creep-
ing; internodes short, semiglossy, to 6 cm diam.,
mm diam.; cataphylls 20-30 cm long, 2
E D-shaped, persisting semi-intact with
brownish fibers at upper nodes; petioles 32-52 ст
long, 1.5-2 cm diam., subterete, somewhat spongy,
obtusely flattened adaxially, surface semiglossy,
short-lineate, usually drying brown to blackened;
blades ovate to ovate-cordate, subcoriaceous,
slightly bicolorous, acute to acuminate at apex, cor-
date at base, 25-48 cm long, 18.5-40 cm wide
(1.2-1.97 times longer than wide), (0.6-0.98 times
the petiole length), about equal in length to petiole,
upper surface somewhat silvery, semiglossy, lower
surface drying dark brown to blackish; anterior lobe
8 cm long, 18.7-38 cm wide (2.1-3.1(3.6)
times longer than posterior lobes); posterior lobes
rounded, 7-17 cm long, 8-17 cm wide; sinus usu-
ally hippocrepiform, 12 cm deep; midrib flat,
slightly paler than surface above, convex, paler
than surface below; basal veins 4-7(8) per side,
with all free to base, third and higher order veins
coalesced 1.54 cm long; posterior rib naked for
1.5 cm; primary lateral veins 3-6 per side, depart-
ing midrib at a 35—40° angle, + straight to the
margins, sunken above, convex and paler than sur-
face below; minor veins moderately distinct below,
arising from both the midrib and primary lateral
veins. INFLORESCENCES 1 per axil; peduncle
2.5-7 cm long, 8-10 mm diam., whitish streaked;
spathe 10-17 ст long (2.4-3.4(4) times longer
tan peduncle), constricted slightly above the tube,
+ ellipsoid; spathe green, short-lineate outside,
reddish purple throughout inside; spathe tube 5 cm
long; spadix sessile; 10-13 cm long; pistillate por-
tion 4-4.5 cm long, 1.7-2.3 cm diam. at base, 2-
2.7 cm diam. midway, 1.7-1.9 cm diam. near apex;
staminate portion 8-8.5 cm long; fertile staminate
Portion 1.8-2 cm diam., sterile staminate portion
usually much broader than pistillate portion at an-
thesis, weakly tapered toward apex, scarcely or not
at all constricted above the sterile portion, 1.4-1.8
cm diam. midway, 11-12 mm diam. 1 cm from
арех, bluntly acute at apex; sterile staminate por-
tion not very obvious, ca. 1.3 cm long, abruptly
Contracted at base, as broad as or narrower than
the pistillate portion; pistils 6.2 mm long, 3.4 mm
iam.; ; ovary 6—7-locular, 5 mm long, 3 mm diam.,
with axile placentation; locules 5 mm long, 1 mm
diam.; ovules 13-20 per locule, somewhat trans-
parent, 1 mm long, much longer than funicle; fu-
nicle 0.2 mm long, adnate to lower part of partition,
style 1 mm long, 3 mm diam., similar to style type
D; style apex + rounded; style boss broad, pro-
nounced; stigma 1-1.1 mm diam.; the androecium
truncate, oblong, prismatic, margins irregularly 4—
5-sided, 1 mm long, 2–2.5 mm diam. at apex. IN-
FRUCTESCENCE with pistillate spadix 4.5-5 cm
long, 3—4 cm diam.; berries 1 cm long, 4 mm diam.;
seeds 6-8 per locule, oblong-ellipsoid, 2.2-2.5 mm
long, 0.7–0.9 mm diam., sticky.
The flowering phenology in Philodendron purul-
hense is poorly known. No flowering collections
have been made, and only one post-anthesis col-
lection is known (July), but immature fruiting col-
lections are known from March, July, September,
October, and November. This might suggest that the
species flowers throughout at least a substantial
part of the rainy season and perhaps also in the dry
season (December through April).
Philodendron purulhense ranges from southern
Mexico (Chiapas) to Guatemala (Alta Verapaz) and
Honduras (Cortés and Olancho), at 1360 to 1870
m elevation in Tropical Lower Montane wet forest
and Premontane wet forest life zones.
Philodendron purulhense is a member of P. sect.
Philodendron subsect. Philodendron ser. Fibrosa.
This species is characterized by its short inter-
nodes; sharply D-shaped to two-ribbed cataphylls
persisting semi-intact; subterete petioles (about as
long as the blades), dark brown to blackish-drying;
ovate-cordate blades with basal veins free or weak-
ly coalesced and scarcely or not at all naked on
the usually hippocrepiform sinus; solitary, short-pe-
dunculate inflorescence, more or less elliptic green
spathe; and ovaries with 13-20 ovules per locule.
Philodendron purulhense may be confused with
P. advena because leaves of the latter species dry
a similar color. Philodendron advena differs in hav-
ing long internodes, deciduous cataphylls, more or
less sagittate blades, and one to three (four) ovules
per locule.
pesci specimens examined. GUATEMALA. Alta
rapaz: Tactic, Río Frío, 1400-1500 m, Standley 90510
(P: Tactic-Tamahú, 1500-1600 m, Lf — (F);
Río Carchá, Cobán-San Pedro Carchá, ca. 1360 m, Stan-
89902 (F). Baja Ver: : Highway СА. 14 to Coban,
un 41189 урне Biotopo del Quetzal, WNW of Purulhá,
600 m, Stevens et al. 25446 (MO); Mpio. Purulhá,
Д Progreso-Cobán, 1620-1720 m, 15?13'N, 90^12'W,
oat & Hannon 63765 (CAS, CM, K, b MEXU, MO,
US, S USCC). HONDURAS. Cortés: acional Cus-
uco, Río de Cusuco, ca. 22 km W of San Pedro Sula, 1500
m, 15%30'N, 88^13'W, Evans 1490 (EAP, МО). Ocote-
peque: Cordillera de Celaque, 3 mi. N of Belén Gualcho,
$m
506
Annals of the
Missouri Botanical Garden
1870 m, 14?30'06"N, 88°48'02"W, Davidse et al. 35340
(MO). MEXICO. Chiapas: Mpio. Jitotol, 5 km SE of Ji-
55077 (MO); Mpio. La Trinitaria, 4 km guna, near
Dos Lagos, 1300 m, Breedlove 38810 CAS; Mpio. Las
Margaritas, 12 km E of Tziscao, 1200-1300 m, Davidse
et al. 29865 (MO).
Philodendron radiatum Schott, Oesterr. Bot.
lotype lost). Schott ic . 2623 (neotype, here
designated). Figures 1, 324, 331—335.
Epiphytic or hemiepiphytic, rarely terrestrial;
stem appressed-climbing, creeping if terrestrial, sap
clear to orange, watery, sticky, leaf scars conspicu-
ous, 2—4(7.5) cm long, 2.5-6(7) cm wide; internodes
thick, 3—7(12) cm long, (1)3-8 cm diam., about as
long as broad or longer than broad, dark green to
gray-green, sometimes scurfy and light brownish tan,
transverally lined (raised 2 mm); roots 4 mm diam.,
with swollen nodes and branched tips; cataphylls to
38 cm long, soft, bluntly to sharply 2-ribbed, rarely
unribbed, green, sometimes conspicuously reddish-
lineate, deciduous; petioles 108 cm long, (2-
3)4-17 mm diam., terete or subterete to obtusely
flattened adaxially, dark green, surface dark greenish
or to greenish red-lineate; xir persisting, some-
times moderately spongy; blades + triangular-ovate
in outline, (15)27-101 cm ig а 1)25-90 cm wide
(ca. 0.8–2.1 times longer than wide), (0.5—1.7 times
longer than petiole), broadest at point of petiole at-
tachment, thinly coriaceous, weakly to moderately
bicolorous, semiglossy, acuminate to long-acuminate
at apex, upper surface dark green, lower surface
slightly paler, usually drying yellow-brown to reddish
brown on both surfaces; margins weakly incised-lo-
bate to usually deeply incised-lobate to within 1-8
cm of midrib, rarely entire and sagittate; apex often
acute, sometimes acuminate, very short acuminate
or + rounded (the acumen tightly inrolled if present,
1-8 mm long), lobed-cordate at base; basal segments
pinnatifid, lateral segments entire, sinuate or the
lowermost of the anterior lobe pinnately lobed with
1-2 lobes on each side, final divisions linear-lan-
ceolate in shape (0.9-2.1 times longer than wide),
segment apex obtuse to broadly obtuse; interlobal
sinuses 0.7-0.97 the length of the lobes; basal sinus
hippocrepiform to obovate or closed, 3-15 cm deep;
midrib slightly raised to convex, weakly reddish
green-lineate, concolorous to paler than surface
above, convex to prominently raised to round-raised
below; basal veins 4—5(8) per side, directed into the
segments of the posterior lobe, 0-1 free to base,
(124 coalesced to З cm long, naked 1-6 cm; pos-
terior rib absent; primary lateral veins 3-8 per side,
departing midrib at a 50-60” angle, straight to mar-
gin, weakly raised above, raised to convex and paler
than surface below; minor veins moderately visible,
paler and slightly raised below, arising from both the
midrib and primary lateral veins. INFLORES-
CENCES erect, 1-3(4) per axil; peduncle 2-12 cm
pne (3)5-10(19) mm diam.; spathe erect, 11-25
cm long (1.3—6.3 times longer than peduncle), ob-
scurely striate, margins paler; spathe blade cuspi-
date at apex, pale green to yellowish green,
glossy, sparsely purplish-dotted outside, pink to dull
red or pale brownish (post-anthesis) inside; spathe
tube dark purple-violet or dark green, sometimes
tinged purple-violet outside, 4 cm diam., dark рш-
ple-violet inside; spadix sessile to weakly stipitate,
cylindrical to weakly tapered; staminate portion
creamy white, protruding forward at anthesis, point-
ed at apex, 10-17 cm long; pistillate portion whitish,
weakly obovoid, 3.5—5.2 cm long (1 cm shorter on
back side), 1.1-1.8 mm diam. at apex, 1.5-1.9 mm
diam. at middle, (7)17 mm wide at base; staminate
portion 6.3-8.1 cm long; fertile staminate portion ta-
pered, 1-1.9 cm diam. at base, 1.2-1.6 cm diam. at
middle, 0.9-1.4 cm diam. ca. 1 cm from apex,
broadest at the base, slightly constricted above the
base; sterile staminate portion as or slightly be^
than the pistillate portion, white with tan ring aro
apex, 0.7-2 cm diam.; pistils (1.6)4—5(8) mm v
(1)2.8-3.2(5.2) mm diam.; ovary (7)8-locular,
(1.6)3.3-4(6.7) mm long, 1.5-3(5.2) mm diam., with
axile or sub-basal placentation; locules (0.6)1.9-
3.4(6.5) mm long, 0.5-0.8(1) mm diam.; оуше sac
1.8(3.5) mm long; ovules 8 per locule, 1 1–2-вепаје,
contained within translucent, gelatinous ovule sac,
0.1-0.3 mm long, longer than funicle; funicle UE
0.2 mm long, style 0.3-0.5 mm long, 1.2-3 mm
"em similar to style type B (rarely C); style ape*
flat to weakly rounded, sometimes domed; stigma
usually hemispheroid, sometimes subdiscoid, pink,
2-1.5 mm diam., 0.6-0.7 mm high, covering entire
e apex; the androecium truncate, oblong, pris-
matic, margins irregularly 3—5-sided, sometimes
weakly scalloped, 1 mm long, (0.7)2.5-2.7 mm er
at apex; thecae cylindrical, 0.3 mm wide, + par
to one another and contiguous; sterile staminate
flowers usually truncate, sometimes clavate or irreg-
ularly 5-6-sided, 1.1-2.9 mm long, 0.6-1 B mm
wide. INFRUCTESCENCE 11-17 cm long, 4.5 cm
diam.; pistillate spadix 6-8 cm long, 2. 53.5 ст
diam.; berries white, somewhat translucent, 4 per
locule, oblong, sticky.
semi-
Philodendron radiatum ranges from Mexico por
Luis Potosí to Chiapas and on both Atlantic т
Pacific slopes in Chiapas) to Colombia (Antioqu!
from sea level to usually no more than 700 m
a e an os on A Jl i RR i a SSS aS rd E uc em EAD SEE qug ES 1. -“"– A .____- ____- Ӯ
——————— A И И EE EE
Volume 84, Number 3
1997
Croat 507
Philodendron Subgenus Philodendron
ly 1250 to 1860 m; most collections are from below
100 m) elevation. It is perhaps more widespread in
South America than collections indicate (only two
are known). The plants are very large and difficult
to collect and may have been overlooked in other
areas.
This species is highly variable morphologically
and ecologically versatile as well. In Mexico, it oc-
curs in “Selva Alta Perennifolia,” “Selva Mediana,”
and “Selva Baja Caducifolia” in mesic areas on the
Atlantic slope as well as in “Selva Mediana Sub-
perennifolia” and “Selva Baja Subperennifolia” in
the drier Yucatán Peninsula. On the Pacific slope
it also occurs in areas of “Bosque Pino-Encino.” In
Central America, this taxon occurs principally in
Tropical moist forest but also in Premontane wet for-
est and drier parts of Tropical wet forest.
Philodendron radiatum is a member of P. sect.
Polytomium. Philodendron radiatum has two vari-
eties, the typical variety having more deeply divid-
ed leaf blades with the segments often incised al-
most to the midrib and variety pseudoradiatum with
the blades only weakly incised-lobate (less than
half the distance to the midrib). While the latter
variety is restricted to southwestern Chiapas, even
populations of the typical variety on the western
slope of Central America have less deeply incised-
lobate blades than those on the Atlantic slope.
There is also considerable clinal variation in the
width of the divisions throughout Central America,
with plants in Mexico having, on average, broader
pinnae (averaging 4.5 cm wide) than those in Pan-
ama, for example, where the pinnae average only
about 3 cm wide. Leaves of Mexican populations of
P. radiatum also consistently lack free basal veins,
whereas elsewhere in Central America such veins
are present. In western Mexico and Guatemala,
ants are also more likely to have longer inter-
nodes and to occur more frequently on rocks or
creeping over the ground than plants on the more
mesic Atlantic slope of Central America.
KEY TO THE VARIETIES OF P. RADIATUM
la. Blades deeply lobed, the divisions near the mid-
id of the blade extending more than E i way
о the HEC cv e се r. radiatum
Ib. eds ткы lobed, the divisions ел ће
middle of the blade extending less than e
way to the midrib __ we peudoraiatom em
Philodendron radiatum Schott var. radiatum
Philodendron augustinum K. Koch,
nd. sp. Hort. berol.
in A. Braun et al.,
ppe 853: 1854.
TYPE: Central America. Without кан locality (ћо-
i B? lost). Schott ic. 2605
~
neotype, here des-
Philodendron polytomum Schott, Bonplandia 7: 164
9. TYPE: Mexico. Veracruz: Colipa, Hac. de Sta.
Bärbara, Liebmann s.n. сира C: әуре; К по!
Philodendron impolitum Schott, Prodr. Syst. Aroid. 291.
860. TYPE: Costa Rica, Wendland s.n. (not seen).
Internodes 3-7 cm long, 3-8 cm diam.; cata-
phylls bluntly to sharply 2-ribbed; petioles 33-108
cm long; blades deeply lobed, the divisions near
the middle of the blade extending more than % the
way to the midrib, 21-101 cm long, 25-90 cm
wide; posterior lobes (8)12.5-25 cm long, (9)16-18
cm wide; primary lateral veins 3-8 cm long; basal
veins 4(5) per side, (1)2—4 coalesced to 3 cm long,
naked 1-6 cm. INFLORESCENCES 1-3(4) per
axil; spathe 11-25 cm long; spadix 10-13 cm
long; pistillate portion 3.5-5.2 cm long; pistils
(3)4-5(8) mm long; ovaries with axile placentation;
style similar to style type B (rarely C).
Flowering in Philodendron radiatum var. radia-
tum is documented by few flowering collections
made in February, August, and September. Post-
anthesis collections are common, from every month
of the year, and there are no clearly marked differ-
ences in the phenology of this species in different
parts of Central America. Material was seen at var-
ious stages of development from all parts of Central
America in about equal numbers. Mature fruits are
known only from February and May.
Philodendron radiatum var. radiatum is charac-
terized by its moderately thick, mostly short inter-
nodes; sharply two-ribbed, deciduous cataphylls;
terete petioles; and especially by its deeply incised-
lobate leaf blades that mostly linear to linear-lan-
ceolate segments, themselves often lobate toward
the base of the blades
Philodendron radiatum var. radiatum might be
confused with P. dressleri and P. warszewiczii, Cen-
tral American species with incised-lobate leaf
blades. Philodendron dressleri differs in having
thicker leaf blades, more succulent stems with
shorter internodes, and especially by having blades
that are incised lobate only about midway to the
midrib. In addition, P. dressleri has one inflores-
сепсе per axil and 3—4 locules per ovary, while P.
radiatum has 1—3(4) inflorescences per axil and
7-8 locules per ovary. Philodendron warszewiczii
differs by having thinner blades, drying papyra-
ceous with the minor veins distinctly visible. In ad-
dition, the lateral leaf blade segments usually have
three or more lobes per side. Philodendron radia-
tum, in contrast, has blades drying subcoriaceous
with the minor veins rather indistinct and with the
508
Annals of the
Missouri Botanical Garden
lateral leaf blade segments usually 1-2-lobed per
Philodendron radiatum var. radiatum has a
unique attractant, whose odor profile is made up
almost entirely of compounds unique to it, and is
pollinated by a new species of Cyclocephala
(Schatz, 1990).
Additional specimens examined for P. var. radia-
tum. BELIZE. Honey Camp, Lundell s.n. (US); ao
Gentle 1141 (MICH, NY); Gracie Rock, 154 f
avidse £ Brant 33146 (MO). Cayo: Roaring
Creek, ts 12677 (МО); 1.5 € W of Augustine, Río
Frío, ca. 450 m, Sutton = al. 211 (BM); Hummingbird
Highway, Mile 20, Dwyer & noe 12130 (MO); mig а
bul National Park, Caracol Archaeological Reserv
m, 16745'N, 8907", Ingram & pasting clle 1935
(MO); Río Ma Cal, San Luis—Cuevas, 1050 ft., Croat
23534 (MO); Vaca, Gentle 2552 (MICH). Равзат edge
ghw
tion to San Antonio, ca. 100 ft., e Losa phe So.
lomon Camp, vic. of Richardson Creek and Bladen Branch
, 80-420 m, 16°32-
avidse & ios 32198 (MO); Colum-
bia Forest Station "ein 1.5 mi. S of Mayan Village of San
José, Croa 24335 (МО); Holst 4303 (МО). Жы аң
Alajuela: Cafias—Upala, 10 m N of Bija
Croat 36488 (MO). Heredia: La Selva Field GP Che
yum 2534A (DUKE); 100 m, e (DUKE); Jacobs 2290
(DUKE); 2707 (MO); 10°26
Ortiz 78615 (MO); Río бшсш. near Puerto Viejo, 20 m,
Croat 35680 (MO); Puerto Viejo-Guápiles, 7 km N of
Buenos Aires, 10723'30"N, 83%48'30"W, Croat 68386
(MO); Río Sarapiquí, S of La Virgen, A m, Lent 33 (MO);
La Tirimbina, 220 m, end 821 (WIS). Limón: Barra
del Gaani N side, 0— 10%47'N, 83°35'W, Stevens
24133 (CR, MO); Barra e. Сафина Ни San Juan, La-
na de Atrás, 5 m, 10?48-52'N, 83°3 38'W, Davidse &
etg 31512 (CR, MO); Cerro Coronel, E of Laguna
nto, ди m, 10%41'N, 83°38’ W, Stevens > oe
MO): P que Nacional Tortuguero, Estación Agua Fría,
m, Robles. 13 ha (CR, MO); 40 m, 10°26’N, 83?35' W, 1730
(CR, МО); 600 m S, 4 m, 10?32'N, d W, 1846 (CR,
MO); Cibus: Poveda 1166 (CR ; Cahuita-Limón,
0-10 m, 9°44’N, 83?20'W, Baker & Burger 157 (CR, NY);
aiite rente La Suerte, 29 air km W of
rtuguero, 40 m, 10%30'N, 83°47’ WD avidson & Dona-
i 8729 Mo, ea 8437 (RSA); Río io Colorado, 3.5 air-
120 m, 10*
line km S of Islas Buena Vista, 39'N,
83°40’ idse & Herrera bio ( 0); Barra del
Colorado, 1-5 m, 10?47' E S avidse
Herrera 30882 (MO); Río Pacuare, 50-100 , 10?15'N,
W, Thompson & Rawlins
1175 (CM); Río тома Етса манка below
Cairo, 25 m, Standley & Valerio 48969 (US). EL SAL-
VADOR. ај its Calderón 1245 (US). San Salva-
dor: Tonacatepeque vicinity, Standley 19538 (GH, NY,
US); Calderón 201 (GH, NY, US). GUATEMALA, Alta
H); near Finca
e
Verapaz an u
Sepacuite, Cook & Griggs 7. 4 (US) ym 350 m,
Tuerckheim 8330 (US); бесы Estor, 5 ті. W of
Tucurú, 600 m, Croat 41509 (MO); Tucuri-El Estor, Fin-
ca Argentina, above Papalha, 15 mi. Telemán,
50-650 m, Croat 41550 (МО); ca. 6 km NE of Panzós,
500 m, Croat 41616 (MO). Izabal: Quiriguá vicinity, 75-
225 m, Standley 23939 (US); ca. 7 mi. S of Puerto Barrios,
50 m, Croat 41803 (MO); Río Frío, Cerro San Gil, 75-
150 m, Steyermark 41534 (MO, NY); W of El Estor and
abandoned nickel mine, 1-10 m, Stevens & Martínez
сл
); 15
12702 нЕ: Tikal National Park, ои ;
Quezaltenango: along СА-2, 4 ті. N rnoff to Co-
loba, Croat 32767 (MO): Coatepocue RARE Hwy.
CA-2, 3 mi. 5 of turnoff to Colomba, 600 m, Croat &
Hannon 63414 (MO, US). Sacatepéquez: Volcán Santa
Clara, 1250-2650 m, Steyermark 46621 (MO). HONDU-
RAS. Puerte Sierra, Wilson 321 (NY). Atlántida: Lance-
tilla Valley, near Tela, Standley 52945 (US); 53668 (US);
53985 (US);
from La Ceiba, Blackmore £ Chorley
UNAH); Río чеш Tocoa—La Cieba, 11.3 mi. from Río
Cangreja Bridge a 47
Croat & Hanno
joa, Pun
ا et al. 3079 (MO); Ocote —
Lago de Yojoa, Nelson et al. 5868 (MO , VBD); y Puerto
Cortes-Guatemalan border, 2-3 mi. SW ч Отоа, Croat
42558 bei Jo of Lago de Yojoa, along old fabis l,
са. 2 . from junction with new Highway 1, SW о
Santa hae de Yojoa, 600 m, Croat 42745 (MO). Gracias
a Dios: Río Plátano Biosphere Reserve, Las Marías (Ba atil
Gualaco-San нећак Río Olancho, 7.4 mi. NE of
Esteban, 540 m, 1520'N, 85%42'W, Croat 64365 (K, MO.
NY). Santa Bárbara: Lago Yojoa, Punta Gorda, E
14°52'N, 88°W, езг sapiat et al. 3126 (MEXU, M
NY); 630 m, 14^53'N, 88°W, Liesner 26769 (MO); 700 bee
ibas па (MO); El Novillo, on m, 14
Yoro: Aguán River ips
MEXICO. Location ыа Reko 3690 (US); CH
al 14 (US). Campeche: Mpio. Hopechén, Rancho El p
men, 33-35 km S of Xmaben е road from Hopelchén
pujil, near Xpanzil, 200 m, 20'W, Sande
et al. 9735 МОУ. Huatusco, peas 14 (05); Раша
Engler 197 (BM, GH); Santa Leo 5
Barl (BH). Chiapas: San Manuel, ene km
enque, on road to Ocosingo, ca. 500 m, Ма
ED: Excma Triunfo, 1 mi. N of са 100 de
Croat 43811 (MEXU, MO); Esperanza-Escuintla, vices
16662 (F, MEXU); 17788 (NY: 150 m, 18045 (MEXU):
EE
о eee ç or س
Volume 84, Number 3
1997
Croat 509
Philodendron Subgenus Philodendron
Tapachula—Nueva Alemán, 4 пи. N of Tapachula, 250 m,
Croat 43794 (MO); Escuintla—El Triunfo, ca. 8.5 mi. NE
of Escuintla, 250 m, Croat 43820 (MO); Escuintla—Monte
Ovando, 2.8 km NW of Mid ca. 100 m, Croat 47511
(MO); Acacoyagua, Cerro Ovando, 800—900 m, Croat
78548 (CHIP, MO Hu iria en de Mendoza, 48
m N of Huixtla, ca. 200 m, Croat 47523 (MO); 2 mi. S
oft Chiapas edis joe Hwy. 195, 8 mi. N of cet dg
80 m, Croat 40088 (MO); 5 mi. SE of Palenque, on road
pak, 89-90 mi. SW of Palenque, 350-370 m, Croat 40217
(MO); 73 mi. SE of Palenque, 460 m, Croat 40267 (MO);
Ocozocoautla—A pitpac, 2 mi. N
Croat 40650 (MO); 5 mi. N of Ocozocoau
Croat 40543 (MO); ا de Mendo өрүт pe 15
mi. S of Motozintla de Mendoza, 900 m, Croat 40765
(MO); Mpio. Ixtacomitan, 7 km SW of Ixtacomitan, 250
x
onusco, new unfin
m, 15°31'N, 9
Mpio. Ocosingo, Lacanja—Chanzayab, Palenque—Boca La-
cantum, 340 m, Martinez 15071 (MO); Laguna agrum
12 km N of eh Líbano, trail to Chancala, 980 m
tínez 17029 (MO); Bonampak, 520 m, Breedlove & Almeda
58051 (MO); Mpio. Palenque, 8-9 km 5 of Palenque, 300
m, Breedlove & Strother 46895 (CAS); 25 km S of Pal-
Pl
g 8904 (BH, мо); 8910 (ВН); Тих-
tepec—Oaxaca, 0. Te mi. oS of Valle Nacional, Highway 175,
120 m, Croat 39700 (MO); 3 km 5 of Hidroeléctrica Те-
Carranza & Zamudio 8 (MO); ca. 1.5 km E of La
Boquilla, 270-320 m, d 5 Diaz 4721 (MO). San
Luis Potosi: Highway 85, 6 mi. NW of Tamazunchale,
250 m, Croat 39262 (MO); cial Edwards 626
abasco: 30 mi. E of ie atitlán, Barkley
Gs 36221 (НУ a 10 mi. NW of Cardenas, Bar-
low 26/25 (BH و ا ме. Сагдепаз, “Cowan
2021 (МО). Mpio. on aas f km 1 ón
de Huimanguillo, Cowan & Magaña 3255 (CAS, NY). Ve-
racruz: Huatusco—Puente Nacional, El Mirador, 21 km E
Huatusco, ca. 1200 m
Bunting 8855 (BH); Playa Escondida, N of Sontecomapan,
along Caribbean cliffs, 10-60 m, Gentry et al. 32610
(MO): Córdoba—Veracruz, San José de S aie са. 750 m,
Croat 39612 (MO); Highway 180, 6 mi. E of Coatzacoal-
cos, Croat 40064 (M Mi 5 ; ;
to Martínez de | . 320 m, Moore & Bunting 8946
p Dtto. Papantla, Kelly 126 (BH); Mpio. Coatepec, 5
m by air SE of Tuzamapan, 680 m, 19°21'N, 96750" W,
Nee & Taylor 26026 (NY): Mpio. Cosamaloapan, 10 km
by air NW of Tres Valles, 25 m, 18°17'№, 96°13’ W, Nee
& Taylor 29306 (NY); Мріо. Hidalgotitlán, 1 bs SE of
gustín Melgar, 100 m, 17^15'N, 94° e 29765
(NY); Mpio. Naolinco, 2 km from Concha al det 900
m, Ortega 630 pt Yucatán: Izamal, 20°56'N,
89*01"W, Gaumer 23200 (GH, NY, US); 1413 (GH, MO);
es Schott 766 ctr NICARAGUA. Boaco: Воасо–
slope of Cerro Mombachito, 500—900 m, c
12:24 05; N, 85°32- 337, Stevens & Grijalva 14699
(MO); Hwy. 33, vic. Río Quilan bridge, ca. 300-310 m
12*35'N, 85°32’ W, Siih 9330 (BM, MO). Chuintales:
E Libertad, ca. 17.4 km NE ix cane > at
ford of Río El Bizcocho, 2'N,
85^17'W, Stevens 4093 (BM, МО); 4095 (BM, Mo; ca.
2.8 km above Сиара, 400—500 m, ca. 12°17'N, 85°23'W,
Stevens 3671 (MO, PMA). Estelí: km 167 on Hwy. 1, ca.
15.8 km N of entrance to Estelí, 825—850 m, ca. 13?15'N,
86°22'W, iig 5788 (MO). Jinotega: Río Bocay, Salto
Acatulu, ca. 130 m, 14?13'N, 85?10'W, Stevens et al.
16749 (MO). Madriz: Cerro Quisuca, 1100-1250 m, ca.
13°30'N, 86°31'W, Stevens & Grijalva 16063 (MBM, MO).
anagua: Ciudad Managua, Reparto Bolonia, Guzmán et
al. 1071 (MO). Nueva Segovia: Río San Fernando Valley,
ca. 5.2 km N of San Fernando, 13%2-3'N, 86°19-20'W,
Río San Juan: ag Santa Cruz—
Cafio Santa Crucita, La Palma, 11?2-4'N,
84?24—26'W, Stevens 23427 (MO); Caño pm ear 20
km NE of El Castillo, 200 m, Neill £ Vincelli 3620 (MO).
Zelaya: La Barra de Punta Gorda, 0-2 m, 11%30'N,
83746", Moreno 13225 (MO); Caño е тошћ
of Caño El Consuelo, ca. 10 m, 11?35'N vla oreno
15027 (MO); 6.5 km al SE de Ым, 5 0-560 m,
13°16'N, 85°24’ W, Moreno 17288 (МО); == Cabezas—
Rosita, ca. km 47, ca. 5.3 km W of Rfo Wawa Ferry, <10
m, 14%06'N, 8335'W, Stevens 8568 (MO); along road to
—5 km from main
m,
(м0) Puerto apo Wawa, Ibo Tingni, drainage of
ño Sung Sung, <10 m, 14°9-11'N, 83°29-31'W, Ste-
vens 10658-a (MO); Auastara vicinity, <10 т '
14^19'N, 83°12-13’W, Stevens 10441 (МО); SW of Blue-
Pipoly 4370 (MO); ca. 100-200 m,
49'W, 4239 (MO); 4162 (MO); са. 100-268 m,
13°39'N, 84?48-49' W, Stevens 7378 (MO); Caño Costa Ri-
quita, ca. 1.8 km SW of Colonia Naciones Unidas, 1
180 m, ca. 11%43'N, 84?18'W, Stevens 5081 (MO); Cerro
Saslaya-San José del Hormiguero, aeger Саћо Sucio
and Loma Mollejones, ca. 3 , 13%45'N, 84°58-
59'W, Stevens 7011 (MO); Cayo eid 0-10 m
11°34'N, 83°39'W, Stevens 20765 (MO); San Juan del
pie pum 22 (MO); Seymour 5906 (MO); Cerro Baká,
E of Río Coperna, 200-320 m, 13°40'N,
84:30" W, ctas 4846 (MO); Estación Experimental El
Recreo, W de Cerro La Ceiba, 12°10'N, 84^18'W, San
dino 1656 Pom Monkey TOM. с. El Pato, ca. 10 т,
11%35'N, 83742", Moreno 12399 (MEXU, MO); 1-5 m,
11°36'N, dede W, ade (МО); 1-5 m, 11?36'N,
83:38'W, Moreno & Sandino 12007 (MO) 0-20 m,
11%35'N, 83*39' W, Stevens 20031 (MO); 1 km S of Mon-
key Point, ca. 5 m, 8339' W, 11%35'N, Moreno & Sandino
12541 (MO); Río Likas, near Silima Lila, 50 m, ca.
14*30'N, 8350'W, Pipoly 4097 (MO); Río dip ане
0.3-1.9 km N of pee 8–
84713", Stevens 8264 (M
100 m, са. 13°42'N,
a Gorda, Atlanta, 10 m, 11°33’N, 84
& Sandino 12808 (MO); Atlanta, 2 km 5 of Carolina del
11%32'N, 84°O1'W, Moreno & Sandino
: а, са.
IPN, 84°16! W, Stevens 8849 (MO); Río Sucio, E "
Bonanza, ca. 140 m, 14°O1'N, 84%34'W, Stevens 12345
(MO); Río Waspuk, 15-100 m, 14^15'N, 84°36' W, Stevens
510
Annals of the
Missouri Botanical Garden
13091 eda PANAMA. Ahorca Lagarto, Cowell 262
). Bocas del Toro: Gualaca-Chiriquí Grande, 4.2 mi.
82%09'W, Croat 66816
(AAU, CM, F, MO); Changuinola-Almirante Railroad,
Milla 7.5, 100 m, Croat & Porter 16433 (MO); Croat
38122 (MO); Chiriquí Lagoon, Water Valley, von Wedel
989 (GH, MO); Isla Colón, Swan Key, 2 km N, Tyson &
Loftin 6307 (FSU, MO); Rio Cricamola, Finca St. Louis—
— Woodson et al. 1901 (GH, MO, NY); Río aed
guinola, ca. 1 km above mouth of Río Teribe, «100
9°21’ AO'N. 82°31'40"W, Croat & Zhu 76446 (CM, MO)
anal Area: Hayes 805 (NY); Gorgona—Gatún, 10-15 m
Pittier 2300 (US); Gatún, Standley 27023 (US); Ft. Sha.
mag Standley 30992 (US); Frijoles, Standley 27467 (US);
t. Randolph, Standley 28626 (US); 66-70 m, Dodge et
т 16870 (MO, U, UC); near Gatún, ae 27203 (US);
vic. Summit Garden, Tyson & Lazor 6121 (FSU, RSA);
Barro Colorado Island, Gattin Lake, Standley 31328 ји
кеге (05); Aa 178a an Elmore X14 (MIC
, US); Croat 7178 (MO); 79%49'46"W, besa
a (MO, NY, SCZ); 6060 al. 4655 (МО, SCZ); Aviles
44 (MO); Shaitück 266 (GH, MO); Navy Pipeline Road,
Smith & Smith 3313 (US); 0.25 km NW of Río Macho
bridge, 10 km NW of Gamboa, 125 m, Nee 7863 (MO,
RSA, TEX). Chiriquí: Gualaca-Chiriquí Grande, 13.6 mi.
N of Continental Divide, 120 m, 8°57'N, 80%56'W, Croat
74934 (MO). Coclé: 27 km N of Penonomé, on road to
Coclesito, at Continental Divide, 1500 ft., Hammel 1648
O); Llano Grande-Coclesito, 12 mi. from Llano
m. 8° 47'N, 8028", Churchill et al. 3987
(RSA). сом ca. 8 km E of Piña, 50-100 m, 9%17'N,
807, Thompson 4815 (CM, MO); Río Indio-Miguel de la
Borda, 0 m, Croat 36927 (MO); near Nuevo Tonosí, 2 mi.
m Portobelo, on road to Nombre de Dios, «100 m
Croat 33527 veris Santa Rita Ridge, Transisthmian High-
way, ca. 10 mi. from di Porter et al. 4738 (MO); ca. 7
mi. from Tr ићи way, са. 650 ћ., Wilbur et al.
15083 (MO); Santa Rita Tra T: des (NY). Panamá:
ca. 20 km from Inter-Am N of Cerro Azul,
Mori et al. 3786 (MO); Río pets Valley, El Llano—Cartí,
8 km from Pan-American Highw.
, 0-50 m, 9%20'N, 78°13’ W, Herrera ан
МО); Мене = El Llano-Cartí Road, 9 mi. N of m
highway, 350 m, 9°20'N, 79°W, Croat & Zhu 76566 (MO,
SEL). V eraguas: Santa Fe-Río Calovébora, 0.6 mi. be-
yond agricultural school near Santa Fe, 735 m, Croat
32982 (MO).
~
Philodendron radiatum var. pseudoradiatum
(Matuda) Croat, stat. et comb. nov. Basionym
Philodendron pseudoradiatum pice Revista
Soc. Mex. Hist. Nat. 1: 96, fig. 8. 1950. TYPE
Mexico. Chiapas: Mun. Escuintla: Turquía,
Salto de Agua, 8 Sep. 1947, Matuda 17787
(holotype, si нин, DS, NY, UC). Fig-
ures 8, 327-3
Internodes 2-12 cm long, 1-2.5 cm diam.; cata-
phylls unribbed; petioles 28-78 cm long; hades
shallowly lobed, the divisions near the middle of the
blade extending rni rm Y the way to midrib,
11-60 cm lo m wide; posterior lobes
(8)12.5-25 cm Aie = 2-18 ст wide; basal
veins 5(7-8) ed side, with 0—1 free to base, poste-
rib abse ary lateral veins 3—5 per side.
INFLORESCENCES T per axil; spathe 12-18 cm
long; spadix 12.8-17 cm long; pistillate portion 4-
4.5 cm long; pistils 1.6-2.2 mm long; ovary with
sub-basal placentation; style similar to style type B.
The flowering phenology of Philodendron radia-
tum var. pseudoradiatum is still poorly known.
Flowering collections are known only from January
and September, but the flowering pattern is prob-
ably just poorly documented rather than bimodal.
Philodendron radiatum var. pseudoradiatum is
endemic to Mexico, where it is known only from
southwestern Chiapas, at 200 to 350 m elevation
in “Selva Alta Perennifolia.”
Philodendron radiatum var. pseudoradiatum is
distinguished by its incised-lobate blades with the
lobes divided less than halfway to the midrib. The
typical variety generally has the blades incised
more than three-fourths of the way to the midrib.
This taxon might be confused with P. dressleri,
which also has leaf blades divided only about half-
way to the midrib. That species differs in having
much thicker stems (typically to 6 cm diam),
sharply 2-ribbed cataphylls, and by occurring in
much drier areas where it loses its leaves during
the dry periods. In contrast, P. radiatum var. pseu-
doradiatum has rather slender stems, unribbed cat-
aphylls and occurs in mesic areas
Until recently, Philodendron idani var. pseu-
doradiatum was represented by only two collec-
tions, Matuda 17787 and 18718, from the same
locality. Several collections are noteworthy. Matuda
18045 consists of two sheets, one of which has a
leaf lobed in much the same manner as the type,
while the other has a leaf lobed to more than two-
thirds of the way to the midrib. This may be an
indication that the type of P. radiatum var. pseu-
doradiatum is merely an aberration in the degree
of lobing for P. radiatum
Another noteworthy polation is Croat & Han-
non 63381 from Sierra de Soconusco, northwest of
Mapastepec. It is perhaps also P. pseudoradiatum
but has smaller and narrower leaves. It looks much
like a hybrid between P. radiatum var. pseudora-
diatum and some other species, but if so, it is not
clear which other species might be involved in the
putative hybrid plant.
Additional specimens examined. МЕХІ ICO. NEN
Escuintla, Salto de Agua, 350 m, Matuda 1 8718 (
ished road to Tuxtla Gutiérrez, 200 m, 1531 М, dede х
Croat & Hannon 63381 (CAS, СМ, Е, MEXU, MO,
EOE > 7 7 RR س «NN REUS — — — —— > —? nca digi. ENS —
Volume 84, Number 3
1997
Croat 511
Philodendron Subgenus Philodendron
Philodendron vits bees sp. nov.
: Panama. Panam o Jefe, al
road just below the summit, oa 15'N, 79°30" w
17 June 1976, Croat & Zhu 76211 (holotype,
MO—4619415; isotypes, AAU, B, CAS, CM,
COL, CR, DUKE, F, GH, HUA, K, M, MEXU,
NY, P PMA, RSA, 5, SEL, TEX, US, VEN).
Figures 341-345.
Planta ipee. raro hemiepiphytica; serene 1-4
ст longa, 1.5-2.7 cm diam.; cataphylla 12-27 m longa,
acute 2-costata, beni semi-intacta nodis UM.
bus; petiolus U-formatus, obtuse sulcatus, 10-23 cm lon-
gus, 3-6 mm diam., crassior quam latus; lamina (22)30—
49 em longa, 9. 8-21 ст lata, elliptica vel oblongo-ellip-
р ongus; spatha
6.5-10 ст linens lamina spathae extus saepe rubella pler-
umque atriviridi, intus pallide viridi vel marronina; tu
spathae extus pallido-lineato, intus pallide viridi vel mar-
ronino; pistilla (4)5—7(8)-locularia; loculi 14—18-ovulati;
c e.
Terrestrial or rarely hemiepiphytic; stem ap-
pressed-climbing or creeping, leaf scars incon-
spicuous, obscured by cataphyll fibers; inter-
nodes short, semiglossy, 1-4 ст long, 1.5-2.7 c
diam., longer than broad, dark green to gray-
green to reddish, obscured by cataphyll fibers;
roots many, thin, wiry, descending, greenish
brown; cataphylls 12-27 cm long, sharply
2-ribbed (unribbed to weakly to sharply 2-ribbed
in Colombia), pale red to purple-violet, drying
dark brown, sharply D-shaped, persisting semi-
intact at upper nodes, as fibers below; petioles
10-27 cm long, 3-6 mm diam., U-shaped, thick-
er than broad, firm, dark green to reddish, tinged
purple, somewhat flattened to obtusely or narrow-
ly sulcate adaxially; geniculum pale green to red-
dish violet, 1 cm long, 1.5 cm diam.; blades е!-
liptic to oblong-elliptic, sometimes oblanceolate,
rarely elliptic, coriaceous to subcoriaceous, con-
spicuously bicolorous, sometimes acuminate at
apex (the acumen inrolled, 1-3 mm long), acute
to narrowly rounded, sometimes attenuate at
base, (22)30—49 cm long, 9.5-21 cm wide (1.8–
3.5(4.5) times longer than broad), (1.6–3.2 times
longer than petiole), about twice as long as wide,
upper surface dark green, matte to semiglossy,
lower surface moderately paler, semiglossy to
glossy; midrib narrowly sunken, paler than sur-
face above, narrowly convex, thicker than broad,
matte, paler than surface and sometimes reddish
green below; basal veins lacking or sometimes 1—
2 per side, free to base; primary lateral veins 8—
14(16) per side, departing midrib at a 40—55°
angle, sunken to narrowly sunken, concolorous to
paler than surface above, convex and darker than
surface below; interprimary veins few, sunken to
narrowly sunken, concolorous to paler than sur-
face above, raised and darker than surface below;
minor veins few, moderately obscure above,
darker than surface below, arising from midrib
only, sometimes prominulous and weakly undu-
late on drying. INFLORESCENCES 1-2 per axil;
peduncle 2.5-6 cm long, 2.5-5 mm diam., red-
dish, prominently green streaked; spathe 6.5-10
cm long (1.3-3.3(4.6) times longer than pedun-
cle), red to maroon or violet-purple or sometimes
green heavily tinged red, sometimes greenish
white throughout; spathe blade frequently dark
green (red in Colombia) outside, pale green to
maroon inside; spathe tube short pale-lineate
outside, 3—4 cm long, 2 cm diam., pale green to
maroon inside; spadix = tapered, 6.8–8 cm long,
broadest at the base; pistillate portion white to
pale green, cylindrical, 2-3.7 mm long, 1-1.5
mm diam. at apex, 11–7.2 mm diam. at middle,
9–11(17) mm wide at base; staminate portion
3.9—4.9(6) cm long; fertile staminate portion
cream, cylindrical to somewhat tapered, 10 mm
diam. at base, 8 mm diam. at middle, 8 mm diam.
са. 1 ст from apex, broadest usually at base,
narrower than the pistillate or sterile portion;
sterile staminate portion as broad as the pistillate
portion, 10 mm diam.; pistils 2.5 mm long, 1.6—
1.9 mm diam., tinged ren ovary (4)5-7(8)-
locular, 1.4 mm long, 1.6 mm diam., with axile
placentation; locules 1.4 mm Whe 0.5 mm diam.;
ovules 14—18 per locule, contained within a ge-
latinous matrix, 2-seriate, 0.3-0.5 mm long,
longer than funicle; funicle 0.2-0.4 mm long, ad-
nate to lower part of partition, style 0.4-0.6 mm
long, 1.6-1.8 mm diam., similar to style type B;
style apex flat to weakly rounded; stigma usually
subdiscoid, 1.1-1.3 mm diam., 0.2-0.5 mm high,
covering entire style apex; the androecium trun-
cate, prismatic, oblong, margins irregularly 4–6-
sided, 0.7 mm long, 1.2-1.5 mm diam. at apex;
thecae oblong to cylindrical, 0.5 mm wide, +
parallel to one another and contiguous; sterile
staminate flowers clavate, blunt, irregularly 4—6-
sided, 2.3 mm long, 2 mm wide. INFRUCTES-
CENCE with pistillate spadix 44.5 cm long,
1.5-2.5 cm diam.; berries white (immature), 3.5
mm long, 2.6 mm diam., 12-14 per locule; seeds
7-9 per locule, whitish, oblong, 1.8 mm long,
sharply tapered on one end.
Philodendron roseospathum ranges from Panama
to Colombia. In Panama, this taxon is restricted to
both sides of the Panama Canal, ranging from Ver-
512
Annals of the
Missouri Botanical Garden
aguas to San Blas, mostly in Premontane rain forest
or less frequently in Tropical wet forest life zones,
at 350 to 1000 m elevation. In Colombia, it has
been collected only in tropical pluvial forest at 100
Philodendron roseospathum is a member of P.
sect. Philodendron subsect. Canniphyllum. The
typical variety is recognized by its generally erect,
frequently terrestrial habit, short internodes, an
rosulate cluster of leaves with moderately short,
bluntly sulcate, thicker-than-broad petioles. The
dark green, elliptic-oblanceolate leaf blades are
narrowly rounded to acute at the base. It is es-
pecially recognizable by its masses of weathered
persistent cataphylls and clusters of inflorescences
with reddish spathes (hence the name “roseospa-
. This species is unusual for the genus in
that it is generally terrestrial. It has prospered in
cultivation and is deemed an important horticul-
tural ee
ndron књ appears to be most
inde related to P. cuneatum Engl. from the Pa-
cific slope of Colombia, but the latter species dif-
fers in having acutely sulcate, C-shaped petioles
and smaller, more narrowly pedunculate inflores-
cences with white spathe blades. It is also related
to several other undescribed species from the Am-
azon basin.
Philodendron roseospathum is here divided into
two varieties, with variety angustilaminatum Croat
differing from the typical variety in having some-
what longer petioles and proportionately much nar-
rower blades (4.5 times longer than broad vs. an
average of 2.5 times longer than broad for the typ-
ical variety). See P. roseospathum var. angustilam-
inatum for additional differences
KEY TO THE VARIETIES OF P. ROSEOSPATHUM
Blades oblong, ca. 4.5 times longer than
wide; Panama (Colón), EN m elevation .
angustilaminatum
Blades + elliptic, ca. 2.5 ids longer than
wide; — a to Colombia, 400—900 m el-
atio
evatión < o0 ee a var. roseospathum
—
e
—
T
Philodendron roseospathum Croat var. roseo-
spathum
Flowering in Philodendron roseospathum var. ro-
seospathum occurs in the late dry season and early
rainy season (March through July) with post-anthe-
sis collections known from March through Septem-
ber and immature fruits from July to December. The
few South American collections concur with this
pattern. Flowering collections are known from Co-
lombia in July.
In Central America, P. roseospathum var. roseo-
spathum is common locally in cloud forests along
the Continental Divide as far west as Veraguas and
as far east as the El Llano-Cartí Road. It is cer-
tainly to be expected in Darién Province.
For differences with P. var. angustilaminatum,
see the preceeding key.
Additional specimens examined for P. roseospathum
var. roseospathum MA. Coclé: Continental Di-
vide ridge, Coclesito Road, Hammel 2540 (CR, MO);
El Copé region, 7 km N of Copé, Alto Calvario, 800 m,
Folsom & Mauseth 7948 (MO); 750-900 m, Croat
44738 (F, MO); 710—800 m, 8?39'N, 80?36'W, 68763
(MO); 900-1000 m, 8?39'N, 80736", Croat 75057
Ok El Valle region, La Mesa, N of El Valle de Antón,
m, 8?38'N, 80°09'W, Croat 67130 (CAS, СМ,
k MEXU, MO, NY, PMA, TEX); 860-900 m, 37346
(F, = 830-900 m, 8°36'N, 80°07'W, 74810 (CM,
LE, MO, WIS); ca. 800 m, 25435 (F, MO); Luteyn &
пе 1652 (MO); 850 m, 8°37'N, 80706", s
vers et al. 3513 (MO); 2700 ft., Sytsma et al. 4350 (CM,
MO); РАДНЕ m, 838'N, 80°7.5'W,
9908 (MO); Cerro Gaital,
80°07'W, Knapp 5311 (MO); 860 m, 8°37'N, 80°08'W,
Croat & Zhu 76734 (AAU, CAS, CM, MEXU, MO); N
slope and summit of Cerro Pilón, 900-1173 po Croat
22951 (MO). Panamá: El Llano—Cartf Road, 4 mi. be-
yond the highway, 500 m, Croat 49135 (MO); Ae. Jefe
region, 750-850 m, 9?15'N, 79°30'W, McPherson
11166 gi 11197 (CM, MO); 1000 m, Croat 49088
(MO); ca. 1000 m, Mori et al. 3795 (MO); ca. 950 m,
ca. 9:15' N, 79°30’ X. McPherson 7107 (B, is MO, US):
00-3000 ft., mel 3704 (MO); 750-800 m,
9°14'N, 79°22' W, en 67089 (МО); vic. jov de Pa-
cora, 800-1000 is Croat 22672 e L, MO, WIS); 3-
3.5 mi. NE of Altos de Pacora, 750 m, 9°15'N,
7925'W, Croat 68635 (MO); La тета
Kennedy et al. 3374 (US); md ". Weil 3158
у 8 (МО); 6 mi.
N
ле
79°W, Croat 69282 (MO); Mile 10.9, 450 m, 9*18'N.
79759", Croat 75123A (MO); Cerro Obu, er m,
400—500 m, de Nevers et al. 8054 (MO). Veraguas:
Santa Fe region, Santa Fe-Río I Luis, past Бо
Agricola Alto de Piedra, ca. 5 mi. N of school, 670 m,
833'N, 81%08'W, Croat etri (cn. MO, RSA); Santa
с Ради School, 570
Orquideas,” Sector Venados, 900 m
Cogollo et al. 3344 (MO). Chocó: Quibdó-Is stmina, polo
ity of Quibdó, <100 m, 6°28'N, 76°36'W, Croat t&
52233 (MO); Quibdó-Bolívar, 117-118 km E of Quibdó vi
465 m, 5°44’N, 76°28'W, Croat 57515 (CHOCO, s ,
COL, JAUM, MO, NY, PMA); Río Baudó, Рис a |
22048 (COL); ia ue at Km 208.5, 9 km pr
Tutunendo, ca. 9 km E of Quibdó, <100 m, 539
76°40'W, Croat pied
Volume 84, Number 3
1997
Croat 513
Philodendron Subgenus Philodendron
Philodendron roseospathum Croat var. angus-
tilaminatum Croat, var. nov. TYPE: Panama.
Colón: Cerro Bruja as approached from Río
Escandaloso, 915 m, 18 May 1978, Hammel
3133 (holotype, MO-2639732). Figure 349.
rnodia brevia; cataphylla song petiolus 24–27 с
ак lamina oblonga, 42—46 cm longa, 10 cm u
circa duplo longior quam petiolus; peduióulüs 3.8 cm lon
gus; spatha 7 ст longa, omnino viridalba; pistilla (4)5-
locularia; loculi 6—8- secs
Internodes short; cataphylls not seen; petioles
24—27 cm long; blades oblong, acuminate at apex,
rounded at base, 42-46 cm long, 9.5-10 cm wide
(4.2—4.8 times longer than wide), (1.75 times longer
than petiole); basal veins 2 per side, free to base;
primary lateral veins 10 per side. INFLORES-
CENCES with peduncle 3.8 cm long; spathe 7 cm
long greenish white throughout; spathe tube 3 cm
long; spadix 6.3 cm long; pistillate portion 2 cm
long in front; staminate portion 4.3 cm long; pistils
1.7 mm long; ovary 4(5)-locular; ovules 6-8 pe
locule, 1-seriate, ca. 0.3 mm s style similar to
style type D. Berries unkno
Flowering in Philodendron o ЈЕ уаг. ап-
gustilaminatum probably occurs in the wet season,
with the only flowering collection from May.
Philodendron roseospathum var. angustilamina-
tum is known only from the type locality near the
Canal Area in Colón Province, at 915 m elevation
in Premontane rain forest.
This variety is distinguished by its long-petio-
late, oblong leaf blades with rounded bases and
greenish white spathes. It is distinguished from the
typical variety by its proportionately much narrower
blades (4.5 times longer than wide in P. roseospa-
thum var. angustilaminatum vs. about 2.5 times
longer than wide in var. roseospathum) and propor-
tionately longer petioles (only 1.7 times longer than
petiole vs. 1.6-3.2 times longer in var. roseospa-
thum). In addition, P. roseospathum var. angusti-
laminatum has only 6-8 ovules per locule (vs. 16–
18 for P. roseospathum var. roseospathum).
It is unusual that P. roseospathum var. angusti-
laminatum has style type D while the autonymic
variety has style type B. Perhaps it is an artifact of
the poorly preserved nature of the material.
One collection, Croat 57592, from near Quere-
mal in Valle Department, at 1300 m in an area of
either Premontane moist forest or Tropical Lower
Montane wet forest, may also represent this variety
but has unribbed cataphylls. Croat 57036, a sterile
collection from Cotopaxi Province, Ecuador, also
unribbed cataphylls and may represent the
same taxon
Philodendron rothschuhianum "di Croat &
G : 659.
71): 124. 1920. TYPE: Nicaragua. Matagalpa:
Matagalpa, 1000 m, 12%55'N, 85?55'W, Roths-
chuh 229 (holotype, B). Figures 4, 9, 31, 336—
340, 347, 348.
Hemiepiphytic to epiphytic or rarely terrestrial;
stem appressed-climbing, creeping, sap watery, leaf
scars conspicuous, 2 cm long, 2.5 cm wide; inter-
nodes short, semiglossy, 1.8-3 cm long, 1.5-3 cm
diam., about as long as broad, gray-green, epi-
dermis fissured; roots scurfy, few per node; cata-
phylls fleshy, soft, 16-23 cm long, unribbed then
sharply 2-ribbed at apex, or bluntly 2-ribbed
throughout or weakly 1-ribbed throughout, green,
short dark striate, semiglossy, deciduous, emargin-
ate with subapical apiculum at union of ribs at
apex. LEAVES erect-spreading to spreading; peti-
oles 33-71 cm long, 3-13(20) mm diam., subter-
ete, moderately spongy, medium green, obtusely
l-ribbed near apex adaxially, surface weakly glossy,
densely green striate; sheath inconspicuous; blades
deeply 3-lobed, subcoriaceous, weakly bicolorous,
gradually acuminate, long to very long acuminate
at apex (the acumen inrolled, 1–2.5 mm long), =
hastate at base, 22—42 cm long, 25-57 cm wide
(0.6-1 times longer than wide), (0.5-1 times the
petiole length), about two-thirds as long as petiole,
upper surface semiglossy, lower surface weakly
glossy, slightly paler; sinus hippocrepiform; median
lobes 16—39.5 cm long, 7-20 cm wide; lateral lobes
narrowly ovate, 14-28 cm long, 5-17 cm wide, di-
rected toward apex, broadly confluent 2-5 cm with
medial lobe, the margins undulate; midrib sunken
to narrowly concave, paler than surface above, al-
most round-raised, paler than surface below; basal
veins 10—15 per side, the remainder arranged in a
* regular manner in the posterior lobe, coalesced
in part throughout much of their length into a cen-
tral rib; posterior rib usually not naked, sometimes
naked for 1 cm, rarely to 2.5 cm; primary lateral
veins 5-12 per side, departing midrib at a 50—65°
angle, narrowly sunken, concolorous above, convex
below; interprimary veins sunken and concolorous
above, raised, almost as conspicuous as primary
lateral veins below; minor veins darker than surface
below, arising from both the midrib and primary
lateral veins. INFLORESCENCES erect, 2—3(9) per
axil; peduncle 5.2-21 cm long, 5-10 mm diam.,
medium green, densely lineate; spathe (6.5)8—
14(16) cm long, (0.6–2.2 times longer than pedun-
cle), markedly constricted above the tube; spathe
514
Annals of the
Missouri Botanical Garden
blade pale green to greenish white to —
green outside, to ca. 5 cm wide when open, ca.
cm diam. at constriction, greenish white to creamy
white inside; spathe tube ellipsoid, 4—7 cm long,
pale green to yellowish green outside, greenish to
greenish white to creamy white inside; spadix very
short stipitate; somewhat protruding forward at an-
thesis, 10.5 cm long, constricted at base of fertile
staminate portion; pistillate portion pale lime-
green, 3.7 cm long in front, 2.5 cm long in back,
1-1.3 cm diam. at apex, 1.2-1.4 cm diam. at mid-
dle, 1.2 cm wide at base; staminate portion 6.4—9.3
cm long; fertile staminate portion creamy white,
clavate, 1 cm diam. at base, 1.3 cm diam. at mid-
dle, 1 cm diam. ca. 1 cm from apex, usually broad-
er than the pistillate portion; sterile staminate por-
tion broader than fertile or pistillate portion,
creamy white, 1.6-1.9 cm diam. at base, 1.3-1.4
cm. diam. at apex; pistils 1.6–2.6 cm long, 1.2 mm
diam.; ovary 5—7-locular, 0.8 mm long, 1.1 mm
diam., with sub-basal placentation; locules 0.8 mm
long, 0.3-0.4 mm diam.; ovule sac 0.6 mm long;
ovules 1 per locule, contained within translucent,
gelatinous ovule sac, 0.3-0.5 mm long, longer than
funicle; funicle 0.1-0.3 mm long (can be pulled
free to base), style 0.6 mm long, 1.2 mm diam.,
similar to style type B; style apex sloping to round-
ed; stigma brush-like, hemispheroid, 0.7-1 mm
diam., 0.3-0.6 mm high, covering entire style apex;
apex drying button-like, with or without radial ridg-
es from center; the androecium truncate, oblong, +
prismatic, margins irregularly 4—6-sided, 1-1.6 mm
long, 1.5-1.9 mm diam. at apex; thecae oblong, 0.4
mm wide, = parallel to one another, nearly contig-
uous; sterile staminate flowers blunt, sometimes
clavate or prismatic or irregularly 5—6-sided ог 3—
6-sided, 1.7-2.3 mm long, 1.6 mm wide. Berries
pale green or pale yellow-green to greenish white,
4 mm long, 2.5 mm diam.; seeds 1—3, medium
green, (1.2)2.1-2.3 mm long, 0.6–0.8 mm diam.,
within thin envelope. JUVENILE plants with peti-
oles sheathed to about midway; blades with poste-
rior lobes hastate, acuminate at apex, promptly au-
riculate on posterior margin, broadly confluent with
anterior margin on anterior margin.
e in Philodendron rothschuhianum oc-
curs during the dry season and early rainy season.
Most аба have been made in March, but
flowering occurs as late as August. Post-anthesis
collections have been made from January through
November (except October), but are particularly
abundant from March through May. Immature fruit-
ing collections have been made from April through
September (also December), but mature fruits are
known from only August.
Philodendron rothschuhianum ranges from Hon-
duras (Gracias a Dios) and from Nicaragua (Rfo San
Juan, Zelaya, Jinotega, and Matagalpa) to Panama
(Bocas del Toro, Veraguas, and Coclé), principally
on the Atlantic slope, from sea level to 1450 (most-
ly less than 1000) m elevation. It occurs in Pre-
montane wet forest, Tropical wet forest, and less fre-
quently in wetter parts of Tropical wet forest and
Premontane rain forest life zones.
Philodendron rothschuhianum is a member of P.
sect. Tritomophyllum. This species is distinguished
by its short internodes; unribbed, deciduous cata-
phylls; subterete, moderately spongy petioles; and
especially by its deeply three-lobed blades about
two-thirds as long as the petioles with the lateral
lobes broadly confluent with the medial lobe and
the sinus hippocrepiform and naked along the pos-
terior rib for a short distance from the petiole.
асар rothschuhianum is most frequently
confused with P. tripartitum, which differs in having
less auriculate Laci} lobes that are scarcely or not at
all naked along the posterior rib. In addition, P. tri-
partitum has a much more slender, scarcely constrict-
ed, spathe tube reddish within (vs. a markedly inflat-
ed, uniformly greenish tube in P. rothschuhianum).
In terms of leaf shape, P. rothschuhianum is clos-
est to P. cotobrusense. The latter differs in medial
lobes with 18—19 pairs of primary lateral veins (vs.
5-12 pairs for P. rothschuhianum).
The style apex is apparently variable in this spe-
cies, with Croat 66772 and Neill 1569 having type
B styles while Croat 35657 has style type D. The
latter has the style apex prolonged into a distinct
boss separated from the rest of the style apex by à
distinct neck.
Additional specimens examined. COSTA RICA. Ala-
juela: Quebrada Guillermina, on N side of Volcán Arenal,
500 m, 10° jet 84°42’ ex е et al. 3415 (Е, U); Nar-
ке = 8 km is d 600 m, Croat
46942 (MO); Finca Los Aia ‚ 11 mi. NW of Zar-
cero, ca. 850 m, pier 43567 (MO), 43637 (MO); У
teverde Biological Reserve, Rio Pefias Blancas, 7
Bello 1980 (CR, INB); 850-900 m, 10°20'N, M
Haber & Bello 6836 (MO); 850 m, 10°19'N, 8474.
Haber & Cruz 8409 (MO); Finca de Tomás Guindon, 900
m, 10°19'N, 84243'W, Bello 763 (CR, MO); Río mere
road to Colonia Virgen del Socorro, ca. т 10°5.5'
мя Y Grayum & Hammel 5516 ‚ СЕ, MO); A
Burger & Antonio 11101 (CR, s 11097 (F); e
m, ront 68301 (MO); Vara Blanca-Puerto Viejo. p mi.
of San Miguel, 380 m, Croat 35657 (CR. oe ау
Ораја, 3 km NNE of Bijagua, 450 m, 10°45 N85 ..
Burger & Baker 9881 (CR, F); 8 km N of M
2 eng 36502 (CR, MO); 13.8 km N of Bijagua.
50 m, Croat 36438 (MO); 36402 (CR, MO): Mi
e 1.8-2.7 km S of Río Canalete, 100 m 550
36402 (MO); 13 km W of Fortuna, Río Tabacón, a: ub
m, 10°29’N, 84%43'W, Liesner et al. 15250 (МО); 1
g^
Volume 84, Number 3
1997
Croat 515
Philodendron Subgenus Philodendron
NW of Arenal by air, 700 m, 10%34.5'N, 84°54’ W, Liesner
et al. 15088 (MO); 15065 (MO); San Ramón, Bittner &
Venschott 1941 (CR); Nilsson 460 dd Cartago: 4 =
SE of Pejibaye, 700 m, 9?48'N, 83°42’W, Liesner et al.
14330 (MO); Turrialba, etal dee (US). Guanacaste:
Parque Nacional Guanacaste ee смара 600 m
11°02’N, 85%25'18"W, xd
redia:
E
8026 (F); Tirimbina, 700 ft., Proctor 32148 (LL); La Selva
Field Station, 100 m, pere 2826 (DUKE); Folsom 9874
(DUKE); 50-80 m, 10°26'N, 84°01'W, Grayum & Cha-
varria 8300 (MO); Puerto Viejo de са ара Grayum
2120 (DUKE); с :
5756 (CR, US); Volcán Barva, Río Peje-Río Sardinalito,
700-950 m, 10%17.5'N, 84%04.5'W, Grayum 6981 (CR,
MO); 700-800 = 10°17.5'N, 84°04.5'W, 6720 (CR, m
Limón: ca. 1 mi. NE of Bribrí, ca. 40 mi. SW of Lim
above Río Catarala, 50-100 m, Croat 43245 (МО); 7 не
SW of Bribrí, 100-250 т, Gómez et al. 20320 (MO); Río
Pacuare, TS m, 10%05'N, 83°20" W, Burger & Liesner
6895 (F, MO); vic. of Guápiles, 300-500 m, Standley
37469 (US); 2 km W of Río Toro Amarillo, W of Guápiles,
275 m, 10°13'N, 83°50'W, Thompson & Rawlins 1216
(CM); Hacienda Tapezco—Hacienda La Suerte, 29 air km
о
rt a
m, 10°N, 83°05'W, Thompson & Rawlins 1176 (CM); Cer-
ro Coronel, E of Rio Zapote, along and above new road,
within 1 km of Río Colorado, 10-40 m, 10?40'N, 83%40'W,
Stevens 24277 (CR, MO); E of Río fepe 10-100 m,
10%40'N, 84740", Stevens 23974 (CR, MO); E of La-
na Danto, 20-170 m, 10?41'N, 8: ?38'W, Stevens
24453 (CR, MO); Parque Nacional Tortuguero, Puesto
Cuatro Esquinas, 4 m, 10?32'N, 83?30'W, Robles 1837
(CR, MO); Barra del Colorado, 0-2 m, 10?47'N,
83735'W, Stevens 24073 (CR, MO); Río Colorado, Barra
del Colorado, 1—5 i 10?47'40"N, 83?35'30"W, Davidse
MO); 2 airline km SSE of Islas rn
0 m, 10?40'N, т ^^
31035 (MO); Río Reventazó
Cairo, ca. 25 m, Standley & "Valerio 49027 (US); Parque
La Amistad, Fila Tsiurábeta, between Río Urén and Río
ri, 800 m, 9?27'30"N, 83°W, A. Chacón it v E
я hills above Santa Elena, 3
teverde, 1450 m, 10?20'N, 84°50’ W, — & чы 5067
(МО). 8ап r S of San José, Greenman & Greenman
5353 (MO); Ванд Carrillo National Park, Fila Carrillo,
m, . 21149 (CR, MO); 600-700 m, Croat
78777 (CR, INB), HONDURAS. Gracias a Díos: Ahuas
Bila, 200 W of Puerto Lempira, 100 m, Nelson &
Cruz 06 (CM. MO, NY, UNAH). NICARAGUA. Chon-
13°51'N, 85722' W, Stevens et
alpa: Macizos de Pefias Blancas, SE side, drainage of
Quebrada El Quebradón, slopes NW of Hacienda en
s tián, 800-1100 m, 13%14-15'N, 85738' W, Steve:
(MO); summit of El Toma Road, Меш 1569
(HNMN, MO); 10.5 km NW of Matagalpa, ca.
12°57'N, 85°51'W, Moreno 10233 (MO); El ا а 10
km de Matagalpa, 800—820 m, 12%59'N, 8 Alte Mo-
reno 6607 (MO); N of Cerro Musún, near W: s, Ara-
quistain & Moreno 2741 (LE, МО); NW of Сато Мо.
пеаг Раујо, m, Araquistain & Moreno 2572
OR 800-1200 m, 2510 (MO); 2495 (CAS, HNMN,
MO); 300—600 m, 2471A (MO); Río Bilampí, NW of Cer-
ro Musún, 4 km SW of Wanawás, 200-500 m, he
01'N, 85%14'W, Araquistain & Moreno ¿da (HN
MO, US); Comerca Wanawás, 180-200 m asia
85°13'W, Araquistain & Moreno 2398 (HNMN, K, MO);
near Cerro Musún, 8 km from Población Wanawás, Palán
er , 500-800 m, 12?59'-13^N, 85^14'W, Araquis-
n «€ Moreno 2355 (HNMN, MO, NY); Río Bilampf
3
Valley,
500-800 m, Neill 1800
Chontalefio, 20 km NE of El Castillo (Río Indio water-
shed), 200 m, Neill 3367 (MO); Neill & Vincelli 3541
(BM, MO); Río Indio end 6 km upstream from the
junction with Сапо La Pimienta, 300—600
(MO); Río Sábalo, Buenos phe 70 m, 11?02'N,
eds 28'W, Moreno 25595 (MO). Zelaya: ca. 1.5 km =
Las Esperanza de Las Quebradas, 300-350 m
13°38'N, 85°02'W, Stevens & кү 19308 (МО); Tou
baika-El Empalme, ca. 3.9 km SE of El Empalme, ca.
vi 2 ca. аха 84°24'W, pad 12930 (MO); po
a Nueva América, ca. f main road, 5
vens d MOL. Roéita-Puerto Cabezas, ca. 15.7 e
SW of Río Kukalaya, «100 m, ca. 13%58'N, 84°12'W,
Stevens 8500 (MO); Cerro El Inocente, near Cafio Ma-
jagua, ca. 800-1000 m, са. 13745"М, 85^W, Stevens 6805
(MO); ipn i qui ca. 31.4 km pagg id E ca.
8.9 km beyond Rosa Grande La Balsama, —
vens 7456 Mo): Saslaya-San José "n eos
ero, from Loma Mollejones eastward, ca. 200-400 m, ca.
13?44—45'N, 84?57—58' W, Stevens 7049 (BM, МО); Саћо
Majagua, ca. 750—850 m, са. 13745" М, 85°00-01 зида Ste-
vens ње (МО); Cafio Majagua—Cafio Sucio, са.
13°45'N, 84?59'-85?W, Stevens 6821 en
PMA): vic. TE La Рета. ca. 13?45'N, 84759" W, Pipoly
6225 (MO); Cerro La Pimienta-El Hormiguero, ca
1000 m, ca. 13%45'N, atai W, ond 6012 е Cerro
La Pimienta, number О m, ca. 13?45'N,
84759", Pipoly S (мо) са. 3 gi above “a ria,
<200 m, Pipoly 3794 (MO);
Risco de Oro, ca UP m, ду rk 5043 (MO); 10 km NE
of Siuna, along Cafio Madriguera, 250 m, Neill 3754
(BM, MO); 4 km NE of Siuna, road to El Dos, 300 m,
13?45'N, 84^45' W, Moreno & Robleto sr (MO); l km
W of El Naranjo, 200-210 m, 13°34'N, 85717, owe
& Robleto 20616 (MO, oy a of a El Hor
‚ ca. 900-1000 m, N, 84^59'50"W, Grijalva
473 (MO); Río lyas, Quebrada й Toro, 260—280 m, Vin-
celli 370 (MO); near San Juan del Norte, C. Smith 5 (F).
ANAMA. Bocas de roin Gualaca—Chiriquí Grande,
ca. 10 km SW of Chiriquí Grande, ca. 300 m, 8°52'N,
82^10'W, Thompson 4936 (CM); 4.2 mi. S of рн
Grande, ca. 0 т, 855'N, 82°09' W, Сөн С Жыш
Fortuna Lake area, Continental in
-Pherson & Aranda 10185
i m, 8°4
7950'04"W, MPs 10546
MO, US); ca. 300 m, ca. 8°50’N,
82715 W, ЖР 8522 (К, MO, US); 6.6 mi. N of
bridge over Fortuna Lake, 780 m, 8°45'N, 82°18'W,
516
Annals of the
Missouri Botanical Garden
Croat 66772 (MO); 1.2 mi. N of Divide, 910 m, 8° 44°N,
82°17'W, Croat 60462 (MO, PM i
of Divide, 770 a 8°45'N,
ios Río паран Finca St. ae s-K {
l. 1919 (MO). Coclé: El Copé region,
4. € mi. N of El Cons, 750 m, 8738'N, 80°35’ W, Thompson
4760 (CM, MO); 5-6 mi. N of El Copé, 600-800 m
8°38'N, 8°35'W, Croat & Zhu 77224 (CM, MO); 680-770
m, 8°39'N, 80°36’W, Croat 74828 (CM, МО); Alto Cal-
vario, Croat 68848 (MO). Veraguas: 15.6 km NW o
Santa Fe, 450-550 m, Croat 27704 (MO); 350-400 m,
27385 (MO)
=
Philodendron sagittifolium Liebm., Vidensk.
Meddel. Dansk Naturhist. Foren. Kjgbenhavn
1849: 17. 1849. TYPE: Mexico. Veracruz: Río
Nautla at Pital, Apr. 1841, Liebmann s.n. (ho-
lotype, C). Figures 346, 350-356
Philodendron ен опит, Liebm., ја вола "m
urhist.
ro ide Соно, Liebmann s.n. (holotype, С; 180-
Philodendron tanyphyllum Schott, Prod. Syst. Aroid.: 272.
E: Mexico. Liebmann s.n. (W destroyed).
Den ic. wp (neotype, here gros У).
agp capte sanguineum Regel, Ind.
. 1868. TYPE: Mexico. ert unknown: Kar-
nn s.n. (holotype, LE? no тена extant); t. 621
in Regel, es 18. 1869 (neotype, here des-
ignated).
Philodendron lancigerum Standl. & L. O. Williams, Ceiba
51. TYPE: Costa Rica. Puntarenas: vic.
Palmar Norte, Rio Térraba, 30 m, Allen 5612 (ho-
99728 В (ориуре;
designated, MO-3319112; ocios B, GR,
K, NY, PMA
Роне tuxtlanum G S. Bunting, Gentes Herb. 9.
` i : Mexico. Veracruz: vic. Santiago
Tuxtla, Bunting & Davies 162 (holotype, US).
Usually hemiepiphytic, rarely terrestrial or epi-
lithic; stem appressed-climbing, parchment-white,
sap turning blackish, slimy; internodes usually te-
rete, weakly angular, sometimes obscurely flattened
on one side or closely and acutely ribbed, semiglossy
to matte, 1-20 cm long, 1.5—4 ст ark green,
becoming usually gray-green to о sometimes
pale yellow-green, epidermis somewhat ridged
(“wrinkled”), sometimes fissured transversely; roots
reddish brown, 6-10 mm long, 3-5 mm diam., aris-
ing from and along the node on one side; cataphylls
18-39 cm long, usually weakly 1-ribbed, sometimes
unribbed or weakly to sharply 2-ribbed or sharply
l-ribbed (ribs to 1 cm high), soft, green, sometimes
reddish to pinkish, sometimes sparsely green-spot-
ted, purple-maroon or darker striate near base, dry-
ing reddish brown, margins sometimes prominently
and thinly raised, usually soon deciduous, rarely
persisting as a rotting mass, rounded at apex; peti-
oles 20-65(91) cm long, 3-15 mm
subterete, moderately spongy to firm, m
drying greenish brown, obtusely Poi usually
obtusely and narrowly sulcate, rarely obtusely and
broadly sulcate adaxially, surface minutely an
densely short purple- or occasionally white-striate,
sometimes dark green or violet-purple blotched,
sometimes smooth to irregularly ribbed and folded;
sheath subtending an inflorescence, ee cm long;
blades ovate to ovate-triangular, semiglossy, mod-
erately coriaceous, weakly to moderately hien
acuminate to narrowly acuminate, sometimes short-
acuminate at apex, prominently cordate to + sagit-
tate at base, 30—72 cm long, 15-39 cm wide (1.85-
2 times longer than wide), (0.6—1.6 times longer than
petiole), margins somewhat hyaline, weakly revolute,
upper surface medium green, drying brownish green
to greenish brown, semiglossy, lower surface pale
yellow-green, sometimes reddish, drying usually red-
dish brown, sometimes yellowish brown, weakly
glossy to matte; anterior lobe 24—61 cm long, 13-
34.5 cm wide (2—5.1(5.5—6.4) times longer than pos-
terior lobes); posterior lobes usually narrowly round-
ed, 6–20.5 cm long, 5-17.6 cm wide; sinus =
V-shaped to almost closed, 6-15 cm deep; midrib
flat to weakly raised, paler than surface, drying
broadly raised and + concolorous above, convex to
narrowly raised, sometimes maroon-spotted or with
white flecks, concolorous to darker than surface, dry-
ing broadly raised and reddish below; basal veins 3-
5(6) per side, with 0—1(2) free to base, third and
higher order veins sometimes coalesced 2.5-4.5 cm,
a few additional veins sometimes coalesced to 6.5
cm; posterior rib not at all naked or only briefly so;
primary lateral veins 4—9 per side, departing mi
at a 60—70° angle, weakly curved to the margins, but
usually turned prominently up just before the mar-
gin, rather prominently downturned just before the
midrib, round-raised to flat to obtusely sunken and
paler than surface above, convex, concolorous to
darker than surface below; interprimary veins weakly
sunken, concolorous above, flat, darker than surface
nating with dark,
INFLORESCENCES erect, 1-3 per axil; `
4—15 cm long, 1.6-1.8 cm diam., somewhat flatten
to terete, green, sometimes tinged reddish, drying
greenish, densely short and broad striate; spathe
8-22 cm long, 2-3 cm diam. (0.8-2.2 times i
than peduncle), weakly to obscurely constrict
above the tube, semiglossy, usually green, sometimes
TE mer mi eee
Volume 84, Number 3
1997
517
Philodendron Subgenus Philodendron
plum-red, often purple-spotted, densely short pale
lineate throughout, weakly so near apex, blunt to
narrowly cuspidate-acuminate to prominently acu-
minate, frequently tinged purplish violet at base;
spathe blade green to pale yellow-green, 8-11 cm
long (opening 3—4.3 cm wide, sometimes opening to
near the base), greenish white, weakly tinged red-
purple in throat to pale yellow-green or white inside,
sometimes reddish throughout in age; resin canals
appearing as continuous lines, red-purple to orange
in color; spathe tube green, sometimes moderately
to heavily tinged red-purple to red (B & K red-pur-
ple 3/7.5), with sparse, dark purplish spots (mostly
medially) throughout outside, 4—7 cm long, 2.5-3.5
cm diam., red to reddish purple (B & K red-purple
3/7.5), white striate inside; spadix stipitate to 7 mm
long; tapered toward apex, (8)9—16 cm long, broadest
below the middle or near the base, usually protrud-
ing somewhat forward at anthesis but not curved;
pistillate portion pale green (anthesis) to greenish
white to yellow-green (post-anthesis), uniformly wide
throughout or weakly tapered toward both ends, 2.5—
6.2 cm long in front, 24.7 cm long in back, 1-1.2
cm diam. at apex, 1.1-1.7(2.1) cm diam. at middle,
1.1-1.7 cm wide at base; staminate portion 6.3—
10.3(13) cm long; fertile staminate portion usually
creamy white, sometimes pale green to pinkish, 9—
13 mm diam. at base, 11-13 mm diam. at middle,
8-10 mm diam. ca. 1 cm from apex, broadest at base
ог middle, about as broad as the pistillate and sterile
portions; sterile staminate portion often broader than
the pistillate portion, white, (0.9-1)1.2-1.9 cm
diam.; pistils (0.9)1.8—4.4 mm long, (0.9)1.3-2.3 mm
diam.; ovary 6-9-locular, 1-1.7(3) mm long, 1.3-2.3
mm diam., with sub-basal placentation; locules 1—
1.7(3) mm long, 0.2-0.4 mm diam.; ovule sac
(0.6)0.8-1.2 mm long; ovules usually 24, rarely 5—
8 per locule, l-seriate (2-seriate, if 4 or more
ovules), usually contained within translucent, gelat-
inous ovule sac, sometimes contained within gelati-
nous matrix (no true envelope), 0.1-0.5 mm long,
usually longer than funicle; funicle 0.1—0.3 mm long
(can be pulled free to base), sometimes adnate to
lower part of partition, style (0.4)0.6-0.8 mm long,
(0.8)1.2-1.6 mm diam., similar to style type B; style
apex flat to sloping; stigma discoid or subdiscoid,
truncate, (0. YA 5 mm diam., (0.1)0.3-0.5 mm
igh, cov tyle apex, times shallowly
depressed at кы the по повећа truncate, pris-
matic, oblong, margins irregularly 4—6-sided to
weakly ovate, ca. 1 mm long, 1.6-2 mm diam. at
apex; thecae oblong, 0.4-0.6 mm wide, + parallel
to one another, sometimes contiguous; sterile sta-
minate flowers blunt, irregularly 4—6-sided, some-
times clavate or prismatic, 1.4-2.1 mm long, 1.4-2
mm wide. INFRUCTESCENCE pink, green at base
with reddish spots outside; berries pale yellowish,
rarely orange, with stigmas reddish brown, 7 mm
long; seeds 2-3 per locule, drying pale brown to
tannish brown, narrowly ellipsoid to oblong-ellip-
soid, 1.4—2 mm long, 0.7-0.9 mm diam., with faint
striations.
Flowering in Philodendron sagittifolium occurs
almost throughout the year, principally after the on-
set of the dry season and continuing throughout
much of the rainy season. There is a slight geo-
graphical shift, with flowering beginning about one
month earlier in Mexico and Guatemala (January
through August, less frequently in September and
October) and continuing somewhat longer in Pan-
ama (February through September, but also rarely
in December). Fruits apparently mature in about
two months’ time, but mature fruits have seldom
been collected, only from January and July.
Philodendron sagittifolium ranges from Mexico
(Veracruz) to Colombia (and probably also to Ven-
ezuela), from sea level to 1800 m elevation. It is
probably the most morphologically variable, and
one of the most ecologically versatile, species in
Central America. In Mexico, this species occurs in
“Selva Alta Perennifolia,” “Selva Mediana Subper-
ennifolia,” “Selva Baja Caducifolia,” and “Bosque
Caducifolio.” In the remainder of Central America,
it occurs principally in Tropical moist forest and
Premontane wet forest but also in Premontane rain
forest and Tropical wet forest life zones.
Philodendron sagittifolium is a member of Р
sect. Calostigma subsect. Macrobelium ser. Macro-
belium. This species, though highly variable in
most regards, can be characterized by its ap-
pressed-climbing habit; short, stout internodes;
sharply two-ribbed, deciduous cataphylls; obtusely
flattened, firm petioles usually spotted with violet-
purple; and ovate-triangular, moderately coriaceous
blades with the posterior rib not at all or only brief-
ly naked along the sinus. Also characteristic is the
externally green, frequently purple-spotted spathe,
which is reddish purple on the tube within.
In Mexico and Guatemala, Philodendron sagit-
tifolium is most easily confused with P. advena and
P. purulhense, differing from both in having the
blades somewhat triangular and drying reddish
brown rather than generally ovate and drying black-
ened. See P. advena for additional discussion.
In Panama, P. sagittifolium may sometimes be
confused with P. annulatum, which differs in hav-
ing the petioles somewhat spongy with a purple dis-
tal ring, blades typically ovate-oblong, and spathes
commonly white on the blade portion. It can be
518
Annals of the
Missouri Botanical Garden
confused at some dy of development with sterile
specimens of P. баке
Considering the highly variable nature of this
species there are many noteworthy collections, only
a few of which can be discussed here.
Whitefoord & Eddy 222 from Panama has the
lower blade surfaces drying yellowish brown rather
than reddish brown and spathe solid plum-red out-
side. Also noteworthy is Hammel et al. 14598,
which reports fruit color to be іа
А few collections from Puntarenas Province,
Costa ea (Croat 57243, uem Grayum & Ham-
, differ in having the primary lateral
veins m rather than darker on the lower surface.
Sterile specimens from Cocos island (W. Klawe s.n.,
Foster 4177) differ in having the minor veins mi-
nutely raised on the upper surface. These speci-
mens may prove to represent distinct species.
A large number of sterile and ostensibly juvenile
collections from Nicaragua are of Sgen identity.
ne series, including Pipoly 3826, ;
Stevens 7628, 12673, 12739, may dien prove
to be P. bakeri. Two other collections (Pipoly 5190,
5194) might prove to be still another species.
Croat 60804, from the coastal cordillera of Ven-
ezuela, reported as Р cf. sagittifolium (Croat &
mbert, 1986) is either this species or a very close
relative.
When Regel described P. sanguineum Regel, he
cited no specimens or country of origin, but Krause
(1913) cited a collection he had prepared from the
Berlin Botanical Garden of a cultivated Karwinsky
collection from Córdoba in Veracruz State, Mexico.
While no such collection still exists, Karwinsky
collected in Mexico only a few years (1840—1843)
before Regel described P. sanguineum. Thus it is
possible that both Regel and Engler could have
been dealing with the same material originally col-
lected by Karwinsky at Córdoba.
Another synonym that deserves mention is P.
lancigerum Standl. & L. O. Williams, correspond-
ing to a narrow-leaved form of P. sagittifolium re-
stricted to the Pacific lowlands of Costa Rica. Ep-
itypification was necessary because the type
specimen (Allen 5612) consists of only an inflores-
cence.
Additional specimens examined. BELIZE. Cayo
iocis fee Аа Bartlett 13052 (MICH), Blanes:
Wiley 460 (MO); Water Hole, near Vaca,
Cd 2447 (MICH): Río Ma Cal, San Luis-Cuevas, 1050
ft, Croat 23497 MON 1050 ft., 23539 (MO). cra
Punta Gorda, Cosminsky 227 (F); Richardson Creek, lower
part affluent of Bladen Branch, lower part of Maya Moun-
tains, 100-250 m, 88°46-48'W, 16°33'N, Davidse &
p 32334 (MO); Софији road, Gentle 6118 (LL); San
Antonio, Gentle 5510 (LL); Punta Gorda, Cosminsky 226
(F); Columbia River Forest Reserve, SW Maya Mountains,
trail between Gloria Camp and Edwards Camp to the 5,
16°22'N, 8910'W, Holst 4499 (MO). COSTA RICA. Al-
I : mi. N of San Miguel, 380 m, Croat 35642 (CR,
MO); 8.9 mi. NW of San Ramón, 1100 m, 10°10'30"N,
84^30' W Croat 68070 (L, MO); Río Sarapigü£, road to
Colonia Virgen del Socorro, 830 m, 10°1’6N, 84^11'W,
83°48'W, 3322 (K, MO); 31
9*57'N, 83?36'W, Thompson
~
—
zZ
eo cz
YT
i
uw
A
=
=
E
“<
e
~
~
S
S,
- р
+ =
Ф
~
сл
©
B
9°20'N, 83241" W, Lent 2967 (CR, F). Heredia: Mpa Salvi
Field Station, Hammel 10244 (MO); Grayum 2306 (MO);
Hammel 964] (DUKE); La Zona Protectora, SSE of Mag-
en Schatz 722 (CR, DUKE, MO); 4 mi. N of Vara Blan-
350 m, Croat 35579 (MO). Limón: vic. of Moin, 0
m, ane .04W, Croat 61206 (MO); Parque Nacional Tor-
uguero, 2 km S of Tortuguero, З m, 10°31’N, 83°30'W,
Robles 2166 (CR, MO); 2178 (CR, MO). Puntarenas: Pal-
mar Sur—Piedras Blancas, 20 m, Croat 32917 (MO); Que-
brada Aparicio-Quebrada Aguabuena, Rincón de Osa,
200-400 m, 8°42'N, 8 А W, Grayum а а!. 4017 (СЕ,
MO); Rincón de Osa, region to W of airstrip, 40—100 m,
Utley & "Utley 1119 Ў Зал ma ~ Coto Brus—Ciudad
Neily, роо m 1 W, Hammel 14159
(CR, MO); E ia Agua ari na, get 842'50'N,
8331'42"W, Herera 3959 (INB, МО); Las к sas—Las
Juntas de Coto Brus, ca. 900 m, 8°52’N, 82°57'W, Gra-
yum & Hammel 5692 (INB, MO); Parque Nacional Cor-
covado, Sirena, 1-10 m, 8?28'N, 83?35' W, Kernan & Phil-
lips 1028 (MO); 1-50 m, Kernan & Phillips 1089 (MO);
Kernan 461 (CR, MO), 574 (CR, MO); ds Aguilar 3396
(INB); Río Claro, 5 m, Kernan 783 (CR, MO); Las Cruces
Botanical Garden-Río Jaba, ca. 3 km SW of San Vito de
oto Brus, ca. 1050-1200 m, 8°47’ N 82°58’ W, Grayum
€
(MO); gp near TV mission er, n
8°49'N, 82°58'W, Croat 57243 (MO), 57231 (MO); Finca
Loma o. 1 mi. SW of Cañas Gordas, 1150 m, Croat
22287 (F, MO), 22257 (MO); hills above Palmar Norte,
naa m, Croat 35141 (MO); Punta Catedral, ca. 7 km
E of Quepos, 20-70 m, 922.5'N, 84097, Grayum &
pia 5903 (MO); Punta Quepos (3 km S of Puerto Que-
pos), O m, 9°24’N, 84°10'W, Grayum 6613 (CR, MO,
PMA, US); road to Rincón de Osa, 16.5 km W of Inter-
American Highway, 280 m, 8°45'N, 83722", Grayum el
al. 7550 (INB, MO); Reserva Forestal Golfo Dulce, 5 :
Rincón de Osa, 150 m, 8°37'N, 83°28’ W, Hammel & R
bles 16728 (CR, MO); 150-200 m, 16759 (CR, MO); 450
m, С. Herrera 3953 (CR, INB); ca. 0.7 km N of Golfito-
Villa Briceño road, 160—260 m, 8?40'30"N, 83°12'W, Gra-
yum & Hammel 10066 (CR, MO); Cantón de Golfito, Ji-
ménez, Alto de la Carbonera, road to Cerro de Osa, 200-
350 e e 30"N, 83°19'W, A. Chacón 1069 (CR, p
MO); 3 NW of center of Golfito, 30 m, 811'N
mia, pia 67632 (CM, CR, MEXU, MO, WIS); Pie-
Blancas-Rincón Road, Mile 3.7, 90-105 m, Croat
pol (CR, MO); Cocos Island, valley behind Bahía le:
lesias, Foster 4177 (F, МО); Klawe s.n. (POM). San Jose
San Isidro de El General-Dominical, SW of San спевова
Volume 84, Number 3
Croat 519
Philodendron Subgenus Philodendron
4.8 mi. from Rfo Pacuare, 1000 m, Croat 35254 (MO); 9
mi. SW of Rio Pacuare, 680 m, 35343 (MO); ca. 0.5 mi.
above turnoff to Canaán at Rivas, 900 m, Croat 43430
(CR, MO); 43415 (MO); умун Carrillo, Fila Cañon del
Río Sucio 450-700 m Chacón & Herrera 1705 (CR,
MO); Carara—El до de Turrubares, 200-310 т,
9%45'15'N, — 30"W, Grayum 10451 M INB, MO);
7. La greja, Santa Rosa де purisc m,
9?42' '50"N, 84093 30"W, Morales 1468 (CR, MO); Acosta,
Valle del cid € m, Mere & González
4581 (CR, INB); T: 1100-1200 m,
Croat v (INB, MO): 78959 у TNB. "мој; 990-1100
m, Croat & Hannon 79110 (MO); Parque Nacional, sector
Esquinas, vic. Fila Gamba, 200-300 m, Croat & Н
79294 (INB, MO). GUATEMALA. Eastern пуну of Ver.
apaz and Chiquimula, Watson 106 (GH). Alta Verapaz:
E of Tactic, 1300 m, Steyermark 43993 (F, US); near Chir-
riacté, on the Petén Highway, ca. 900 m, Standley 92189
(F); Chapultepec Farm, 62 km from Cobán on Sebol Road,
Contreras 4823 (LL); Río Carchá, Cobán-San Pedro Car-
há, ca. 1360 m, caro 89891 (F). Baja Verapaz:
greso-Cobán, Hwy. CA-14, 17 mi.
Hw wy. 17, 1620-1720 m, 15?13'N,
uisa m,
dley 62029 (F). Izabal: Escobas—Santo cde s, Santo To-
más Bay, 0-2 m, Steyermark 39331 (F); between Milla
49.5 and ridge 6 mi. from Izabal, Montaña del Mico, 65—
ai 3283 (NY); Sierra de las Minas, 13 km E of Doña
240 m, ca. 15?]4'N, 89?20'W, Harmon & Fuentes
5857 (UMO); Río Frío, 75-150 m, Steyermark 41533 (F,
LL). Petén: Canchacan, near San Luis, Lundell 16325
: La Shuya, SW o
16202 m, Stanley 67882 e HONDURAS. Mont. de la
Flor, 3300 ft., von Hagen & von — 1304 (F, NY).
Mida t dando Grande, ca. 10 km SW of La Ceiba,
15%42'N, 86751", Door & Mejía 26011
> 26352 (МО); 80-180 m, 15?42'N, 86751" У, Liesner
26179 (MO). Comayagua: El Achote, above Siguate-
o. 1350 m, Yuncker et al. 5894 (F, GH, K, MO, NY,
U). El Paraiso: El Junquillo-El Robledal, Sierra el Chile,
ta ~ чку чүү (EAP, Е, NY). Olancho: =
of Catacamas, m, Sta
Poe “ee vin 9875 (TEFH, US); Río Olancho, San
Esteban-Bonito Oriental, 14.8 mi. NE of San Esteban,
, Croat & Hannon 64295 (F, K, MO, NY) Mpio.
La Unión, 6 mi. E of La Unión along roa
950 m, 15°03'N, 86°35’W, Davidse et al. 35428 (MO).
Yoro: Cordillera Nombre de Dios, El hare José
де Texfguat, Cerro A ales, 830 m,
87°37'W, Davidse et al. 34524 (EAD, MO, mw. 34524
TE T
Ej
Я
~
5
N
Р
—
e
~
1 , 87°35'W, Liesner
26581 (MO). MEXICO. Patria, Engler 198 (GH, P). Chia-
:
mel et al. 15677 (MO): centies 69 mi. SW
of Palenque, 450 m, Croat 40197 (F, MO), 6 mi. N of
алея 1000 т, Croat 40581 (В, MEXU, МО); Pal-
que—Ocosingo, Hwy. shi
m, stun 10337 (MO): 13 m
al, m,
17°29'N, 92°05'W, Hammel et al. 15641 (MO); 18-20 km
N of Ocozocoautla, 800 m, Breedlove 25202 (DS); 3 mi.
N of Pueblo Nuevo Solistahuacán, 1700 m, Breedlove &
Smith 32436 (DS); 6700 ft, Thorne & Lathrop 40267
A). Morelos: Cuernavaca, Barranca del Salto de San
Antonio, Moore & заразен 8823 (ВН); ena р»
1417 (К, P). са: Latani, Dtto. Choa
17°24'N, 95°48’ W, gore Џез & Reko 900 (ECON, GH)
Sierra de Juarez, Tuxtepec—Oaxaca, ieh 44 е y ті.
above bridge at Valle Nacional, са. 5
ting 8902 (BH, CM); 8892 (BH): 660 n " vss 39752
(МО); 39757 (МО); 43917 (CM, MO); 1400 m, Croat
48000 (L, MEXU, МО); 705 m, 17%44'N, 96°19'W, Croat
& Hannon 65537 (B, MO); 55 km S of Tuxtepec, 630 m
17°37'N, 96°20’ W, Hammel & Merello 15482 (MO, NY);
Uxpanapa region, Esmeralda—Rfo Verde, 100 m, 17°10'N,
94?45'W, Croat
m о m,
17%09'N, 94°37'W, Hammel & Иле» 15556 (МО);
15555 (МО); Uxpanapa, 14 km E of Campamento La La-
guna (Poblado D.S.), 150 m, 17%05'N, 94°35'%, Hammel
& Merello 15578 (MO); Río Uxpanapa, Matuda 38666
(MEXU); Ixtlán, Mpio. Comaltepec, Puerto Antonio, ca.
1300 m, 17%45'N, eee Luna 436 (MO); Highway
a Hermosa, 1255 m, 17°35'N,
n ity, on high-
e & Bunting = (ВН, МО); 8863 (ВН, МО);
El ا 8857 (BH, MO); Fortin de las Flores (Posada
Loma), 850 m, Nevling & Gómez-Pompa 287 (cultivated,
XAL); Cátomáco-Acayucah; ca. 8 mi. S of Catemaco, near
Zapoapan, ca. 400 m, Moore £ Bunting 8925 (BH); For-
tín, Cervecería Moctezuma, 1000-1150 m, Croat 39415
(CM, F, MO); 900 m, Croat 44038 (MO); 5.7-6 mi. viciis
Catemaco, road to Sonte cog rae ca. m,
tepfo, 0 m, 18°38'N,
Bunting 8941 (BH); Play
95°05'W, Hammel & Mer ello eps (BR, CM, MO); Co-
atzacoalcos River, Fortuño, 30-50 m, Williams 8702 (V);
cn de Biología Trioucak “Los Tuxtlas," ca. 31 km
by road N of Catemaco, along road to Montepío, 250—300
, Grayum & Sleeper 8355 (МО); 300
m, 1844, 85°10’ W, Hammel & Merello 15490 (CM,
MO); 170-200 m, Gentry et al. 32488 (MO); 250 m, Ibarra
589 (MO); Mpio. Coatepec, Tuzamapan—Jalcomulco, Cerro
de Achichuca, 700 m, 19°23'N, 96°48'W, Castillo € Tap-
ia 678 (F, MO); Mpio. Hidalgotitlán, toward La Laguna,
160 m, 17°16.6'N, 94°33.4'W, Valdivia 744 (XAL); Hi-
dalgotitlán, 150 m, Valdivia 1351 (XAL); near La Es-
cuadra, 160 m, 17?19'N, 94°38.5'W, Valdivia 1330
(XAL); Mpio. J. de Ferrer, Cerro de Villa Rica, near Plan
de la Flor, 1250 m, 19°48'N, 96°46'W, Castillo et al. 1715
(F); Mpio. Jesús Carranza, 2 km N del Poblado 2, Ejido
F. J. Mina, 120 т, 17°16'N, 94°40'W, Vásquez & Navar-
rete 2383 (MO); Mpio. Pajapan, emen San Martín Paja-
Paj ^18'20"N,
Montepío, М оЁ Ѕап гаки Tuxtla, 150-200 m, 18°42'N,
95°10'W, Croat & Hannon 63158 (MO, NY, US); Mpio.
рана: Río Tablazos, ehe ена MBS 6 km
SSW of Tlapacoyan, 19°56'N, 97°13'W, Nee et al. 26090
520
Annals of the
Missouri Botanical Garden
(F, MO, NY); Vallé de Córdova, Bourgeau 2296 (K, P).
NICARAGUA. Estelí: El Zacatón (Plan > са. 1300
МО); Laguna de
0). ата са.
1450-1520 m, 13°01'N, 85°56'W, Stev
20435 (MO), Las Camelias—La ари 1100-1150 m,
13?05—06'N, 85°53-54'W, Stevens et al. 15327 (MO); N
slope of Volcán Yalí, 1200-1400 m, 13°15'N, 86710",
Stevens et al. 15091 (MO); Matagalpa—Jinotega Road, Km
146, 1 1400 m, Moreno 571 ; Laguna Miraflores,
ca. 26.1 km Hwy. 1 at Estelí, 1250-1300 m,
13%15'N, 86%15'W, 243 (MO); Las Ca-
vens & Grijalva 15322 (MO); Macizos de Pefias Blancas,
vic. of Finca of Manuel Estrada (El Cielo), 1200-1400 m
са. 13?15'N, 85741", Stevens 11665 (MO); 1500-1650
m, ca. 13°15-16'N, 85°41'W, Stevens 11430 (MO); Río
Bocay, Salto Kayaska, 190-340 m, са. 13°51'N, 85°22'W,
pace et al. jes m (MO). Matagalpa: m У Arenal,
road to Sanatorio de Aranjuez, 1400 13°02'N,
85°55’ W, Davidse e et al. 30444 (MO); 13901 Х, 8554 'W,
Moreno 9583 (MO); Fuente Pura, km 142, 1400-1450 m
13°N, 85?55'W, Moreno 17017 (MO); бай to Cerro La
Carlota, 2 km from Tuma, 1040-1100 m, 12%58'N,
85752'W, Moreno 15659 (MO); Cerro Picacho, behind La
Selva Negra Hotel, 1200-1540 m, 13°N, 85%55'W, Dav-
idse et al. 30311 (MO). Nueva Segovia: Río San Fernan-
do Valley, Cerro El Peñascal, 800-1125 m, 13°2-3'N,
20
m,
Stevens 12739 (MO); road to Mina Nueva América, ca. 10
m N of main road, Stevens 12673 (MO); ca. 11.3 km N
ог main road, Pipoly 5283 (MO); са. 6.3 km 5 of bridge
at pin Yolania, ca. 200-300 m, са. 11?36-37'N,
dis Stevens 4824 (MO); Cerro El Hormiguero, W
e, 1100-1183 m, ca. 13°44'N, 85°W, Pipoly 5198
(MO) 5194 (MO); 5190 etek Cerro La Pimienta number
a. 900-980 m, ca. 13° € W, 5107 (МО);
irte (MO); ca. 13 km ie ve Kururia, on road to San
Jerónimo, <200 m Sici po (NO, 1 km S of Monkey
Point, ca. 5 m, 11°35’ Moreno & Sandino
12531 (MO); Monkey adi p km NW, 1-5 m, 11%36'N,
83°38'W, Moreno & Sandino 11957 (MO); Río Kisalaya,
near Tala Has and Puente Mango, m, са. 14%41'N
84°03’ W, Stevens 7628 (BM, MO); Río Mico, near El Re-
creo, ca. 30 m, Standley 19404 (F); 19525 (EAP, F),
ded (F); Río Pis Че, 0.5—1. : km from Plantel El Salto,
road to Bonanza, ca. 140 m, ca. 14%03'N, 84°37'%, хав
vens aer (MO); Río Puma p Atlanta, ca. 10
11 , 84701", Moreno & ен bee MOS Was
ا Cabezas, Río Tron <200
14743'N, 84°06'W, Pipoly 4055 (MO). PANAMA. воа
del Toro: Gualaca-Chiriquí Grande, 4.2 mi. E of Chiri-
ми Grande, ca. 0 m, 8°55'N, 82°09’ W, Croat 66814 (MO);
mi. SE of Punta Peña, 120 m, 8?57'N, 8°56’ W, Croat
pes (AAU, CR, MO, NY); Fortuna Dam area, N of For-
tuna Dam, 650-700 m, 8%45'N, 82%15'W, Морено
11131 (МО); ge Grande Гоага. 1.6 mi. N
tinental Divide, 790 m, 8°45'N, 82°17'W, Croat &
Zhu 76453A дин 76456A (CM, МО); 10 mi. N Divide,
,
1 mi. N from highway, 130 m, 8°46'N, 82°11'W, Hammel
=" al. 14598 (MO); 4.3 km N of Divide, 590 m, 8°46’N,
2714", Croat & Grayum 60210 (CM, MO); 3. 2 mi. N of
end 700 m, 8°45'N, 82°15'W, Croat & Grayum Y n
(CAS, CR, F, KYO, M, MEXU, MO, PMA). Canal Area
Frijoles-Monte Lirio, 30 m, Killip 12133 (US); hills N of
Frijoles, Standley 27598 (US); Pipeline Road, 14 mi. NW
Gamboa gate, Croat 12363 (MO); Barro Colorado Is-
land, Croat 6472 (F, MO, RSA, SCZ); Aviles 25 (F); 46
F); Сашп Lake, Standley 31398 (US); Croat 6531 (F, MO,
SCZ), 6334 (MO), 5052 (MO), 10901 (MO), 8290 (MO),
10859 (MO), 9018 pods 8999 (MO), 10076 (MO); Sil-
vestre s.n. (MO); Croat 6023 (MO); Summit Gardens, Croat
59 139 (мо), hee MO) 10768 (MO), gera m
quí: Finc e Agua, 1300 m 1'N, 8
чийи 1 590 (МО); ‘Catia Gordon, ^ m E ~ см
Согдав, са. 1000 т, =, 22304 (MO); 8 mi. У of Puerto
rmuelles, in vic. of San Bartolo Limite, ca. 600 m, Croat
22022 (MO), 22026 (MO); Cerro erent above San Fé-
ix, 15-18 mi. N of Pan-American Highway, 800-1000 m,
Croat 33179 (CM, MO); Gualaca-Chiriquí Grande, For-
tuna Dam area, 7.2 mi. beyond Los Planes de Hornito,
1165-1200 m, 8°44'N, 82°14'W, Croat 67843 (CM, MO);
5.9 mi. NW of Los Planes de Hornito, 1225 ee 8°45'N,
82°14'W, Croat 67795 (CR, MO, PMA); 1400 m, Croat
48719 (MO); 48697 — 48681 (MO); Sara АЫ
тапде, 8 mi. Los Planes de Hornito, trail to Río
poner 1010-1130 m, 8?44' N, 82°14’ icon Croat 67918
(CAS, COL; L S MO, PMA, US); 10.1 mi. NW of
8
o
=
om
~
Plc de Hornito, 1250 m, 82917, °45'N, Croat
50040 (CM, MO); rar er Los Chorros—Quebrada Hon-
da, N of Fortuna e, 8°45'N, 82°14’ W, Chur-
о т,
chill & pos a. (MO); 4.5-5 km N of dam over
Fortuna Lake, 1100-1135 m, 8?43'N, 82717", Croat &
Grayum 60056 = CAS, F, L, MBM, МО, Р, PMA); Cerro
Colorado, 2.3 km above Chame, 1000 m, Croat 37067
је Coclé: Penonomé—Coclecito, Río Cascajal, 5.7 mi.
N of Llano Grande, 210 m, 840'N, 80°26’ W, Croat 67541
(CR, MO); El Copé region, 9.4 km above El Copé, 750-
900 m, Croat 44726 (MO); Alto Calvario, 710—800 m
Croat 68716 (EAP, G, HNMN, MO, TEX); 8 239'N,
80736", 68849 (MO); 580-740 m, 8”38'N, 80°36 6",
Croat 67522 (MO, US); El Valle region, La Mesa, above
> le de Antón, 860-900 т, Croat 37407 (MO); 870
, 850'N, 80%07'W, Hoover 1320 (MO); 800-900 m,
8%38'N, 80°09'W, Croat 67152 (B, CAS, CM, COL,
DUKE, F, K, L, MEXU, MO, NY, PMA, QCA, TEX, VDB,
WIS); 800 m, 8°36'N, 80%07'W, Croat & Zhu 76691 (МО);
Cerro Gaital, Churchill 3907 (MO). Colón: E
Portobelo, Río Piedras Lumber Road, 250 m, 9*22'30"N,
ded 1'30"W, Croat 75159 (MO); Río beth to de la
a, 0 m, Croat 36906 (MO); Miguel de la Borda, Croat
9856 | (E, MO, т» "s US); Santa ius Ridge Road, 10.6 km
from highway, ca. 380 m, Croat 34345 (MO); Mile 6.5.
370 m, 9°21’ 15", 79744" W, te & Zhu 76964 (МО);
along Río Guanche ca. 2 km E of bridge on ~ main
e 9°39' ү,
Croat 75176 (М
slopes оп W side, 60 m, 7%57'N, 7146" W, Croat
68958 sente 68893 [o ned Cana-Río Escucha Ruido,
above Cana Gold Mine 1430 m, Croat 37755 (MO):
37818 (мо; 37827 мој А of El Real, Alturas де Nique,
near Cana mine, along old Camino Real toward Colombia,
m, 7%45'N, 77%40'W, McPherson 11536 (МО);
mi. from P. Cocalito, Whiteford &
Eddy 222 (BM, МО); Parque Nacional del Darién, Pana-
ma-Colombia border, Río Pucuro, Cerro Tacarcuna, са. 6
— ө өөөчөӨӨЧч A РГ" "ГП ~ -СК[ЙЙЙЙЩ,,Щшыъҥъ ни m
Volume 84, Number 3
1997
Croat 521
Philodendron Subgenus Philodendron
km N of Cerro Mali, 1300-1500 m, 8?09.5'N, 7715 W,
Higo, N slope, 2400 ft., Ноу 4227 (МО). "we Вам
Azuero Peninsula, Jobero-JAMo Pedregal, 300—700 m,
Croat 34479 (MO); Río Guanico valley, 600 m, 7°18'N,
80°30'W, McPherson 9245 (MO, NY, SCZ). Panamá: Cer-
ro Azul, Croat 17281 (MO); El Llano—Carti Road, ca.
16-18 km N of Pan-American Highway, 400 m, Tyson &
Nee 7352 (MO) Mile 5-9, 200-250 m, 9?15-16'N,
78°59'N, Thompson 4624 (CM); Mile 10, 330 m, Croat
33823 (CM, F, MO, TEX, WIS); Mile 7, 460 m, 75109
(MO); Km v 2, 360—400 m, 25096 (MO); Mile 8.2, 450
m, 914'N, , Knapp & Hufi 4412 (MO); Mile 12,
200—500 m, Owen 695 (MO); Mile 5-9, 200—250 m,
9*15-16'N, 78*59'W, Thompson 4636 (CM); Cerro Cam-
N, 79*56' W, Croat ина (AAU, МО);
74771 (MO, PMA); 17177 (MO); са. 850 m, 8°40’N
79°50'W, Thompson 4577 (CM); 700 m m, 79*55'W, 8^40'N,
pe et al. 1174 (MO); — д (МО); са. 1 пи.
rom Inter-American Highway, 0 m, Croat 35983
eni күзө (МО); 800 m, BAIN, ce W, 74770 (MO);
efe region, near summit 750—800 m, 9*14'N,
79°29! W, Croat 67087 (F, MO); 800-1000 m, Gentry 2892
(GH, MO); 770 m, 9°15'N, 79°29’W, Croat & Zhu 76608
cora, 11.1-11.6 mi. b d
m, 9?43'N, 78° '69' W, ‘Croat 60506 (MO); Cerro Brew-
ster, 800-850 0 m, 918'N, 79°16’ W, de Nevers et al. 6287
(MO); El Lano-Carí Roa d, vic. Nusagandi, 450 m
9°18'N, 79*59'W, Croat 75150 (CM, MO, PMA); 350 i
9°20'N, 79°W, Croat & Zhu 77012 (CM, MO). Veraguas:
Dist. Montijo, Isla de Coiba, 7?37'N, 81744", Galdames
Santa Fe region, 7
RSA); Cerro
2284
81°47'W, Galdames 2256 (MO, P MA);
km W of Santa Fe, 2900 ft., Nee 11192 (MO,
me to 1200 m, Witherspoon et al. 8873 (MO); Santa Fe- *
fo San Luis, at Río Segundo Brazo, 480 m, 8°33'N,
81°08'W, Croat. 66886 (MO, PMA); 66916 (MO, PMA),
66920 (MO).
Philodendron scalarinerve Croat & Grayum, sp.
nov. TYPE: Panama. Panamá: road past and
3—3.5 mi. NE of Altos de Pacora, 7.8—8.2 mi.
above Pan-American Hwy., 11.1-11.6 mi. be-
yond Lago Cerro Azul, 700—750 m, 915'N,
79*25'W, Croat 68692 (holotype, MO-
3585744; isotypes, B, CAS, COL, CR, F, K,
NY, PMA, US). Figures 357-363.
Planta hemiepiphytica; internodia 1-1.5(3) ст longa,
1.5-3.5(4) ст diam.; cataphylla usque 22 cm longa, in-
lentia dederis petiolus subteres, 20-75 cm longus,
0.7-1.5 em diam.; lamina oblong-elliptica vel leniter
cordata 33-67 em longa, 11.4—40 cm lata, cum nervis
conspicue аи inter nervos minores; inflores-
centia 1-4; pedunculus 6-20 cm longus, subteres; spatha
16–21.5 ст longa, pallide viridis; lamina spathae sem-
initida, extus striata не ж intus viridi diluta; tubo
spathae hebetato, extus bo basi, intus viridalbo;
al
pistila 4–6(7–8)- oros: rad (10)12-14-ovulati.
Appressed-climbing hemiepiphyte; stems mod-
erately short; leaf scars conspicuous, 2—4 cm wide;
internodes 1-1.5(3) cm long, 1.5-3.5(4) cm diam.,
about as long as broad; roots moderately few per
node, short, drying dark brown, matte, longitudi-
nally ridged; cataphylls moderately thick, to 22 cm
long, unribbed to bluntly l-ribbed, rarely sharply
2-ribbed, drying dark brown, persisting semi-intact,
eventually fibrous at upper nodes. LEAVES erect-
spreading, clustered at or near stem apex; petioles
20-75 cm long, 0.7-1.5 cm diam., subterete, stiff,
green, obtusely flattened, sulcate adaxially; sheath-
ing 1-2.5 cm long; geniculum scarcely thicker than
petiole, 6–7.8 cm long slightly paler than petiole;
blades oblong-elliptic to narrowly ovate, moderate-
ly coriaceous, acuminate at apex (the acumen
sometimes inrolled, to 2 mm long), obtuse to sub-
cordate at base, 33-67 cm long, 11.4—40 cm wide
(1.24 times longer than wide), (0.82-2.11 times
longer than petiole), broadest at the middle, mar-
gins weakly undulate, upper surface dark green,
semiglossy, lower surface much paler, matte to se-
miglossy, obscurely dark-punctate at 10X magni-
fication; midrib convex to narrowly raised, some-
times paler than surface above, convex to thicker
than broad and concolorous below; basal veins 1—
4 per side, with all free to base; primary lateral
veins 5-18 per side, departing midrib at a 60—75*
angle, arcuate-ascending to the margins, sunken
ve; minor veins arising from the midrib only;
with scalariform *cross-veins" weakly visible when
fresh, conspicuously raised above on drying, barely
visible below. INFLORESCENCES spreading, pen-
dent, shorter than leaves, 1—4 per axil; peduncle
6-20 cm long, subterete, white-streaked near apex,
spongy; spathe 16-21.5 cm long (0.6-1.4 times
longer than peduncle), pale green, acuminate at
apex, obtuse at base, convolute to about the middle
at anthesis; spathe blade semiglossy, white-streaked
at apex outside, light green inside; spathe t
matte, white-streaked at base outside, 5-7 cm long,
2–3.5 cm diam., greenish white inside; spadix ses-
sile, erect, (7.6)9—12 cm long, broadest at the base
or at the middle or below the middle; pistillate por-
tion light green, slightly tapered above middle or
toward both ends, (1.7)2.6—4.3(7.8) cm long, 0.8—
1.3 cm diam. at apex, 0.8-1.4 cm diam. at middle,
0.9-1.4 cm wide at base; staminate portion 4.8-8.7
cm long; fertile staminate portion (9)12 mm diam.
at base, (9)12 mm diam. at middle, (6)10 mm diam.
522
Annals of the
Missouri Botanical Garden
ca. 1 cm from apex, broadest at the base, slightly
narrower than the pistillate portion, slightly narrow-
er than the sterile portion; sterile staminate portion
as broad as the pistillate portion, 1.2-1.4 cm diam.;
ponk (1.4)2.3-2.8(3.2) mm long, (1)1.4-1.9 mm
diam.; ovary 4—6(7-8)-locular, (0.7)1.5-1.8(3) mm
ibi: (1.1)1.5-1.9 тт diam., with axile placenta-
tion; locules (0.7)1.5—1.8(3) mm long, 0.4–0.6 mm
diam.; ovules (10)12-14 per locule, 2-seriate,
(0.1)0.3—0.4 mm long, longer than funicle; funicle
0.1-0.2(0.3) mm long, adnate to lower part of par-
tition, style 0.5(0.7) mm long, 1.1-1.4(2.2) mm
diam
lobed, 0.9-1.2(1.4-1.7) mm
(0.1)0.2-0.3 mm high, covering entire style apex,
inserted on center of style apex, sometimes medi-
ally depressed; the androecium truncate, Е
sometimes weakly oblong to oblong, т
sided, sometimes scalloped, 0.608-11 mm long,
1-1.8 mm diam. at apex; thecae oblong, 0.3-0.5
mm wide, + parallel to one another, contiguous or
nearly contiguous; pollen ellipsoidal, <0.1 mm
long, <0.1 mm diam.; sterile staminate flowers ir-
regularly 4—6-sided, clavate or weakly rounded,
blunt, rarely irregularly 3-5-sided or depressed
with a furrow, (0.8-1.2)1.7 mm long, 0.9-1.5(1.5-
2 mm) wide
Flowering in Philodendron scalarinerve occurs
principally in the early rainy season (June through
August) but also in March. Post-anthesis inflores-
cences have been collected primarily from June
through September, but one such Costa Rican col-
lection was made in January and one South Amer-
ican collection in December. Immature fruiting col-
lections are known from July through December.
Philodendron scalarinerve ranges from Costa
Rica to Ecuador, from sea level to 1325 m in Trop-
ical wet forest, Premontane rain forest, Tropical
Lower Montane rain forest, and Tropical rain forest
(Colombia).
hilodendron scalarinerve is a member of P. sect.
Philodendron subsect. Philodendron ser. Fibrosa.
This species is distinguished by its short inter-
nodes, bluntly 1-ribbed to unribbed cataphylls per-
sisting as pale fibers with small fragments of thin,
pale epidermis, terete to subterete petioles, narrow-
ly ovate to oblong-ovate blades drying dark with the
"cross-veins" (tertiary veins that extend between
the minor veins) numerous and conspicuous on
both surfaces and uniformly greenish spathes.
Additional specimens examined. COSTA RICA. He-
redia: Finca El irm N of Río Sarapiquf, ania de
Sarapiquí, са. 100 m, 10%27'N, 84°04’ W, Grayum & Ray
5564 (CR, MO); Cerros — ca. 2-2.5 ki N of Chi-
10°28’N, 84704", Gra-
yum et al. pe d MO). ки: Cerro Coronel, 5 of
m Е ren eN
NY, PMA, QCA). no Fortu
riquí Valley, vic. IRHE facilities, 1100-1200 m, 8%45'N,
82718", Croat 66593 (AAU, CAS, CM, COL, DUKE, С,
GH, HUA, K, L, MEXU, MO, NY, PMA, QCA, TEX, US,
VDB, VEN); Lago Fortuna, йон trail to Rfo Hornito,
8°45'N, 82°18'W, Croat & Zhu 76419 (MEXU, MO, SCZ,
US). Coclé: El Copé region, 5—6 mi. N of El Copé, 600-
800 m, 8°38'N, 8°35'W, Croat & Zhu 77222 (HUA, МО,
MY, QCNE); ca. 0.5 mi. N of Continental Divide at Alto
Calvario, ca. 5.5 mi. N of El Copé, 800 m, p ies
80°36’ W, Croat 75081 (MO, US); Alto Calvario, ca.
N of El Copé, sq - bid ti ?36' W, Croat cates
rata 80 Hag W, Mc-
Pherson 12856 (BR, en "s CR PMA); m, Croat
49196 (CAS, L, MO); 49174 era 49154 но 68722
(DUKE, F, MO, PMA); 750-900 m, 44737 (СМ, МО);
44720 (MO, PMA); El Valle region, La Mesa, N of El Valle
de Antón, f 80°09'W, Croat 67203
(CR, MO, her QCA, US); 25436 (F, MO); 880 m, 37569
(F, MO); ca. 1000 m, Kennedy & Dressler 1759 (MO); Lu-
teyn & урне 1717 (DUKE); Cerro Gaital, 800–900 m,
8°37'N, 80°07’ W, McPherson 11196 (MO, PMA). Colón:
Santa Rita Ridge Road, ca. 22 km from у 500 seño
925'N, 7940", Hammel et al. 14478 (MO). P.
Vertiente Pacifica, 150-200 m, 9?13.5'N, 78°15’ W. "Her.
rera. 1604 (K, MO, PMA, beter US); Cerro Jefe region,
= ра km Кот highway, са. 600 т, "-— 35919 дине
po Tres, 3 mi. NE s eag de Pac
eps 22704 (CM, MO, ; 700 m, үзү 27072 (E
MO), 27093 (MO); ca. 1 m e Alto de Pacora, 2600
& Kress 13420 hee El Llano-Cartí, 56
. N of Inter-American Highway at El Llano, 350-375
m, i Ch) 34805A (F, MO, NY, QCA, У WIS) Mile 6.8, 350
m, Croat 49112 (MO); Mile 7, 9^1 9'N, 79°59'W, Croat
75103 (CM, MO). San Blas: Nusagandi, 300 m, P15'N,
19*W, McPherson 11066 (MO, PMA); је Urgandi-Cerro
Óbu, on trail, 100—300 m, 9°23'N, 7848'W, de Nevers el
al. 8019 (MO, PMA); Río Diablo, 40 m, ges 78734" W.
Herrera et al. 1714 (MEXU, MO, PMA, STRI, US). Ver-
aguas: Santa Fe region, slopes of Cerro Tute, NW of Santa
Fe, 1250-1350 m, Croat 48971 (F, MO, W); Santa Fe-
Río San Luis, 8 mi. N of school, 450 m, 8”33'N, 81708 W.
Croat 66959 (CM, ENCB, MBM, MO, PMA, QCA); Alto
Piedra-Calovébora, Río Dos Bocas Valley, 350-400 m,
Croat 27367 (F, MO, PMA,
COLOMBIA. Chocó: Medellín-Quibdó, 78 km W of
Bolívar, 466 m, Croat 49286 (MO); Km 175-176, 117-
118 km E of Quibdó, 465 m, 5°44'№, ds 6°28'W, Croat
57485 (CHOCO, COL, JAUM, К, MO, US); Quibdó-lst-
mina, Km 14, Msn vea 6731 W, Croat & Cogollo
52215 (MO); S of ncherita, Km 31-32, «50 m.
STN, u W, Psi 51372 (CHOCO. COL, JN
MO); Quibdó-Lloro, vic. Río a. 150 m, 529N,
76735'W, Croat 55979 бм JAUN, мо, nice Rio al
Rio Митђа, upriver from 80-12 5°37'N,
76°25'W, Juncosn 1457 (MO); ууч етен 12 km 5 of
Volume 84, Number 3
1997
Croat 523
Philodendron Subgenus Philodendron
Quibdó, 60 m, 5°38' №, 76°40'W, Croat 56261A (CM, МО);
Serranía de Baudó, Las Animas—Pato (Río Pato), 4 km S
of Pato, 150 m, е 76 а ы Croat 56151 (МО); Je-
quedó, 41 km s Animas, ca. 10 km E of Río Pato,
220 m, Gentry & Rentería 241 12 (МО
78°08’ W, Betancur et al. 4778 (MO).
region, Buenaventura-Río Calim
Tomar, 50 m, 4%02'N, 77%07'W, Creed 61278 (AAU, B
CAS, CM, COL, CR, F, K, L, MO, NY, QCA, US); 61380
(СМ, MO); road to Málaga, 6 km S of main road, 50-8
m, 3°56'N, 77%07'30"W, Croat 69417 (AAU, COL, ad
СВ, MEXU, MO, NY, QCA); 100 m, 3?55'N, 77°W, Mon-
salve 892 (MO); Buenaventura-Málaga, Km 65-66, 40—
m, 4^10'N, 77?12'W, Croat 71055 (MO); at Km 17.5,
3°57'N, 77%01'W, Croat & Bay 75630 (MO); Km 28, W
of Em Man road, 3°59'N, 77°03'W, Bay 269
(MO); Km 35.2, 100 m, 4°N, 77°03’W, Croat & Bay 75759
(MO); Km 44 , «100 m, gg Ne А roat & Watt
70199 (COL, "мо, NY, U 2.4, 140 m, 4%03'N,
S; K
7705", Croat & Bay 75724 alo: 75728 (MO); 11 km
NW of Cali-Buenaventura Highway, 3756' 30"М, 77%01'W,
Croat 69321 (CM, MO). ECUADOR. Carchi: Chical,
1200-1250 m, 0%56'N, 78711", Thompson & Rawlins
761 (CM, MO).
Philodendron schottianum H. Wendl. ex Schott,
Oesterr. Bot. Z. 15: 72. 1865. TYPE: Costa
Rica. Wendland s.n. (holotype, W? lost). Schott
ic. 2735-36 ор. here designated, W).
Figures 2, 17, 365-371
Usually epiphytic or hemiepiphytic; stem ap-
pressed-climbing, grayish green, sap strongl
thyme-scented; internodes short, thick, semiglossy,
4–7.5 ст diam., broader than long, pale green to
gray, epidermis thin, yellow, fragmented, without
fissures; roots moderately few per node, drying dark
rown to ca. 5 mm diam., epidermis semiglossy,
flaking; cataphylls 16—46 cm long, sharply 2-ribbed
(ribs to 1.5 cm high), usually tinged red, drying
brownish yellow, often glossy (as if surface is shel-
lacked), broadly concave to broadly D-shaped
adaxially, persisting semi-intact, finally as a dense
mass of whitish fibers; margins acute; petioles
35-83 ст long, 1–2.2 cm diam., subterete to
D-shaped, firm to moderately spongy, medium
green, drying yellowish brown, obtusely flattened
with obtuse medial rib toward apex adaxially, sur-
face semiglossy and obtusely striate; blades ovate,
subcoriaceous, semiglossy, moderately bicolorous,
acuminate to abruptly acuminate at apex (the acu-
men strongly inrolled, 2-8 mm long), cordate to
sagittate at base, 30-77 cm long, (17.5)23-64 cm
wide (1-1.7(2.7) times longer than wide), (0.6-1.2
times longer than petiole); upper surface dark
green, semiglossy to subvelvety-matte; lower sur-
face semiglossy ог гагеју matte, moderately рајег;
anterior lobe 23-57 cm long, 24.5-64 cm wide
1.6-2.7 times longer than posterior lobes); poste-
rior lobes 8.5-28 cm long, 10-29.6 cm wide, ob-
tuse to broadly obtuse; sinus usually spathulate, 8—
20 cm deep; midrib flat to broadly convex, paler
than surface above, convex to narrowly rounded,
concolorous or slightly darker than surface below;
basal veins (1)6-7(8-9) per side, with 0—1(2) free
to base, most of the remainder coalesced 1-5.5 ст,
2 coalesced to 11 cm; posterior rib dede nus
1-3 cm long; primary lateral veins 3-8 per side,
departing midrib at a usually 40—70° ilte quilt-
ed-sunken to sunken, paler than surface above,
convex and slightly paler than surface below; ter-
tiary veins visible and darker than surface below;
minor veins conspicuous, arising from both the
midrib and primary lateral veins, moderately prom-
inulous on drying, alternating with secretory ducts
perpendicular or more frequently oblique, some-
times branching; “cross-veins” weakly parted. IN-
FLORESCENCES erect to erect-spreading, (1)2-3
per axil; peduncle 2-9 cm long, 8-14 cm diam.,
pinkish red, white striate, especially toward apex;
spathe 10-17 cm long (1.9-7 times longer than
peduncle), weakly constricted, oblong-ellipsoid;
spathe blade light green outside, cream, pale li-
neate in upper one-half inside; spathe tube green,
tinged red outside, 6-9 cm long, red with conspic-
uous resin canals inside; spadix tapered to some-
what ovate, weakly protruding, 8.8-14.4 cm long;
pistillate portion weakly ovoid, whitish, (2)3—4.9
cm long, 2–2.4 cm diam. throughout, 1.4–1.5 ст
diam. at apex, 1.4—1.9 cm diam. at middle, 1.4—
1.8 cm wide at base; staminate portion 5.7-10.8
cm long; fertile staminate portion bluntly tapered
at apex, 1.1-1.4 cm diam. at base, 1.2-2.3 cm
diam. at middle, 1(1.7) cm diam. ca. 1 cm from
apex, broadest at upper two-thirds, broader than the
pistillate portion, slightly narrower than the sterile
portion; sterile staminate portion broader than the
pistillate portion, 1.4-2.5 cm diam.; pistils (1)3—
3.9 mm long, 1.5-1.9(3.8) mm diam., margins
broadly rounded and slightly raised above the apex;
ovary 4—6-locular, 1.9-2.5 mm long, 1.3-1.9(3.8)
mm diam., with axile placentation; locules 1.9-2.5
mm long, 0.4-0.7 mm diam.; ovule sac not present
or to 1.9 mm long; ovules 10-14(18) per locule,
2-seriate, rarely contained within translucent or
transparent envelope, 0.1 mm long, longer than
funicle; funicle 0.1–0.3 mm long, adnate to lower
part of partition, style 0.8-1.2 mm long, 1.5-
1.9(3.6) mm diam., similar to style type B; style
apex barely raised, button-like, broadly concave,
medial apex with a whitened margin, raised and
apparently like type D on drying; stigma subdis-
coid, truncate, 1.4-1.6 mm diam., 0.3-0.4 mm
high, covering + entire style apex, sometimes de-
524
Annals of the
Missouri Botanical Garden
pressed shallowly and medially; the androecium
truncate, oblong, prismatic, margins irregularly
5-sided, sometimes scalloped, 1 mm long, 1.8-2.1
mm diam. at apex; thecae + oblong, 0.4–0.5 mm
wide, + parallel to one another; sterile staminate
flowers blunt, irregularly 4—6-sided, prismatic, 2.5—
3.5 mm long, (1.4)1.8-2.2 mm wide. INFRUC-
TESCENCE with pistillate spadix 5—6 cm long, 3.5
cm diam.; berries white, 1.1 cm long, 44.6 mm
diam.; seeds 1.4 mm long, 0.5 mm diam., cream-
colored.
Flowering specimens of Philodendron schottian-
um have been collected only from March and June,
but post-anthesis collections have been made from
March through August. Immature fruiting collec-
tions have been collected from January, May, Sep-
tember, and November. The immature January
fruiting collection is a clear indication that the spe-
cies must flower much earlier in the dry season
than March (as indicated above). Perhaps it flowers
throughout the dry season. In the cloud forest
regions where this species occurs the dry season
would not be very severe.
Philodendron schottianum ranges from Costa
Rica to Panama at (490)730 to 2250 m in Premon-
tane rain forest and Tropical Lower Montane wet
forest life zones. In Panama, this species ranges no
further east than Veraguas (Cerro Tute), except for
a disjunct occurrence on Cerro Jefe in Panamá
Province. All Costa Rican collections are from the
northern slopes of the Cordillera Central in Alajue-
la, Heredia, and San José, and the northern end of
the Cordillera de Talamanca in the Tapantí region
of Cartago Province. It is to be expected throughout
much of the Cordillera de Talamanca
Philodendron schottianum is a sine жм of P. sect.
Philodendron subsect. Philodendron ser. Fibrosa.
This species is distinguished by its short, thick in-
ternodes; sharply 2-ribbed cataphylls persisting as a
dense mass of fibers (frequently with patches of
glossy, yellowish to orange-brown epidermis); ob-
tusely to sharply D-shaped petioles drying somewhat
grayish or rarely yellowish and glossy; large, broadly
ovate blades with a deep, usually spathulate sinus,
usually pale-drying primary lateral veins; and rather
conspicuous secretory canals between the veins; 2—
З short-pedunculate inflorescences per axil; and
d constricted, externally green spathes red on
the tube within.
In Costa Rica and at higher elevations in Pana-
ma, such as on Cerro Colorado and on Cerro Pate
de Macho (1000 to 2200 m), the petioles are sub-
terete or obtusely flattened adaxially. At lower el-
evations in Bocas del Toro, Veraguas, and Coclé,
the petioles become D-shaped to sharply D-shaped
with erect margins, and at the lowest elevations
they are nearly always wing-margin
In Bocas del Toro Province, |i at middle
elevations and in mesic situations, this species is
most easily confused with P. findens, which also has
sharply D-shaped petioles. Philodendron findens
also has spathes which, like those of P. schottian-
um, are barely constricted midway. In rare situa-
tions where the blades of P. findens do not promptly
tear into narrow segments, P. findens can be distin-
guished from P. schottianum by having primary lat-
eral veins of the lower surface drying darker than
the surface.
In central Panama, P. schottianum can also be
confused with P. llanense. Both P. schottianum and
P. llanense occur in the Cerro Jefe region, though
P. schottianum has been collected there only once,
northeast of Altos de Pacora. This collection (Croat
68691) exhibits most of the diagnostic features of
P. schottianum, especially the persistent yellowish,
semi-intact cataphylls, and the acutely D-shaped
petioles with an obtuse medial rib (unknown in P
llanense), but has a blade shape midway between
that of P. schottianum and P. llanense (blade length/
width ratio 1.5 vs. an average of 1.4 for P. llanense
and 1.65 for P. schottianum). Philodendron llanense
differs in having at most obtusely flattened petioles
and in lacking the conspicuous. yellowish cata-
phylls of P. schottianum. llanense also
rarely occurs above 500 m (to 950 m), whereas Р.
schottianum only rarely occurs to as low as 500 m.
hilodendron schottianum can be confused with
P. thalassicum and P. alticola, especially in Costa
Rica. Philodendron thalassicum differs in having
bluish green leaf blades that are glaucous beneath
and sharply D-shaped petioles drying somewhat
blackened rather than merely obtusely flattened
and light yellow-brown, as in P. schottianum. Philo-
dendron alticola differs in usually having narrower
leaf blades (usually more than 1.8 times longer than
wide) and stigma пок exserted as minute funnels
on the dried sti
Philodendron dodioni may also be confused
with P. schottianum. The former differs in occurring
usually at lower elevations principally in tropical
wet forest and premontane wet forest, and in having
larger leaf blades, more long-pedunculate inflores-
ences, and longer spathes with a normal constric-
tion above the spathe tube.
One collection, Grayum 7333, differs from more
typical collections in having blades reportedly
matte on both surfaces and lacking promin
minor veins and conspicuous secretory ducts (80
evident in other material of this species where they
=== — ———— — НЬ он $” По IO, ct
Volume 84, Number 3
1997
roat 525
Philodendron Subgenus Philodendron
alternate with the minor veins). In addition, this
specimen has darker-drying petioles and more frag-
mentary old cataphylls as well as intact cataphylls
(perhaps juvenile?) drying brown (rather than the
typical brownish yellow) and a spathe with a nar-
rowly acuminate portion extending a full 5 cm be-
yond the end of the spadix. On other specimens,
the spadix ends only about 1 cm or less short of
the end of the spathe
Hammel et al. 14705 is unusual in lacking cat-
aphylls, suggesting that they might have been de-
ciduous (or forcibly removed during preparation).
ари“ prins — COSTA RICA. Ala-
i. W of San Ramón, ca. 800 m,
ea р g (MO); Re pee y Alto Tm 1900 m,
Lent 1845 (CR, F); Río Gorrión—Río m,
10°12'N, 84°19'W, 2996 (CR, F); Rro Cariblanco Canyon,
ariblanco,
°12'W, Grayum et ~ 61 86 (СЕ,
МО, NY); Río се road to Colonia Virgen del So-
corro, 830 m, 10°16'N, 84°11'W, Croat 68341 (CR, MO).
Cartago: 31 km from San José, SE of CR-2, 1750 m,
Harmon & Е, uentes 6081 (UMO); Tapantí Watershed Pre-
ве с . SW of Paraíso, ses او m, 9°43’N,
83°47'W, Croat 47047 (MO); 1500-1 nb & Gra-
yum 68293 (B, K, MO); 1500-1700 m, ii 36110 (MO)
(MO). Heredia: 3 mi. S of inp
7333 (CR, MO, US); Volcán Barva, Finca Montreal, Río
Voleán-Río San Fernando, 1740 m, 10?12'39"N,
84%06'45"W, Boyle et al. 1155 (MO, NY); Volcán Barva,
along Río Vueltas, 1 m, 10%06'N, 84%04'W, Burger &
Сетту 9042 (F, US); i fe Nacional Braulio Carrillo,
000 m, 10°16’ /38"N, 84%04'57"W, Boyle 964 (CM, MO,
9: 1223 (CR, МО); 1990 m, 10?11'03"N, 84?06'27"W,
8 (CR, MO); 1750 m, 10%03'40"N, 84°01’ ha pened
(CR. MO, NY). Puntarenas: Monteverde Reserv
m, са. 10°17'N, 84?48'W, Croat 61194 (CM, ja uS
а Cordillera de Talamanca, 730 m,
of Las
Utley & Utley 5259 (DUKE); Cerro de la Muerte, ca. 2000
m, Croat 32859 (MO); Braulio Carrillo National Park,
1000-1500 m, 10%05'N, 83?57'W, че 61226 (М О).
PANAMA. Восав del Toro: Cerro ~
ТЕ
& Zhu 76518 к DUKE, MO, WIS: са Cerro
Rio San Félix, 2000 m, Croat 48460 (MO, TEX); 18.5 mi.
N of Río San Félix, 1660 m, 8°30’N, 81746'W, Croat
74991 (AAU, CAS, CM, COL, СЕ, С, K, MEXU, MO,
QCA, US); Cerro Hornito, above Los Planes de Hornito
1750-1900 m, 8°41’N, 82^10' MA Croat 67979 (F, MEXU,
MO, PMA); Cerro Pate Macho, NE of Водице ea и
m, 8°46'N, 82725'W, Croat 66395 (F, MO); 1 aia, m,
48560 (MO); 1900-2000 m, 66505 (B, смге СОГ, СК,
ap F, L, MEXU, МО, ОСА, TEX, US, VDB, VEN);
а. 2200 m, Croat 48545 (MO); 1650-2000 m, 8?50'N,
82°25/ W, McPherson 11315 (MO, PMA); Cerro Punta,
above town, 2250 m, Croat 48601 (MO); Fortuna Dam
area, Gualaca-Chiriquí Grande, Río Chiriquí, 9.1 mi. be-
yond Los es aF ga Ys 1300 gh vato 48758 (MO);
Quebrada Are m, 845'N, 6'W, Hammel et
al. 14705 (MO). Co ae Е] Valle анти JB Mesa, N of El
Valle de Antón, E edge of Cerro Gaital, m,
8°37'N, 80%08'W, Croat 67234 (MO, PMA); 900—930 m,
37433 (F, MO); El Copé area, on western slope, just S of
the old saw-works area, 700 m, 8”38'N, 80?35'W, Croat
> Zhu 76795 (MO). Panamá: Cerro Jefe region, 3-3.5
NE of Altos de Pacora, 7.8-8.2 mi. above Pan-
Merlin Highway, 700-750 m, 915'N, 79*25'W, ез
68691 (AAU, CAS, К, МО, NY). Veraguas: Santa Fe
e—Calovébora, beyond Escuela Agrícola
d Río Tercero Brazo, m, 8°31'N,
Primero Brazo, 490 m, 8°33'N, 81°08’ W, 66879 (CM, CR,
HUA, IBE, JBGP, MO, PMA, QCA, US); ca. 3 mi. N of
the school, ca. 700 m, Croat 49002 (MO); 1.7 mi. past
the school, 570 m, 8*33'N, 81708", Croat & Zhu 76832
(MO); god of Cerro Tute, near Escuela Agrícola Alto
ана ag
m, 8°33’N, 81*08'W, Croat 67002
(CAS, CR, L, MO, NY 1 PMA).
Philodendron smithii Engl., Bot. Jahrb. Syst. 26:
540. 1899. TYPE: Guatemala. Izabal: Río
Dulce at Livingston, sea level, 5°49’N,
88°45'W, J. D. Smith 1535 (lectotype, here
ааг us add K). Figures 364,
2E 924949469
Usually Teese sometimes terrestrial or
epilithic; stem usually appressed-climbing, leaf
scars conspicuous, 7-15 mm long, 7-16 mm wide;
internodes glossy, (1)2.2-3 cm long, 2-3 cm diam.,
usually longer than broad or about as long as broad,
or gray-green to dark green, epidermis drying
yellow-brown with loose flakes, fissured conspicu-
ously; cataphylls soft, ече 12-20 cm long,
bluntly to sharply 2 d, green, dark green
short-lineate, и at apex, drying tannish to
yellowish, deciduous, intact; petioles 21.5-63 cm
long, 4-16 mm diam., subterete, spongy, medium
green, obtusely flattened with obtuse angle toward
apex adaxially, surface dark short-lineate, dark with
a dark purple-black ring around apex; blades nar-
rowly ovate, subcoriaceous, glossy, moderately bi-
colorous, acuminate to long-acuminate at apex (the
acumen inrolled, 1—4 mm long), cordate at base,
26.5-53 cm long, 11-31 cm wide (1.2-2.2 times
Annals of the
Missouri Botanical Garden
longer than wide), (0.7—1.4 times longer than pet-
iole), margins hyaline, convex is lower half of
blade, upper surface dark green, drying dark
gray-green to yellow-brown, lower surface light
greenish, usually drying yellow-green to yellow-
brown, sometimes olive-green; anterior lobe 18—41
cm long, 18-30 cm wide, (1.8-2.9 times longer
than posterior lobes); posterior lobes (3.2)7.5—16
cm long, 8.7-14.2 cm wide, obtuse to broadly
rounded; midrib flat to broadly convex, paler than
surface above, prominently raised, concolorous be-
low; basal veins (3)4—5(6) per side, with 0-1 free
to base, 1-2 coalesced (1.8)2.5—5.5(7.5) cm; pos-
terior rib not at all naked or briefly so, usually 1—
1.5 cm (rarely to 2 cm); primary lateral veins 3-6
per side, darker than surface below, departing mid-
rib at a 50—70° angle, + straight, eventually weakly
arcuate to the margins, sunken above, convex be-
low; interprimary veins weakly raised, darker than
surface below; minor veins numerous, sparsely vis-
ible, darker than surface below, arising from both
the midrib and primary lateral veins. INFLORES-
CENCES erect, 2 per axil; peduncle 9-22 cm long,
(1)9-12 mm diam., medium green, moderately
spongy; spathe + erect, 8.8-19.8 ст long, (0.5—
1.4 times longer than peduncle), prominently con-
stricted at anthesis; spathe blade yellowish green
to green, glossy outside, 7 cm long, 1.7 cm diam.,
whitish green, minutely white-dotted, sometimes or-
ange-striate inside; spathe tube ellipsoid, green
outside, 8.5 cm long, 2.5 cm diam., light red to
maroon (B & K red-purple 2/10) inside; spadix
stipitate 6-15 mm long; gradually to bluntly ta-
pered to apex, 9.3-16.9 cm long; pistillate portion
pale green to greenish yellow, 3.3-5.2 ст long in
front, 2.2-2.5 ст long in back, 1.4 cm diam. at
apex, 1.3-1.6 cm diam. at middle, 1.6 cm wide at
base; staminate portion 6.5-11.3 ст long; fertile
staminate portion creamy white, 9-12 mm diam. at
base, 1-1.2 cm diam. at middle, 5-7 mm diam. ca.
1 ст from apex, broadest at the middle, narrower
than the pistillate and sterile portions; sterile sta-
minate portion broader than the pistillate portion,
creamy white, (9)1.1-1.5 cm diam.; pistils 1.8-2.3
mm long, 1.1 mm diam., creamy white; ovary 6-8-
locular, 0.9-1.2 mm long, 1 mm diam., usually with
basal, somtimes sub-basal placentation; locules
0.9-1.1 mm long, 0.3-0.4 mm diam.; ovule sac
0.9(1.2) mm long; ovules 1 per locule, contained
within transparent ovule sac, 0.3-0.4 mm long,
about as long as funicle; funicle 0.3-0.4 mm long
(can be pulled free to base), style 0.6-0.8 mm long,
1.1 mm diam., similar to style type B; style apex
= sloping; stigma subdiscoid, greenish yellow, 1
mm diam., 0.3 mm high, covering entire style apex;
the androecium irregularly 5—6-sided, sometimes
oblong or quadrangular, 1.6–1.9 mm long, 0.9-1.2
mm diam. at apex; thecae oblong, 0.4–0.5 mm
wide, + parallel to one another and nearly contig-
uous; sterile staminate flowers blunt, irregularly 4-
6-sided, sometimes prismatic, (1)1.6-2 mm long,
(1.2)1.7-1.8 mm wide. Berries white or sometimes
yellowish; seeds 1 per locule, tan, oblong, 2 mm
long, 0.8 mm diam.
Flowering in Philodendron smithii occurs prin-
cipally in the dry season, and flowers have been
collected from December to April (but also July).
Post-anthesis collections are common from Febru-
ary through July but with a few in October, Novem-
ber, and December. The latter, especially those in
October and November, are the most aberrant,
since even if this species is cued to flower by the
onset of the dry season (which appears to be the
case), they would appear to have flowered too early.
This may be explained by the fact that the dry sea-
son starts much earlier in Mexico and Guatemala
than in Costa Rica and Panama.
Philodendron smithii ranges from Mexico (mostly
in Chiapas, Tabasco, and Quintana Roo) to Gua-
temala, Honduras, and Nicaragua on the Atlantic
slope at 40 to 1630 (mostly less than 600) m ele-
vation. It is known from “Selva Alta Perennifolia”
and “Selva Mediana Subperennifolia” in Mexico as
well as from Tropical moist forest and Subtropical
wet forest in Guatemala.
Philodendron smithii is a member of P. sect. Ca-
lostigma subsect. Glossophyllum ser. Ovata. This
species is characterized by its conspicuously fis-
sured tan internodes, which are about as long as to
longer than broad; sharply two-ribbed, deciduous
cataphylls; spongy subterete petioles; usually
blackish, narrowly ovate-cordate leaf blades; long-
pedunculate inflorescences; and externally green
spathes that are maroon inside the tube.
Philodendron smithii is not easily confused with
any other species in Central America. It is most
similar to P. panamense, which is endemic to Pan-
ama, and shares with that species similarly dark-
drying ovate-triangular, long-petiolate blades and
long-pedunculate inflorescences. Philodendron
panamense differs in having much shorter inter
nodes (shorter than broad on flowering plants).
weakly one-ribbed cataphylls, moderately firm pet-
ioles, and spathes green on the tube outside with a
white blade and greenish white throughout within
(vs. green throughout outside and maroon on the
tube within for P. зтићи).
Philodendron smithii is also similar to P. stra"
minicaule and P. wilburii, both from Costa Rica and
A И س AAA (q € Station El Recreo, 30 m, 12?10'N, 84°18’ W, Dav-
idse et al. 30768 (MO); Río Prinzapolka, ca. 2 km S of
Wanî, ca. 0-100 m, 13°42' №, 84^50' W, Pipoly 4733 (MO),
4738 ; (MO), 4747 pd Río Punta Gorda, Atlanta, 10 m,
11°34' №, 84?01'W, Moreno & Sandino T S me =
pam-Puerto Cabezas Río Troncera,
14°43'N, 84^06' W, Pipoly 4042 yee Río Mila сие
Wilike region, 100 т, 13%05'N
MO). PANAMA. Bocas del паши
of cial Divide, 700 m,
t & Grayum 60281 (MO); Chan-
guinola—Almir, e Mi lla 7.5, Croat & Porter 16241 (MO);
Valle del Silencio, Tp Changuinola, ca. m above
mouth of Río Teribe, <100 m, 9721'40"N, 82°31'40"W,
Croat & Zhu 76431 (МО). Canal Area: Frijoles, 25-30
m, Pittier 3755 (US); Barro Colorado Island, Shattuck 182
, MO); Ebinger 99 (MO); Croat 11838 (MO);
Schmalzel 797 (MO); Dwyer et al. 8474 (MO); Croat 16535
(MO); 15067 (MO); 11006 (MO); 10741 (MO, UC); 10198
(MO, SCZ); 10085 (MO); 6214 (MO); 6294 (MO); 8605
(MO); 8040 (MO); 6861 (MO); 6495 (MO); 4001 (MO);
Summit Garden, Croat 17058 вани 10284 (МО, =
~
^
33015 (MO); 10915 (MO). Chiriquí: Volcán-Serrano, 7
mi. N of Volcán, 8°50’N, sip W. Cro менү (МО); уїс.
El Hato del Volcán at Las Lagunas, 3 mi. from Nueva
«fi
California, 1380 m, 8°46’N, 82 3°40" ^W. "ian 67707 (MO,
yo
40'N, 82? 13' W, Croat 67739 (CM,
MO); 2.8 mi. e Los Planes, 1200 m, 48817 (MO);
Quebrada La Mina, 2.3 mi. N of Los Planes, 800 m,
8°41'N, 82°13'W, Croat & Zhu 76364 (CM, MO); along
Río Colorado, 1200-1400 m, 82%43'W, 8*50'N, Hamilton
& Krager 3757 (MO, RSA); vic. Boquete, са. 1450 m,
Croat 26744 (MO). Coclé: La Mesa, N of El Valle de
Antón, 800-900 m, 8?38'N, 80°09’W, Croat 67210 (CM,
MO); Croat 13358 (MO); ca. 1 mi. N of El Valle, Gentry
& Dwyer 3591 (MO); 900 m, Gentry 7433 (F, GH, MO).
A trail beyond La Mesa towards Los Llano
4066 (MO). Colón: Santa Rita Ridge, 26 km
sisthmian Hi 9°26'N, 79*57' W, Knapp et al.
9°25'N, 79°40'W,
of >
803.5" N, 7717 W, Ha u “г ed 16293 (МО); western
slopes of Сен Рите, 17 km N of El Real, Его Регіѕепісо,
100 m, 801'N, 77°40W, Croat & Zhu 77187 (МО); ме.
Cana, 500-600 m, Croat 37641 (MO); Alturas de Nique
пало. 900-1250 m, 7°45’N, 77°40'W, McPherson 11603
(MO); Cerro Sapo, 800 ft., Hammel 1330 (MO); E base of
Cerro Sapo along stream (Río Celorio?), 1300 ft., 1131
(MO); 18 km SE of Jaque, 25 m, Garwood et al. 98 (BM,
CM, MO, PMA); N of Punta Guayabo Grande, NW of En-
senada El Guayabo ridges, 0-200 m, 7'24'М, 78% o d
Knapp & Mallet 3160 (MO, US); Río Coasi, Cerro Coa
Duke 15554 (US); Río Cocalito, SE of Jacque, 7187,
77°58'W, Whitefoord & Eddy 223 (BM, MEXU, МО).
Гем: : Juan Díaz, Standley 30618 (US); Cerro Jefe re-
п, 750-800 m, 9°14’N, 79°22’ W, Croat 67082 (F, MO):
67084 (MO); 4.6 km beyond peak on road to Altos de
Pacora, са. 600 m, Croat 35900 (MO); at Altos de Расога,
750 m, 9°15'N, 7929'W, Croat & Zhu 76606 (MO); 3-
3.5 mi. NE of Altos de Pacora, 700-750 та, 9°15'N,
79°25'W, Croat 68636 (CM, MO); Campo Tres, 5 km NE
of Altos de Pacora, Busey 835 (MO); Lago Cerro Azul,
500 m, Sullivan 82 (MO); Cerro Campana, 2500 ft., Ham-
mel 3767 (MO, SEL); Río Majé-Quebrada Brava, 60 m.
Croat 34630 (MO); El Llano-Cartí Road, 7-12 km from
Volume 84, Number 3
1997
Croat 546
Philodendron Subgenus Philodendron
rege r 360-400 m, Croat 25122 (MO);
a. 16-18, 4
78°50! W, Thompson 4704 (CM, MO); Serranía de Majé,
along Río Ipetí, near confluence with Río Ambrosio, 100
m, 8°57'N, 79°32'W, Churchill & de Nevers 4480 (MO,
PMA, RSA). San Blas: Nusagandi, along El Llano—Carti
oad, 11.6 mi. N of highway, 450 m, 9?18'N, 79°59’ W,
Croa 75135 (СМ, МО, PMA). Veraguas: Santa Fe—Río
uis, vic. Río Tercero Brazo, 700 m, 833'N,
81°08" W, Croat 66925 (MO, PMA); Rio Dos Bocas, ca. 5—
8 km N of Alto Piedra, 730-770 m, Croat 25915 (MO);
ca. 500 m, Croat 34242 (MO); vic. Escuela Agricola—Ca-
00 m, 81%07'W, 8°32'N, Hamilton et al.
127
Philodendron chis Croat, sp. nov. TYPE: Pan-
~
MO -S6146T^ isotypes, AAU, B, CAS, CM
CR, DUKE, F, GH, K, M, MEXU, NY, P, PMA,
RSA, US, VEN, W). Figures 29, 425—430
Planta plerumque hemiepiphytica, interdum terrestris;
internodia 2-6 cm longa, 3.54 cm diam.; cataphylla
(10)21-33 em longa, incostata vel leniter 1-costata, saepe
rubella, persistentia epit sat emum decidua; petiol-
us teres vel subter m longus, 8-15 mm diam.;
lamina ovata, coriacea, ыйа. 32-70 cm longa, 24—50 ст
i.
3-
8(11) mm diam.; spatha 11.5-19 cm longa, lamina spathae
extus viridi, s ii usa marronina aut rubella, intus alba vel
oe тн tubo spathae extus suffuso marronino aut
По, intus atrimarronina suffuso in laminam; pistilla
(96-809). нта loculi (4)5—7-ovulati.
Usually hemiepiphytic, appressed-climber,
sometimes terrestrial; internodes short, stout, se-
miglossy, smooth, 2-6 cm long, 3. cm diam.,
dark green; roots elongate, fine, branched; cata-
n (10)21-33 cm long, unribbed to weakly
-ribbed, sometimes sharply 2-ribbed near base
is unribbed near apex, frequently reddish, fre-
quently striate, emarginate at apex, persisting semi-
intact at upper nodes, eventually deciduous, emar-
ginate at apex; petioles 39-90 cm long, 8-15 mm
diam., terete to subterete, surface densely pale-stri-
ate, glossy; sheath 10.5-11 cm long; blades ovate,
coriaceous to moderately coriaceous, glossy to se-
miglossy, conspicuously bicolorous, acuminate at
apex (the acumen sometimes inrolled and —
cordate at base, 32-70 cm long, 24-50 cm wide
(1-1.8 times longer than wide and averaging 1.4
times), (0.4—1.1 times longer than petiole), aver-
aging about % as long as petiole, upper surface
dark green, drying dark brown to blackish Bo
lower surface slightly paler, drying dark yellow-
rown to greenish brown; anterior lobe 25—52.5 cm
long, 20-50 cm wide (2-4.8 times longer than than
posterior lobes); posterior lobes broadly rounded to
broadly obtuse, 8-20 cm long, 11-22 cm wide; si-
nus parabolic, sometimes spatulate, (3)4-12 cm
deep; midrib broadly convex to almost flat, con-
spicuously paler than surface above, convex to
luntly acute, darker than surface below; basal
veins 5—9 per side, with (0)1(2) free to base, num-
bers 3—4 coalesced 1.5—3 cm; posterior rib usually
scarcely or not at all naked, rarely naked for 3 cm;
primary lateral veins 4—10 per side, departing mid-
rib at a 60—70° angle, moderately straight to the
margins, narrowly sunken above, convex and some-
what paler than surface below; interprimary veins
sunken, usually concolorous above, raised, darker
than surface below; minor veins moderately distinct
below, arising from both the midrib and primary
lateral veins. INFLORESCENCES 4–5 per axil; pe-
duncle 3-12 cm long, 3-8(11) mm diam., green,
finely striate; spathe 11.5-19 cm long (1.
times longer than peduncle); spathe blade green to
white, lightly to heavily tinged maroon or reddish,
finely striate-speckled, margins clear outside, white
to pale green inside; spathe tube tinged maroon or
reddish, finely striate-speckled outside, 4.5-10 cm
long, dark maroon, suffused into blade inside,
weakly constricted above the tube; spadix white
throughout, 10.7-15.6 cm long; pu portion
3.4—5.2 cm long in front, 2.4—3.1 cm long in back,
broadest midway and gradually t toward both
ends, 1.2 cm diam. at apex, 1.2-1.4 cm diam. at
middle; staminate portion 7.7—9.4 cm long; fertile
staminate portion white, barely constricted above
the sterile staminate portion, barely tapered toward
apex, 1-1.2 cm diam. at base, 1-1.2 cm diam. at
middle, 6—7 mm diam. ca. 1 cm from apex; sterile
staminate portion 9-10 mm diam.; pistils 1.5-2.7
mm long, 0.8-1.6 mm diam., white; ovary (5)6—
8(9)-locular, with sub-basal placentation; locules
(0.8)1.1-1.3 mm long, 0.3-0.6 mm diam.; ovules
(4)5—7 per locule, (0.1)0.3—0.4 mm long, 2-seriate,
contained within transparent ovule sac; style simi-
lar to style type B; stigma 0.8-1.2 mm diam., 0.2—
0.3 mm high. INFRUCTESCENCE with pistillate
spadix 6.5 cm long, 4 cm diam.
Flowering in Philodendron tysonii occurs mostly in
June and July but also in September. In reality, flow-
ering probably occurs from January through Septem-
ber or perhaps October, since post-anthesis collec-
tions have been made nearly throughout the year
except February. Immature fruiting collections have
been made in March, September, and November.
Philodendron tysonii is known only from Pana-
ma, but is expected to be found in the Cordillera
550
Annals of the
Missouri Botanical Garden
de Talamanca of Costa Rica. It ranges from 600 to
1500 m elevation in Premontane rain forest and
Tropical Lower Montane rain forest life zones.
Philodendron tysonii is a member of P. sect. Ca-
lostigma subsect. Macrobelium ser. Reticulata. This
species is characterized by из stout, short (usually
3-5 cm diam.) internodes; unribbed or weakly one-
ribbed, frequently reddish cataphylls persisting
semi-intact (often becoming fibrous) at the uppe
nodes; terete to subterete petioles; апа
ovate-cordate blades averaging 1.4 times longer
than wide and about three-fourths as long as the
petioles, with 6–9 basal veins and a weakly devel-
oped posterior rib, which is rarely naked at the
~“
8.
Philodendron tysonii is perhaps most easily con-
fused with P. antonioanum, with which it may oc-
cur. The latter species differs in having sharply
2-ribbed cataphylls, blades drying gray-green with
aquia ‘cross-veins” and 4 to 6 pairs of pri-
lateral veins, and short-pedunculate inflores-
cences with spathe tubes dark magenta on both sur-
faces and barely constricted above. In contrast, P.
tysonii has usually bluntly к ае.
blades drying + blackened and s h above, 6–
10 primary lateral veins, more айыы редип-
culate inflorescences, and prominently constricted
spathes merely tinged red on the tube outside. In
addition, the two species differ in the nature of their
ovules, with P. tysonii having (5)6—8(9)-locular ova-
ries with (4)5-7 ovules per locule borne in an en-
velope with sub-basal placentation, while P. anto-
nioanum has 5-locular ovaries with about 30 ovules
per locule, with parietal placentation and not borne
in an envelope.
This species may also be confused with P. sag-
utifolium (especially broader-leaved forms), which
shares blades with mostly free basal veins with pos-
terior ribs short or lacking and scarcely or not at
all naked along the sinus. The latter species differs
in having narrower stems (rarely more than 3 cm
diam.), green, sharply 2-ribbed deciduous cata-
phylls (vs. persistent in P. tysonii), and blades usu-
ally somewhat reddish brown and proportionately
longer (averaging about 2 times longer than broad)
with the posterior lobes usually narrowly rounded.
In addition, the juvenile blades of P. sagittifolium
are elongated, usually oblong to narrowly triangu-
lar, and usually up to 2.5 times longer than wide
(vs. ovate to narrowly ovate, less than 2.5 times
longer than wide, and with broadly rounded pos-
terior lobes in P. tysonii).
Additional repa сое РАМАМА. Вос
del Того: Fort Dam area, Gualaca—Chiriquí "тона
just N of the и ан Divide. 850-950 m, 8°46'N,
82°17'W, Hammel et al. 14700 (MO); ca. 8.5 mi. from
bridge over Fortuna Lake, 600 m, 8°46'N, 82?16' W, Croat
67005 (CAS, CM, L, MO, PMA, TEX); Cerro Colorado,
9.2 mi. W of Chame, 1450-1480 m, 8?35'N, 81°50'W,
Croat 69017 (DUKE, IBE, MEXU, MO, NY, PMA, QCA,
RSA, WIS); 7 mi. from Chame, ca. а. 18785
81°45! W, McPherson 8810 (MO); 5.3 mi. N Py center uf
Fortuna Dam, Continental Divide Trail, £ 844", 82°17'W,
Croat & Zhu 76346 (MO); near Lago Fortuna, along trail
to Río Hornito, 8°45’N, 82?18'W, Croat & Zhu 76374
(MO, PMA). Chiriqui: Cerro Colorado, above San Félix,
7 mi. N of Pan-American Highway, 1200-1500 m,
Croat 33148 (MO); 24 mi. N of Río San Félix, 1430-1500
m, 48486 (MO); 34.1 km N of Río San Félix, 1390 m,
37258 (МО); 28 mi. above Río San Félix, 1200-1500 m,
33269 (MO); from Chame to ca. 9 mi. along road, 1100—
1750 m, 8°35'N, 81%54'W, Hammel & Trainer 15008
(MO, NY); Fortuna Dam area, Fortuna-Chiriquí Grande,
Continental Divide, 1170 m, 8?44'N, 81717'М, Croat
66711 (B, BR, COL, CR, K, MO, NY, PMA, US); Hammel
& Kress 13490 (DUKE); Continental Divide trail, 8°44' №,
82°17'W, Croat & Zhu 76330 (K, MO, NY, SCZ, US); trail
o Río Hornito, 1100-1350 m, 8°45’N, 82715", Mc-
Pherson 11660 (MO); Croat & Zhu 76417 (AAU, CM,
ENCB, GB, IBE, M, MO, NY, PMA, TEX, WIS); conflu-
ence of Río Hornito and Río Chiriquí, 1050-1100 m, ca.
8°44'N, 82713", Stevens 18376 (MO); 18377 (MO); de
IRHE facilities, 1100-1200 m, 8?45'N, 82*08'W, Cro
66589 (AAU, COL, DUKE, F, K, MBM, МЕХ. MO, Р,
PMA, US, VDB); 1300 т, Croat А00 (СМ, МО); Que-
brada Los Chorros—Quebrada Fra of reservoir, 1100
m, 8°45’ N, 82°13’ W, Churchill & Churchill 6081 (MO);
83 m i. beyond Los Planes de Hornito, 1260 т, 8°44’N,
82°16! W, Croat 49943 (MO); 1300 m, '845'N, 82°17’ W,
49821 (MO); 1400 m, 48704 (AAU, MEXU, MO, U, UC,
W). Coclé: El Copé region, Alto Calvario, Continental
850 m, 67577 (AAU, CM, F, G, L, MEXU, MO, NY, PMA,
TEX, о 930 т, 49203 (МО); 650-850 m, Folsom 6217
(MO). Panamá: Cerro Jefe region, vic. of summit, 9°15'N,
79°30'W, Croat & Zhu 76210 (CM, MO); Witherspoon &
Witherspoon 8531 (MO); ca. 1000 m, Croat 49084 (CAS,
MA); 1000 m, 9%15'N, 79%30'W, Knapp 2240
(MO); La Eneida, 1000 m, Luteyn & Kennedy 1775
(DUKE); 0.8 mi. beyond turnoff to Altos de Pacora, 770
m, 9?15'N, 79°29'W, Croat & Zhu 76614 (CM, МО, WIS);
3-3.5 mi. NE of Altos de Pacora, 700-750 m, 915",
79°25'W, Croat 68688 (AAU, CM, Е, C, MEXU, MO,
PMA, TEX, US); 800-1000 m, 22671 (F, MO); 2400 ft.,
Antonio 4753 (MO); Croat 67059 (AAU, B, BR, C, CAS,
M, COL, CR, DUKE, EAP, ENCB, F, HNMN, НЏА,
IBE, JAUM, JBGP, K, L, M, MO, NY, PMA, QCA, US,
USCG, WIS); жое: m, 9°15'N, 79°30’W, McDonagh
et al. 36 (BM m, Mori et al. 3737 (МО); са.
950 m, ca. "o N. = W. McPherson 7109 (MO); Cer-
efe, ca. 1000 m, Luteyn 3209 (DUKE);
mi. from highway, 350 m, Croat 49120 X Veraguas:
Santa Fe region, Alto Piedra-Calorébora, 3: 5600 ft..
Hammel 4604 (MO); Río Tercero Brazo, beyond Alto Pied-
ra, Croat 27328 (MO); between Río Tercero Brazo and
summit of hill leading down to dis Atlantic lowlands, 700
m, 8°33'N, 81*08'W, Croat 66926 (B, CAS, F, K, MO,
PMA, US); Parque Nacional Cerro Tute, near Alto Piedra,
ا M — س ы. ес — — — — —————— ا
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
0.5 mi. N of Alto Piedra, 1250-1350 m, Croat 48955 (F,
MO); 800-1030 m, Croat & Zhu 76911 (CM, MO).
Philodendron ubigantupense Croat, sp. nov.
TYPE: Panama. San Blas: on mainland oppo-
site Isla Miria Ubigantupo, along trail to Di-
gole, to 20 m, 9°26'N, 78°54'W, 19 July 1987,
H. Herrera 294 сае MO-3585503; iso-
type, PMA). Figure 436.
Planta hemiepiphytica; internodia usque to 7.5 cm lon-
ga, 1 ст diam.; catap ylla decidua; petiolus subteres, 12—
с in sicco m diam.; lamina oblonga,
anguste ae hone 27-32 cm longa, 8.5-9 cm lata,
idis; nervis lateralibus I grece inflores-
centia 1; sp А 8 ст longus, in sicco 5 mm diam.;
spatha - 3 cm longa, viridis; pistilla (7)8-locularia; кой
1-ovulat
Hemiepiphytic; stem scandent; internodes to 7.5
cm long, 1 cm diam., slender, drying 7 mm diam.,
gray-brown, irregularly and closely fissured, some-
what flaking; roots several per node, drying mod-
erately smooth, reddish brown, semiglossy; cata-
phylls lanceolate, deciduous; petioles 12-13 cm
long, drying 5-6 mm diam., subterete, drying dark
gray-brown, minutely striate, sheathed to 1.5 cm;
blades oblong, acuminate at apex (the acumen
tightly inrolled, 5 mm long), narrowly cordulate at
base, 27-32 cm long, 8.5-9 cm wide (3-3.8 times
longer than wide), (2.2-2.5 times longer than pet-
iole), about as long as petiole; upper surface dark
green, drying weakly glossy, gray-green, lower sur-
face slightly paler, semiglossy, drying greenish gray;
sinus to 1.3 cm deep; midrib broadly convex above,
convex and paler below, drying faintly striate; pri-
mary lateral veins almost totally obscure above an
below; many minor veins, all free to base, arising
from the midrib only, drying obscure above, close
and weakly raised below, spreading at a 55-75"
angle, + straight to the margin. INFLORES-
CENCES solitary; peduncle 8 cm long, drying 5
mm diam.; spathe green, 9.3 cm long, (1.2 times
longer than peduncle); spathe tube weakly con-
stricted, ca. 4.5 cm long; spadix bluntly pointed at
apex, 10.8 cm long; pistillate portion 4 cm long in
front, 3.7 cm long in back, 8 mm diam. at apex,
drying 1 cm diam. at middle; staminate portion 8.1
cm long; fertile staminate portion 8.5 mm diam. at
middle, 6.5 mm diam. ca. 1 cm from apex; sterile
staminate portion 8 mm diam. at apex, 9 mm diam.
at base; pistils 1.5 mm long; ovary (7)8-locular, 1.2
mm diam., with sub-basal placentation; locules 1.1
mm long, 0.2-0.3 mm diam.; ovule sac ca. 1 mm
long; ovules 1 per locule, contained within trans-
parent ovule sac, ca. 0.3 mm long; funicle ca. 0.3
mm long (can be pulled free to base), style 0.9 mm
diam., similar to style type D; style apex sloping to
weakly rounded; style boss moderately broad and
fairly pronounced; the androecium truncate, some-
times prismatic and weakly clavate, margins irreg-
ularly 4—5-sided, 0.8-1.5 mm long; thecae oblong,
0.2-0.3 mm wide, + parallel to one another; sterile
staminate flowers clavate to prismatic, irregularly
4—5-sided, 1.3-1.7 mm long.
The flowering phenology of Philodendron ubi-
gantupense is uncertain owing to its rarity, but since
it has small inflorescences and was in flower in
July, it probably both flowers and fruits in the wet
season.
Philodendron ubigantupense is endemic to Pan-
ama, known only from the type locality along the
Atlantic coast, at or near sea level in a Premontane
wet forest life zone.
Philodendron ubigantupense is a member of Р.
sect. Calostigma subsect. Glossophyllum ser. Glos-
sophyllum. This species is characterized by its
scandent habit, long, slender internodes, subterete
petioles and especially by its oblong, gray-green
drying blades over twice as long as the petioles,
weakly cordulate leaf bases, and almost totally ob-
scure primary lateral veins.
Philodendron ubigantupense is perhaps most
similar to P. folsomii, which has blades of similar
size and color. The latter species differs in having
short internodes and blades merely rounded at the
base and with 8—10 distinct primary lateral veins.
In addition, P. folsomii has 6—7 locules per ov
that might be confused with P. ubigantupense, but
it differs in having proportionately broader blades
(to 2.8 times longer than broad) with up to five,
obvious primary lateral veins.
Philodendron utleyanum, Croat, sp. nov. TYPE:
Panama. Colón: Santa Rita Ridge, about 7 mi.
from Transisthmian Highway, ca. 200 m, 21
Dec. 1971, Wilbur et al. 15068 (layne,
DUKE 226389). Figures 431, 432.
Planta ES internodia 1—3.5 cm longa, 1.5—
2 cm diam., in sicco pallide — cataphylla usque
27 cm longa, аы 5 -costata, sicco obscure rubri-
brunnea, decidua; petiolus atia atribrunneus, obtuse
subcomplanatus adaxialiter versus apicem, 19— cm
longus, in sicco 6-7 cm diam.; lamina oblongo- óblincéo-
lata, 32-39 ст longa, 10-14 cm lata, aliquantum inequi-
laterala et rotundata vel truncata aut leniter cordulata
pedunculus usque 9 cm longus, in sicco atribrunneus;
spatha usque 11 cm longa, pallide viridis, extus suffusa
Annals of the
Missouri Botanical Garden
subrosea, intus alba, tubo us 5 cm longa; pistilla 4—
5-locularia; loculi 1-ovulat
Hemiepiphytic; stem appressed-climbing; inter-
nodes drying moderately glossy, irregularly and
acutely ribbed with a few flat smooth intervening
areas, 1-3.5 cm long, 1.5-2 cm diam., drying light
brown; roots several per node, short, drying brown,
to 3 cm diam.; cataphylls to 27 cm long, weakly
2-ribbed, but sharply 2-ribbed toward the apex, red
tinged toward the base, drying dull reddish brown,
deciduous; petioles 19-21.5 cm long, drying 6-7
cm diam., subterete, dark brown, obtusely some-
what flattened toward the apex adaxially, surface
smooth to weakly ribbed; blades somewhat inequi-
lateral, oblong-oblanceolate, subcoriaceous, semi-
glossy, acuminate at apex, somewhat inequilateral
and rounded to truncate to weakly cordulate at
base, 32-39 cm long, 10-14 cm wide, broadest
above the middle, 1.3-1.8 times longer than peti-
oles, upper surface dark green, drying gray-brown,
lower surface often somewhat reddish, drying me-
dium brown; posterior lobes to ca. 2 cm long;
midrib drying weakly and obtusely raised, slightly
paler than surface above, drying convex, drying
light brown with a distinct pale border along its
edges and paler than surface below; basal veins 2—
3 per side, and with all free to base; primary lateral
veins 8-9 per side, departing midrib at а 65-80”
angle in the lower one-half, to 45° angle toward
apex, weakly arcuate to the margins, drying incon-
spicuous except near the midrib, paler than surface
above, weakly convex and paler than surface, some-
times undulate below; minor veins drying moder-
ately inconspicuous and close, weakly undulate,
arising from the midrib only, the surface and minor
veins minutely and densely granular at 10X mag-
nification. INFLORESCENCES 1 per axil; pedun-
cle to 9 cm long, drying 4.5 mm diam., dark brown;
spathe moderately coriaceous, to 11 cm long, ca.
2 cm diam., pale green with pinkish tinge, drying
dark brown throughout, white within; spathe tube 5
cm long, 1.8-2 cm diam.; spadix sessile, 10 cm
long.; ne portion 5 cm long, drying to 1.7 cm
ase, 1.5 cm diam. midway, 1 cm at
apex; siaminate portion 4.7 cm long, too deterio-
rated for detailed studies; pistils 2.5-3 mm long,
1.5-2 mm diam., drying light brown, smooth except
warty near apex; style type not studied; stigma 0.4—
0.6 m diam., button-like on drying with a medial
nipple and 4–5 radiate arms, sometimes with frag-
ments of the dried stigma apron-like still attached;
locules 4—5; ovaries with sub-basal placentation, 1
ovule per locule.
Flowering in Philodendron utleyanum is poorly
known. The single collection with immature fruits
was made in December. This species probably flow-
ers in the rainy season.
Philodendron utleyanum is endemic to Panama,
known only by the type specimen from Colón Prov-
ince in a region of Premontane wet forest at 200 m
elevation.
Philodendron utleyanum is a member of P. sect.
Calostigma subsect. Glossophyllum ser. Glossophyl-
lum. This species is distinguished by its short tan
internodes; sharply 2-ribbed, deciduous cataphylls;
subterete petioles about three-fourths as long as the
blades; and oblong-oblanceolate, brown-drying
blades broadest above the middle and rounded to
weakly subcordate at the base
In terms of coloration and texture, P. utleyanum
appears closest to P. sagittifolium, but in terms of
blade shape it is closest to P. pseudauriculatum.
The former differs in having the blades broadest at
the base and decidedly cordate, the latter in having
dark gray-green-drying leaves, more or less spongy,
proportionately shorter petioles, and the spadix
clearly demarcated from the peduncles by a marked
color contrast. Philodendron utleyanum lacks any
clear limit between the peduncle and the spathe.
This species is named in honor of John Utley, a
participant in the Duke University expedition that
collected the type. Utley has been responsible for
collecting many interesting and important Araceae
during his trips to Central America and during his
tenure as a Peace Corps volunteer in Costa Rica.
Philodendron verapazense Croat, sp. nov.
TYPE: Guatemala. Alta Verapaz: 7 mi. on road
to Oxec, along road turning N off Hwy. 7E be-
tween Tucurú and El Estor, ca. 6 km NE of
Panzós, 700 m, ca. 15°28'N, 89°04’W, Croat
41656 (holotype, MO-2743518; isotypes,
GUAT, US). Figures 433-435.
Planta hemiepiphytica; internodia longiora quam lata,
conspicue multisulcata; cataphylla 14—17 cm longa, acute
2-costata, decidua; petiolus acute D-formatus complanatus
adaxialiter, 29-44 longus, in sicco 2-5 mm diam.; lamina
longe cordata vel sagittata basi, 26-34 cm longa, 11. 5-14
em lata; nervis lateralibus 1 3-4 utroque; inflorescentia 1;
pedunculus 3-14 ст longus, 4-5 mm diam.; spatha 10-
18 cm longa, extus rubriviolacea; tubo spathae intus саг-
mesino; pistilla 4—6-loculari; loculi 2-ovulati.
Hemiepiphytic; stem appressed-climbing or
sometimes scandent; pre-adult internodes to 20 cm
long, 1.5-3 cm long; adult internodes green becom-
ing tinged purple, finally tan, conspicuously
many-sulcate except above each petiole, longer
than broad; roots few per node, 10-30 cm long,
Volume 84, Number 3
1997
Croat 553
Philodendron Subgenus Philodendron
drying dark brown, weakly glossy, 2.3 mm diam.;
cataphylls 14—17 cm long, sharply 2-ribbed, green
tinged red, deciduous; petioles 29-44 cm long,
drying 2-5 mm diam., sharply D-shaped, flattened
adaxially, with adaxial margins sharp; blades nar-
rowly ovate-triangular-cordate, subcoriaceous, long-
acuminate at apex (the acumen sometimes apicu-
late), long-cordate to sagittate at base, 34 cm
long, 11.5-14 cm wide (2-3 times longer than
wide), (0.7—1.1 times longer than petiole); anterior
lobe (13)21-26 cm long, (5.4)13-14 cm wide (2.9-
4.3 times longer than posterior lobes); posterior
lobes 6-7.5 cm long, (2.3)6-7.1 cm wide; sinus
hippocrepiform, sometimes spathulate, 7-7.5 cm
deep; midrib broadly convex and paler above, con-
vex and darker below, drying somewhat acute; basal
veins 3(4) per side, with 0(1) free to base, the re-
mainder coalesced 2-2.5 cm; posterior rib naked
for most or all of its length; primary lateral veins
34 per side, departing midrib at a 55-65” angle,
+ straight to weakly arcuate to the margins; minor
veins arising from the midrib only. INFLORES-
CENCES 1 per axil; peduncle 3-14 ст long, 4-5
mm diam., green or tinged with reddish violet;
spathe 10-18 cm long ((0.7)2-3.7 times longer
than peduncle), reddish violet outside; spathe blade
acuminate; spathe tube cylindrical, 8-9 cm long,
1-1.5 cm diam., crimson-red inside; spadix stipi-
tate to 3.5 cm long; acute at apex, 15—16.5 cm long,
constricted ca. 4 cm above base of sterile staminate
portion; pistillate portion cream, 3.2 cm long in
front, 1.7 cm long in back, 1.5-2.6 cm wide at
base; staminate portion 8.3-9 cm long; fertile sta-
minate portion cream, tapered to apex, 7-10 mm
diam. at base and midway, 3—4 mm diam. ca. 1 cm
from apex; sterile staminate portion 1.8-2.2 mm
diam.; pistils 1.9-2.3 mm long; ovary 4—6-locular,
l mm diam., with sub-basal placentation; locules
0.9 mm long, ca. 0.3 mm diam.; ovule sac 0.4-1
mm long; ovules 2 per locule, contained within
transparent, gelatinous ovule sac, longer than fu-
nicle; funicle 0.2-0.3 mm long (can be pulled free
to base), style 0.8 mm diam., similar to style type
D; style apex somewhat rounded to flat, depressions
surrounding stylar canal exits, style boss broad,
very shallow; the androecium truncate, prismatic,
= oblong, margins irregularly 5—6-sided, 0.8-1.1
mm long; thecae oblong, 0.4—0.5 mm wide, + раг-
allel to one another, very nearly contiguous; sterile
staminate flowers prismatic to weakly clavate, ir-
regularly 4—6-sided, mostly 5-sided, 1.3-1.9 mm
ong.
Flowering in Philodendron verapazense occurs in
April and May, with a single, post-anthesis collec-
tion known from J y.
Philodendron verapazense ranges from southern
Mexico (Chiapas) to Guatemala (Alta Verapaz), at
700 to 1525 m elevation in *Bosque Pino-Encino"
in Mexico and Tropical moist forest in Guatemala.
Philodendron verapazense is a member of P. sect.
Calostigma subsect. Macrobelium ser. Macrobel-
ium. This species is recognized by its scandent
habit with the internodes longer than broad and
conspicuously pluri-sulcate, as well as by its sharp-
y two-ribbed, deciduous cataphylls, sharply D-
shaped petioles, narrowly ovate-triangular-cordate
blades, solitary inflorescences, red to reddish violet
spathe tubes, and greenish spathe blades.
Philodendron verapazense is probably most sim-
ilar to P. sagittifolium especially in the shape and
coloration of the blades, the sharply two-ribbed, de-
ciduous cataphylls, and the D-shaped petioles. It
differs from the latter species principally in its
deeply sulcate stems, sh D-shaped, shorter
(less than 50 ст long) petioles, and leaf blades with
3—4 pairs of primary lateral veins (vs. subterete,
— petioles, which are greater than 50 cm long,
4—8 pairs of primary lateral veins for P. sag-
ittfolium).
—
Additional specimens examined. GUATEMALA. Alta
Verapaz: near Chirriacté, on the Petén Highway, ca. 900
m, Standley 91978 (F). Izabal: vic. EXMIBAL camp 2,
W of Lake Izabal, 400—500 m, Jones & Facey 3354
hi Los Lagos, 3 mi. NW of Rancho
San José, 5000 ft., Carlson 1846 (Е).
Philodendron verrucosum L. Mathieu ex Schott,
Syn. Aroid. 85. 1856. TYPE: Locality un-
known (holotype, W? lost?). Schott ic. 2757—
2759 (neotype, here designated). Figures 437—
440, 445.
Philodendron daguense Linden € André, Ill. Hort. 18:
cde t 79. 1871. TYPE: Colombia. Valle: Río Dagua,
is s.n. (holotype, K
Philodendron pilatonense Engl., Bot. Jahrb. Syst. 37: 129.
5. TYPE: Ecuador. Pichincha: Río Pilatón, May
pem Sodiro s.n. (holotype, B; isotypes, у МО).
Philodendron discolor K. Krause, Noti zbl. >
lin-Dahlem 9: 273. 1925. :
Jauja, in the valley of Río Masamerich (Río Ponta-
chuela) confluent of Río Pangoa, above Rasthütte
Calabaza, 1 1 m, 11°30’S, 7 May 1913, We-
berbauer 6663 steely B; isotype, F).
Usually hemiepiphytic, rarely terrestrial or epi-
phytic; stem appressed-climbing, densely scaly or
setose, trichomes green to greenish white; pre-adult
internodes to 20 cm long, 1-2 cm diam.; adult in-
ternodes smooth, scurfy, minutely wrinkled, semi-
glossy to matte, 3-10 cm long, 2—6 cm diam., about
as long as broad or longer than broad, gray to
brown, roots moderately numerous, mostly short,
554
Annals of the
Missouri Botanical Garden
covered at least on one side with fine root hairs,
drying dark brown, semiglossy, faintly appressed-
scaly; cataphylls 10-30 cm long, unribbed, some-
times bluntly 1-ribbed, green or reddish, densely
scaly or setose, persisting as a tangled network of
fibers. LEAVES erect-spreading; petioles 33-
65(90) cm long, 3 cm diam. at base, 1.5 cm diam.
at apex, subterete, purplish violet to brownish, sur-
face densely scaly, the scales of two distinct types,
short, broad, often lacerate scales interspersed with
long acicular scales, the latter erect, or spreading
then erect; blades broadly ovate-cordate, thinly co-
riaceous, moderately bicolorous, acuminate to nar-
row acuminate at apex (the acumen mostly inrolled,
1-2.5 mm long), cordate at base, 28-75 cm long,
19-60 cm wide (1-1.7 times longer than wide),
(0.7—1.5 times longer than petiole), broadest below
point of petiole attachment, upper surface dark
green or bronze-green (blackish green on new
leaves), velvety to matte, drying dark brown, yel-
low-green or gray-green, lower surface pale green
tinged red-purple between secondary veins (weakly
glossy and purple-violet on new leaves), matte, dry-
ing yellow-brown to reddish brown; anterior lobe
9.6-51 cm long, 10-59 cm wide (1.9-3 times lon-
ger than posterior lobes); posterior lobes 8-21 cm
long, 5-28 cm wide, broadly rounded to obtuse;
sinus hippocrepiform to obovate or closed; midrib
flat to sunken, slightly paler than surface above,
convex, matte, darker than surface below; basal
veins 6-8 per side, with (0)1-2 free to base, num-
bers 3—4 coalesced 1-5 cm; posterior rib not at all
naked on small blades or naked for 0.5-6 cm; pri-
mary lateral veins 3-6(8) per side, departing midrib
at a 5° angle, sunken, paler than surface
above, round-raised to convex, darker than surface
below; minor veins distinct and darker than surface
below, arising from E the midrib and primary
lateral veins; “cross-veins” conspicuous, in part
raised below. INFLORESCENCES 1—4 per axil; pe-
duncles 5-25 cm long, 1-2 cm diam., medium
green to purplish, usually 0.66-1.75 times longer
than the spathe; spathe densely scaly or setose,
10-22 cm long, 4 cm diam.; spathe blade medium
green, whitish or reddish green outside, white to
pinkish inside; spathe tube reddish green, medium
green, or dull purple-violet outside, red or pale red-
dish (dark green post-anthesis) inside; spadix 14.6
cm long; pistillate portion cylindrical to weakly cla-
vate, 1.8—4.6 cm long, 1 cm diam. at apex, 8 mm
wide at base; staminate portion 9.4 cm long; fertile
staminate portion cream; pistils 2.6-3.3 mm long,
1.4—1.7 mm diam.; ovary 4—5-locular, 1.9-2.5 mm
long, 1.4-1.7 mm diam., with axile placentation;
locules 1.9-2.5 mm long, 0.5-0.7 mm diam.; ovule
sac 1.8-2.2 mm long; ovules 20-24(34) per locule,
2-seriate, contained within translucent, gelatinous
ovule sac, 0.1–0.2 mm long, as long as or longer
than funicle; funicle to 0.2 mm long, adnate to low-
er part of partition, style 0.5-0.6 mm long, 1.4-1.7
mm diam., similar to style type B; style apex + flat;
stigma + lobed, 1.4 mm diam., 0.2-0.3 mm high,
covering entire style apex; the androecium trun-
cate, prismatic, oblong, margins + 4—6-sided, 0.8-
0.9 mm long, 2-2.5 mm diam. at apex; thecae ob-
long, 0.3-0.4 mm wide, + parallel to one another;
sterile staminate flowers blunt, irregularly 4—5-sid-
ed, sometimes clavate or prismatic, 1.6-1.8 mm
long, 1.4-1.7 and 1.9-3.4 mm wide. INFRUC-
TESCENCE with ripe berries white.
Flowering in Philodendron verrucosum occurs
from April through June (also September and No-
vember) in Central America. The species probably
also flowers earlier in the dry season, perhaps as
early as February, as suggested by the fact that a
post-anthesis collection was made in March (and
even in February in Colombia). Post-anthesis col-
lections have also been made from April through
July (but also in December). Immature fruits have
been seen from July, October, and December. The
flowering collections from September and October,
as well as the immature fruits from October and
December, appear to reflect bimodal flowering. On
the other hand, no flowers were seen from July and
no flowers or fruits from August.
Philodendron verrucosum ranges from Costa Rica
to Peru at 200 to 1500 (mostly above 500) m ele-
vation, mostly in Premontane rain forest and Trop-
ical Lower Montane rain forest but also in Tropical
wet forest. In Central America, it occurs principally
on the Atlantic slope or near the Continental Divide
in Costa Rica and Panama but also on the Pacific
slope in southwestern Costa Rica. In South Amer-
ica, it ranges along the Pacific slope of the Andes
to as far south as El Oro and Cotopaxi Provinces
in Ecuador, but also occurs on the eastern slopes
of the Andes in Napo and Morona-Santiago as well
as in Peru in the departments of San Martin and
Junin at 750 to 1850 m elevation.
Philodendron verrucosum is a member of P. sect.
Philodendron subsect. Achyropodium. This species
is characterized by its short internodes, subterete
petioles (about as long as the blades), broadly
ovate-cordate blades velvety on the upper surface,
and especially by the densely scaly or setose ves-
titure of the stems, cataphylls, petioles, and inflo-
rescences.
Philodendron verrucosum might be confused with
a few other species that have scaly parts, including
Volume 84, Number 3
1997
Croat 555
Philodendron Subgenus Philodendron
P. hammelii, P. malesevichiae, P. squamipetiolatum,
and especially P. squamicaule. (See under the in-
dividual species for discussion of the differences.)
Philodendron squamipetiolatum differs by its lon-
ger, more slender internodes and much smaller leaf
blades (ca. 30 cm long on flowering plants) with a
semi-glossy upper surface. It also differs in having
solitary inflorescences (vs. usually more than one
per axil in P. verrucosum). Philodendron hammelii
differs in its much smaller size, petioles with foli-
aceous (rather than acicular) scales restricted to
near the apex, and glabrous cataphylls, Рен
and spathes. Philodendron malesevichiae
its terrestrial, creeping habit; persistent, dien
tact, glabrous cataphylls; narrowly sulcate petioles
(rather than subterete as in P. verrucosum); and se-
Miibiey blades lacking the conspicuous “cross-
veins" of P. verrucosum.
m
Additional specimens examined. COSTA RICA. Ala-
juela: Salto La Paz, 1350 m, Ferreyra 15788 (USM); San
NP aes ca. 2.3 km N Río Balsa, ca. 1050-1150 m,
sa, 900-1 000 m, 10?12'N, 84731^W, Stevens
13762 (CR, MO); 4—7 km N of Balsa, ca. 750 m, 10^13'N,
°32'W, Liesner & Judziewicz 14763 (MO); 2.5 km N of
—
E
ziewicz 14726 (CR, MO); Finca Los Ensayos, ca. 1
NW of Zarcero, са. 850 m, Croat 43636 (MO); 2 km N t
Angeles Norte de San Ramón, 4000 ft., Luteyn 3691 (MO);
шде, 11.8 km past aptid 1230 m,
^23'W, Hoover 1352 (CR, MO); E of Río San
Rafael, УТ of La Marina, 500 т, 10°23’N, 84^38'W, Bur-
ger & Stolze 5041 (CR, US); Monteverde Biological E
serve, Río Peñas Blancas, 900 m, 10?18'N, 84745",
ber & Bello 8458 ^ MO); 820 m, Bello 2672 (CR)
50-900 m, 4478 (CR, INB); 950 m, Bello & Méndez
2639 (CR, INB); Самба Alfaro Ruiz, Fe Рећа de Zarcero,
Smith 961 (F); Los Chiles-Los Angeles, San Ramón, 500—
200 m, 10%10'N, 84°30'W, gii oe et x 2200 (CR,
МО); E. Schmidt 306 (CR). Cartago: 4000 ft., J. D. Smith
391 (G); Moravia-Quebrada Platanillo, Mia 3-5 km
from Finca Racine, 1200-1300 m, Croat 36623 oe
1200-1400 m, 36795 (MO); Turrialba-Limón, along H
32, ca. 11 mi. NE of Turrialba, 850 m, Croat 43355 (CR.
МО); 31 km S of Siquirres on road to Turrialba (CR-101),
850 m, 957'N, 83°36'W, Tide & Rawlins 1163
(CM); El Muñeco, S of Río Navarro, 1400-1500 m, Stan-
dley & Torres 51 745 (CM, US): Standley 33425 (US);
Finca Navarro, 1350 n Maxo n 636 (NY, US); 10 km 5
of Tapantí, 1600 m, 83 ^46' W, Burger & Burger
8424 (CR, F); 3.5 km SE of Tapantí, 1250 m, Lent 851
(F, NY, US); Río Grande de Orosi, 2 km SE of ама
1350, Lent 107 (F); 14 "i from Turrialba, road to Pavo-
nes, Carlson 3244 (F); Orosi, Standley 39736 (US); La
Estrella, Standley 39547 (US); 4250 ft., J. D. Smith 5963
nay Río Gato, 4 km SE of Pejibaye, 700 m, 948'N
2'W, Liesner 14412 (MO); Tapantí Hydroelectric Re-
ith & Taylor 1009
region, 1.5 km past Río Macho, along small stream along
road to Humo, 1330 m, 9°17’N, 83°50'W, Hoover 1345
(CR, MO); 4250 ft., Cooper ua (CM, GH, US); Tucur-
ue, Las Vueltas, 635—700 m, Tonduz 13321 (US); Re-
ventazón Valley, Juan Viñas, Cook & Doyle 288 (US). He-
redia: 4 mi. N of Vara Blanca, 1350 m, Croat 35609
(MO); = mi. 5 of Cariblanco, 760 m, Croat 35794 (МО);
La Selva Field Station, 280-300 m, 10°21'N, 84%03' W,
ama, 1200 qt дисања 23533 (МО
US); Talamanca, Tsaki, 200 512 (CR, US):
Fila Matama, 850 m, Robles E oe pa (CR, MO).
Puntarenas: Las Cruces Botanical Garden-Río Jaba, ca.
3 km SE of San Vito de Coto Brus, ca. 1050-1200 m,
аи 82°58’ W, oe. 5621 (CR, MO); Monteverde
гуе, 1500 m, 10?17'N, 84°48’W, Hammel & Zu-
odi 13886 (MO). San José: San Isidro del General-
Dominical, 4.8 mi. from Río Pacuare, 1000 m, Croat
35251 (MO); 9 mi. SW of Río Pacuar, 680 m, Croat 35349
Gómez 20564 (M,
m
18096 (F, NY); La Hondura, 1300-1700 m, Standley
37783 (US); Río Claro Valley (Bajo La Hondura), below
La Palma, NE of San Jerónimo, 1000-1200 m, 10°3’N,
83°58'W, Burger et al. 9433 (CR, F, А below La Palma
NE of San Jerónimo, 1000-1200 m, 10?3'N, 83°58'W,
Burger et al. 9386 (F, MO), ое y Gentry 9077 (CR,
); 1500 m, 10%3'N, 83° ger & Stolze 5308 (CR,
F), 8499 (F); Tarrazá, vic. iae guero, 11 200 m,
Croat 78935 ies МО). PANAMA. Bocas del Того:
Cerro Colorado, 9.2 mi. W of Chame, 1450-1480 m,
8°35'N, 81?50'W, Croat 69010 (MO); Fortuna Dam area,
Chiriquí Grande-Gualaca, 21.4 km past Gualaca, 8°32'N,
оба Hoover 1327 (MO); Continental Divide, ca. 1200
m, са. 8°45’N, 82°15'W, McPherson 9033 (CA S, CM, K,
MO): кў, ыр 60374 o Old Bank Island, vic. of Chiriquí
Lagoon, von We 3 (MO). Chiriquí:
Gordas, 1 > : о 4
Сојогадо,
37254 do: d cul Grande, vic. IRHE head-
uarters, 1200 m, 8?45'N, 82°18’W, Croat 66719 (MO);
66558 (ААЏ, MEXU, МО, NY, РМА); road to Finca Lan-
dau, 1100 m, Correa et al. 2141 (F, MO, PMA); NW of
ort m, Cerro una, lower slopes, 1150 m,
7 Vodicka 5576 ~ "behind Vivero Forestal, 12 km N
Planes de Hornito, 1200-1300 m, 8?45'N,
82°12/ w Knapp 4949 (MO); 7 km SE of Fortune Dam,
O'Connor 91-512-004 (MICH); Continental Di-
vide, 1200-1500 m, 8°47'N, 82713", Churchill 5298
(МО); 1200 m, O'Connor Perm -003 (MICH); Continen-
tal Divide trail, 1200-1300 m, 8%45'N, 82^15'W,
Thompson 4954 (CM, MO). Coclé: El Copé region, Alto
Calvario, near Continental Divide, 5 mi. N o opé,
900-1000 m, 8°39'N, 80*36' W, Croat 75058 (MO); 750-
900 m, 44736 (MO); la 80°36'W, 68768 (СМ, F,
MO); El Valle region, La Mesa, Gentry 5632 (MO, PMA):
860-900 m, Croat 37. 7339 (MO); T: (MO); 900 m, Sul-
; NE slopes of Cerro
—
—
Mori et al. 6633 с Lallathin 5096 (MO); Cerro Сайа,
900-1000 m, 8°40'N, 80707", Knapp 5301 (МО);
= m, Knapp & seem 4913 (MO); 5.6 km S of El Valle,
m, 8°50'N, 80°07’ W, Hoover 1319 (MO, NY); ca. 3
= N. и El Valle de Antón, 850 т, Wilbur et al. 15656
(MO). Darién: Cerro Pirre region, 1000-1100 m, Hart-
man 4829 (MO); 9-10 km N of Alto de Nique, 1520-
1560 m, Croat 37871 (MO); Río Tuquesa, vic. of Tyler
556
Annals of the
Missouri Botanical Garden
Kittredge gold mine, ca. 2 air km from ti
Croat 27215 (MO). Panama: El cc apris
00 m, Liesner 689
875 (MO); above Su Lin Motel, Croat 14728 (MO); near
Florida State Univ. Building, Croat 12121 (F, MO, SCZ).
San Blas: Cerro Нађа, vic. of peak, 2500 ft., 923'N,
78°49'W, Sytsma et al. 2699 (MO, US); Cerro Brewster,
850 m, 918'N, 79716", de Nevers et al. 5546 (MO);
Cerro Obu, 400—500 m, de Nevers et al. 8053 (MO); Nu-
sagandi, 310 m, 9?20'N, 79°, Croat € Zhu 76993A
(MO). Ver raguas: Santa Fe region,
fork of road beyond Alto Piedra, 0.6 mi. b
the road, 1300-1350 m, Croat 49060 (MO); Santa Fe-Río
Calovébora, 0.6 mi. beyond Alto Piedra, 735 m, Croat Т
Folsom 33991 (МО); 1.7 mi. past Alto Piedra, 1.5 т
beyond Quebrada Cosilla, 570 m, 8°33'N, 81°08’ W, Сузи
& Zhu 76856 (МО).
Philodendron warszewiezii K. Koch & Bouché,
in A. Braun et al., Append. gen. sp. Hort. be-
rol. 1855: 4. 1855-1856. TYPE: Guatemala.
San José and Florida, Warszewicz s.n. (holo-
type, B? lost?). Guatemala. Santa Rosa: Volcán
Jumaytepeque, 6000 ft., Hyde & Lux 4282
(neotype, here designated, K). Figures 16,
, 446-449.
Usually terrestrial, or on rocks, sometimes epi-
phytic; stem appressed-climbing, thick, succulent,
bare, leaf scars conspicuous, 1.5-2.6 cm long,
1.5-2 cm wide; internodes short near apex, semi-
glossy, glaucous, 3-5(15) cm long, 2.5-7(10)cm
diam., often longer than broad lower down, dark
green to gray-brown; roots pale green to dark
brown, few per node, epidermis drying yellowish;
cataphylls thin, soft, 18-33 cm long, weakly to
sharply 2-ribbed, pale green to whitish, lightly and
densely short-lineate, deciduous intact. LEAVES
often deciduous in dry season; petioles 33—58(80)
cm long, 1-2 cm diam., subterete to C-shaped,
moderately soft, weakly flattened to rounded with
thick, medial rib adaxially, with adaxial margins
sharply raised, surface sharply striate, with a dark
green ring around apex; blades triangular-sagittate
in outline, deeply bipinnately or bipinnatisect to
within 1—4 cm of the midrib, thin, semiglossy, mod-
erately bicolorous, + rounded at apex (the acumen
2 mm long), cordate at base, 31-78 cm long, 30—
62 cm wide (0.9-1.3 times longer than wide), (1—
1.3 times longer than petiole), upper surface mod-
erately glossy, moderately paler, margins sinuate;
median lobe 17-52 cm long, 23-63 cm wide,
rounded to obtuse; lateral segments 1-30 cm long,
11-28 cm wide, pinnately lobed with 3 or more
lobes per side, acute or acuminate; interlobal si-
nuses 0.78-0.94 the length of the blades; midrib
flat, obscurely striate, slightly paler than surface
above, raised, obscurely striate, slightly paler than
surface below; basal veins 2-6 per side, with 0-1
free to base, second and third pair coalesced 2-5
cm, third and higher order veins coalesced 5—7 cm;
posterior rib naked along most of its margin; pri-
mary lateral veins 3-6 per side, departing midrib
at a 55—70° angle, straight to the margins, weakly
raised above, raised, paler than surface below; ter-
tiary veins raised, paler than surface above and be-
low; minor veins visible, darker than surface below,
distinctly visible on drying, arising from both the
midrib and primary lateral veins. INFLORES-
CENCES 1(3) per axil; peduncle (2.5—3)4.5—11(16)
ст long, 1-2 cm diam., dark green, dark short-
lineate; spathe 13.5-30 cm long, 4–7.5 cm diam.
(1.1-3.6(4.7) times longer than peduncle), medium
to dark green throughout, blunt at apex, scarcely
constricted; spathe blade short dark green lineate
outside, 12 cm long, pale green to white inside;
spathe tube weakly short dark green lineate out-
side, 14 cm long, pale green to white or purple or
red inside; spadix sessile; white throughout, 14–
24(30) cm long, constricted weakly above pistillate
portion; pistillate portion 3 cm long in front, 2 cm
long in back, 1.3 cm diam. at apex, 1.7 cm diam.
at middle, 1.6 cm wide at base; staminate portion
8.3-15 cm long; fertile staminate portion 1.6 cm
diam. at base, 1.4 cm diam. at middle, 9 mm diam.
ca. 1 cm from apex; sterile staminate portion 1.3
cm diam.; pistils (1.1)4.1 mm long, 3. mm
diam.; ovary 4-5-locular, 2.6-9.5 mm long, 3.54.5
mm diam., with axile placentation; locules 2.6 mm
long, 0.8-1.1 mm diam.; оуше sac 2.4 mm long;
ovules 3-4 per locule, 0.5 mm long, contained in
thick, translucent matrix; style 1.2-1.6 mm long,
1.2-1.4 mm diam., similar to style type E; style
funnel broad, moderately deep; style apex + flat;
stigma subdiscoid, unlobed, 0.7 mm diam., 0.3 mm
high, lining entire upper surface of funnel; the an-
droecium truncate, prismatic, oblong, margins ir-
regularly 4—6-sided, 3.3 mm long, 1.3 mm diam. а!
apex; thecae oblong, 0.4 mm wide, + parallel to
one another and nearly contiguous; sterile stami-
nate flowers rounded to somewhat clavate, irregu-
larly 4—6-sided, 2.6 mm long, 1.3-1.8 mm wi wide.
INFRUCTESCENCE brownish outside, brownish
inside, to 3 cm wide pistillate spadix 3-7.5 cm
long; berries white, 8-11 mm long; seeds 3—4 per
locule, pale yellow, oblong-ellipsoid, 2-2.5 mm
long, 1-1.2 mm diam., with pale raphide cells.
Flowering اا of Philodendron warszew-
Volume 84, Number 3
1997
Croat 557
Philodendron Subgenus Philodendron
iczii are rare, known only from May. Post-anthesis
collections, concentrated between April and June
(but also January, February, March, July, August,
September, and December), imply that the flower-
ing period is relatively broad, perhaps throughout
the entire dry season and the first half of the rainy
season. Immature fruiting collections are scattered
in a somewhat bimodal pattern, one group in the
mid-rainy season from July through September, the
other group primarily in the early dry season from
December through March (especially December
and January). The immature fruits collected in No-
vember might reflect an early dry season flowering.
Mature fruits are known only from this same period
between December and February at the beginning
of the dry season.
Philodendron warszewiczii ranges from Mexico
(Jalisco and Chiapas) to El Salvador on the Pacific
slope, and to Honduras and Nicaragua at 300 to
900 m elevation, in Tropical dry forest life zones.
In Mexico, the species occurs in “Selva Mediana
Subperennifolia" and *Selva Baja Caducifolia."
hilodendron warszewiczii is a member of P. sect.
Polytomium. This species is distinguished by its
moderately thin, deeply dissected, bipinnatifid leaf
blades and thick, succulent stems often leafless
during the dry season
Philodendron warszewiczii is most similar to P.
radiatum and P. dressleri. It differs from P. radia-
tum in having thinner, more highly divided blades
and thicker stems. Although both species occur in
some of the same forest types in Mexico, P. radia-
tum always occupies more mesic sites within these
zones. In other parts of Central America for which
there are Holdridge Life Zone maps, P. radiatum
is found mostly in Tropical moist forest or wetter life
zones, whereas P. warszewiczii is known from Trop-
ical dry forest areas.
Although Philodendron warszewiczii and P. dres-
sleri occupy equally dry sites, the latter is distin-
guished by occurring further north in Mexico and
in having the leaves divided to no more than half-
way to the midrib (vs. nearly all the way to the
midrib in P. warszewiczii).
This species occurs principally on the Pacific
slope except in Nicaragua (one Honduran collec-
tion is from the Atlantic slope). This is probably
due to the fact that the Continental Divide runs
very near the Pacific Ocean in Nicaragua, and be-
cause there is a second, generally much higher,
more centrally located range of mountains and hills
that creates a rain shadow throughout most of the
Pacific side of the country. Because of this, P. war-
szewiczii inhabits many sites in the central part of
the country well within the Atlantic watershed.
pcm names for this species include: “Gua-
amayo," "Cupapayo," *Mano de Lión," *Copapa-
yo," “Ocopayo,” and “Papaya de Monte” (Standley
& Steyermark, ' 1958b).
Additional specimens examined. EL SALVADOR. S of
San Salvador, Calderón 876 (GH, US). Ahuachapán:
Ahuachapán vicinity, 800-1 m, Standley 19726 (GH,
NY, US). San Salvador: са 416 (СН, МУ, 05); Ѕап
Salvador sar ae 50 m, Standley 19624 (N Y USI
GUATEMALA. Jalapa: Jalapa-San Pedro Pinula, 1400—
1800 m, gine To 77055 (F). Jutiapa: Monjas—El Ргорте-
so, above Ovejero, ca. жь т, eRe 77660 (Е); near
a ca. 900 m 5 (F); Agua Blanca—Amatillo,
50-990 m, Gears А. (F); Jutiapa-La Calera, SE
ài Jutiapa, ca. 850 m, Standley 76087 (F). Santa Rosa:
near Cuilapilla, ca. 900 m, Standley 78051 (F). Zacapa:
een Santa Rosalía
2. ;
Río Lima, Sierra
1700 m, Steyermark s.n. (F). HONDURAS. Río de la Or-
illa, SE of El Zamorano, 900 m, Anderson & Spoehr s.n.
(MO). Comayagua: vic. of Comayagua, ca. 600 m, Stan-
50 m, Standley 17007 (F); near Yuscarán, 960 m
Standley et al. 1234 (F). op are Tegucigalpa, Canta
buqui, 2600 ft., von Hagen & von Hagen 1272 (F, NY);
S d e Occidente #350 m. Valerio 3654 (F); Escuela
Agricola ricana, 800 m, Croat 34814 (cultivated)
(MO); ah pos de Ош, 4 mi. N of Zamorano, Croat
42765 hz US); Río de la Orilla region, SE of El Za-
morano, 900-950 m Stay Же 23205 (F), 23213 (F),
АЗАТ (P) near El гадо, 800 т, Morton 7119 (US).
Olancho: Rfo Olancho, San Franciaco | Зе Ја Мо
14 mi. МЕ of San Francisco de la
14°57'N, T W, Croat & Hannon 64186 (B, MO); К
of main 8.6
Croat & Rais се (СМ, МО). MEXICO. Chiapas:
20-30 mi. SW of El Jocote, on road to Motozintla, 700—
900 m, Croat 40714 (MEXU, MO); Tuxtla Gutiérrez-San
Fernando, Chacona canyon, 850 m, Breedlove gel
(CAS); 22 mi. SE of Comitán, ae 1953 (F); 6-8 km
E of Frontera Comalapa, roa iudad Cuauhtémoc,
1000 m, Breedlove 23428 (MO); 10 km E of Mozotintla,
1110-1150 m, Garcia et al. 1527 (MEXU, BM); vic. Tux-
tla Guttierez, 830 m, Breedlove & Bourell 68505, (CAS);
pio. Tzimol, 15 Eri S Ф Comitán, 1200 т, Breedlove
51028 (CAS). Jalisco: Autlán-Barra de Navidad, at km
291.67, ca. 300 m, Moore & Bunting 8744 (BH); 9 mi. N
of road-junction at W end of Bahía de Pyp 300 m
McVaugh & Koelz dad eee Hwy. 110, 2 mi. NE of
ns & Fairhurst Mus (US); 15-
of Barr “ Navidad. Dressler & Wirth 2671
(UG. US); ip wig Field Station, 100 km 5 of Puerto
allarta, 50 m, 19°30'N, 105°03'W, Gentry 63966 (MO),
Bullock 1 101 (MO); El ion, 100 m, 19°30’N, 105*03' W,
Gentry 74426 (MO); 10 km E of Las Palmas, m,
20^50'N, 105%02'W, Cochrane et al. 12022 (MO); Reserva
Biosfera Sierra de Manantlán, Lagunillas-Juluapan, Cerro
Grande, ca. 1400-1500 m, 19°22'N, 103°56' 30W, Iltis et
al. 29716 (МО); Pacte о Cuixmala, Cumbres 1, Arroyo
Cajones, 19°27'N "307, Lott et г 3268 (МО).
Nayarit: Mirador de Ese ca. 14 mi. N of Te m
450—600 m, Feddema 824 (MICH); 67. mi. S of Com
558
Annals of th
Missouri Botanical Garden
postela, 1000 m, McVaugh 18754 pagos à mi. SE of
Las Varas, road to Mazatlán, ugh 15351
МО);
1218'N, 8530", Stevens & Montiel 1 7206 (МО); Jui
galpa-La Libertad, ca. 17.4 km NE of Río Mayales, 35!
400 m, 12°12'N, 85°17'W, Stevens 4023 (MO). Estelí: a
ке: 5 of Estelí, 1000 m, Neill 1186 (MO); km 163 on Hwy.
. 11.2 km N of entrance to Estelf, ca. 920 m, ca.
EN 86°23'W, Stevens 11213 (MO); Cerro Сисатоп-
a, 800-850 m, 10"15"М, 86722", Moreno 14120 (MO);
Salto de Estanzuela, ca. 5 km of Estelí,
13°02'N, 86720", Guzmán et al. d d (MO);
tanzuela, 6 km e Estelí, 1000 m, 13?01'N,
8672 P М. pene 9751 (MO). ДЕ Jinotega- Бан Ка-
"à el, NW of Ji inotega, , Croat 43020
(MO); ү a Abenas, 1 1-2 mi. E of San Rafael de Norte
Jinotega Road, 1000 m, Croat 42970 (MO); ca. 1 mi. SE
of Yalí, 850 m, Croat 42891 (MO). Madriz: San Lucas-
Hacienda El Volcán, 2 km NE de San Lucas, 800-900 m,
13%25'N, 86735", Araquistain & Moreno 2018 (K, МО);
N of El Sicut, 700-1000
86°08' W, у М ка А & Moreno 2316 (МО,
NY); Quebrada El Nancital, ca. 6.2 km N of Ocotal, Que-
brada El Nancital, 700—760 m, са. 13?41'N, 86°24’W,
Stevens 3040 Vd "d 6.5 km de Ocotal, road to Las
13?40'N, сыф eae 14382
; El Jica “Casas Viejas, . 600 m, 13%44'N,
205’ W, Moreno 13504 perd El pine *El Сабый. м
са. куы» , 13°46 57, Moreno 13589 (MO);
El Jícaro, “Río Gra Ka per o Murra, 500 m
13743'N, 86"W, Moreno 13670 (M О).
~
Philodendron wendlandii Schott, Prod. Syst. Ar-
oid. 221. 1860. TYPE: Costa Rica. Wendland
s.n. (holotype, W? lost). Schott ic. 2079 (neo-
type, here designated). Figures 15, 450, 451,
453.
Epiphytic, sometimes occurring high in the can-
opy, rarely terrestrial, perhaps by accident; stem
short, scurfy, leaf scars inconspicuous, 1.3-2.1 ст
long, 2–2.5 ст wide, obscured by root mass; inter-
nodes short, 0.5-3 cm long, 1.8-3.5 cm diam.,
glaucous, epidermis fissured transversely; roots
reddish green becoming dark brown, е
branched near tip; cataphylls somewhat spon
fleshy, 29-36 cm long, unribbed or sometimes
sharply 2-ribbed, green to reddish green, persisting
semi-intact at upper nodes, eventually fibrous
ES rosulate; petioles 9-30 cm long, 1.54
cm diam., broader than thick, spongy, soft, sharply
flattened adaxially, sharply convex abaxially, with
adaxial margins erect on younger petioles, with
dark green ring at apex; blades oblong-oblanceo-
late, coriaceous, slightly bicolorous, semiglossy,
acuminate, sometimes long-acuminate at apex (the
acumen inrolled, 2-4 mm long), + narrowly round-
ed to obtuse at base, 32-67 cm long, 8.5-22 cm
wide (2.6—4.7 times longer than wide), ((1.7)2.6-
4.2(5.2) times longer than petiole), broadest = near
the middle; midrib flat to convex, broadly convex
near base, convex toward apex, concolorous above,
round-raised to convex, slightly paler than surface
below; basal veins lacking; primary lateral veins 7-
12(16) per side, departing midrib at a (50)60—70*
angle, = straight to the margins, weakly impressed
to sunken above, convex to weakly raised, darker
than surface below; minor veins obscure above, vis-
ible, darker than surface below, arising from midrib
and paralleling primary lateral veins. INFLORES-
ENCES erect, 2 per axil; peduncle 3-10 ст long,
1-1.5 ст diam., pale green; spathe erect to
erect-spreading, coriaceous, 12- em long
((1.5)3.4—3.6(5.3) times longer than peduncle), con-
stricted just above the tube to 2.1 cm diam. at con-
striction; spathe blade oblong-ovate, pale green to
white, sometimes tinged pink, sometimes white-
striate outside, 7-13 cm long, 2.6—7 cm diam. when
open (opening broadly elliptic in face view, white
(rarely rose-red)), pale-spotted with dark green cen-
tral rib inside; spathe tube ellipsoid, pale to me-
dium green, sometimes tinged with red, white-
short-lineate in back outside, (3.5)5.5-7 cm long,
2.6–3.5 cm diam., pale green to white (sometimes
creamy yellow or reddish with darker green central
in minutely pale унш inside; spadix stipitate
o 1.3 mm long, light green; cylindrical, bluntly
ied at apex, (8.5)11-14.4(17.2) cm long, of
nearly uniform width; pistillate portion pale green
to yellowish iex cylindrical, or weakly tapered
toward apex, 2.9—4.6(6.7) cm long, 1-1.2 cm diam.
at apex, 1.1-1.4 cm diam. at middle, 1-1.5 cm
wide at base; staminate portion 8.2-11.7 cm long;
fertile staminate portion creamy white, with resin
droplets, 9-13 mm diam. at base, 9-12 mm diam.
at middle, 6–7(9) mm diam. ca. 1 cm from apex,
broadest at the base or equally broad at base and
middle, as broad as the pistillate portion, as broa
as to slightly narrower than the sterile portion; ster-
ile staminate portion slightly broader than the pis-
tillate portion, white, 1-1.3 cm diam.; pistils 1.7-
2.3 mm long, 1.4—1.7 mm diam.; ovary -6-7-locular,
0.7-1.3 mm long, 1.4-1.7 mm diam., with sub-ba-
sal placentation; locules 0.7-1.3 mm long.
(0.2)0.4—0.5 mm diam.; оуше sac (0.7)1-1.1 mm
long; ovules 2 per locule, translucent and contained
within translucent or transparent ovule sac, 0.3-0. 5
mm long, longer than funicle; funicle 0.2-0.4 mm
long (can be pulled free to base), style 0.6-0.8 mm
—— ET AA
س — ————
eee ل ——
Volume 84, Number 3
1997
Croat 559
Philodendron Subgenus Philodendron
long, 1.2-1.8 mm diam., similar to style type B;
style apex sloping to weakly rounded, with depres-
sions surrounding stylar canal exits; stigma subdis-
coid, truncate, 0.7—1.1 mm diam., 0.2-0.3 mm
high, covering center of style apex; the androecium
truncate, prismatic, margins irregularly 3—6-sided,
1.2 mm long, 0.7-1.4 mm diam. at apex; thecae +
ovate, 0.5 mm wide, = parallel to one another,
sometimes nearly contiguous, sometimes distinctly
contiguous; sterile staminate flowers blunt, irregu-
larly -sided, sometimes clavate or prismatic,
1.4-1.6 mm long, 1.1-2(3.6) mm wide. Berries
bright orange.
Flowering in Philodendron wendlandii occurs
from the mid-dry season to the first part of the rainy
season, with Costa Rican populations having been
found in flower earlier (February and March) than
those in Panama (May and June). Post-anthesis in-
florescences have been collected between February
and July, with an outlying collection from Novem-
ber. Immature fruits have been collected only in
June and August, and mature fruits in September.
Philodendron wendlandii ranges from Nicaragua
to Panama, from 10 to 1250 m elevation in Tropical
wet forest and Premontane rain forest life zones.
Most collections are from below 800 m elevation.
Philodendron wendlandii is a member of P. sect.
Calostigma subsect. Glossophyllum ser. Glossophyl-
lum. This species is distinguished by its true epi-
phytic habit, short stem, and rosulate, oblong-ob-
lanceolate, short-petiolate leaves but especially by
the petioles, which are often broader than thick and
sharply flattened adaxially. Philodendron wendlan-
dii is one of relatively few truly epiphytic, rosulate
“bird’s nest” species in the genus.
Philodendron wendlandii is apparently related to
several species of more or less rosulate habit and
with elongate leaf blades and a green or purple line
of demarcation between the petiole and the blade.
These include Philodendron annulatum, P. auri-
culatum, and P. ligulatum. With the exception of
P. ligulatum var. heraclioanum, all of these taxa
differ from P. wendlandii in having more or less
terete petioles. Philodendron ligulatum var. hera-
clioanum itself differs in being а long-stemmed
vine (vs. rosulate in P. wendlandii), and in having
D-shaped petioles scarcely or not at all wider than
thick (vs. sharply flattened and conspicuously
broader than thick for P. wendlandii) and undulate-
winged margins.
Additional specimens examined. COSTA RICA. Ala-
juela: near La Laguna, 6-8 km S of Villa Quesada, 1200
m, Molina et al. 17 7553 (EAP, F, NY, US); Cafias- Upala,
near Río Zapote, 1.8-2.7 km S of Río Canalete, 100 m,
Croat 36376 (MO); 8 km NE of Villa Quesada, 600 m,
е 46925 (МО); Rio Frio, near Los Chiles, 30-40 т,
1°02'N, 84°44’ W, Holm & Пиз 826 (A, GH). Cartago:
Moravia Quebrada Platanillo, ca. 1250 m, Croat 36683
; aste: El S
m, Standley & Valerio 44785 (US). Heredi
Cariblanco, 760 m, Croat 35833 (MO); 5 of Río Sarapiquí,
opposite Chilamate, 60-100 m, 10%27'N, 84%04'W, Gra-
yum et al. 5311 (MO); La Selva F ial Station, Puerto Viejo
de Sarapiquí, 100-150 m, Croat 44313 (MO); са. 100 m,
d 9086 (F, MO); McDowell 943 (MO); Fi Mon 9596
EX); Hammel 8222 (DUKE). Limón: Finca La Suerte,
Davidson 3265 i i
S :
69732 (CM, MO); Hitoy-Cerere, Quebrada Cunabrí, Baja
Talamanca, 500-800 Gómez 24081 (MO); Amubri,
along Río Lari, 200 m, 9°20) 40"N, 82%89'40"W, Herrera
2993 (AAU, CR, MO, US); Guépiles, 850 ft., J. D. Smith
4977 (US); Cerro Coronel, E of Laguna Danto, 20-170 m,
10%41'N, 83738", Stevens ag (CR, MO); E of Laguna
Danto, SW towards hills E o Zapote, 30-170 m
10°40-41'N, 8338-39
E of Río Zapote, 1 km dion Río
10%40'N, 83°40'W, y жые & Montiel 24313 (CR, MO);
Stevens et al. 24683 (CR, MO); between Río Madre de
Dios and Quebrada Cafiabral, 380—440 m, 10°02'N,
83?26'W, Grayum et al. 8712 (CR); Parque Nacional Tor-
tuguero, NE of La Aurora, Guápiles, along Río Sierpe, 30
m, 10%22'N, 83°31'W, Robles 2254 (CR, MO). San Јове:
Carrillo Station, 700 m, Gómez et al. 21192 cn MO).
NICARAGUA. Río San Juan: Bocas de Sábalo, 70—100
84727", Moreno 26780 (MO). PANAMA.
Dam Area, Gualaca-Chiriquí
Grande, 9.4 mi. N of the Continental Divide, 175 m,
8°46'N, 82°16'W, Croat 66819 (K, MO); 2.8 mi. N of Con-
tinental Divide, 8
Pherson 9653 (L, MO, PMA, US). € a
from Río Escandaloso, 2000 ft., Puer 3205 (MO);
Santa Rita ipn
6.5, 370 m, 9?21'15"N, 79744", Croat & Zhu 76962
(MO) аай: Cerro Jefe region, Campo Tres, 3 mi, МЕ
о E n (MO); 3-3.5
mi. NE of Altos de Pacora, 7. 8.82 above Pan-Amer-
ican Highway, 700—750 m, %15'N, 79°25) W, Croat 68633
(MO); El Llano—Cartí, 10 mi. кп main gate near El Lla-
no, Croat 33715 (CM, MO); M i
m, 920'N, 79°W, eter & Zhu Tap (MO). Veraguas:
anta Fe region,
Alto Piedra, 735 m, aer ad 34141 (M0); умна Fe-Calov-
ébora, 1.7 mi. past Alto Piedra, 1.5 mi. beyond Quebrada
Conlin. 570 m, PE33'N, 81°08'W, Croat & Zhu 76820 (F,
MO).
Philodendron wilburii Croat & Grayum, sp. nov.
TYPE: Costa Rica. Alajuela: Cordillera de Ti-
larán, San Ramón-Bajo Rodríguez, vic. La
types, CR, K, US). Figures 452, 454—458
Planta plerumque hemiepiphytica; caulis scandens; in-
ternodia 3-15 cm longa, 1.3-2 cm diam.; cataphylla 7—
560
Annals of the
Missouri Botanical Garden
m longa, incostata vel obtuse 1-costata, decidua; pe-
] teres, moderate s 10818, obtuse
i 2m
ст lata, cordata vel -— basi: inflorescentia 1—3(4);
pedunculus 3.5—29 ongus, 8-12 mm diam.; spatha
1.5-18 cm longa; eie spatha extus albida vel pallide
viridi aut cremeo-flava, intus albida (sub anthesi) vel cre-
meo-flava; tubo spathae extus virenti, intus rubella vel
atrimarronino; pistilla (5)6—7-loculari; loculi 1-2-ovulati;
baccae pallide vel vivide aurantiacae.
Usually hemiepiphytic, sometimes terrestrial;
stem scandent, semiglossy, sap watery, spicy-scent-
ed, sometimes milky-white; internodes usually long,
glossy to semiglossy, 3-15 cm long, 1.3-2 cm
diam., usually longer than broad, dark green to
ay-green, sometimes drying black, epidermis of-
ten cracking or minutely fissured, brittle, some-
times silver-gray; cataphylls soft, 7-26 cm long,
unribbed to bluntly or sharply 1-ribbed, sometimes
sharply D-shaped and sharply 2-ribbed, whitish or
ates ae glossy, deciduous; petioles 16–55 cm
long, 5-12 mm diam., subterete to terete, moder-
ately spongy or moderately firm, bluntly D-shaped
or obtusely flattened adaxially, surface weakly
glossy to semiglossy, smooth; sheathing 6-8 cm
long; blades ovate-triangular, thinly coriaceous to
subcoriaceous, matte to semiglossy, weakly to mod-
erately bicolorous, acuminate to long-acuminate at
apex (the acumen more or less inrolled), cordate to
sagittate at base, 17-36 cm long (averaging 26 cm),
11-31 cm wide, + equal in length to or longer than
petiole, margins hyaline, upper surface drying
brownish, dark brown, or gray-brown to green, low-
er surface drying light brownish green to reddish
brown or yellowish brown; anterior lobe 11.5-32
cm long, 7-24 cm wide; posterior lobes somewhat
spreading, rounded to obtuse, 4-14 cm long,
(2.7)4.2-13.5 cm wide, directed toward base ог
somewhat outward; sinus hippocrepiform, parabolic
to arcuate or arcuate with petiole decurrent, 1–6(8)
cm deep; midrib broadly convex to shallowly sunk-
en to flat, paler than surface above, convex, paler
than surface, drying darker than surface below; ba-
sal veins 3-5 per side, with 0-1 free to base, 1—
3(4) coalesced (0.7)1.3—4(7) cm; posterior rib na-
ked for 0.5-2.6 cm, sometimes not naked, directed
to the tip of the posterior lobe 1-3.5 cm distant
from the lower margin of the posterior lobe; primary
lateral veins (2)3—4 per side, departing midrib at a
40—60(70^) angle, + straight to the margins,
streaked reddish below, broadly impressed to shal-
lowly and obtusely sunken above, weakly convex
and darker than surface below; minor veins mod-
erately distinct, arising from both the midrib and
primary lateral veins. INFLORESCENCES erect,
1-3(4) per axil; peduncle 3.5-29 cm long, 8-12
mm diam., medium green, glossy, faintly white-stri-
ate near apex; spathe glossy, 7.5-18 cm long,
1.4-2.2 cm diam., weakly constricted midway
above the tube, 1.9-2.2 mm diam. at constriction;
spathe blade creamy-white to pale green or creamy-
yellow, sometimes tinged faintly reddish outside, 6–
7 ст long, 2.2-3.2 cm diam. (opening 6.7-9 cm
long, 3.2 cm wide), whitish (at anthesis) to creamy-
yellow inside, sometimes suffused with red; resin
canals orange and appearing as continuous lines at
least around the throat inside; spathe tube medium
to dark green and light maroon band near base,
sometimes weakly to heavily tinged reddish (B &
K red-purple 5/7.5), white-striate outside, 5—7.5 cm
long, 2-3.6 cm diam., reddish to dark maroon (B
& K red-purple 2/10) inside; spadix sessile; * ta-
pered toward apex, 6.7-11.1 cm long, broadest at
or near the base, protruding strongly or weakly out
of the spathe blade at anthesis; pistillate portion
weakly tapered toward both ends, light green to yel-
lowish green, (2.5)3.4—4.3 cm long in front, 2.9-
3.8 cm long in back, 7-13 mm diam. at apex, 1-
14 mm diam. at middle, 1-12 mm wide at base
staminate portion 5-8.4 cm long; fertile sini
portion white or light green, 10-11 mm diam.
throughout, of nearly uniform diameter throughout
or broadest in upper one-third, 9-13 mm diam. at
base, 10-11 mm diam. at middle, 7-9 mm diam.
ca. 1 ст from apex, broadest usually at the Базе,
narrower than the pistillate portion, narrower ke
the sterile portion; sterile staminate portion
broad as the pistillate portion, (pre-anthesis) white
to light gray, 9-14 mm diam.; pistils 1.1-2.3(3.5)
mm long, 0.7-2 mm diam.; ovary (5)6—7-locular,
0.6–1.5(2) mm long, 0.8-1.3(1.6) mm diam., with
ed basal placentation; locules 0.7—1.1(2) mm long,
.6 mm diam.; ovule sac 0.7-0.9 mm long;
zt 1-2 per locule, 2-seriate, contained within
transparent, gelatinous ovule sac, 0.3-0.5 mm long.
slightly longer than funicle; funicle 0.2-0.3 mm
long (can be pulled free to base), style 0.3-1.2 mm
long, 0.8-1.6 mm diam., similar to style type B;
style apex flat to shallowly rounded or concave:
stigma brushlike to discoid or subdiscoid, purplis
violet or golden кэе nent ie 1-1.5 mm diam.,
0.2-0.4 mm high, covering + entire style apex; the
androecium truncate, prismatic, margins irregul. y
4—6-sided, 1-1.7 mm long, 0.7-1.5(2.7) mm diam.
at apex; thecae oblong, 0.4-0.5 mm wide, + par
allel to one another; sterile staminate flowers blunt,
irregular, dense, 4—6-sided, prismatic, 1. 5-2(2.5-
3.3) mm long, (1.2)1.5-1.6(2) mm wide. Berries
pale to bright orange (white, immature); seeds yel-
low-orange, narrowly ellipsoid.
——— —M ——— ل ——————— ل А —— ————— —— —— — —— —
Volume 84, Number 3
1997
Croat 561
Philodendron Subgenus Philodendron
Philodendron wilburii ranges from northern Cos-
ta Rica to the Canal Area of Panama from near sea
level to 2000 m elevation. This species is highly
versatile ecologically, ranging from Premontane
rain forest, Tropical Lower Montane wet forest, and
Tropical Lower Montane rain forest in the highlands
to Premontane wet forest and Tropical wet forest life
zones in the lowlands.
Philodendron wilburii is a member of P. sect.
Calostigma subsect. Glossophyllum ser. Ovata. This
species comprises somewhat scandent plants with
elongate internodes drying grayish to yellowish
brown and closely fissured; mostly 1-ribbed, decid-
uous cataphylls; subterete petioles mostly longer
than the blades; ovate to ovate-triangular blades of-
ten with the lobes directed somewhat outward; and
1-3 prominently pedunculate white to greenish in-
florescences per axil.
Philodendron wilburii is probably most closely
related to P. cotonense from the mountains of east-
ern Costa Rica. That species is distinguished by its
yellowish-brown-drying, more narrowly triangular
blades averaging over two times longer than wide
and thicker stems (2.5—3 cm diam. vs. up to 2 cm
diam. in P. wilburii). It also has typically longer
peduncles (mostly more than 10 cm) and more
ovules per locule (4—5, vs. 1-2 in P. wilburii). The
two species do not overlap in their distribution, but
do occur in the same life zones.
Philodendron wilburii may be confused with P.
straminicaule from Panama, but that species differs
in having thicker and shorter internodes, l-ribbed:
to sharply 2-ribbed cataphylls, and larger blades
(rarely less than 35 cm long, vs. rarely more than
35 ст for P. wilburii), with frequently more than 6
basal veins and 6 primary lateral veins (vs. no more
than 5 basal veins and mostly no more than 4 pairs
of primary veins for P. wilburii). In addition, P. stra-
minicaule frequently has the peduncle shorter than
the spathe and has greenish white to lavender ber-
ries (vs. having the peduncle rarely shorter than the
spathe and orangish berries in P. wilburii). Philo-
dendron wilburii is also similar to P. smithii, which
differs in having the dried stem epidermis conspic-
uously yellow-brown.
This species is named in honor of Robert Wilbur
(DUKE), who has made major contributions to the
study of Costa Rican plants and was one of the first
to collect this species.
This species is here subdivided into two varie-
ties.
KEY TO THE VARIETIES OF P. WILBURII
la. Peduncles typically equal to or shorter than the
spathe, usually less than 12 cm long; leaf blades
with sinus hippocrepiform to parabolic, usually
drying greenish to yellowish green below; Costa
aj pu Guanacaste, Heredia,
2 - var. wilburti
cm long; leaf blades with
sinus usually arcuate or arcuate with leaf tissue
decurrent on petiole, rarely hippocrepiform, usu-
ally drying reddish brown below; Costa Rica
(Cartago and Puntarenas) and Panama (Vera-
guas), 1400 m, 40—450 m along the Pacific
1
slope ____-. var. longipedunculatum Croat & Grayum
Philodendron wilburii Croat & Grayum var. wil-
..
Usually hemiepiphytic, sometimes terrestrial;
cataphylls 13-26 cm long, unribbed to bluntly
l-ribbed; petioles 16—46 ст long, obtusely flat-
tened adaxially; blades ovate-triangular, cordate to
saggitate at base, 17-36 (averaging 26) cm long,
11-27 (averaging 16) cm wide (1.3-2 times longer
than wide, averaging 1.5 times), (1.3-2 times longer
than petiole); sinus usually hippocrepiform, some-
times shallowly parabolic. INFLORESCENCE 1-
3(4) per axil; peduncle 3.5-14 cm long; spathe
7.5-13.5 cm long; spadix 9-11 cm long; pistillate
portion (2.5)3.4—4.3 cm long.
Flowering in Philodendron wilburii var. wilburii
appears to have a bimodal pattern, with the main
event occurring in the late dry season to the mid-
rainy season (March through September but es-
pecially April through August). The second flow-
ering event occurs in the early dry season, es-
pecially in January. A flowering collection from
November is unusual. Post-anthesis collections also
occur in two clusters, June through September and
November through February. Mature fruits have
been collected only in March.
Philodendron wilburii var. wilburii is largely
known from central Costa Rica to an area of the
frontier with Panama but rarely also to central Pan-
ama (Veraguas), extending from 400 to 2000 m,
mostly at over 1000 m elevation. It is largely known
from central Costa Rica, mostly at elevations of
higher than 1000 m in premontane rain, lower mon-
tane wet, and lower montane rain forests. It has
been collected mostly in the Cordillera de Tilarán,
especially in the San Ramón region and in the
Monteverde Reserve at 1350 to 1700 m (averaging
at more than 1500 m) but to a lesser extent in areas
to the east and west of the Central Valley. It has
been collected only rarely in the Cordilleras de
Guanacaste and Talamanca.
One collection believed to be this species, Croat
76799, is from Veraguas Province near Santa Fe in
Premontane moist forest. Because of the isolated na-
562
Annals of the
Missouri Botanical Garden
ture of this а раје и it se е to represent а
distinct taxon. The unusual in having
unribbed to bluntly 1- ariel Лив i ovate
to ovate-triangular, cordate, mostly greenish-drying
blades; and peduncles 3.5 to 14 cm long (averaging
9.3 cm long), usually equal to or shorter than the
spathe.
Philodendron wilburii var. wilburii might be con-
fused with a lowland form of P. wilburii var. lon-
gipedunculatum from southeastern Costa Rica that
has leaf blades more nearly as broad as long (e.g.,
Grayum 4759). Especially confusing are Grayum
4759 and 9167 from Puntarenas Province in the
region of the Osa Peninsula at 300 to 400 m ele-
vation. Both look much like P wilburii var. wilburii,
and if they prove to represent that taxon, they
would be the only collections from such a low el-
evation and the only ones from southwestern Costa
ica.
Also noteworthy is Croat 61199, a sterile collec-
tion that had its locule and ovary number deter-
mined from а field-collected infructescence. This
collection, from Monteverde in Puntarenas Prov-
ince, Costa Rica, has ovaries with only 5 locules
but 10-14 ovules per locule. It otherwise matches
panies P. wilburii. Perhaps it represents a different
spec
A с, from the Azuero Peninsula in Pan-
ama is somewhat intermediate between P. wilburii
var. wilburii and P. wilburii var. longipedunculatum.
Though it has very long peduncles like P. wilburii
var. longipedunculatum, its blades dry somewhat
blackened with the posterior lobes somewhat more
rounded than normal for P. wilburii var. longipe-
dunculatum.
Addit ional petu examined for P. wilburii var. wil-
AR Lo
ond Río Balsa, ават
т, Utley 366 (DUKE); Sai Rai dio Rodriguez, 940
m, Croat 78833 (CR, INB, MO); 1025-1100 A “78885
(CR, INB, MO); San Ramón-Cataratas, km 11, 850 m
mic 11543 (K); Cantón Alfaro Ruiz, Guadalupe,
50 m, A. Smith 2844 (F); San Luis de Zarcero, 1575
m, ну: Smith NY1023 (Е, NY); 12 km NNW
(CR, MO, WIS); 15 km NNW of San Ramén by eS 2.5
km N of Balsa on road to San Lorenzo, 1050-11 00 m,
10°11'N, 84°30'W, Liesner & Judziewicz 14753 (MO); 17-
20 km of San Ramón, ca. m, 10?13'N
84732", 14690 (MO, WIS); 1 km S of Balsa de San Ra-
món, 1200 m, 10^13'N, 84°31'W, Lent 3514 (F, MO); 9
km SE of Devin 1000 m, 1016'N, MEE Loiselle 177
mi. NW o
rcero, Croat 43634 (CM, MO); San Ra-
món-Balsa, ca. 5.7 km N of рн, over Quebrada Volio,
1100-1150 m, са. 10%08'N, O'W, Stevens 14182
(CAS, MO); 2.3 km N, Río Bala, ca. 1050-1150 m,
10%01'N, 84°30'W, Stevens =ч 98 (К, МО); Reserva For-
estal de San Катбп, са. 10
Lorencito, 850—1100 m, 10°1 18'N, 84.
15252 (MO); 800-1000 m, 10*12'53"N, 84°36'28"%, Her-
rera & Mora 179 (MO); Río San Ed Cerro Azahar, 15
km NW of San Ramón by air, 1400-1500 m, 10%09'30'N,
8434-35'W, Liesner et al. 15486 моу 15487 (СА, МО,
NY, WIS); “er n Fila Volcan Muerte, 000-1
m, 84°32'W, 10°12’N, Barringer & Gómez-Laurito 2559
| ), Barringer У box 3784 (CR. F); Мон Reserve,
Dryer 794 (MO), 997 (Е); 1500-1600 m, 10°17’N,
R, MO, MV); 10°18’N,
84°47’ W, Grayum & Sod 3851 (CR, MO).
нле дааа Platanillo, ca. 1250 m, Croat 36658
MO); SE of Turrialba, ca E of La Suiza, ca. 1200
m, Lellinger & White 1 403 (US) Tapantí Reserve, Nilsson
195 (CR), 215 (CR), 220 (CR), 238 (CR), 254 (CR), 266
(CR), 367 (CR), 647 (CR); 1250-1350 m, Ferreyra 15702
(USM); 1500-1800 m, 9?43'N, 83747", Grayum &
French 5827 (INB, МО); са. 1350 m, 9°47'N, 83*48'W,
Grayum & Sleeper 3694 (MO, US); Utley & Utley 5613
(DUKE): 1 500-1800 m, 9°43'N, 83°47'W, Croat & Gra-
yum 68292 (CR, MO, US); Croat 36122 (MO); ca. 6 km
S of Cartago by air, pro rada of
~
aprio Highway, 1620-1650 °46'N, 83°57'W,
Liesner & Judziewicz 14474 (МО); Аю Patillos, 4 Кт 5
of тирани 1480 m, Lent 1240 (F); Río Grande de Orosi,
ca. 15 km S of Tapantí, 1500 m, 9 "42N, 83 83°47'W, Burger
«e Liesner 6752 (F, NY); La Sierra, 6 km NE of Empalme,
2000 m, er 19762 (MO); Santa Teresa-Río Coliblan-
co, 6-7 km of Pacayas, ca. 1700 m, Luteyn 3246
(DUKE, Ton Quebrada Honda-Río Sombrero, c a. 1-2
km above El Мићесо, ca. 1400 m, 3236 (DUKE, MO); 31
km from San José, SE on CR-2, 1750 m, Harmon 6105
(UMO); Río Grande de Orosi valley, r Río Villegas.
1700 m, Lent 1860 (F); Río Macho, Nilsson "942 (CR), 274
(CR), 312 (CR). Guanacaste: Tilarán, Z.P. Tenorio, 1050
m, G. Rodríguez 24 (CR). peepee Río Peje-Río ак
nalito, Volcán Barva, 700— 10?17'30"N,
84^04'30"W, Grayum 6929 (MO); ~ La Paz Grande, 7.5
km N of Vara Blanca, 1270-1350 m, vw 36051 (MO);
La Selva Field uti, Valerio s.n. (USJ). Limón: Kámuk
massif, between агапа and ү Kámuk ршщ
2300 m, mias ч rs W, Davidse & Herre
29203 (CR, MO). Pun nteverde Reserve, cà.
1350 m, Croat 47139 (MO): 2 yon SW ой 1500-15
m, Ingram & Ferrell-Ingram 1734 (МО); ca. 3.5 km NE
of Monteverde Reserve, ca. 1500 m, Wilbur et al. 1581 6
(MO); Kennedy 545 (F); Haber 2369 (MO); Pounds 321
449 (MO); Gentry et al. 48860 (CR, MO, NY); Ha-
сл
es
-
a
ES
—
zZ
=
ug
О
S
R
ON
|
~
=
Y
83-66-1 (F LAS); Burger & Baker 9649 (ЕМ
таг dime 300-400 m, 8” 98: cda
jo La Нш
for road 222. «
S of San E. E Cartago, 1880 m, Utley & Utley 2948
(F); La Palma, ca. 1600 m, Standley 38300 (US); La Pal-
ma area, 1400 m, 10°12’N, 84°10’W, Burg: 4
6212 (CR, F, MO); ca. 6 km NE of San Jerónimo, Primack
et al. 322 (DUKE); Braulio Carrillo National Park,
Volume 84, Number 3
1997
Croat 563
Philodendron Subgenus Philodendron
1000-1500 m, 10%05'N, 83%57'W, Croat 61227 (CM,
MO); Río гашен Zurquí, Cerro Hondura, 1500—1600
m, 10%04'N, 84°01'W, Grayum & Sleeper 6116 (CR, МО);
Puriscal, Z.P. La pita 500 m, Morales 924 (CR, INB).
PA . Veraguas: vic. Santa Fe, along road between
Santa Fe and Alto Piedra, 1.7 mi. N of Hotel Santa Fe,
470 m, 8°31'N, 81%05'W, Croat & Zhu 76799 (В, CAS,
CR, F, HMNM, К, MO, NY, PMA, SCZ, US).
Philodendron wilburii Croat & Grayum var. lon-
gipedunculatum Croat & Grayum, var. nov.
TYPE: Costa Rica. Puntarenas: vic. of San
Vito de Java, ca. 1 km S of San Vito on road
to Villa Neily, 8°48’N, 82°57'W, 1100 m, 13
June 1987, Croat 66169 (holotype, MO-
3610645; isotypes, CR, K, NY, US). Figures
459—464
Planta plerumque terrestris, interdum ами
caulis scandens; internodia 3-15 em longa, 1.3-1.7 ст
diam.; cataphylla mollia, 7-15 cm wes књ D- x ata,
plerumque acute 2-costata, decidua; petiolus subteres vel
obtuse complanatus adaxialitof, 16-55 cm longa, 5-10
mm diam.; lamina 22-35 cm longa, 16-31 cm lata, trian-
gulari-ovata, lobis posticis angustis, patentibus; inflores-
centia 1-2; pedunculus 11-29.5 cm longus, 5-10 mm
diam. "s subteres; spatha 8.7-18 cm "unge, і saepe | viridis
eburnea
suffusa rubra; t e extus spe vel atriviridi, in-
tus atrima ro, rubriviolaceo
ri ino v ru
aut violaceo; ew (| и loculi 1-2-ovu-
lati; baccae aurantiac
Usually terrestrial, sometimes hemiepiphytic; in-
ternodes 3-15 cm long, 1 m diam.; cata-
phylls soft, 7-15 cm long, sharply D-shaped, usu-
ally sharply 2-ribbed, sometimes unribbed or
sharply 1-ribbed; petioles 16-55 cm long, 5-10
mm diam., obtusely flattened or bluntly D-shaped
adaxially; blades triangular-ovate with spreading
lobes, 22-35 cm long (averaging 28 cm), 16-31 cm
wide (averaging 21 cm; 1.03-1.4 times longer than
wide, averaging 1.3 times), (0.6-1.3 times longer
than petiole); sinus usually arcuate or arcuate with
decurrent petiole. INFLORESCENCES 1 or 2 per
axil; peduncle 11-29.5 cm long, 5-10 mm diam.;
spathe 8.7-18 cm long; spadix 6.7-11.2 cm long;
pistillate portion 2.8—5.5 cm long; ovary (5)6–7(8)-
ocular; ovules 1-2 per locule. Berries bright or-
ange (mature) or white (immature); seeds yellow-
orange, narrowly ellipsoi
The flowering phenology of Philodendron —
to be bimodal or, in any event, unlike the eua
of any other species. All flowering collections have
been made between September and April, in the
late rainy season and throughout the dry season.
However, post-anthesis flowering collections from
July make it obvious that some flowering also takes
place during June or July, perhaps both months.
Other post-anthesis collections are from January
through April and also in September. The only ma-
ture infructescence was collected in March.
Philodendron wilburii var. longipedunculatum is
known principally from eastern Costa Rica and
western Panama with an outlying population at Ta-
pantí in Cartago Province, at 900 to 1400 m ele-
vation, especially on the Atlantic slope but also on
the Pacific slope at lower elevations (40 to 450 m).
The Pacific coastal populations range from Carara
to the Osa and Burica Peninsulas in Puntarenas
Province, with outlying populations on the Azuero
Peninsula and in the Canal Area of Panama.
Philodendron wilburii var. longipedunculatum is
distinguished by the sharply two-ribbed, deciduous
cataphylls; subterete, moderately firm petioles; typ-
ically triangular-ovate blades often drying reddish
to yellowish brown and usually with narrow, spread-
ing lobes; and very long-pedunculate inflores-
cences.
The highland populations differ from those of the
lowlands in having longer posterior lobes, which
are narrower and more prominently directed out-
ward, averaging about 11 cm long (vs. about 8 cm
long for the lowland populations). The lowland pop-
ulations have proportionately narrower blades (ow-
ing to their lack of long, outward-directed lobes),
averaging 1.6 times longer than wide (vs. 1.3 times
for highland populations).
Lowland populations in the vicinity of the Burica
Peninsula have somewhat thicker blades drying
with a minutely pustular or granular lower surface
with the secretory ducts mostly not visible. In con-
trast, blades of highland populations dry darker
rown to almost blackened on the upper surface
and are smooth on the lower surface with the se-
cretory ducts clearly visible and alternating with
the minor veins.
The Azuero population, represented by Croat
34476, has leaf blades that dry somewhat black-
ened and have posterior lobes somewhat less nar-
rowed and less directed outward than those of the
highland Chiriquí populations. The Canal Area
populations, represented by Croat 12351 and Croat
& Zhu 77083, also differ in having proportionately
longer бее than material elsewhere. The latter
collection is also exceptional in having four inflo-
rescences per axil.
Grayum & Sleeper 3435 is a noteworthy collec-
tion from Tapantí, Cartago Province, Costa Rica. It
is unusual both in being from well out of the normal
range of the variety, but also in having narrower
564
Annals of the
Missouri Botanical Garden
leaf blades and shorter peduncles. Perhaps it rep-
resents another taxon.
Additional specimens examined. COSTA RICA. Car-
tago: Tapantf Region, N of Quebrada Casa Blanca, 1300
m, 9°47'N, 83°48’ W, Grayu um & Sleeper 3435 (AAU, CR,
мев - — San Vito-Ciudad Neily, Fila de Cal,
m, 841 М, 8 80°56! 30"W, Grayum et al. 6048
(CR. INB, 4 "MO, PMA); Osa Peninsula, vic. Rincón de
, 250-540 m, 8'42'М, 83°31'W, Croat & Grayum
598 79 (СЕ, MO); Quebrada Banegas—Río Riyito, ca. 7 km
W of Rincón de Osa, 100-300 m, 8°41'N, 83?33'W, Gra-
yum 4089 (CR, MO); Rincón-Agusb uena oad to Rancho
Quemado, 100 m, 8°40’N, 83°31' 6. G. Herrera 4619 (CR,
INB, MO); Finca Loma Linda, 1 mi. of Cañas Gordas,
1140 m, Croat 22310 (F, и к Piedras Blancas, da
Costeña, Fila Cruces, guciana
8°49' 18'N, 83"11'15"W, јава 120650 (CR, on мо);
Quebrada Bonita, to са. 1 km
Carara Reserve, 30-40 m,
4759 (CR, L, MEXU, MO). San p San lik de El
eneral-Dominical, 4.8 mi. S of Río Pacuare, 1000 m
ым 35249 (СЕ, К, MO, PMA, US); ca. 0.5 mi. сч
turnoff to Сападп at Rivas, 900 т, Croat 43429 (CR, МО);
ca. 8 km SW of San Isidro de El General, 1000 m, Wil-
liams et al. 28387 (Е). PANAMA. Bocas del Toro: dd
tuna-Chiriquí Grande, Continental Divide, 1200
8°44'N, 82°17'W, Croat € Grayum pens (COL, CR.
DAV, IBE, K, MO, NY, PMA, pu. i. N а. Conti-
nental Divide, 910 m, 8°44’N, 2°17 "n Croat 60454
(MO, PMA); 4.5-5 km N of dis over Factis Lake,
1100-1135 m, 8743'N, 82717", Croat & Grayum 60026
(B, CAS, CR, F, L, KYO, MEXU, MO, NY, PMA, R, S);
Cerro Colorado, ca. 7.5 mi. from Chame Camp, 1200-
1250 m, ca. 8°35'N, 81745", McPherson 8868 (MO). Ca-
nal Area: Pipeline R. Agua Salud, Croat
12351 (DUKE, F, MEXU, MO, SCZ); 6 mi. N of Gamboa,
Ri be W, Croat & Zhu 77083 (CM,
CR, MO, NY). iriquí: rica Peninsula, San Bartolo
Límite, 18 km W of Puerto a n) 450 Y dune 597
(CM, CR, MO); Rabo de Puerco, 8 km W of Puerto Aru-
muelles, 50-150 m, Croat 21958 (F, МО); po m, 22487
(Б MO); Cerro Colocado 34—35.6 km above Río San pan
90-1410 m, Croat 37250 (F, MO, NY, PMA); 28
ibo San Félix, 1200-1500 m, 33202 (MO); Cualaca-
rtuna, 10 mi. NW of Los Planes de Hornito,
82°17' W, ras, Croat 50093 (L, MO, NY); 5. oi mi. NW
es de Hornito, 1370 m, 843'N, 82*15'W,
Croat 40852 (MO); 7.2 mi. beyond Los Planes de Hornito,
165-1200 m, 8°44'N, 82714", 67836 (CM, F, MO,
PMA); 11.8 mi. N of Los Planes de зора 1400 m,
48696 (MO); near Lago Fortuna,
82°15'W, iier, 9081 (MO. ‚ US);
3.4 km N of Sui Chorro mi. N of center of
bridge over lake, 1205 m, 8°43" N, сем. W, Croat 74955
У eo
Pirre, 800—1,050 m, 7?56'N, 77°45'W, Croat е (МО).
antos: Azuero Peninsula, Jobero-Río Pedregal
— 300-700 m, Croat 34476 (MO, PMA). Pan-
: Río Majé, ca. 4—5 mi. above waterfalls, near new
Sivan Lake, <100 m, Croat 34726 (MO).
Philodendron zhuanum Croat, sp. nov. TYPE:
— Coclé: Alto Calvario, vic. El Copé,
m N of El Copé, 770 m, 8°38'N,
hou W, 12 July 1994, Croat & Zhu 76755
(holotype, MO-4619357-9; isotypes, AAU, В,
CAS, COL, CR, F, GH, K, MEXU, NY, PMA,
QCNE, SCZ, US, VEN). Figures 465—468.
Planta hemiepiphytica; internodia 2-4 cm longa, 3-5.5
diam.; cataphylla 37-50 cm longa, acute 2-costata,
decidua; petiolus 40-63 cm longus, obtuse D-formatus, in
sicco manifeste complanatus usque 3-6 cm latus; lamina
anguste ovato-sagittata, 38-62 cm longa, 27-32 cm lata;
tu
tus viridi, intus чалу VERUS ва Др Зи pistilla 8-
9-locularia; Јосић 1-3-ovulat
Hemiepiphytic, loosely or appressed-climbing;
leaf scars conspicuous, 3 cm long, 2 cm wide; in-
ternodes semiglossy, 2-4 cm long, 3-5.5 cm diam.,
dark green, drying dark brown, longitudinally wrin-
kled; roots drying light brown and weakly glossy,
finely striate, epidermis peeling; cataphylls 37-50
cm long, sharply 2-ribbed, medium green, semi-
glossy, drying dark brown, deciduous; petioles 40–
63 cm long (0.8-1.5 times longer than blades),
bluntly D-shaped, faintly raised medially, moder-
ately firm, medium green, surface dark green short-
lineate, drying dark yellowish brown and glossy,
prominently flattening to 3-6 cm wide and only a
few mm thick, smooth to finely and closely striate;
sheath 3-6 cm long, moderately closed; blades
narrowly ovate-sagittate, moderately coriaceous,
acuminate at apex, conspicuously sagittate at base,
38-62 ст long, 27-32 cm wide (1.8-2 times longer
than wide), margins broadly undulate, much paler
than surface, upper surface dark green, glossy, dry-
ing dark yellowish brown, lower surface weakly
glossy, paler, drying reddish brown to yellowish
green with conspicuous, dark intermittent sec
canals visible; anterior lobe 34—35 cm long, mar
gins broadly convex, paler; posterior lobes 8. 5-18. 5
ст long, 7-13 ст wide, directed downward to
slightly inward, broadly rounded; sinus narrow, spa
thulate to narrowly obovate, (7.5)15-19 cm deep.
1-3 cm wide; major veins drying with sharp ridges.
usually darker than surface; midrib flat, paler than
surface above, narrowly rounded, concolorous ђе-
low; basal veins (3)5-6 per side, with the first free
to base, numbers 3-5 coalesced (2)5-7 cm; poste-
rior rib not at all naked or naked to 2.5 cm; primary
lateral veins 6—7 per side, arising at an acute angle
A a —— —
Моште 84, Митбег 3
1997
Croat 565
Philodendron Subgenus Philodendron
then spreading to a 40-45” angle, weakly arcuate
until near the margin, then turned up abruptly,
weakly sunken above, convex, paler than surface
below; minor veins clearly visible, arising from both
the midrib and primary lateral veins, drying prom-
inulous above, faint and weakly wrinkled below.
INFLORESCENCES 3 per axil; peduncle 9-14 cm
long, 1-1.5 cm diam., somewhat flattened, medium
green, faintly short-lineate; spathe 13-18.5 cm
long, margins paler; spathe blade white, except yel-
low-green medially on back side outside, tinged
magenta on lower one-third inside; spathe tube me-
dium green outside, dark violet-purple inside, spa-
dix 11.5-18 cm long; pistillate portion creamy
green, 3.5—8 cm long, 3-6 ст long in back, 1.5
mm diam. at apex, 1.9 mm diam. at middle, 1.8
mm wide at base; staminate portion 8—10 cm long;
fertile staminate portion slightly narrower than the
pistillate portion; sterile staminate portion barely
distinguishable when fresh, 1.2-1.3 cm diam. at
base, tapered only slightly upward when fresh (dry-
ing to 10 mm diam. at apex); pistils 1.9-2.1 mm
long, 1.2-1.6 mm diam.; ovary 8—9-locular, 0.8 mm
long, with basal placentation; locules 0.8-1.2 mm
long; ovules 1-3 per locule, translucent, contained
within gelatinous matrix (no true envelope), 0.3—
0.4 mm long, equal in length to funicle, style sim-
ilar to style type B; stylar canals emerging directly
gularly 4—6 sided, 0.8-1.2 mm diam.
at apex; ende staminate flowers irregularly round-
ed, 0.8-1 mm e.
Flowering in BEA zhuanum is docu-
mented by a single collection from July, at the be-
ginning of the rainy season. A cultivated plant of
the same collection flowered in November at the
Missouri Botanical Garden
Philodendron zhuanum is endemic to Panama in
Coclé Province in an area of Premontane rain forest
at 700 to 900 m elevation.
Philodendron zhuanum is a member of P sect.
Calostigma subsect. Macrobelium ser. Macrobel-
ium. This species is distinguished by its short,
thick internodes; deciduous, sharply two-ribbed
rini. bluntly D-shaped petioles about as long
e blades or slightly shorter; narrowly ovate
blades with prominent posterior lobes, a narrow si-
nus and prominently visible secretory canals; up to
three inflorescences per axil; and spathes green
outside and whitish inside on the blade, and dark
purple-violet inside on the tube.
Philodendron zhuanum is apparently related to
P. sagittifolium and P. aromaticum, differing from
both in having petioles which on drying become
moderately spongiose, markedly flattened (especial-
ly in the middle portions), dark yellow-brown to
yellowish, and often glossy. It also differs in having
conspicuous secretory canals visible in both living
and dried conditions. While secretory canals are
present in both P. sagittifolium and P. aromaticum,
they are quite inconspicuous.
This species is perhaps most easily confused
with P. aromaticum, which has similar leaf blades
with a naked posterior rib. The latter species occurs
at lower elevations and nearer the coast. In con-
trast, P. zhuanum rarely has the posterior rib naked.
While P. zhuanum has the spathe tube dark purple-
violet inside with the same color bleeding onto the
lower one-third of the blade, the spathe tube of P.
aromaticum is white to weakly maroon within and
the color does not merge onto the lower part of the
blade.
Philodendron zhuanum is named in honor of
Guanghua Zhu, my student, field companion, and
a monographer of the genus Dracontium, who
helped to collect the type and many other aroids
during our fieldwork in Panama.
Additional specimen examined. PANAMA. Coclé:
Alto Calvario, vic. El Copé, ca. 5 mi. N of El Соре,
900—1000 m, Croat 75054 (CM, MO).
NOMEN INCERTAE SEDIS
Philodendron auritum Lindl., J. Hort. Soc. London
This is perhaps an older name for Philodendron
anisotomum but was based on a cultivated plant
collected by Skinner, perhaps in Guatemala, and
no specimen or illustration is known to exist to con-
firm that it is even a Philodendron. Lindley stated
that it had foliage very similar to a Syngonium il-
lustrated by Vellozo (1825) in Flora Fluminensis
(vol. 10, t. 113).
ExcLUDED NAMES
Philodendron armigerum Standl. & L. O. Williams,
Ceiba 3: 107. 1952. — Syngonium armigerum
(Standl. & L. O. Williams) Croat, Ann. Mis-
souri Bot. Gard. 68: 585. 1981.
Philodendron brevinodum Standl. & L. O. Williams,
Ceiba 1: 231. 1951. — Monstera tuberculata
Lundell var. brevinoda (Standl. & L. O. Wil-
liams) Madison, Contr. Gray Herb. 207: 92.
1977.
Philodendron hastiferum Standl. & L. O. Williams,
Ceiba 1: 232. 1951. — Syngonium hastiferum
Annals of the
Missouri Botanical Garden
(Standl. & L. O. Williams) Croat, Ann. Mis-
souri Bot. Gard. 68: 595. 1981.
Literature Cited
Armbruster, W. S. 1984. The role of resin in angios
"mre Ecological and chemical considerations.
pes 1149-1160.
937. The oe of Petén. Publ. Car-
negie Inst. © 478:
1995[1
Bay, D. C. 996]. P И in the aroids. Aro-
ideana 18: 32 né
Berlin, B. & Р. 1969. Basic Color Terms, Their Uni-
versality and Русоа. Univ. California Press, Berke-
ley.
виш: М. 1951. The Cultivated Aroids. Gillick Press,
Berkeley, California.
Blanc, P. 1977a. Contribution à l'étude Aracées. I. Re-
marques sur la croissance monopodiale. Rev. Gén. Bot.
84: 115-126.
xr 7b. Contribution à l'étude Aracées. П. Re-
sur la croissance sympodiale ee Мере
krd Engl., le Philodendron fenzlii Engl.,
Philodendron speciosum Schott. Rev. Сеп. Вог. 84: 3
al:
. 19772. Contribution à l'étude Aracées. П. Re-
marques sur la croissance monopodiale. Rev. Gén. Bot.
84: 115-126
1 Aspects de la ramification chez des Ar-
acées tropicales. Théses du Diplome de Docteur З?"
Cycle, ve or = & М. Curie, Paris.
1980. rvations sur les flagelles des Ara-
сеае. ‘Adansonia Ser 2, 20: 325-3 чм
Bogner, Н. & D. Н. Nicolson. 1. A revised classifi-
cation of Araceae with dichotomous ane Willdenowia
: 35—50.
. J. D. Meeuse & J. R. Klima. 1971.
f blooming in Sauromatum guttatum
Schott by darkness. Canad. J. Bot. 49: 1025-1031.
Bunting, G. S. 1963a. Studies in Araceae. Ann. Missouri
Bot. Gard. 50: 23-28.
3b. A reconsideration of Philodendron hed-
eraceum. ош ска 2–6
8 5: Commentary i on Mexican Araceae. Gentes
Herb. 9: 289-3 382.
1968. Vegetative anatomy х i Adr
scandens complex. Gentes Herb. 1
. 1979. Sinopsis de las кы. d cali
Rev. i = (Maracay) 10: 139—
w taxa of Venezuelan Atinede: Phyto-
logia d зыт
. New taxa of Venezuelan Araceae—II.
Phytologi 64: par
. Ara p. 600–6 n: Ј. Steyermark,
E "e & B. Holst Pb abe, Flora of the Venezuelan
Guayana. Vol. 2: Acanth
n, St. Louis.
Floral AMAT of the Pothoideae
and Monsteroideae (Araceae). Ph.D. Dissertation, Mi-
ami و Oxford, Ohio.
"— Phenological behavior of habit a
n Barro Colorado Island (Panama Сила
Zone). пано 7: 270-277.
ога of "i Colorado Island (Panama
Canal m Stanford Uni ss, Stanford.
———. 9. The учим: of ehe Pp. 291-308
n K. Larsen & L. B. Holm-Nielsen (editors), Tropical
те Academic Press, Lon
—— owering behavior of P pA ge-
nus Anthurium (Araceae). Amer. J. B 7: 888—904.
981119 ed А revision of ace (Araceae).
Ann. verti Bot. Gard. 68: 565—651.
——— 1983 ^ revision of Anthurium d of
P
rt 1. Mexico and Mid-
"D
=
. Pp. 234-236 i
es (editor? Costa Rican fund History. Univ. Chi-
o Press, Chicago.
1985a. Collecting and preparing specimens of
Áraceae. Ann. Missouri Bot. Gard. 72: 252-258.
1986a. A revision of Anthurium (Araceae) for
Mexico and Central America. Part II. Pa
Syst. Bot. Missouri Bot. Gard. 14.
1986b. The distribution of Anthurium ee
in Media: Middle America and Panama. Selbyana 9:
ope
апата. Monogr.
————. 1988[1990]. Ecology and life forms of Araceae.
m 11: 4-55.
—. 1990[1992]. A comparison of aroid classification
edt Aroideana 13: 44—63.
1991. A revision of Anthurium sect. Pachyneu-
rium (Araceae). Ann. Missouri Bot. Gard. 78: 539-855.
2. Species diversity of Araceae in Colombia:
: preliminary survey. Ann. Missouri Bot. Gard. 79: 17-
G. S. Bunting. 1979. Зори of Ап-
thurium descriptions. Aroideana 2:
& N. 86. The аб of Venezuela.
Aroideana eee =
Crosby, M. R. à x Muscorum—A compu m
muscological deans: Bull. Brit. Bryol. Soc. 48: 2
26.
—& R. Magill. 1986. TROPICOS: The botanical
database at the т Botanical Garden.
Missouri
Botanical еа uis
Cullen, J. 197 E preliminary survey of ptyxix (verna-
tion) in the preci s Roy. Bot. Gard. Edin-
burgh 37: 161-214.
1. A review of the fossil record of
Dahlgren, R. M. T. & H.
و A Comparativa Study. Academic Press,
Davidse, G., M. Sousa S. & S. Kna 1995. In: С. Dav-
idse, M. Sousa S. & A. O. Chace (editors), Flora Me-
soamericana, Vol. 1, Psilotaceae-Salviniaceae. Univer-
sidad Nacional Анбар de México, Mexico 1.
Missouri Botanical Garden, St. Louis; The Natural His-
tory Museum, London
Dilcher, D. L. & С. P. Daghlian. 1977. Investigations of
angiosperms from the Eocene of southeastern No
America: Philodendron leaf remains. Amer. J. Вог.
926-534.
Dodson, C. & A. Gentry. 1978. Flora of the Rio Palenque
AVECH Center: Los Ríos Province, Ecuador. Selbyana
41: 1—628.
, ——— & F. Valverde. 1985. Flora de Juaneche.
Ediciones del tere del Ecuador, Quito
Dugand, A. . Revaluación de Philodendron heder-
aceum seg como transferencia de Arum hederaceum.
Caldasia 3: 445—452.
Endlicher, S. 1837. Genera Plantarum 1(3). Vienna d
Endress, P. K. 1982. Syncarpy and alternative modes
Volume 84, Number 3
1997
Croat 567
Philodendron Subgenus Philodendron
escaping disadvantages a apocarpy in primitive angio-
1: 4
A. 1 ur Morphologie der Araceae. Bot. Zei-
tung (Berlin) 34: 81-90. 9
1877. Vergleichende ig suchungen = r
бардан Verhältnisse der Araceae. И.
i йы und Sprossverhültnisse der Meca Noki
Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 39:
Sri
. Araceae. Pp. 25-244 in: C. F. P. von Mar-
tius “төн Flora brasiliensis 32A). n бышы Leip-
zig.
. 1879. Araceae. In: A. & С. De Candolle (edi-
tors), Moográphie Phanerogamarum 2: 1—681. Paris.
—— 18 Beitráge zur Kenntnis der Araceae. IX.
16. Revision се Gattung Philodendron Schott. Bot.
Jahrb. а 6: 509—564.
ers a. Beiträge zur Kenntnis der Araceae. X.
18. ier novae. Bot. Jahrb. Syst. 37: 110-143.
P . Araceae—Pothoideae. In: A. Engler (ed-
itor), ipis Pflanzenreich IV.23B (Heft 21): 1-330. W.
Engelmann, Leipzig and Berlin.
—— ———. 1908. Additamentum ad Araceas—Pothoideas.
Pp. 1-3 in: A. Engler (editor), Das Pflanzenreich IV.23B
(Heft 37). W. Engelmann, Leipzig and Berlin.
911. Araceae-Lasioideae. /n: A. Engler (edi-
tor), Das Pflanzenreich IV.23B (Heft 21): 1-310.
—"FuIA, Xi Ме cena n et, a
dreae-H
Engler oaa Das Pflanzenreich 1V.23 (Heft 55): ү
gelmann, Leipzig and Berlin
915. Araceac-Philodendroideae- Anubiadae-
i^ eot PEU In: А. e (editor), Da
Wise prin IV.23De (Heft 64): 1
a. Araceae, pars Mae et Index familiae
generalis. p A. Engler (editor), Das Pflanzenreich
IV.23A е 74): 1-71.
20b. Araceae—Aroideae und Pistioideae. In:
A Eger (editor), Das Pflanzenreich IV.23F (Heft 73):
ES K. Krause. 1908. Araceae—Monsteroideae. Pp.
4-138. In: A. дё ә (editor), Das Pflanzenreich IV.23
(Heft ae
О. Araceae—Colocasioideae. In: A.
Engler Game P5 Pilanxcoreich IV. (Heft 71e): 1–
139
Eyde, R. H., D. H. Nicolson & P. Sherwin. 1967. A sur-
vey of floral anatomy in Araceae. Amer. J. Bot. 54: 478—
497
Faegri, K. & L. van der Pijl. 1979. The Principles of
Pollination Ecology. Pergamon Press, London.
Flores, G., L. Jiménez, X. Ma drigal, R. Moncayo & F.
Takaaki, 1971. Mapa Doe de Vegetación de la
Repüblica México. “i 200,000. Secretaria de Recursos
Hidraulicos, México.
Foster, M. B. 1949. My flower has a temperature! Natl.
3.
1985. Patterns of endothecial wall thick-
enings in Araceae: Subfamilies череда, Lasioideae,
and Philodendroideae. Bot. Gaz 1533:
UN Patterns of samen paros in the
Araceae. ташы Ј. Во
al vods ни v Ariidae Bot. Gaz.
147: 478-495,
- 1987a. Systematic occurrence of sclerotic hy-
podermis in roots of Araceae. Amer. J. Bot. 74: 891—
1987b. The structure of ovular and placental
trichomes of Araceae Ба. Gas 148: 198-208.
1987c
y of resin canals in roots
of нац ~ ane 148; 360–371.
988. Systematic occurrence of anastomosing la-
бейи in Araceae. Вог. Gaz. 149: 71—81.
& Р. В. Tomlinson. 1980. Preliminary observa-
tions on the vascular system = stems of certain Ara-
ceae. Pp. 105-116, pl. 1-9. In: C. D. Brickell, D. Е
Cutler & M. Gregory cir), Petaloid Monocotyle-
dons. Academic Press,
ide patterns in stems of
Vas
Araceae: већину Philodendroideae. Bot. Gaz. 142:
50-563.
& ———. 1984. Patterns ip ge vasculature in
Philodendron. Amer. J. Bot. 71: -1443.
— ——, M. Chung & Y. Hur. deer ginem DNA
phylogeny of am A a оно in Р. J. Rudall, Р.
J. Cribb, D. Е Cut J. Humphries (editors),
Monocotyledons: Systematics & rr Royal Bo-
tanic Gardens,
Gentry, A. H. 1982. Evidence for phytogeographic pat-
ы, as evidence (ог a Сћосб refuge. Pp. 112-136 in
G. T. Prance (editor), Biological Diversification in the
Tropics. Columbia Univ. Press, New York.
Goebel, K. & W. Sandt. 1930. Untersuchungen an Luf-
twurzeln. Bot. Abh. 17: 1—124.
Gottsberger, G
Philodendron species. Ber. Deutsc
41
Pollination strategies in Brazilian
h. Bot. Ges. 97: 391—
———, 1986. Warmeentwicklung von Philodendron-
Bluten. Naturwiss. Rundschau (Stuttgart) 39: 350-351.
1990. Flowers and beetles in South American
tropics. Ber. Deutsch. Bot. Ges. 103: 360-365
— & A. Amaral, Jr. 1984. Pollination strategies in
Brazilian Philodendron species. Ber. Deutsch. Bot. Ges.
97: 391-410.
Si Iberbauer-Gottsberger. 1991. Olfactory
and у al attraction of Erioscelis emarginata (Cyclo-
e Dynastinae) to inflorescences of Philo-
dendron selloum €" Biotropica 23: 23-28.
Grayum, M. H. 1984. Palynology and Phenology of the
Araceae. Ph. = Dissertation, University of Massachu-
setts, Amhers'
1985. Бани and ecological significance
of a storage in pollen of the Araceae. Amer. J. Bot.
72: 1565-1577
arene between pollination biology
n.
А € and phylogeny of Araceae. Ann.
Missouri Bot. Gard. 7 8-697.
1991. Кы, embryology of the Araceae.
Bot. Rev. 57: 167-203.
992a. Comparative external pollen s ond
ture of the Araceae and putatively = taxa. Mon
Syst. Bot. Missouri Bot. Gard. 43
New species of 68788 og
Te (Araceae) from rea and Pacific
uth America. Phytologia 73: 30—
. 1996. Revision of din subgenus Pter-
bmüchum (Araceae) for Pacific and Caribbean Tropical
America. Syst. Bot. Monogr. 47: 1-233.
568
Annals of the
Missouri Botanical Garden
Gregor, H.-J. & J. Bogner. 1984. Fossile Araceen onm
leuropas und ihre rezenten Vergleichsformen. Doc
menta Naturae 19: 1–12.
Greuter, W., F. R. Barrie, H. M. Burdet, W. G. Chaloner,
o pias ge of Botanical Nomenclature (Tokyo
egnum
Hay, P & D Mabberley. 1991. Transference of func-
tion and the origin of aroids: Their significance in early
angiosperm evolution. Bot. Jahrb. Syst. 113: 339-428.
Henry, M. F. & E. J. Nyns. 1975. Cyanide insensitive
respiration. An ene mitochondrial pathway. Sub-
ell. Biochem
Herk, A. W. Hs van. ра Die chemischen Vorgänge
im Sauromatum-Kolben. I. Rec. Trav. Bot. Néerl. 34:
69-156.
Die chemischen Vorgiinge im Sauroma-
—. 1937b.
tum- kolben II. Proc. Kon. Ned. Akad. Wetensch. 40:
607-6
937c. Die chemischen Vorgünge im Sauroma-
tum- Kolben. III. Proc. Kon. Ned. Akad. Wetensch. 40:
709-719
Hermann, Р 1698. Pp. 69–95 in Paradisus batavus. Lei-
en.
Holmes, J. W. 1969. On the absolute fall of sea-level
during the Quaternary. Palaeogeogr. Palaeoclimatol. Pa-
laeoecol. 6: 237-239,
Holdridge, L. R. 1967. Life zone ecology. Tropical Sci-
ence Center, ps p Costa Rica.
‚ W. C. Grenke, W. H. Hatheway, T. Liang & J. A
Tosi, Tr. 1971. jen Environments in Tropical Zones.
Pergamon Press, Oxford.
Hotta, M. 1970. A system of the family Araceae in aom
and adjacent areas. Mem. Fac. Sci. Kyoto Univ., Ser.
Biol. 4: 72-96.
D 71. A Glossary of Botanic Terms.
Fourth Ed. Duckworth, London.
60
31. Enumeratio Syst Plan-
240, Pl. 152. Selectarum Stirpium
Americanárum Паша. Ex officina Kransiana, Vienna.
. 1797. Plantarum Rariorum Horti Семе
Schoenbrunnensis. Vienna
James, W. & H. Beevers. 1950. The respiration of Arum
spadix. New So 49: 353-374.
9. The botany of San José Island (Gulf
of Panama). pai 8: 1-306.
: Ena press. Vol. 10. Araceae. In: D. F. Cut
ry (editors), Anatomy of the Mile.
dons. Clarendon Press, Oxford.
Kelly, N. Epiphytes and climbers of a Jamaican
e a Хаи distribution, life forms and life his-
s. J. ree 12: 223-241.
кылы, R. M. Heat production and temperature
regulations in eastern skunk cabbage. Science 186:
746-747.
Koch, K. 1853. Araceae. P. 14 in Index Seminum in horto
botanico berolinensi Coluntur, Appendix. Berlin.
Kraus, G. 1884. Uber die Blütenwürme bei Arum ilali-
cum. a Naturf. Ges. Halle 16: 746-747.
Рио aus den Tropen. Ann.
Jard. = oe 8: 217-275.
Krause, K. 19 Araceae Philodendroideae-Philoden-
dreae-Philodendrin n: A, er & K. Krause (ed-
itors), Das Pench IV23Db (Heft 60): 1-143. W.
Engelmann, Leipz
Kunth, C. S. 1841. Araceae. Pp. in Enumeratio
Plantarum, Vol. III. J. G. Md "bata & Tubingen.
1910. Untersuchungen über die Bliitenwárme
1916. Die
ihre Blütenbiologische Deutung. Ber. Deutsch. Bot
CREE vhs 3
u, M. Über de Wurzeln der Araceen. Bot.
Jahrb. а 9: 1-38.
Linnaeus, С. 1754. Genera plantarum, ed. 5. Stockholm.
Madison, M. 1977. A revison of Monstera (Araceae).
Contr. Gray Herb. 207: 3-100.
978. Genera of Araceae in the northern Andes.
Fas ка 1; 31-53.
Matuda, E. 1954. Las Araceae Mexicanas. мн Inst.
iol. Univ. M Auton. Mexico, Bot. 25:
2 Nuevas plantas de Mexico. se Inst.
Biol. Univ. ye Auton. Mexico, Bot. 32: 143-155.
Mayo, S. J. 1986. Systematics of Philodendron Schott
(Araceae) with Special Reference to Inflorescence Char-
an
ução grafia d
género Philodendron Schott Пена PR не Bot. Brasil.
1(2) Sup
cial structure in
100: 139-
of gynoec ا
Philodendron EN J. Linn. Soc. Bot.
aa ee story and infrageneric pr ees. of
ШОХ esos Kew Bull. 45: 37—
e, . A revision of Philodendron Ei Me-
RIRA Азы ey Kew Bull. 46: 601-681.
————, J. Bogner & P. C. Boyce. 1995. The Arales. Pp.
277-286 in P. J. Rudall, P. J. Cribb, D. F. Cutler & C.
J. Humphries (editors), Monocotyledons: Systematics &
Evolution. Royal Botanic a , Kew.
& "The pny of Ara-
e. Royal B Botanical сна A In
йан ееиѕе, В. J. D. 1966. The voodoo lily. Sei. pu 215:
19 75. Thermogenic respiration in aroids. An-
В кь. Pl. Sci. 26: 117- 126.
. The physiology of some ree ous
: A. Ri JANE (editor, The Pollination of
Flowers br Insects. Academic Press, London
. G. Buggeln. 1969. Time, space, ‘Tight and
darkness in the metabolic flare-up of the spadix of Sau-
romatum appendix. Acta Bot. Neerl. 18: 159- 1.
Moodie, С. Е. E. 1976. Heat production and pollination
in Araceae. Canad. J. Bot. 54: 545-546.
Кару, K. A., D. K. Odell & R. 5. Seymour. 1972. Tem
perature он by the inflorescence of родни
Science 178: 1195-1197.
Nicolson, D. H. 1960. A i review of classifications in
the Araceae. Baileya 8: 62-67.
Report of de General Committee: 6. Tax-
on 43: 279-281.
Plumier, C. 1756. Plantarum americanum fascicularis
rimus. Fascicle 2. t. 51.
Pohl, F. 1932. Anatomische und ókologische Untersu-
chungen am Blütenstünde von — sein
Schott, mit besonderer Berücksichti arzkan
üle und der Beschaffenheit der Pollenkittstoffe. Planta
Porch. 0. 1911. Die Anatomie der Nahr und Haftwur-
zeln von Philodendron selloum С. Koch. Denkschr.
Kaiserl. venu Wiss., Math.-Naturwiss. Kl. 79: 390-
451, tt. 244
e ———— ا 0
Field Mus. ne Hist., Bot. Ser. 18: 131-1
-—— 19 Wi
Volume 84, Number 3
1997
Croat 569
Philodendron Subgenus Philodendron
Putz, F. E. & N. M. Holbrook. 1986. Notes on the natural
history of hemiepiphytes. Selbyana 9: 61—69.
Raskin, I. 1992. Salic
Physiol. 99: Sasman
ylate, a new plant hormone. Pl.
er & T R. Melander. 1989. deci
tion of heat send n the inflorescences of an aru
lily by завесе ме acid. Proc. Natl. Acad. Sci.
U.S.A. 86: 2 18.
———, А. E xen R. Melander & B. J. D. Meeuse.
1987. Salicylic POM А natural inducer of heat produc-
tion in Arum lilies. Science 237: 1601-1602.
D. I. Axelrod. 1974. Angiosperm bioge-
ography Se - continental movements. Ann. Missouri
Bot. Gard. 61: 539-673.
Ray, T. S. 1986. Growth correlations within the segment
in the sima Amer. J. Bot. 73: 993-1001.
. Leaf types in the Araceae. Amer. J. Bot.
74: E
1987b. Diversity of shoot organization in the
Anche. Amer. J. Bot. 74: 1373-1387.
ج i. xig Survey of shoot organization in the Ara-
r. J. Bot. 75: 56-84.
Risas А 1971. Morphologische Untersuchungen
ur Wuchsform von Philodendron. Bot. Jahrb. Syst. 90:
27-649.
Schatz, G. 1990. Chapter 7. Some aspects of pollination
bi rid in р American forests. Pp. 69-84 in
adley (editors), Reproductive йе бин
of Това Е = der Parthenon Publ. Group, Park
Ridge, New
Schott, H. W. 1629.1 Fi iir Liebhaber der Botanik. Wiener
Z. Kunst 1829: 7
1
832. hae raceae. Pp. 16-22 in H. Schott
Endlicher, Mu botanica. Typis Caroli Gerold,
ienna.
856 6. Synopsis aroidearum complectens, Enu-
merationem а icum generum et specierum hujus
ordinus. Typis Congregationis Mechitharisticae, Vienna.
pem 1858 arum. Typis Caroli Ueber-
reuter. Prostat ne apud Ed Hólzel, Vien
chaos aroidearu m. Typis
Congregationis Mechithari cae, Vienna.
Seymour, S5 G А. чет АЫ & М. С. Barnhart.
983. Respiration and heat production by the inflores-
cence of Philodendron selloum Koch. Planta 157: 336–
343.
a С. — & С. hs — 1984. Re-
rmogeniesis in Philo-
: 229-
: «d. r. 1928. Sys — € Ana-
tomie der Monokotyledonen. Heft 13: 1
Sprengel, K. P. J. In: ud desc
vegetabilium. Ed. 16, 3. Gótting
Standley, P. C. 1937. Flora of bus Rica, Part 1. Publ.
Systema
raceae. In: R. Е. Woodso akan
Schery (e edt Flora d Panama. Ann. Missouri Bot.
ard.
—— & J. A Send. 1958a. Studies of Central
oe a Publ. Field Mus. Nat. Hist., Bot.
Ser. 2
58b. Araceae. Pp. 304-363 in:
Flora of Guatemala: Part 1. Fieldiana, Bot. 24.
Takhtajan, A. 1969. Flowering Plants: Origin and Dis-
persal. Smithsonian Institution Press, Washington, D.C.
Tieghem, P. van. 1907 ee sur la енн од des
Aroidees. Ann. oma „ ser. 9, 5: 312-320.
Torre, P. as prada de la чийет де
Рапата. ia del Atlas de Panamá, Atlas de Pan-
am
аи ТОР 1700. i pt rei herbariae, editio
altera. 1: 158-162. Par
1898. місце ا Borrtraegero, Ber-
Urdentlich, A., R. A. Linzer & I. Raskin. 1991. Alter-
native respiration ape heat evolution in plants. Pl.
Physiol. S 1545-1550.
Usher, G. BA ee of Botany. D. Van Nostrand,
UN New
Van der Pijl, L. 1937. Biological and physiological ob-
servations on the inflorescence of Amorphophallus. Re-
cueil Trav. Bot. Néerl. 34: 157-167.
Vellozo, J. M. da Conceição. 1825 [1829]. Florae Flu-
miensis, Archivos do Museo Nacional Flumine Janu-
ario. [Text corresponding to Vols. 1-8 of the plates.]
PA EMI jag des plantes nouvelles
peu connues. Crapelet, Pari
Walker, D. B., J. Gy Lb. рет & M. J. DeNiro. 1983.
Direct respiration of lipids during heat production in
inflorescence of Philodendron selloum. Бейне 220:
9–421
41 і
Warming, E. 1867. Nogle lagttagelser over Varmeudvik-
linger hos en Aroidee, Philodendron lundii. Vidensk.
Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1867(8—
11): 127-144. t
1883.
Tro pische Fragmente. I. Die она
von Philodendron bipinnatifidum Schott. Bot. Jahrb
à ` Ueber Наћ- und Nahrwurzeln
bei Kieran и und Epiphyten. Ann. Jard. Bot. Bui-
tenzorg 12: 1—7
еа c E. lins In: Caroli a Linné Species Plan-
rum. . .Editio qua
You e H. J. 1986. r pollination of Dieffenbachia
gs (Araceae). Amer. J. Bot. 73: 931-944.
7. Aroid observations: Philodendron roths-
ie | no
1988a. Differential importance of beetle species
pollinating s ei longispatha (Araceae). Ecol-
: 832
1988b. Neighborhood size in a beetle-pollinated
tropical aroid: rs of low density and asynchronous
ering. Oecologia 76: 461—466.
1990. Chapter 11. Pollination and reproductive
biology of an understory neotropical aroid. In: K. S.
Bawa & M. Hadley (editors), Reproductive Ecology of
Tropical Forest Plants. Parthenon Publ. Group, Park
Ridge, New Jersey.
Zavada, М. 5. 1983. С phology of monocot
llen and evolutionary sein of apertures and wall
structures. Bot. Rev. (Lancaster) 49: 331-379.
570
Annals of the
Missouri Botanical Garden
Philodendron Subg. Philodendron.
of Central American
Appendix 1. Geographic Distribution
BRAZIL PERU) BOLIVIA! CUBA | TRINIDA|
GU
Volume 84, Number 3 Croat
1997 Philodendron Subgenus Philodendron
á ~ т
ЈЕ
Е
< |= = =
3
=
o
ш
Ф T тт
Ё
iJ — T
i
1
о
2
=> + اا bl paio = سس —
Es
BI
A is Ds pages a
cutres ИРА АИ al |
E ; 5
3 3 HE
É JHE E
P 5| |= Ый ре 28) le
E o5 S ооо |g 3 58| (E
E [LIBE нн ТИЕШ
* || {ШЕ TERBHEEEBEEEEEEE
571
Missouri Botanical Garden
Annals of the
572
aum E it E NE NM 55 забаазмл |
UN E E иш E ___ EOL |еаоаавапе 9 S303dS VU
8 ИШ шш = = E E
2 E. -. EE Р 5 Je SL (9 пт uu S 8 aa. 14802 NOXV.L
"m UM SU v |^ AN unuenuz
maa= зн нл ш ш unjejounpadiBuo| ел
ПШ um Uu UB UNS NN шпајм зел пара
mim = | | |. пи 7 IIzolwezsiem
L | Fu ng NUN US ES UN NN шпаоотшел
ИШ ЫШ = DN E 13 7 asuazedelaa
q poo. шпиеќәрп
am ЕЕ ЕЕ. eT E ^ REESE asuadnyuebiqn
mE UE хп uU m UM поз
| TF UU пш ш ш ш unyyedin
nm EE S ES E NM unoissejeuy
E Ur Ib mg nm E S S UM anua)
| nm EE S S NM NUM ejneorons
mE ШШ ЫШ 3 — 1 unsiouiqns
| ЫШЫ um H US US жш ш unionis
YE Rk Бил |» ШШ ш ejeaiuuens
ZEE mm NE US ШШ me Mu wine|onadiwenbs
[DH гаш иш D шл .» Nm s eneonuenbs
|] E ИШ ШО ИШ ИШ E ! aesnos
muuu ШШ E E NM D uu г nyus
во o EX EN unuemouos
= ПШ EE ШШ zm Nm Em enous
1 a mnm E E NM ш UM r штцојдубез
TJ n m = um mm eus. unueiqnuosuyoi]
HL] d à. |. | | = | E Е unjeuiue|nsnBue зел
рагавы ^. ош шаш ви . '08$01 Јел umujedsoaso!
mum um NE y m uw rom unjeipeiopnasd јел
Ex Um (ОШ E mm пш ш 71 uumeıpeı зел Uungerpei
zum mm : mz [Im L asuayjrund
амы yana vanoa (nad пуна no noa UM Nd мо Ма ON NA PUOH МЯ AUS NH 2198 NH WO NH XW
13
"ропштиој "т xipuaddy
573
Philodendron Subgenus Philodendron
Croat
Volume 84, Number 3
1997
euou | ѕәџәѕ2 G0 | 050 | 060 01 01 0L axe foei orf 9 | 3 | [aepo] vd | uinaajinpue|
jeuou seuesg-2| (0 | oro | oro | o2 | 02 02 axe [ozs|082]| S |¢ ајәргоо Add | .. seb
| dsuenauou | seuas-2| £020 | oco | 020 9E ZE єє | epa [062 091 | 6 | 2 |: aepo Wd | _ шпш5зциәлбә
euou | seues-2| 4070 | 050 | 0v0 01 01-6 qns fosi 06: | 8 |, әјәроо! Add | __ _ азивипџој
| одеш jeb-euou | j| EFE | 0€0 020 | qns 090 | 090 | ¿ | 9 је э! боодо 555. NWOS|O}
auou seues-| S0€0 | oso | 00 22 st | (z2 02-51 axe [019091 | 9 | c | 8iepio2| dd . Suepuy
, 8u0u seues-|| SUED | 050 | 00 | Р рг |(emejqns [061 | 02! | o1 | в [a |2 | аеро NAO шпашбгшај
| xew 1аб-аиои sauas 2-1| $0 0S0 | oso | + be | ans ро тр, | = | фер:оз | NAO штәрпиерә
|. BUOY 0 aiu г DIO | | med mp моја | 2 |3 | аәроо WWO _пәАмр
| Чед | sauas-2 0 эъ | 0Р0 3 PE axe |052 052 | S | + [8 |2 аеро 103 | џејззар
| хцеш jab-auou | | POE 0Р0 DE0 | qns fozo[ozo| 9 | s [8 |? | аеро 059 asue[eoiunuop
| dsuen seuesz-|| 10 MA | оа | | 5 «qns __ [082 | 001 | 6 1 Биојао sol wnKydoyoio
HERR 0€ au | mU 02 02 Jd axe [020 | 022] S аюрюэ, ddd didi . ADO
це, | ОРО | 00 21 21-9 qns _ [062] 062 | Ol | 6 [8 | 2 [pucogns) 3W9 пџвгојеоод dsqns nuospinep
п | wb | Gi у (01) 8-Р qns [052 021! | el IDE Эмо | miospwep dsqns ишовриор
RA m 0 | o 02 02 је __| әрә ЫЕ ШЕЕ» | Suono Эа | E An
Мо) 091 081 01 г. (| ee [006 005 | ; | Э. 1р:сэ9п8 IWO | __ штује 5! 55940
peo! 000 | te | ib qns 1090] 090] 9 | 3 | sep 099 | _ eee
E 0t0 | 020 Е qns fori orf: ајар:оо L| _ esuesniqoo2
TEES | | qns _ [092 | 092 | 9 Мо | бһиодо 950 | _
¡ab 133 — tee: 22 02 22-02 axe _ у ајер:о2 | Idd | NOSE
| xew C saues-z ozo | 020 і 2 ¿+ ape |081 081 | < ајәр:оо | у | X eesuapgiojoo
d : 20 ozo | 020 4 2 2 qns Joell oct [ ; empio2| ОО пуамај
euou seues-|| 20-10 020 0L0 2 3 21-8 exe fori] ori] «c ajeno! Od asusoduyo
| xew јеб-аиоџ oer 90 090 | 090 pl pl pL 99 axe fore! ore s аеро | Jdd зоџапбш
КИР T мра 12-60 012 060 8l el ferse | exe [ог oso | ; аро dg | _ ајпезгаџиг)а
L MM 80 90 90 | 2 2 qns [090 | 090 | ‹ бело 5959 | _ __ 8su8i8jswelq
| euou 0-0 РО о | bl 9 Р1—9 axe – [090 | 090 | ‹ ayepioo| Sd | — unyedsmag
| хдеш 185-euou РО ОР'0 0Р0 21 9 21-9 qns fosil ogi | + TES T
| 8uou polo F0 | 00 02 02 02 92 axe [осо | oco | ‹ ayepı02 | чая
| — dsuey 13. DES | oco 9 9-р qns _ [002 | 002 | $ ајерлоз | дур
| jab-dsuen _ 5020 | 050 | 020 А (Qu | ans [oot ovo [ £ биојдо | 555
. dsuea . 3euesz-l| bozo | 00 | 020 p pE _ | qns 031 | 020 uojqog| 55902 | O
| Чзиәдозиәд зеџез2-1| €0 | oco | 0Е0 Е Ez qns __ [081 | 081 | ol aepo | NINO | sat.
| | euou 3euest2| рй "m I s DE S2 0Е-2 | ape [ooz|002| s | 3 |g |2 | аеро ddd шпивомојџе
dsuen ^ | заџаз-|| 1-0 021 0E 0 5 _(+1 | ans _ [092 | 060 i Buoigo| WAS
| __|еб-5ивд seues-|| _ РО oro | 00 S Sy) | ms [оге | ozz ¿ редоне| 181 |____ l
| dsuen E 90 090 090 г (2i qns _ [02] | 060 ¿ радон: | ML unqo¡ysnbue
аиоџ seuas-2| 9070 | 090 | OO 8l 21 81—21 axe [002 0/1 ‹ йр | Jdd | __- ом
[| auu 7] 02 8L 02-81 axe fors] obs ‹ 8j9pJ02| vdd =>
¡SUBA 020 Р bl qns PEST с ayepuoo
мошамоо unu XEM uw ganool NOILWL xew u xew | ши [33 3av18 1295 мономзаотна
3dO13AN3 | HNN HION31 3MAO | emooysemeo | ;s3in^o | -маомла | нумал s3T001la1 | ЕЕ] |
| злпло | | amoo Ja fas | |
| | | | | | о BY] gum eR |
| | | | QOCHETEM RENE S|H| | |
Missouri Botanical Garden
Annals of the
574
| о | ob [Ору | ewe [05206119 | v» је |2 рој ddd
“pt | OL [rii | ame [оос oof 9 | v ја 5 |piooans| ddd
BE e ae: rz | ans [oes 00: e | 2 [8 |» | аер:оз | NAO
Cri o | «ws __[080 | ово [| 9 | 3 |е pegort| HL
e 1-9 e3 | ame folllollf r | а чојдо| 2d
t | Mi а! | axe [oriri ort | 2 | g чојдо| Dd | _wnyjedsoeso: ‘Jen wnyedsoasos
Е E € | ans _ [090 | 090 | 8 | а [2 |докзош 104 штереорпега лел шәрә;
—€— — 8 { 8 axe [059 061 | 7 OJa э |чонзош | 10d - Ro. ~ лел WNyeIped
02 El 02-81 | axe [005 009 | 2 [а ajepı02 | ddd
02 st | 02-51 | 902 [029] 029] S а |^ | өер Sd
02 02 o | ewe |061 | ost | 9 8 ВЕНЕ
b (zi | qns ое ort] 8 g | | Buoıqo| 550 | X штщгүпошпәрпәз
Е 6 qns fost | ozo] 8 | х ја |^ arpo NAO А:
02 | 02 02 axe [02102119 |s ја азарлоз | Jdd
ЈЕ 02 1£-02 axe |о0е| 00119 | v ја ewpio29| ddd | - uie
4 4 cs ES apı09| JIWO |
qns JOEL | 0с у s E Сй зч шыл
qns [ово ogof 7 | 9 [8 |^ |рюзапз 095
£ (£)2-L qns [osz|000|] 2 | 53 [а |^ әәро ии _____ [Ls
@ | 02 02 exe fog | ос! | 9 | s [8 |) әәроә vd
[ i1 | ws AAA | 3 [О |? |редон fut
oz | 9 [Oæ eme [089] 051 | 8 | ә ја ә epo заа)
spass) 2 qns 00L/| 001 4 9 g |^ р:оодпѕ 995 штело тел unje|nbi|
> > 2- | ада 091 | 091 | = = о џојдо 559 | Wnueoipelay лел шпјејпбу
| qns __|орг | ово | 8 | 9 [a v биодо 990 | ___ штејпби i геге
‘ 9+ (2-1) qns [002 | 021 з [а ло ргозапѕ 3^5
FL | OL 1-01 epe [091 | 011 s [a ejapioo| ddd |
, [ b | ewe [080 080 b ја ајерлоз | WWO
82 | 8l 82-81 | ape [021] 021 ғ [8 ajepi09| Jdd
ee speesg | axe [026] 260 ¿ [а | 2 | aepo uno
хијеш TRE “| g 2 qns __ [02 | OF L ғ [а |^ appo | OVA
[ A | E | = wi | M Тш y ја биојдо | 555 упр
| [SUBA L р-С(0 qns [021 060 з [а o ргозапѕ! 09 аешајац
Ж БК = A fi аферлоо | Sd штрлезлко ‘10 лел шпазеарац
| өцоџ z oz | so | әрә |009) 009 | s | s [а |^ аерсо Sd | әрим гәл шпазеларац
| и D 2 | ог | o | ewe [оов оов] 9 | у [а |^ | epics] sy uneaerepeu ел wneomepey
[ uou e | et | pee. | әрә [оду се: | e | s |а |2 | epics) idd | |
E LIR T axe fooi ool] 9 | 9 fg |! aepo] va |
auou) 86 dsuen coke 20 eu | RE ТЕ ТЕЕ ШШШ ајар:оз | WIND |
ep LI 00 1 PW SEIT! бојао, 990 | yT
auou 2145 22-91 ајхе net 001 3 | و [8/3 | epics! dd
NOLLIGONOO | Д WN шу аппоол NOLLVL 5 хен | ст = | um 33 1 | заула 1385
| 3do13^ |dO13AN3 r 9jn201/s9[n^O IS31nAO “-N30V1d H. ШОКЕ S301 - 1 | ЕТЕ)
| ЮИ CONE GET
vu NONE I C TE ONERE NU E ПЕ са CIL : _
E б | {| S|H|
'penuguo) > xipueddy
LO
~
LO
| |. UntuÁADOJSW = IVA ee
c | | umwuoWod = 109) unyjeqomew TIT әшбцѕо[е0 = NANO
o | Em E TUS T wnupodoiAysy uojpuepo|lud = vd
5 | aub wubuajsousjos uojpuepolud = Sd
S T | ч аы: “dd
8 E
£ Em. E “euibysojeg = 655 [esoo
wine3Auoed шпцадоюеу ешбцѕојео =д р |
8 uos | ејејпоцен шпједолој ешбцѕојед = NO) 1 apt ону
S дәр:ооэ шпцедооуј _ вшбузоје) BSH i. + ae о UOIPUAPOII = Idd
2 ___зеџе5 uomesqng иоцоәЅ apo) _ lo l1 2 Pe S PS
f
-
Ф ПЯ bz0 bS6 | РОД | eBeıare |
5 bb bb 9 9р juno3 D
> 91 TI БЕР | pee wns pusana, = ||
|- BUA =A cH
Ф Ре TES МАВАРА + 9 T E ___|_ Једи =3
= 8 хщәшјаб | заџез-|| р0-с0 00 DE0 3 €-1 seq |021 080 | € | э | әјәроо ANO | OO unuenyg
Ој | ¡eb-dsuen | saues-1 (9002 0-20 | 090 020 2 2-1 qns [051 oro | + 3 1м) expioo| 050 мтепоџпрефбиој тәл шта
об jab-dsuey _ |seues-|| 4060 | 050 0Е'0 4 2-1 qns [002 | 020 | 2 3 23 ајер:о2 | 099 шпал JGA типом
| 15u04 10 зиед EE | 60 | o) д 2 qns _ [осі | 0/0 У [а 2 | биодо 990| ирџәриәм
хщеш [SUBA YOIY »euasz-|| FO 090 050 PE axe [ose | 092 2 о |gorsou| 103 |__________паоимаглем
jab gdsuey | sauas-2| 20-10 020 010 ФЕ oz | (2e)p2-02 | ewe [092 061 2 aJepi0o| vd штзозпшал
аб is dsuey РО РО РО 2 qns |060 060 : ayepı02 | WWO a
| | qns | биојдо | 555
sa РЕ Oe BE £0 | qns fori] one биојдо | 555 M utm
dsuen sauas-2 p | РЕ a 10 L eo |- Dig d b Est) | ans [pet] 080 юрю ыр | иод
хщеш eb-euou | 2 2-1 qns [оса | 060 Мо | редове | ld | — & unda
| BE E ES 7 22-91 axe foe | 002 aepo] Idd unaisse[eui
БЕТИ 7 | pl ral #121 axe [065 | 091 ajepi09| ddd | enua
| dsuey 10 suey | П qans [оро оро | s | > [а |? | аергоз | 055 8|neamns
Lo m је. | al 91 axe [oro|oro]|] v | > | 2 |дор-вюш WWO winsiouigns
Buou — | 82 02 82-02 axe joe! 08: | 9 | S ayepı02 Idd MTS
| dsuey Аҳоцѕ | | 3 to | we не пыр з | 5 ajepio9| 099 ајтеошшәдѕ
co Г euou заиәѕ-с-21 20-10 | 020 | o10 DE 02 oaz | ape [021] 001 | s | v раозап5 | wq unjoyadiwenbs
5 auou sauas2-1) 20 020 | ozo ez | 02 82-02 | ape [021 | 001 fp |> афер:оз | vd | атәошепЬѕ
8 e6 dsuey suen] 20-0 | 020 | 00 z ¿ 2 (qns) језедј бр | oro | e | 9 ајар:оз | WWO | 8esnos
E dsuey Р'0-Е 0 0Р0 0£ 0 | (ans) језод| 01) 060 | 8 | 7 aepı09| 05 "gius
= NOLLIGNOD DNVHHV (шш) хеш unu xen UA 31noo1 МОУ хер | ши pen | uw [33 | 3av18 1295 монамзаотна
i 3dO13AN3 HI5N31 HION31 3INAO | amosan | /S31NAO | -мзомла | Hi5Nail sa3105201la1 | | БЕЈ |
5 311۸0 | | | 310201 # luna | E
o | | | | avi | |
E | |
5 | | | | E 1-1 S/H! посна 1
22
‘penuyuoy) >р xipuaddy
576
Annals of the
Missouri Botanical Garden
Appen
subg. Philodend;
ron in Уве America with number
species in Central Ameri
SECTION BAURSIA
мека 1. Philodendron sect. Baursia (Rchb. ex ем
ngl. in Mart. Е!. bras. 3(2): 134. 1878
SECTION PHILOPSAMMOS
Section 2. Philodendron sect. сосен С. 5. Вип-
ting, Phytologia 60(5): 306. 1086. None
SECTION PHILODENDRON
Section 3. de c cra (Philodendrum) Scho
Subsection 1. Mac see (Schott) Engl., gU
Fl. Bras. 3(2): 139.
Subsection 2. Camilla end Mayo, pii ;
Linn. Soc. 100. 168. 1989 ______ 3 sp., 1 subsp.
пази“ 3. plaspodim (Schott) Engl., Martius,
Fl. Bras. 3(2): 137. 5s
Subsection 4. Poropdin (Schott) Engl.,
Fl. Bras. 3(2): 138. 1878 on
Subsection 5. Sole enosterigma (Klotzsch ex Schott)
Engl., Martius, Fl. Bras. 3(2): 139. 1878
sp.
draw
Non
3 8p,3 om
— D Fhuodenaron. Lue sp.
Seri Phi
ilodendron. grin
par impolite ser. NOV у: sp
Series 3. Velveta ser. nov 1 sp.
Series 4. Fibrosa ser. nov ......... 15 sp
Series 5. Albisuccosa ser. поу _ ls
sp.
Subsection 7. Achyropodium 1 (Schott) Engl., smsen
Fl. Bras. 3(2): 139. 1878 sp.
SECTION CALOSTIGMA
Section 4, Calostigma (Schott) Engl., pos Fl. Bras
sp., 4 subsp.
dix 3. Sectional Composition of Philodendron
ral of
Subsection 1. psi an (Schott) Engl., Martius,
Fl. Bras. 3(2): 143. 1878.
ies 1 т
Series 2. Ecordata ser. nov... 5 sp., 2 si
Series З. Reticulata ser. пу -l-e
e achycaulia ser. nov ___________
Subsection 2. сЗа и (Schott) Croat, Ek
nov.
Series 1. Glossophyllum Schott .......... 15 spp., 3
P
Series 2. Ovata ser. nov ..........- subsp
Subsection 3. и ur pu $ Bat
Јаћг. 26: 535. 1 2 sp
Sergi et 4. asa Mayo, Bot. J. Linn. Su
00: 168. 19
meis EN Er (Engl.) Mayo, Bot. $ Lim
Soc. 100: 168 ne
SECTION TRITOMOPHYLLUM
Section 5. Уш ай (Schott) Engl., — e
Bras. 3(2): 144.
SECTION SCHIZOPHYLLUM
Section 6. rite iei (Schott) Engl., Martius, Fl.
Bras. 3(2): 144. 1878 None
SECTION POLYTOMIUM
Section 7. Polytomium (Schott) Engl., Маи. Fl. Bras.
3(2): 145. 1878 р., 2 taxa
SECTION MACROGYNIUM
Bot. Jahrb. 26: ge
Section 8. Macrogynium Engl.,
1899
SECTION CAMPTOGYNIUM
on 9. Camptogynium K. Krause, in Engler, Das
Pflanzenreich IV. 23Db (Heft 60): 3, 127. 1913 Pw
one
— —
— —À——— —
пина AS AY << nnd
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
577
Appendix 4. Phenological Patterns of Central American Philodendron subg. Philodendron.
Pattern
Flowering
All Year| Wet
PHILODENDRON
advena
1
Wet->Dry| Dry
Dry-> Wet
BiModal | Unknown
1
| غ d| 2
578 Annals of the
Missouri Botanical Garden
Appendix 4. Continued.
Flowering Pattern
PHILODENDRON All Year] Wet [Wet>Dry] Dry | Dry? Wet | BiModal | Unknown)
jefense 1
1
i 1
lenti 1
الم ا —
—
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
579
Appendix 4. Continued.
Flowering F Pattern
| All Year c ES IL UM ACIE S ME
4.85%) 28 4.85%
580 Annals of the
Missouri Botanical Garden
E c
Figures 14. —1 (top L). Art radiatum, ME m entitious roots. Cultivated at ees dioi E
(top R). P. schottianum (Coat d 6395), showing stems with transverse fissures and coarse striations ex of node.
ing
3 (bottom L). P. malesevichiae, showin tric ae like scales и 76707). —A (bottom m. E lid show
stems that dry with minute cracks and scurfy condition (Croat 57199).
Volume 84, Number 3
1997
Figures 5-8. —5 (top L). Philodendron conforme (Venezuela), showing stems with transverse fissures at points of
stress, seca scar, and scar of inflorescence (Croat 59355). —6 (top R). P. straminicaule, dried stem cts con-
spicuous ribbing caused by drying (Croat & Grayum 6005 7). —7 (bottom 1 ^ P. findens, dried stem showing interruption
of conspicuous epidermis by puis ole base (Croat 66811). —8 (bottom R). P. radiatum Schott var. ao rine ne
showing stems with cracking and protruding epidermis (Croat & Hannon 63381).
Annals
hoatn laps RIP Garden
Figures 9-12. —9 (top L). и rothschuhianum, showing i grams cataphyll scars and petiole scars
(Croat 57199) (photó: P d ich). — 2 op R). P davidsonii (Davidson 3956), showing stem with conspicuous cataphy ll
scars and petiole scars. —11 (bottom ie ? те galophyllum (Croat 54252), stem with intravaginal squamulae occurring
just above the cat ses" scar. —12 (bottom R). P. auriculatum (Croat 59730), stem with spreading дый roots showing
spinelike root branch buds.
Volume 84, Number 3 Croat
1997 Philodendron Subgenus Philodendron
Figures dis 13, 14. Philodendron lazorii (Croat 69833). —13 (top L). Showing clasping roots closely appressed
to tree. —14 (top R). Showing clasping roots and a much риба feeder root. —15 (bottom L). P. wendlandii, showing
e intact cataphylls held in place contiguous petioles (Croat 69732). —16 (bottom R). P. warszewiczii,
showing weakly 2-ribbed, short-lineate cataphyll (Croat & жые: 64186).
Annals о
Missouri di Garden
Figures 17— —1 (top L). Philodendron schottianum, showing bluntly 2-ribbed cataphyll (Croat & Zhu 76 s кле:
—18 (top R). " megalophyllum, , Showing petioles which are iid sulcate with broadly rounded margins (C^
). P. dav
rt
2524). —19 (botte n, showing sweet, viscid droplets and combination of e
—20 (bottom T. ds auriculatum, showing dark green ring
marcating petiole and blade and free basal veins (Croat & Grayum 59730)
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
Figures 21-24. —21 (top L). Philodendron malesevichiae, showing snapped petiole with slender strands of latex
(Croat & Bay 75810). —22 (top R). P. giganteum (Croat 68568), showing midrib, primary lateral veins, and basal
veins (the lowermost primarily fused to pr реке rior rib). —23 (bottom L). P. auriculatum, showing quilte d primary
lateral veins (Croat & Grayum 59730). (bottom R). P. megalophyllum, showing arrangement of veins: midrib,
primary lateral veins with lowermost branc mod (these secondary veins), and interprimary veins (Croat 5.3901).
Annals of the
Missouri Botanical Garden
Figures 25-2
— 225 (top L). | eom malesevichiae, Pete lowermost ney lateral veins and basal veins (these
P
loose ly coalesc ad toward the base) (Croa (p hoto: P. ! vich). —26 (top R). P m ii mum per minor
veins arising from the midrib and primary y latera veins он. 53901) 27, 28. р | mali (Croat и $ H
(bottom L). Fresh spadix with pistillate e staminate flowers in middle, and a portion o
flowers at base e rtile
staminate spadix at apex. —28 (bottom R). Older spadix Pasan the ا à staminate spadix and white sterile pee
portion.
Volume 84, Number 3 Croat 587
1997 Philodendron Subgenus Philodendron
Figures 20-32. —29 (top. L). — tysonii, showing spadix with sterile male flowers having been consumed by
beetle pollinators (Croat 67577). —30 (top R). P. fre m polle n grain (Grayum nui 36, DUKE). Photo: M. H. Grayum.
01 (bot ttom L). ` rothsc rer Main == strands of pollen emerging from anthers (Cro 35657). —32 (bottom R). Insect
Visitors to. newly ning spadix of P. anisotomum. gem beetles shown (Erioscelis color mbio » le 'gitimate pollinators;
smaller insects ( New la. order Hemiptera) are frequent visitors to Philodendron же play no “sone dera in their biology. Photo:
1. Your no
Annals of the
Missouri Botanical Garden
re
Figures 33-36. —-33 (top L). еу hebetatum, showing — beginning to break up to expose —
berries (Croat & Monsalve 61396 4 (to › В). P findens, S lowing 8 r left by faller — and coarsely 5
s ; апд
peduncle apex (Croat 38218). a5 d L). P pseudauriculatum, ни mature in ve cences with spathe
; with
staminate portion of spadix fallen (Croat & Zhu 76251). —36 (bottom R). P. brenesii, donem infructescence
exfoliating spathe and partially consumed berries (Croat 35519)
Volume 84, Number 3 Croat
1997 Philodendron Subgenus Philodendron
> ~ =
Figures 37-40. Philodendron advena. —81 (top 1). (Croat & Наппоп шо) dern blade. —38 (top R). (Croat &
> > зе ataphylls and an elongated petiole
ath. —39 (bottom E. e ) "tescences нан ying sp: already n). —A0 ion
R). (Croat & Hannon pa He nearing maturity (
adaxial side of spadix is longer than the abaxial side.
590 Annals of the
Missouri Botanical Garden
Figures 41—44. 41—43. Philodendron albisuccus. 41. 42 (top L & R). (Croat 37851). —43 (bottom L). (Croat 68940).
th: (bottom R). P. anisotomum (Croat & Hannon 64364 64).
Volume 84, Number 3 591
Croat
Philodendron Subgenus Philodendron
Figures 45-48. Р. alticola (Croat 74906). —45 (top L). Stem with post-anthesis inflorescence. —46 (top R). Juvenile
and pre-adult leaves. —47, 48 (bottom L & R). Adult leaves.
Annals of the
Missouri Botanical Garden
ж e
xc WEA k
Figures 49-52. Philodendron ыг. с — 49 (top L). (Croat i 162) Habit, in cultivation. —50 (top Em Pss
displae ed from trees, with inflore ces (Croat & Pis 64522). 51, 52. (Croat 61162). —51 (bottom L). In
ence at anthesis. —52 етта R. Habit i in itis
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
Figures 53-56. Ph ilodendron anisotomum. —53 (top L). Habit in cultivation (Berlin Botanical Garden (031-60-
74-83). —54 (top R). Stem with anchor roots spreading from stem. —55 (bottom L). Leaf in cultivation (Croat &
ns 64364). —56 (bottom R). Plant with inflorescence in cultivation (Selby 80-1668, from Mexico, Chiapas, Ocos-
go).
Annals of the
Missouri Botanical Garden
and
Figures 57-60. Philodendron annulatum. 57-59. (Croat 74803). —57 (top L). Habit. —58 (top R). benc wing
inflorescences. —59 (bottom L). Leaf with petiole showing purple ring at apex. —60 (bottom R). vds 74855) Sho
stem with cataphyll, unopened на and bracteole.
Volume 84, Number 3
Croat
Philodendron Subgenus Philodendron
Figures 61-64. Philodendron antonioanum (Croat & Zhu 76909). —61 (top L). Habit. —62 (top R). Leaf. —63
(bottom L ). Inflorescences partially obscured by cataphylls. —64 (bottom R). Leaves, both lower and upper surfaces.
Annals of the
Missouri Botanical Garden
Figures 65-68. —65 oy, L). Philodendron auriculatum, habit (cultivated by John Banta). 66—68 aromaticum.
66, 67. -(C roat 68382). —66 (top R). Habit. —67 (bottom L). Stem and inflorescences. —68 (bottom R). "Cea 68423)
Open inflorescence.
Volume 84, Number 3 597
1997
Croat
Philodendron Subgenus Philodendron
Figures 69-72. Philodendron auriculatum. —69 (top L). (Croat 59730) Habit in cultivation. 10 (top R). (Croat
& Grayum 35205) Open inflorescence. 71, 72. (Croat 32956) (photo: P. Malesevich). —71 (bottom L). Open inflores-
cence showing protruded spadix. —72 (bottom R). Sharply 2-ribbed cataphyll.
'(Surue[4 релиођ Aq pejeAn[no) пдец 1509 4 “(y wonoq) 97— "qeu 26882 12015) штпшоозир q “(| uonoq) с/— (29119 72015) xipeds ayeurues оцу jo uongiojoostp оцу
pue Aeme mə ayjeds цим xipeds 3utMous *soouooso10pgut srsaqque-1sod ‘wngozysnsup q “(y до) р— "qeu ‘(2777p 10047) оиәаро молригропца "(| do) e1— 92-6) вәли
Missouri Botanical Garden
Ф
<.
—
~
о
Y
o
|
c
<
Volume 84, Number 3
1997
599
Croat
Philodendron Subgenus Philodendron
DLE) HERBARIL'
N? 845556
STANFORD DIYESIN
Figures 77-80. 78. Philodendron bakeri. —77 (top L). Plant hand-held (Croat 37508). —78 (top I Habit,
showing abaxial odd 'es of blades (Croat 67456). —
79, 80 (bottom L & R). P. breedlovei (Breedlove 34518
Annals of the
Missouri Botanical Garden
» af #247
PA! ys 224^.
"wa eol "e,
2 f f uil
D» — 1 je».
e , и
= Е 4 А А By S : ^ У X.
ich). 82.
Figures 81—84. Philodendron basii (Croat 45442). —81 (top L). Habit (cultivated at MO; photo: P. Malesevich)
"me БАС: ey :Josed
3 (Cultivated at SEL). —82 (top R). Leaf, abaxial surface. —83 (bottom L). Stem with intact cataphylls and clos
petiole sheaths. —84 (bottom R). Stem with inflorescence.
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
Figures 85-88. —85 (top L). Philodendron breedlovei, open inflorescence (Breedlove 35181) (photo: D. Breedlove).
. P. brenesii, habit. —86 (to op R). (Croat 67578). —87 (bottom L). (Croat 68084). —88 (bottom R). Apex of stem
with bases of petioles, cataphyll, and unopened a ane (Croat 78806).
Annals of the
Missouri Botanical Garden
602
[[£ydejeo олциг цим wog "(T шопод) T6— иден “(Y doi) 06— “(98S8S 12010)
Tihs
Ap RCE AL
"(GPSS v 12 si2aoN| 2q) asuasaismaiq а “(Y WONG) 26— '9опооволојш p: ausdoun pue
umwipdsia24Q Y “16 ‘06 '(1а92р 12015) Wey “11sau21q иолригропца (т do) 68— "76-68 so1n3t4
Br
Del. Thomas
NORON brewster
PHILODE
21299 19015) uado mə зопаоволојиј “(y шоцод) 96— '(peoejdsrp 1ue[d) идең ‘(7 шоцод) có— '(peoejdsip јивја) уувоце əjonəd роједиојо шолу Suse әоиәоѕәлори
pue 51001 8шлоцоце Чим 921] [ешѕ 0} роцовце ways Surwous jug[q (y dor) рјб— ичен (q doi) gó— “(18592 NYZ 29 10017) "C6-£6 'гјутполомипад иоіриәрорца || '96-g6 SANA
N | |
5
o
©
то
2
=
а
о
3
Е
Ф
[92]
2
5
N
5
5
©
©
©
E
Aa
Volume 84, Number 3
Annals of the
Missouri Botanical Garden
Figures 97-100. 97 (top L). Philodendron brunneicaule, leaf blade = Leger: (Croat & Zhu 76581). 98-
98 f
100. P. chiriquense Cree 69068). —98 (top R). Leaf blade adaxial su 99 (bottom L). Stem apex showing
inflorescences emerging from cataphyll fibers. —100 (bottom R). "irit with tube portion cut open.
Volume 84, Number 3 Croat 605
1997 Philodendron Subgenus Philodendron
Figures 101-104. —101 (top L). Philodendron chirripoense (Burger & Liesner 7139). —102 (top R). Р. clewellii
(Gentry & Clewell 7028). 103, 104. as сс еее —103 (bottom 1). Нађи (Croat 75039). —104 (bottom R). Plant
hand-held with inflorescences (Croat
Annals of the
606
Missouri Botanical Garden
“asuanbi1149 q `(T шопо) 701— (68052 72015) ичен (q 901) 901— (89125 7204) sa
>
(Z9Z9
эчәәвәлоууш цим U31 “(1 d01) GOT
9 10047)) (01) ^W 1L Aq рјәҷ зое) asuau0102 ү] “(y uonoq) 901— (89069 10047)) qeu
‘asuapD10]09 иолригротца '901 “601 '301—201 $931 4
Croat 607
Volume 84, Number 3
1997
Philodendron Subgenus Philodendron
(рррдр 19015) Uqey umymdsisspa2 q “(y wonoq) 211— '(622рр 12015) s22u22s210jut pauadoun pue зојоцод pedeys-q Ҷим Wag
(1 шопо) TTI— '(622рр 17019) (uado зио) soouo: ose1ogur jo лојепјо “(y do) 011— '(62109 19047) дең (1 doi) 601— “asuado> иолригропца '111–601 '211–601 = 210314
Annals of the
Missouri Botanical Garden
; : 44729). 114.
Figures 113-116. 113-115. Philodendron copense. —113 (top L). Leaf blade adaxial surface (Croat 44749). : dd
115. (Croat 68765). —114 (top R). Leaf blade abaxial surface. — 115 (bottom L). Inflorescence emerging from саіар")
fibers. —116 (bottom R). P. correae, habit (Croat & Zhu 76395)
Volume 84, Number 3
Croat
Philodendron Subgenus Philodendron
мом
BOTANICAL |
MERBAS
2 344!
Figures 11 120, 117-119. dator correae (Croat 66748). —117 (top L). Habit. —118 (top R). Habit with
inflorescence and infructescence. 9 (bottom L). Plant with mature infructescence and fully r petioles. —
).
120 (bottom R). P. cotobrusense (Gra: rayum & rese 5689
Annals of the
Missouri Botanical Garden
Figures 121-124. 121-123. Philodendron cotonense. —121 (top L). Habit (Croat 66169). 122, 123. (Croat 66504).
—122 (top R). Habit showing abaxial surfaces. —123 (bottom L). Stem with inflorescences. —124 (bottom R)
crassispathum, habit (Croat 33150).
Volume 84, Number 3
1997
Figures ин 125. 126. Philodendron cretosum (Croat & Zhu 76661). —125 (top L). Habit. —126 (ор | g^
with persistent cataphyll fibers, obtusely sulcate petioles, and an unopened inflorescence. 127, 12
амай Маны 33150). —127 (bottom L). Habit. —128 (bottom R). Inflorescence with spathe tube cut open.
(ESOS 10047)) воопооволоу уши peuedoun Чм jo ходу AM шоцод) ctl = КРАДЕ: У 1D047)) попела по ul иче E uopnoq) Igi-- "umun40]D20q "dsqns nuospiapp
(VZEZZS 19045) (y do) 0£1— “1602 морао (1 do) 671— `чоцюлцүпә ur qe “тиоѕртарр ‘dsqns пиоѕртарр иолригропца ^O€l '621 “ZEL-6Z1 sandiq
Missouri Botanical Garden
Ф
t
©
2
©
€
c
<
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
Жаки 133-136. 133-135. Phi € uem subsp. bocatoranum (Croat 38177). —133 (top L). Habit in
cultivation. —134 (top R). Open inflorescence. —135 (bottom L). Leaf blade abaxial surface. —136 (bottom R). P.
7).
aet мдер, бедий. ореп fia ence был, 709
Annals of the
Missouri Botanical Garden
Figures 137-140. Philodendron dodsonii. 137. 138. (Croat 72982). —137 (top L). Habit. —138 (top R). Apex of
stem with petioles, cataphylls, and unopened inflorescence. —139 (bottom L). Unopened inflorescence (Croat & Hannon
79114). —140 (bottom R). Inflorescence on abaxial blade surface showing normal open nature.
462129 19015) qe ‘tunsori q (у ш X]) РРГ— (69664 10017) 5
9pe[q jes] (1 шопод) epp—
Philodendron Subgenus Philodendron
Croat
со
—
Ф
о
E
3
=
ae
со
Ф
Е
=
о
>
N
о
о
=
(пошта за :ojoud) аошлола ешвива ‘реоу те7-оиовү [¥ шолу oyd PAPAJO лом '92u95s3]onagut әлеу “Y uionoq) Өрт 9»uooso10gut peusdoun ҷим плода jo ходу “(1 мшоцод) 171
— ‘sa шолу pooe[dsip зивја “(y doi) gp[— (066/9 19045) “LET “OPT “(80022 742 X 12045) чеч peor uo ичец emey “(1 do) epp— “um Aydoyonop иолригротца — "вуј-буј Seana
Missouri Botanical Garden
Ф
=
=
—
о
2
©
E
E
<
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
617
Figures 149-152, —149 (top L). Philodendron crassispathum, stem with unope ora inflorescence (С mi 33150). —150
yal 82
7%
(top В). Р. dodsonii, habit with inflorescences arising from elongated petiole sheaths (Croa À
dolic нам yllum, habit (Croat 2.
2908). —152 (bottom R). P dressleri, leaf blade abaxial uis е (Croat e
51 oe D.P
Annals of the
Missouri Botanical Garden
MISSOURI
TANICAL GARDEN
шт
N2 2395111
ligt
de smaller
(atemat
BELIZE
District of Cayo
White; terrestrial %
— 155 (bottom L). Ё. dwyeri
icalense (Croat 35268)
Pielodendron dertei Croat
53, 154 (top L € R). Philodendron domir
Figures 153-15€ |
(Dwyer & Liesner 12 —156 (bottom R). P. edenudatum, habit (Croat & Zhu 771
Croat
Philodendron Subgenus Philodendron
Volume 84, Number 3
1997
e,
if
184
ul
Pr.
os
^4
Pag
A а
E
k
JF PP
И
Figures 157-160. doces dressleri. 157, 158. cow 0, —157 (top 1). Нађи. —158 (top К). Stem with
unopened inflorescences. Cultivated at Joseph Fon s. —159 (bottom L). Habit. —160 dob R). Apex
of stem showing persistent, ا you at uppermost hic d petiole bases with closed sheath.
Annals of the
Missouri Botanical Garden
oe r 164. ree MAU 161, 162. (Croat & Zhu 77087). —161 (top L). Leaf blade adaxial
162 (top R). Stem apex. 163, 164. (Croat 33988). —163 (bottom L). Blade abaxial surface, showing purplis
spots o on п petiole x^ lower midrib кы Р. Malesevich). —164 (bottom В). Inflorescence in cultivation in final stages
of closure
Philodendron Subgenus Philodendron
Croat
Volume 84, Number 3
E
e
M
"d
=
—
са
a
S
$
2
N
N
су)
m
~
©
=
O
=
2
с
=
=
|
2.
e
=
ид
о
pura
s (Croat & Zhu 77029), juvenile foliage. —168 (bottom R). Juvenile
(bottom L). Cluster of inflorescence
Е igures 165-168. Philodendron ferrugineum.
roat 75116).
75155). —167
foliage (C
Annals of the
Missouri Botanical Garden
Figures 169-172. 169-171. Philodendron кан = 169 д LE oe young blade ees shredding occurs
ў roat 67153). —170 (top К). Showing div ide 1 blades which rally become pinnate (Croat 67919). — 171 (bottom
Showing winged petiole (Croat & Zhu 76502). —172 uei R). P. folsomii (McPherson 1361! 9).
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
623
25098).
4 (top R). (Croat
í
|
4, 175. Cluster of inflorescences with one open.
176 (bottom R). P. folsomii (McPherson 13619).
tł. Philodendron ferrugineum. —173 (top L). Habit (Croat 34349). 1
-175 (bottom L). P. findens (Croat 38218). —
Dy LE
46 ]
Figures 173-1
624 Annals of the
Missouri Botanical Garden
gures xig cs dis 178 ise sisi findens (Croat 38218). —177 (top L). Apex of stem with uno pened
Fia ences, —1 p R). Open inflorescence. —179 (bottom L). P. fortunense, habit н 67921). "" 180 (bottom
R). Р po PS lg ка. with open обола ence (hand-held) (Croat 3975 3).
625
Philodendron Subgenus Philodendron
Croat
Volume 84, Number 3
1997
{#1299 19017)) зооизоволојј
#шмоцв јео ‘(1 шопод) 281— ‘(H1299 20027) идең “(Y до) zer— «
п 91njgurui JO лојепјо цим ходе шә] `{ шопо) р81— “(1Z28p 19015) ојоцод ровшм рив зовјдпв [erxeqe
£62
0000€ 19045) (coy Ne Aq P[?u-pueu) зең ‘(q do) 181— “asus
m
d SR
L]
+»
&
-
ито] uospuəpojiyd ‘ÞSI-I8I sandy
Annals of the
626
Missouri Botanical Garden
а
ib
РРА
зооиз: әвәли:
pue вәтоцәа padeys-( Чим вәлвә цим ичер “(y шоцод) 881— `чәрлесу үготиво 142021 Jo AVISIOATU Гү
9u іе po1eA^nn?) (SZ6S
110110q > Wd dr 191 ‘ogg "wmunssipuDadpaf 4 ‘881—981 '(86052 19015) чедо mə ouo цим soouooso1ogut jo 1o1sn[o “wneuldnisaf иолригропца "(| do) сат— “881
(OZSTT 1047)
apaun) +1
сој sanat]
Volume 84, Number 3 Croat 627
Philodendron Subgenus Philodendron
Figures 189-192. 189-191. Philodendron fragrantissimum. — 189 (lop L). Habit (Croat 11526). —190 wp on
53912).
Apex of stem with persistent cataphyll fibers (Croat 53912). —191 (bottom L). Juvenile leaves (Croat 9003).
бон В). Р. gigas, stem with petiole bases and intact cataphyll fibers (Cr roat 33680).
Missouri Botanical Garden
Ф
=
—
يها
о
=
©
=
=
<
629
Philodendron Subgenus Philodendron
Croat
Volume 84, Number 3
1997
(вјешәзепе)
"9sof ues '(*w's 421/2404) °[[02) цәрлтесу [eoruejog yrun aui 1e parean¡no ‘wnafi~npunjs ‘dsqns штаајтрио а y “(y wonoq) 002— '(ејопзопод) (69029 19945) әәиәәѕәлори uad()
(1 tuonoq) 661— 'xiejy-9pmng орәдоу Aq pareanino (y y 7 doi) 861 '261 "w"mupogpmurmo *dsqns umaafimpupj2 а '661–161 “unsafimpuo]2 иолригропца 000—161 $910314
Annals of the
Missouri Botanical Garden
Figures 201-204. 201, 20
2. Philodendron glanduliferum var. glanduliferum. —201 (top L). (Croat 43909) Leaves
) — 202 (top R). Showing weathered cataphyll fibers (Croat 39753). 203, 204. P. grandipes. gr
). Habit (displaced) (Croat & Watt 70200). —204 (bottom R). Apex of stem with intact cataphylls anc
unopened inflorescences (Croat 61323) (Colombia. Valle: Bajo Calima).
Volume 84, Number 3
1997
Figures 205-208. —205 (top L). Мани gigas, blade adaxial surface with unopened inflorescence (Cre
Zhu 76988). —20€ 6 (to ор R). Р. тапар, оре
L&R 5543 x
R). P. granulare (Croat & Porter 1
Croat 631
Philodendron Subgenus Philodendron
»
inflorescence (Croat 33648) (photo: P. Malesevich). —207, 208 "eoi
Annals
ent ا Garden
ew 209-212. Pico sete Sh ins —209 (top L). o blade adaxial surface (Croat 66814). —210 (top R).
er of unopened inflorescences (Croat & Zhu hs dina 1 (bottom L). Blade adaxial surfaces (Croat 669 969). —
212 аи R). Apex of stem with ipli inflorescence; iden ЈА адар sulcate base (Croat 74840).
‘(P9992 NYZ x mos 9) | зафриоа y (y wonoq) 912— 'ичен
(20222 "uz » 10015) (т uonoq) ст2— ‘avu soyur uad() (y doi) ppz— qey '(1 doy) 212— ‘(6299 19015) 112 ‘ета т ‘SIZ : —Ü
~
| |
A
Philodendron Subgenus Philodendron
7)
; Wh
Volume 84, Number 3
Annals of the
634
Missouri Botanical Garden
SIDUVBOSIIOPYUT JO 19ISN][O 1 woyoq) BOIS
fc
(22222 1001) UONBAR NO ut IDUIISIAO ји usado “(y uonoq) ozz— “(OST EZ 10047)
1499 1001) ен “(Y doi) 817— "шттлодоц а "022-817 (1992 јешшон) njourumy иолригропца *(1 doy) /17— `0@@—!.1@ so1n314
~
i
|
|
|
On ridge W of Бе!
эс.
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
Figures 221-224. aee y aperit 221—223. P. hederaceum var. hederaceum. 221, 222. Habit. —221 (top
L). (Croat сар — 222 (top К). (Croat 69834). —223 (bottom L). Cultivated at Kiev Botanical Garden (juvenile form
with velvety lea ves). ar а R). 7 jiu iii var. kirkbridei, close-up of stem showing minutely warty surface
and anchor roots (Croat 75108).
Annals of the
Missouri Botanical Garden
А Figures 225-228. 225, 226. Philodendron hebetatum. —225 (top L). Leaf blade adaxial surface (Croat 73150). —
226 (top R). Open inflorescence (Croat 69231). 227, 228. P. heleniae (Croat & Zhu 76738). —227 (bottom L). Habit-
—228 (bottom R). Нађи with inflorescences (displaced from tree).
Volume 84, Number 3 Croat
1997 Philodendron Subgenus Philodendron
Figures 229-222 229. 230. Philodendron heleniae. —229 (top 1. + а luster of inflorescences (one open) (Croat & >
76738). — 230 (top R). Open inflorescence with pre rotruding spadix (Croat 61275) (Colombia. Valle: Bajo ( — 231, 232.
P. jacquinii. —231 (bottom L). Habit (Croat 69835). —232 (bottom R. Habit with inflorescences (Croat 124.
Моште 84, Митбег 3 Croat 639
1997 Philodendron Subgenus Philodendron
Figure s 237-240. —237 (top L). Philodendron jacquinii, сето infructescences with fallen spathe (Croat 14911).
—238, 239 (top R & bottom L). P. jefense (McPherson 10038). —240 (bottom R). P. knappiae, habit (Croat 67982).
Missouri Botanical Garden
Annals of the
640
(Z86Z9 10017) зовј
so[A1s pesuo[oad Surwous uado nə ваопзаоволојиј “(Y doi) zpz
ns [BIXE
pins [et
pe
ЛА
орв
44 147 Y 19015) 99€
[ч ‘aviddpuy q (у шопо) рус — “(08009 19015) пдец “ununisiabpol
jns ¡eixepe ореја “(| doi) ppz— "питборг uospuapop
d C) wonoq) eyz— "(gerc 10047)
Md стуб lvo үүс-1Ус samiy
Volume 84, Number 3
Croat
Philodendron Subgenus Philodendron
vě LA a
Figures 245-248. Philodendron Ara —245 (top L). ( en o - rs (Croat 60080). —246 (top R). Blade
ad: m surface (Grayum 3216). —247 (bottom L). Habit (Croat 60450) ) (bottom R). Open inflorescence (Croat
(98144 142 Y 10017) sooepns [erxeqe pue [erxepe Zutmoys sape[q Jeq “(y шоцод) 922— (22869 12015) ичен (J uonoq) eez— uozo] 4 "987 ‘660 “(28629 017) "qeu
готадриу q “(y doi) pez— “(62129 1D047)) sa2uoose1ogut Jo лодапјо pue s19qy [[Aydeyeo juaysisued цим ходе ways “wnunisiabpol uospuapopiyg “(1 do) eez— — "'9ez-eez sam
Philodendron Subgenus Philodendron
Croat
Volume 84, Number 3
Annals of the
Missouri Botanical Garden
W^ i \ е. ~~
Figures 257-260. 257, 258. Philodendron lazorii. —257 А. ор L). Leaf blade adaxial surfaces (Croat & Zhu 77126).
—258 (top R). Stem apex with partially intact cataphyll fibers (Croat 69833). 259, 260. P. lentii. —259 (bottom |).
7).
Нађи, displaced from tree (Croat 50041). —260 (bottom Stem with open inflorescence (Croat 6664
645
Philodendron Subgenus Philodendron
Croat
Volume 84, Number 3
1997
(16899 10017) aseq әјоцәа 9]eo[ns цим WS BULMOUS “(y uonoq) poz— ЧЕН “(1 шо11094) 292
— "sajonad pedeus-q pue soorjins [erxeqe SULMOYs S9AE2] “(Y do}) 292—— (8£€49 10017) '292 '292 UÜTF999 19015) пен `7 doy) 1[92— шә] uoipuapopqd “POE [92 soundly
же 4
(8297 мору) eqn) ayieds uodo-1no pue xt peds Surpnajoad Чим оопооволо ји uodo ‘әѕиәитү d “(y wuonoq) 8392— "xipeds Burpnajyoad Ҷим оомооволо YU]
(т woneq) ,oz— зен “(y do) 99z— '(с616р 10047) штаојт ту лел штотгау а '29% ‘997 (29129 12015) wqeu “mua иолригропца “(y doi) сос—— “B9Z-SOZ зәл
issouri Botanical Garden
Ф
=
—
c
o
2
с
c
c
<
M
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
Figures 269-272. 269-271. Philodendron дин var. ligulatum. —269 (top L). Нађи (Croat 67318). —270 (top R).
Stem with open inflorescence (Croat 34302). 1 (bottom L). Habit (plant displaced from tree) (Croat & Zhu 76954); note
dark ring separating petiole from Made 012 ideas R). P. ligulatum var. ovatum, habit (Croat & Zhu 76754).
Annals of the
Missouri Botanical Garden
Figures 273-276. —273 (top 1). Pepa ee var. ovatum, habit (Croat & Zhu 76888). 274-276. P.
ae var. heraclioanum (Croat & Zhu 77098). —274 (top R). Habit showing blades spotted on lower surface
275 (bottom L). Leaf blade adaxial surface = sharply D-shaped petiole with narrow wing and dark ring а! pet
apex. —276 (bottom R). Unopened inflorescence, stem, and D- shaped petiole.
“ole
Volume 84, Number 3 Croat 649
Philodendron Subgenus Philodendron
¢ Comarca ће ñan
it d шы س a
Tia е. х forest та зао ip A а E E rad
гаје“ sky
По ا eae he Ur. ут 4
T : Lm n en. m ‘olay hess
‚үө
- as me dom: petiole ама petoncto
E га «bis sent ta им de
Figures 27 do 277. 218. Philodendron llanense. —277 (top L). Leaf rag adaxial surface (C ht е
278 (top R). Cluster of en ned inflorescences (Croat & Zhu 76993, 4). — 280 (bottom L & R). P. madronense
(Hammel & Ме 'Pherson 14526
Annals of the
Missouri Botanical Garden
y
Lum DIR оним
Figures 281-284. Philodendron malesevichiae (Croat 74818). —281 (top L). Habit. —282 (top R). Stem with intact,
. ] T
— cataphylls. —283 (bottom L). Petiole with conspicuous, trichome-like glands. —284 (bottom R). ope
тпогезсепсе. 7
Volume 84, Number 3
Croat
Philodendron Subgenus Philodendron
Figures 285-288. Philodendron mexicanum. 285, 286. (Croat & Bay 65778). —285 (top L). Habit, displaced from
tree. — 9286 (top R). Stem close up. 287, Cultivated by Monroe Birdsey (photo: K. Upton). —287 (bottom L).
Inflore 'scence. —288 (bottom R). Open spathe.
Annals of the
Missouri Botanical Garden
‘MISSOURI
BOTANICAL GARDEN
на 3398570-
Figures 289-292. 289, 290. Philodendron microstictum (Croat & Hannon 79210). —289 (top L). Stem, leaf blades,
inflorescence. —290 (top R). Cataphyll, anchor roots. 291, 292. P. morii. —291 (bottom L). (Liesner 567). —292
30).
(bottom R). (Hammel & McPherson 1453
Volume 84, Number 3 653
1997
Croat
Philodendron Subgenus Philodendron
MISSOURI
BOTANICAL GARDI
HERBARIUM
N2 241670
Philedesdren pique dorm
1
AACLAT or CENTRAL (TR
wet c UN
Fiiledeudras quen
Figures 293-296. Philodendron niqueanum. 293-295. (Croat 37942). 293, 294 (top L & R). Adult leaves. —295
(bottom L). Juvenile leaves. —296 (bottom R). Juvenile leaves in cultivation (Croat 37886) (photo: P. Malesevich).
Annals of the
Missouri Botanical Garden
игез E 300. 297-299. Philodendron panamense (Croat 55184). 7 (top L). Leaf blade adaxial su
298, 2 › К & bottom L). on n inflorescences and sharply 2-low- ~ и слон те — 300 (bottom R). 3 pseu-
dauric if ss habit (Croat 37595).
Volume 84, Number 3 655
Croat
1997 Philodendron Subgenus Philodendron
HAAN OF SPICAS SUPPORTED BY A GRANT твом THE улей FOUNDATION, ST. 40 MISSOURI
BOTANICAL GARDEN
HERBARIUM
No 2416713
АЉАСКА OF CENTRAL AMERICA
1
«Ф, Philedewdven pirms
Philodeudron frs
á E
: E js i
ё t >
{ T
A $
Figures 301-304. Philodendron ритепзе. 301, 302. (Croat 37944). —301 (top L). Adult leaf. —302 (top R). Stem
with detached fibrous cataphyll and inflorescence. 303, 304. (Croat 68952). —303 (bottom L). Apical half of blade
with loose inflorescences. —304 (bottom R). Basal half of blade.
(92882 10017) оопзоволо ит pauado Чим ued “(q шоцод) 802— “39M шолу paoe[dsıp “ичен xu uonoq) 7 0€— (сово 10047))
Missouri Botanical Garden
Ф
£
-
~—
о
2
d
c
С
<
657
Philodendron Subgenus Philodendron
Croat
Volume 84, Number 3
1997
ишш
“BEE
NS 18 ројелујпо) (26201 19015) s[¡4ydejeo yuajstsiod рив ѕәәиәәѕәлорш JO лојепјо цим SWS Jo ходу (у
—.
'(suapame)
1 шопо) zI£— “(1859 10047) uqe (1 wonoq) 112— mj0421d 4 ‘ZIE
цотләвә[е Y :o1oud) saseq әјоцәа цим шәдѕ Jo dn-aso[) “(y doi) OLE— ‘uqe ‘(q do1) 602— (ос 12015) штттомпорпога иолригропца “OLE “60€ "CI€-60€ вәлпйї 4
658 Annals of the
Missouri Botanical Garden
Figures 313-316. 313, 314. Philodendron pnm ulatum. Nec 3 (top L). Habit in c repr with leaves, in-
florescence, and soak 2-ribbed cataphylls (Croat 33526). —31: (top ^g (C roat 486964). 3 316. P. pterotum. —
315 (bottom L). Habit (Croat 6581). —316 было R). ree м ence (Croat 10265).
Моште 84, Митбег 3 Croat 659
Philodendron Subgenus Philodendron
Figures Mg 320. — purpureoviride. 317, 318. (Croat 59857). —317 (top L). Leaf blade adaxial surface.
-318 (top R). Habit. — (bottom L). Opened in florescence. Cultivated at Wilson Botanical Garden (photo: D.
Be ath). — 320 (bottom R). pu roat & Zhu 76677) Habit with cataphyll recurling and beginning to recurl.
660 Annals of the
Missouri Botanical Garden
5”
n Fo
B
Ady
B
ay
As
af
Figures 321-324. 321-323. a: purulhense. —321 (top L). Habit (Croat 41752). —322 (top R). Le
dre 1
blade adaxial surface (Croat & Hannon 6376 65). —323 (bottom L). Stem M with unopened inflorescence (Croa
41189). —324 (bottom R). P. radiatum, pra at Wilson Tropical Garden
Croat 661
Volume 84, Number 3
1997
Philodendron Subgenus Philodendron
‘uedo jno әоиәәѕәлоуи цим рипола uo јивја “(y шопод) 822— “UBH (T uonoq) /22— '(18229 uouung 2р 19045) umqpippaopnosd лвл
UNIDIPDA 4 ^9ce ‘LZE "pieu-pueu шеја (y doi) 9zg— ‘әәвупѕ ¡erxepe ape[q je *(T doi) ezeg— '(29229 uouung a 10047)) огиоутита иолригропца "oce “STE “BTE-STE волта у]
662 Annals of the
Missouri Botanical Garden
2
б 2 3
ә".
{
4
"d
"
PA ~“
eil...
: Figures 329-332. 329, 330. Philodendron radiatum var. pseudoradiatum (Croat & Hannon 63381). —329 (top L).
Closed inflorescence. —330 (top R). Leaf blade on ground with a displaced post-antheis spadix. 331, 332. P. radiatum
var. radiatum. —331 (bottom L). Habit (Croat 32767). —332 (bottom R). Pre-adult leaf blades (Croat 66075) (photo:
P. Malesevich).
663
Philodendron Subgenus Philodendron
Croat
Volume 84, Number 3
1997
ЧТЕбР2 19015) wnuniynyssyi04 q “(y шоцод) 9о22— '(0909 12015) (Pns perxepe ZuLMoYs *pooe[dsrp јеој 20) ичен (т uronoq) сее— 'npunmy
ur овпоц зәрец пәрлесу peoruejog unossijy əy} је paramo “(y doi) рее— “(91899 12015) (1 doi) сее— -^wnrpippa “лел штупрол иогриәрорца "SEE-ELE "Ogg-ggg SAMBA
Annals of the
Missouri Botanical Garden
Figures 337-340. Philodendron beim —337 (top = Adult к Tortuguero National Park (not col-
bod: pris M. H. Grayum). —338 (top R). (Cr at & Zhu 77224) Juvenile leaves. —339 (bottom L). Habit in
cultivation at the Missouri Botanical Garden (Croat 35 57): pt (bottom R). Stem with inflorescences (Croat 27 704).
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
Figures 341-344. Philodendron roseospathum var. rose ospathum. —341 (top L). Habit in с "ultivation showing leaves
and intact cataphylls (Croat 35931). 342-344. (Croa 7515). —342 бер В). Нађи. —343 (bottom L). Stem with intact
cataphylls and inflorescences. —344 (bottom R). гай п inflorescenc
“(92309 'UNÁDAO 29 10047)) WAS jo xode је ѕәоиәоѕәлоуи Jo 19jsn[o YUM 11918 “(Y шоцод) вре— '(126р2 12015) їчен (1 woyoq) Lpg— “UNUDIYNYISYIOL d “BPS ‘LYE
(26929 12015) wqeu “unmofimáos q (y doi) 9pg— ‘(Ames “A :o10ud “ECE-Z6-TAS) SUSPIRO Aqjag e parean[no читугодкогвол uospuapopy ‘(1 do) eye— ‘gpg-gpg soma
Missouri Botanical Garden
Ф
=
—
-—
o
2
©
c
c
«
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
ARACEAE
pal Jodenáren
Figures 349-352. —349 (top L). Philodendron roseospathum var. angustilaminatum (Hammel 3133). 350-352. P.
sagittifolium. —350 (top R). Leaf blade adaxial surface (Croat 67918). —351 (bottom L). Apex of stem with petiole
bases, cataphylls, and inflorescences (Croat £ Grayum 60259). —352 (bottom R). Open чың scence (Croat 69731).
Annals
sawed мы Garden
ж
Figures 353-356. creia sagittifolium. —353 (top L). Leaf blade adaxial surface (Croat 74939). —354 (top
R). Plant with ope n inflorescence (Croat 33179). —355 (bottom L). Inflorescence with open spathe and protruding
spadix (Croat 69 356 (bottom R). Leaf blade adaxial surface (Croat 60506).
Volume 84, Number 3 669
Croat
Philodendron Subgenus Philodendron
: $ =
Figures 357-360. Philodendron scalarinerve. —357 (top L). Habit, plant displaced on pam (Croat je '
358 (top К). Stem apex with petiole bases and persistent cataphyll fibers (Croat 68723). —359 M L). Plant with
inflorescence, displaced on ground (Croat 66959). — 360 (bottom R). Open inflorescence (Croat 67203).
оррер 19045) зоопзоволојш stsayjue-jsod Арҷӯцѕ pue stsoyjue-oid qm орвју Jea paoe¡dstp “Bug mmus q (үү шоо) v9e— "(26989 10017)
ѕәоиәовәлорш JO ISSN) (71 шопо) 292— (22289 20017) ичен “(Y doi) 292— "(61792 NYZ Y 12045) идең (1 dor) [og— "әмәшытюүрә иолригропца “E9E-19€ "ӘЄ-19Є so1n313
Missouri Botanical Garden
Ф
&
=
~
[e]
2
©
c
c
<
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
=.
ES
Figures 365-368. Philodendron schottianum. —365 (ор, 1). Leaf blade adaxial surface (Croat 74991). 366-368
(Croat & Zhu 76518). —366 (tc op R). Le eaf blade adaxial surfac -367 (bottom L). Stem apex with p» intact
cataphyll fibers and a cluster of inflorescences. —368 ако В). С баб г of inflorescences (one partially open)
Annals of the
Missouri Botanical Garden
Figures 369-372.
369-371. Philodendron schottianum. —369 (top L). vd to adaxial surface Cron n
—370 bl b Stem apex with partially ieu mposed cataphyll fibers and im e inflorescences (Croat 7 R). P
371 (bottom L). Open inflorescence eme g from persistent pace Teng Сы 35620). —372 (bottom J.
smithii, blade adaxial surfaces (Croat & Hana 64588).
'(£ 1069 12015) qey 'атооштир у y ` !£— “eg 19015) 1 o[ruoAnf читтјоподтиттр«
d “(1 uonoq) сле— (96297 nyZ 29 70015) qey 'omponupnbs q “(y do) рле— (расрдо uouupng o 10015) 1 uospuapopy “(1 do) £LE—
Philodendron Subgenus Philodendron
Croat
Volume 84, Number 3
Annals of the
Missouri Botanical Garden
|
т.
”
+
т
a
~“
а.
се: о i vis Philodendron smithii. —377
588).
inflorescence
64: p R).
20450). mede: Fs chis m Deadlies 25142).
Tine; spathe green wi
Pup areas on the * atsido) DR
inside marosn-porple ы
(Croat 40079). 379, 380. Р. зоиза
“У HiP аи. Prawns дк CHIAPAS,
boin ie ale Philodendron
МА | basally
Брзрћуте.
Flowers, am i $
51. EE ty stre:
P deus Inga and E Y
(top L). Stem apex, herren on ground (Croat & Hannon
. —379 (bottom L). (Davidse et. al.
Annals of the
Missouri Botanical Garden
: : R).
Figures 385-388. Philodendron squamipetiolatum. 385-387. (Croat 74857). —385 (top L). Habit. —386 (top )
Leaf blade adaxial surfaces, —387 (bottom
зе : - inflo-
L). Stem with scaly petioles. —388 (bottom R). Stem with leaf and in
rescence (Croat 68767).
Volume 84, Number 3 677
1997
Croat
Philodendron Subgenus Philodendron
Figures 389-392. Philodendron straminicaule. —389 (top L). Habit (Croat 67920). —390, 391 (top R & bottom
L). (Croat 69013) Open inflorescences. —392 (bottom R). Habit (Croat 66584).
Annals of the
Missouri Botanical Garden
Figures 393-396. 393-395. igre 7 eri (Croat 59130). —393 (top L). Leaf blade adaxial hey s
394 (top R). Plant with internodes broader than long, persistent, intact cataphylls and inflorescences, one M ena nii
iip, L). Open inflorescence. —396 е пот des P. subincisum, flowering plant in cultivation, Cornell Univ
(Moore & Bunting 8956) (photo: С. S. Bur
{29182 12015) пен “(y шопод) дор— “(6PZ09 unini р 10015) Ways peqqu Зшмоц цию ‘(1 tuonoq) 66g— moams q “OOP “66€ 's92usoss10gut Jo
19I5N[9 pue srunapida yeyur “шщ цим зАудејео цим xade wars (y doi) g6¢— зең “(1 do) L6£— (11292 туў Y 10015) шту иогриәрорца “86€ “LEE “OOP-LOE Samiq
Philodendron Subgenus Philodendron
c
о
—
О
Volume 84, Number 3
Annals
и س Garden
Figures 401-404. 401, 402 (top L & R). Philodendron subincisum (Moore & Bunting 8956). —403 (bottom L). Р.
sulcicaule, leaf blade adaxial surface (Croat 75167). —404 (bottom R). P. tenue, leaf blade adaxial surface ill 3608).
"(10799 19015) jns әре јео [егхеде ‘wnoisspppy) 4 “(y шопод) 901 "зопаоволојш pauador us
“(y doi) одр— (S9EOL 1104 Y 10017) >} ур “OOF “(809€ 1112) uado auo 'saousoso10put jo IANO цим xode шојс “(тт doi) eot
Philodendron Subgenus Philodendron
Croat
Volume 84, Number 3
Annals of the
682
Missouri Botanical Garden
10047)) sope[q пру “(Y шопод) zpp— “($098 10017) sepe[q opruoAnf pue зовјтв [erxepe ореја jeop mpy ‘(q шоцод
epe[q Jeo] (y do) Отр— “(10P99 12015) soousose1ogut jo злојепјо pue sioqy [¡Aydejeo цим xede ways ("ү d01) 60
(8019S
) Fip "штод d ‘CLP 1 LP (161 19 70017) IMHNS jetxeqe
== “UNIISSDIDYI џолригроу ја ‘OL? ‘60F Тр бор so1n3t Д
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
~ a
os adi АТА
Figures 413-416. 413, 414. Philodendron tenue (Croat 74634) (Venezuela. Zulia: La des M P "wg
blade adaxial surface. —414 (top R). Stem apex with cataphyll fibers and inflorescences. —415 (bottom D. P. th
assicum, leaf ували 79049). “416 (bottom R). P. tripartitum, habit (Croat 43438)
"(67195 10049) sooegms [erxepe орвја увә “(y WONG) OZP— (92102 10019) зоовјтв qerxepe opeqq јео (1
шопод) бр — -9oueoserogur подо “(y doi) gpp— идең “(1 doi) 7 pp— "штелофу воштеду 18 poreantny (20612 12045) ‘BT у “unyod иогриәрорца одру Samt
Missouri Botanical Garden
®
А ==
—
м
о
2
©
| =
c
<
Volume 84, Number 3 685
1997
Croat
Philodendron Subgenus Philodendron
Figures 421-424. Philodendron tripartitum. —421 (top L). Cultivated at the Lyon Arboretum (#80-890), habit. —
422 (top R). Leaf blade abaxial surface (Croat 70136). —423 (bottom L). Stem with open spathe (Croat 59160). —
424 (bottom R). Habit (Croat 56108).
Annals of the
Missouri Botanical Garden
Figures 425-428. Philodendron tysonii. —425 (top
surfaces (Croat 67844). 7
with cluster of infloresce
L). Habit (Croat 68688). —426 (top R). Leaf blade adaxial
—42/ (bottom L). Leaf blade adaxial surfaces (Croat 66711) —428 (bottom R). Stem apex
nces, one spathe open (Croat & Zhu 76346).
Volume 84, Number 3
1997
Croat
Philodendron Subgenus Philodendron
Figures 429-432. 429, 430. Philodendron tysonii, —429 (top L). wr blade surfaces (Croat 66589), —430 ) (top
R). un "scences emerging from cataphyll fibers (Croat 67577). —431, 432 (bottom L & R). P. utleyanum (Wilbur et
al.
Annals of the
Missouri Botanical Garden
Figures 425-428. Philodendron tysonit. —425 (top L). Habit (Croat 68688). —426 (top R). Leaf blade adaxial
surfaces (Croat 67844). —427 (bottom L). Leaf blade adaxial surfaces (Croat 66711) —428 (bottom R). Stem apex
76346).
with cluster of inflorescences, one spathe open (Croat & Zhu 7
Volume 84, Number 3
Croat
Philodendron Subgenus Philodendron
Figures 429-432. 429, 430. Philodendron tysonii. —429 (top L). Leaf blade surfaces (Croat 66589). —430 ) (top
R). узе ences emerging from cataphyll fibers (Croat 67577). —431, 432 (bottom L € R). P. utleyanum (Wilbur et
al. 15(
688 Annals of the
Missouri Botanical Garden
AS
PLANTS OF MEXICO
PE oko drm PPS EI
Beterised by Themes B. Croat, 2
Miser Botesical Carden (NO)
liama, ad, e gi
beg eros eed:
Cathet f Kes йл. ГА
and.
—438 (top es Leaf showing abaxial surface and petiolar scales. Costa Rica. Heredia: Braulio ( "pie Park (not
collected). —439 (bottom L). Leaf showing abaxial surface with varie vines c diviti (Croat 71413). —440 (bottom
R). Stem with увели“ 'ences (Croat 66558).
690
Annals
Missou
of the
ri Botanical Garden
|». ESO ut "y,
— 442 (top R). Habit. 443, 444. (Croat 34814) Cultivated at Escuela
sung leaves at beginnin
1
. Same plant with full-sized leaves and
~
(Croat 40714). —441 (top L). Leaf blade adaxial surface.
9стр9 UQUUDH 29 10015) sooua: оволојш YIM шоу “(y шоцод) дрр— "(рл / 0p 19045) UBH "(T wonoq) Ltr
врио грвојвошон чәшесу "| Aq peieAn[no 91qey “(y doi) opp— EMISSION а “Bb -9pp (85509 72015) 1PY ‘wnsoanssaa иолригроп ца `( dor) срр— ‘Sbb-Crp sonst
691
Philodendron Subgenus Philodendron
Croat
со
~
Ф
à
E
3
=
E
со
Ф
Е
=
O
>
Annals of the
Missouri Botanical Garden
ee 449-452. — 449 (top L). Philodendron warszewic zil
, 451. P a Costa Rica. —450 (
open mm ences and sharply flattened petioles (
(Croat 6808:
: pone inflorescences (Croat & Hannon 64136 )
top R). Habit. Heredia: La Selva (not collected). —451 (bottom L). den
Croat 34141). бајк (bottom R). P. wilburii var. wilburii, habi
Annals of the
Missouri Botanical Garden
ies us
Figures 457—460. 457. 458. dere wilburii var. wilburii (Croat 68083). —457 (top L). Plant with =
inflorescence. —458 8 (top R). Open inflorescence. 459, 460. P
—459 (bottom L). Habit. —460 (bottom R). pes
ed
F. wilburii var. blas (Croat & Zhu 77083).
displaced with closed inflorescence.
Volume 84, Number 3
1997
Figures 461—464. Philodendron wilburii var. longipedunculatum. — 461 (top L). Blade adaxial surface (Croat
66565). —462 (top R). Habit (Croat & Grayum 59879). 463, 464. (Croat & Grayum 60372). —463 (bottom L). Habit.
—464 (bottom R). Stem with inflorescences
Annals of the
Missouri Botanical Garden
\ \
Figures 465—468. Philodendron zhuanum (Croat & Zhu 76755). —465 (top L). Habit. —466, 467 (top R & bottom
.). Leaf blade adaxial surfaces. —468 (bottom R). Open inflorescence.
- " _— — ——— MM MM HM — سسس KK NS TN? —— M аа
A AAA O Пи
Volume 84, Number 3 Croat 697
1997 Philodendron Subgenus Philodendron
Figu
views virides
area not show
Type B
Type D
Style types of Central American Philodendron subg. Philodendron. Semidiagrammatic longitudinal
vii with Soe views (right) showing pr Tene route of stylar canals (based on Mayo, 1989); е
yo Style Туре В. Philodendron advena. —Мауо Style Туре С. Philodendien warszewiczii. —May
Style Type D. Philodendro on ferrugineum. —Mayo Style Type E. Philodendron granulare.
698 Annals of the
Missouri Botanical Garden
Index Homalomena Schott ex Schott & Endl. 312, 322, 323,
Orus 314
glao 315, 923
Alocasia jee © Don 325
mbrosina F. Bas
314
Annona 1. Schott .
370
- 312, 315, 319, 344, 345, 346, 347,
350, 369, 374, 375, 377
349,
sect. Pachy 312
surinamense di
Anubias Schott 323, 360
Arisaema Mart. 314
Arisarum Mill. 314
m L. 314
grandifolium Jacq.
hederaceum Jacq.
380
314, 458, 459, 460
314
lingulatum 1
tripartitum Jac 546
Asterostigma F. E. 1 Fischer & C. A. Mey. ______ 323
Baursia [without rank 380
ubg. Eubaursia T. Post & Kuntze ___________ 380
gnera Nicols 323
Bucephalandra Schott 322
Caladium Vent 314
baue Е ВАН lla с с. 380
HON O OR Hook A A ete 444
поа DOM, ааа 546
Nw Ren ume ) WUM o o c а RUE: 543
шы ло e: Sid Pon аана а ee 314
Cab res [without rank] _.__--____ 324, 331
Conr Schot “~ n s 323. 360
E 370, 374 ا
бс ылыш ы а 370, 508
amblyopsis Bates hi SS 361
mplata Bates oi сло с iS 374
conspicua Sharp .— .. о 361
gravis Bates а 361
Каша Enduüdi . . .. . .— .. — 361
ligyrina Bates ш 361
gaia Burmeister 0 __ 361
melanos Bates a n 361
nigerrima Bates — — __- о 361, 370
scens Баба a s ыг I 361
rustica (Oliver) a o o 361
sexpunciata Casteinau ._____________ 361
Сутборекцит Benth. ос 370
longispatha Engl, c ` 374
олы о 314, 565
Dracunculus Mill. л chee ы eee 314
Elopium H. W. Schott .. 20 380
surinamense (Miq. ) О ЕАК ол 380
Erro AL 370
colombica Endtüdi TS 361, 587
proba Shans | | TS 360, 361
Euphilodendron insane mn] E 32
Furtadoa M. Нона у. _- e 323, 360
с М E. L Wed t MU 322
Gear N. E. BE ___- - -_-- о o s 323
Gorgonidium Sel Maan БАНИ л SANA 323
Сутлонасћу R Bee 314
324, pou p
Mangonia Sch 323
тагы Т те И о) 324
Microcasia Becc. 322
Monster 324, 343, 351, 353
Adan
alero TERN var. brevinoda و & L. O. Wil-
jams 319, 565
esee Crueg. 323, 360
370, 587
Mere on Schott 323, 360
Orontium L 314
Peltandra R 329 300
Philodendron Schott Pus 312, 31:3, 314,315, 316. on
313,319. 320; 321, 322, 323, 324, 325. 326. TEI.
329, 336, 340, 341, 342, 343, 344, 345, 346, 347,
348, 349, 350, 351, 352, 353, 356, 357, 358, 359,
360, 361, 362, 364, 365, 366, 367, 368, 369, 370,
371, 372, 373, 374, 375, 377, 378, 379, 380, 394,
395, 436, 459, 461, 488, 546, 565, 576, 587
subg. Meconostigma (Schott Engl. .. 315, 317, 322,
324, 325,326, 344, 345, 346, 348, 351, 358, 360,
361, 363, 365, 367, Ыга 373, 375, 3
subg. Philodendron Schot 311, 312, 314341,
8, 319, 320, 321, 392, 323, 324, 325, 326, 328,
372, 376, 377, 378, 379, 380, ex 395, 576, 697
subg. Pteromischum Scho 2, 314, 315, 318,
319, 322, 324, 325, 326, 328, Ят 343, 344, 346,
351, 356, 358, 359, 360, 361, 363, 364, 367, 368,
369, 370, 373, ple: 375, 0, 380,
sect. Baursia (Rchb. ex Schott) , 318,
26, 327, 328, 334, ко 338, Pm 366, 368, 516
sect. Calostigma (Schott) Pfeiffer 17, 324,
26, 327, 328, 330, 331, 332, 335, 336, 338, 339,
347, 351, 364, 365, na 369, 370, 392, 397, 401,
, 405, 410, 414, 419, 420, 422, 424,
426, 429, 433, 435, 437, 438, 439, 442, 452, 453,
, , 478, 480, 490, 491, 492, 493,
497, 500, 517, 526, 528, 533, 7 538, 550, 551,
552, 553, 559, 561, 565, 575, 576
sect. Camptogynium К. Krause 312, 317, 318, 326.
sect. Cardiobelium а зоре Engl.
sect. Fruticosa Grayum a us 3
sect. ‘ea Xd. 312, 317, 318, 326, 332.
35, 337, 338, 347, 369, 470, 575, 576
. 817, 318, 326,
sect. Macrolonchium (Schott) Engl. .
9, 3
sect. Mec eg biases! Ени. 2. 316, 348, 363
sect. Oligocarpidiu 332
sect. Oligospermium er
sect. penne Schott 312, 315, 316, 324, 326,
328, 330, 331, 332, 333, 335, 338, 342, 347, 351,
364, 365, 366, 370, 394, 395, 402, 409, 412, 415.
417, 418, 421, 428, 431, 441, 444, 445, 447, 448,
450, 455, 456, 459, 473, 477, 485, 488, 495, 496.
502, 503, 505, 512, 522, 524, 530, 532, 535, 540,
542, 554, 575
sect. Philopsammos G. S. Bunting 312, 327, 328,
576
,
5 , s
sect. Platypodium (Schott) Engl. _____--------- 329
а ан ie ae i e III мани “авио c
— —
me 423, 487, 514, 549, 575, 576
subsect. Achyropodium (Schott) Engl. ........
55, 488, 530, 532, 554, 575, fit
— Belocardium age Engl i. 332, 3
-- 335, 336, 337, 339, 351,
subsect. Bulaoana May
vie Canniphy um (Schott) Mayo ____ 328, 331,
334, 428, 512, 575, 576
subsect. унту атм Ке 330, 394
subsect. Doratop. 3
yuu,
subsect. Eubelium (Schott) E Роса a 0
subsect. Eucardium (Engl.) "а те di 236, 339, 575,
576
subsect. Glossophyllum Spore Croat. i 327, 333,
334, 335, 339, 350, 369, 397, 406, 407, 414, 422,
424, 433, 435, 442, 452, 468, 480, 491, 492, 500,
526, 533, 538, 551, 552, 559, 561, 575, 576
subsect. Macrobelium (Schott) Engl. _ 332, 333, 334,
338, 339, 351, 392, 401, 404, 408, 410, 420, 426,
429, 437, 438, 439, 453, 472, 475, e "We 493,
516, 529, 537, 550, 553, 565, 575
subsect. Macrolonchium (Schott) Engl. ...... yet 329,
5,575, 576
subsect. Oligocarpidium (Engl.) Mayo -...... 385, 339,
419, 466, 575, 576
subsect. Philodendron Schott __ 329, 330, 331, 339,
395, 402, 409, = 447, 450, 455, 473, 477, 485,
495, 496, 505, 522, 524, 535, et 542, 575, 576
subsect. Playelatum. emp
T oiga hott) кай, > ` 329, 331, 415,
1, 441, 444, a,
Es. Psoropodium (Schott) Engl. .. 329, 330, 338,
576
subsect. Solenosterigma (Klotzsch ex Schott) Engl.
29, 330, 331, 335, 338, 412, 459, 503, 575, 576
ser. Albisuccosa Croat ____ 331, 339, 394, 575, 576
ser. Ecordata Croat _ 333, 339, 369, 410, 426, 429,
478, 493, 575, 576
ser. Fibrosa Croat __ 331, 339, 395, 402, 409, 417,
1, 450, 473, 477, 485, 495, 496, 505, 522, 524,
540, 575, 576
ser. Glossophyllum Croat ... 334, 339, 397, 405, 407,
415, 422, 433, 442, 452, 468, 480, 492, 500, 551,
ser. Macrobelium (Schott) Croat 333, 339, 392, 401,
0, 437, 438, 439, 453, 475, 490, 497, 517,
529, 537, 553, 565, 576
ser. Ovata Croat 334, 335, 339, 424, 435, 491,
333, 339, 472, 550, 575; 576
330, 336, 339, 447, 575, 576
Volume 84, Number 3 699
1997 Philodendron Subgenus Philodendron
sect. еен Engl. _ 317, 318, 324, 326, 328, grex Achyropodium Sehott _____________ 316, 331
0, 364 grex Baursia Rchb. ex Schott __________ 315, 326
sect. poe (Schott) Engl. ......... 312, 317, 318, grex Belocardium Schott... 316, 332, 334
20; 228; 332: 337, 338 344 851 436, 507,557, grex Canniphyllum Schot _____________ 315, 328
, 576 grex Cardiobellium Sehott шшш 316, 330
sect. Preromischum (Schott) Engl. ___ 315, 316, 317, grex Cardiophylacium Schott
26, 327,348 grex Doratophyllum Schott
sect. Schizophyllum (Schott) Engl. ..... 312; 317, 318, rex Eubelium Schott
326, 337,338, 576 grex Glossophyllum Schott -.................. 315, 332, 334
sect. Schizoplacium (Schott) Engl. ............... 317,351 grex Imbea
sect. pat ipa че (Schott) Engl. __ 312, 317, 318, grex Macrobelium Schott ...................... 316, 332, 459
6, 332, 335, 336, 337, 338, 347, 351, 365, 397, grex Macrolonchium Schott ......................... 316, 328
grex Psoropodium Schott 315, 329
grex Pteromischum Schott .................-.....-.-.. 315, 316
grex Schizophyllum Schot... 316, 337
grex Solenosterigma Klotzsch _______ 315, 329, 459
grex Sphincterostigma Schott... , 316
grex Tritomophyllum Schott z-a- he 336
Gruppe Achyropodium Schott
Gruppe Belocardium Schott ......................... 317, 332
Gruppe Cardiobelium Schott... 317, 330
Gruppe Doratophyllum Schott ______ 335, 336, 351
Gruppe Eucardium Engl. 336
Gruppe Macrobelium Schott 317
Gruppe a IM ааа а ай 317, 351
Gruppe Platypodium
Gru ruppe Solenosterigma LED Schott _
317, 335
pecies:
accrescens N. W. Simmonds
acrea rause
а Schott
acuminatissimum Engl. .....
m Schot
327
316, 332, 461
30 339
acutifolium K. A um 334
1 334
advena Schott... 317, 318, 319, pos 332, 333, 334,
339, 344, 349, 353, 357, 3 363, 366, 368,
369, 370, 371, 376, 379, 390, pera 505, 517,
529, 570, 573, 571, 589, 598, 697
affine Hemsl. 4 {
albisuccus Croat ........ 311, 331, 339, 353, 355, 357,
376, 386, 393—394, 570, 573, : )
alticola Croat & Grayum . 311, 322, 331, 339, 355,
356, 358, 360, 370, 386, pw 425, 524, 534,
570, 573, 577, 591
amplectans А. C. Sm.
andreanum Devansaye .........................
angustialatum Engl. .
angustilobum Croat & Grayum .. 311, 322, 336, 338,
345, 350, 354, 355, 357, 358, 359, 362, 379,
396—397, 490, 570, 573, 571, 592, 598
413
331, 336, 447
334
318, 507
337
316, 317, 318, 319, 320, 336,
338, 345, 351, 352, 353, 355, 351, 358, 361, 369,
370, 379, 383, 397—400, 544, 565, 570, 573, 511,
587, 590, 593, 598
annulatum Croat € Grayum ...... 311, 322, 333, 339,
348, 349, 350, 351, 353, 355, 357, 370, 383, 384,
386, 387, 388, 400—402, 517, 559, 570, 573, 511,
594
antonioanum Croat ... 311, 331, 339, 347, 348, 351,
700
Annals of the
Missouri Botanical Garden
353, 356, 357, 358, 359, 360, 365, 376, 402—403,
550, 570, 573, 577, 595
apocarpum ккө.
arcuatum К. Kraus
armigerum Standl. & І. O. Williams ____
aromaticum Croat & Grayum _ 311, 322, 333, "339,
353, 355, 357, 358, 359, 377, 390, 402—404, 453,
565, 570, 573, 579,
318, 320, 469
331, 412
asperatum K. Koch 329
augustinum K. тыз $17
aurantitfolium Schot 315, 916
veru Standl. & L O. Williams ___ 318, 319
4, 339, 346, 348, 350, 351, 352, 355, 358, 359,
T 369, 377, 385, 404—406, 434, 500, 559, 570,
973, 577, 582, 584, 585, 596, 597
auritum Lindl. 565
bahiense Engl.
bakeri Croat € Grayum __ 311, 321, 322, 334. 349,
349, 350, 351, 355, 356, 357, 366, 369, 376, 383,
1,492, 518; 570; 575. 1% 599
barrosoanum g S. Bunting ____ 5, 336, 337, 490
basii Matuda __ 318, 319, 320, s "330, 342, 352,
353; 355, p 8957; 376, 377, 387, 407—408, 570,
578.977.
К. Kruse.
billietiae Стоа
RAHSA R Schott ex Endl. _
blanchetianum Schott
brandtianum K. Krause 329
bre sgt Cmat — SELL I2 FFl, E 357, 358,
60, 376, 377, 388, 389, 390, 409, 437, 529, 570,
e 571, 599, 601
ama рар. 318, 319, 320, 321, 333, 339, a
ears 361, 365, 369, 370, 390, 409-412
0, 573, uir 588, 601, 602
"uet ul SORO. 00 0
brevinodum Standl. A: L. O. Williams .
brevispathum Schott _ 316, 317, 319, 330, 331, 337,
338, 343, 345, 357, 361, 369, 378, 379, 388,
412-413, 470, 570, 573, 577, 602
subsp. holmquistii (G. S. Bunting) G. S. Bunting
412,
3
361, 371, 372, 373
328
329, 444.
brewsterense Croat __ 311, 312, 334, 339, 349, 352,
354, 355, E ku 376, 383, 413-414, 570, 573,
577, 413-4
brunneicaule Qr & GET um .. 311, 322, 329, 338,
345, 350, 351, 352, 353, 354, 355, 357, 358, 378,
388, 389, 414—416, 421, 570, 573, 577, 603, 604
bulaoanum т A 335, 337, 339
334
te ЕН
nr ИН 27, 328, 361
subsp. po ere 5. Bunting -. 327
canaimae G. S. Bunt ing .
mn и 327
chiriquense Croat ...... 311, 327, 331, 339, 344, 345,
347, 349, 353, PN 357, 376, 386, 416—417, 421,
570, 573, 577, 604, 606
chirripoense Croat & Grym . 311, 312, 321, 325.
1, 344, 349, 352, 354, 355, 357, 359, 365, 376,
377, 383, 417-418, 491, 570, 573, 577, 605
clematis Wright in Griseb. i 329, 444
clewellii Croat .. 311, 335, 339, 351, 355, 357, "359,
389, 390, 418-419, 570, 573, 577, 605
coerülescóns ЕШШ, LS ушш 318, 328
chult. 455
coloradense Ci. do 311, 333, 339, 353, 376, 388,
390, 419—420, 572, 511, 605, 606
conforme G. S. Buntin 581
consanguineum Schot 9, 332
— sees 311, 329, 338, 344, 349, 350, би 354,
5, 356, 357, 358, 359, 365, 385, 386, 415, 417,
ш 496, 570, 573, 577, 607,
corcovadense Кит 344
cordatum hort. non Vell.
correae Croat __ 311, 334, 339, 350, 351, 354, 355,
357, 361, 364, 367, = 383, 401, 421—422, 443,
480, 570, dud 577,
cotobrusense Сто Мера 311, 312, 322,336;
338, 352, 353, 355, 357, 376, 377, 382, 422–423,
487, 514, 544, 545, 570, ve ni
cotonense ска: & Grayum ___ 335, 339, 345,
350, 356, 357, 367, pind 388, po 423-425, 561,
570, 573, 577, 606, 6
craspedodromum R. E. Seba:
crassinervium Lindl
crassispathum Croat & Grayum 311, 319, 321, 322,
333, 339, 350, 353, 355, 357, 358, 350, 363, 369,
370, 387, = 414, 425—427, 570, 573, 577, 607,
610, 611,
cretosum Croat $ Grayum 311,322, 328, 331, 339,
349, 353, 355, prin 365, 384, 394, 427-428, 570,
573, 571, 611, 6
cuneatum Engl. __ 328, 512
cuspidatum K. Koch & Bouché _______ 459, 460
cyrtocoleum F. Diels
monum Liebm.s 55 2 АЛ. 316, 317, 320, 516
dagilla Soha 6, 397, 398
daguense Linden & André 553
davidsonii Croat ____ 318, 319, 320, 333, 339, 345,
349, 350, 351, 353, 357, 358, 359, 360, 362, 363,
366, 384, 385, 428-429
subsp. davidsonii Croat _
573, 577, 612, 613
subsp. bocatoranum Croat ___ 311, 351, 357, 365,
5, 429, 430, 570, 573, 577, 584, 612, dee
2
348, 355, 429—430, 570,
iatum Sch 332, 335, 469
discolor: K: Krause 20 ae а UR
distantilobum K. Kraus 337
e
dodsonii Croat & Grayum 311, 322, 331, 335, 339,
345, 347, 352, 355, 357, 359, 360, 376, 378, 379,
386, 387, 430-432, 435, 453, 524, 570, 573, 577,
614, 615, 617
dolichophyllum Croat ____ 311, 322, 334, 339, 351,
394. 355, di 360, 370, 384, 432—434, 570, 573,
577, 616,
dolosum ы E 009 9 Оа К 329
dominicalense Croat & Grayum 311, 335, 339, 355,
376, 377, 387, 570, 573, 577, 431, 434—435, ere
dressleri G. S. Ванта SN 318, 319, 320, 321, Ed
344, 347, 351, 353, 354, 355, 357, 365, 376, 311.
382, 435—436, 507, 510, 557, 570, 573, 577, 617,
619
dunstervilleorum G. S. Bunting _________- 321
dwyeri Croat ___ 311, ke "30, 333, 339, 352, 353,
Зэт. se: 364, 376, 377, 390, 436-437, 57 10, 513,
577,6
Misi ^ E. Schult. var. dyscarpium I Ac BN 321
nN. dia аы. 2. з... ТТТ
A av
"о НЕ ج >
Volume 84, Number 3 701
1997 Philodendron Subgenus Philodendron
сене Trig: E о A AA 539 Harlow E-M: Johns <<< 15055 318, 320, 460
edenudatum сом 311, 333, 339, 345, 350, 354, hastiferum Standl. & L O. Williams ____ 319, 565
357, 360, 376, 388, 437-438, 453, 454, 570, 573,
577, 618, 620
effusilobum Uu C o RS dH 335, 336
MOT TMB ESA ДОПА И A wise 372
Haphéglosioides Schott. DE SES 334
ele, UR I ts Ne ada uot andi. 337
duc Enak AAA 327
engléitunum Steyerm., ¿O 32
епапзопа 1. M. Johnst. L2 saa 318, 320, 469
rnet Engel A uL ПЕ. 329, 415
ërubestens Linden <- o din. пе оне 332
Sender Ri Magara Ло Сани и ак 328, 337
PME i. usus 317, 318, 543, 545
333, 339, 344, 350, 351,
353, 355, 359, 360, 2d. 369, 370, 376, 378, 390,
438—440, 453, 486, 570, 573, 577, 621, 623, 626,
97
оокат Poepp 20 1145s 328, 334, 345
fi Croat € Grayum __ 311, 319, 322, 329, 338,
345, 349, 351, 352, 354, 355, 356; 357 358-359,
360, 369, 370, 382, 385, е 502, 524, 570,
573, 577, 581, "ps er
folsomii Croat _ 311, ois Чеч 858: 355, 357;
e Croat 311;
352, 353, 357, 389, 443-444, 5415573; 577, 624,
fragrantissimum (Ноок.) С. Don ___ 320, 328, 329,
, 341, 344, 349, 357, 359, 360, 363, 365, 369,
376, 378, 379, 385, 444—446, 571, 573, 577, 587,
26,
Fragrantissimum (Hook.) Kunth _____ 316, 317, 318
fraternum Schott ________ 330
330, 339, 585
gigas Croat ___ 311, 330, 331, 336, 339, 344, 346,
= 349, 351, 352, 353, 355, 356, 357, 359, 360,
386, 446-447, 571, 573, 577, 627, 628, 631
пи Машаа ___ 3672 518. 319; 320, 321,
338, 342, es 348, 350, 351, 352, 355, 357, 371,
379, 447-449, 488, 573,
subsp. inibi —. 331, 379, 387, 448, 571,
577, 629, 630
subsp. camiloanum Croat __________ 448, 629
rcg André ____ "€ 444
eren ы ч Вагтозо 351
gracile Schott ______ 315;.316,.317,:318, Mes 539
cando (Jacq) Schott = 315, 330, 380
Krause 317, 318, 319, 329, 331, 338,
em 342, 345, 347, 348, 349, 350, 353, 354, 357,
359, 360, 361, 363, 369, 374, 376, 378, 379, 380,
387, 449—451, 473, E 574, 577, 630, 631, 633
granulare Croat ____ 34, 339, 353, 355, 357,
364, 368, 369, 370, jet. 451-452, 571, 574, 571,
631,
graveolens ae
grayumii dar 311, 322,338, 349,345; 351, 353,
re, 355, 356, 357, 360, 361, 370, 388, 419, 420,
1, 438, 440, 452-454, 571, 574, 577, 632, 633
331
malense Engl. 317
guttiferum Kunth 317, 319
hammelii Croat 311, 312, 331, 338, 342, 345, 350,
357, 376, 387, 454-455, 488, 532, 555, 571, 574,
77, 634
hebetatum Croat ........ 311, 330, 339, 347, 351, 352,
353, 355, 357, 359, 360, 362, 368, 378, 379, 455,
536, 542, 571, 574, 577, 588, 634, 636
hederaceum (Jacq.) Schott 314, 315, 316, 317, 318,
319, 320, 329, 330, 332, 335, 338, 341, 343, 345,
353, 357, 358, 360, 363, 364, 365, 369, 370, 376,
377, 378, 379, 385, 388, 389, 458-460, 470, 504,
5
var. hederaceum .. 317, ЗАЗ, € 378, 460, 460—
463, 464, 571, Sad 577, 63
т. hederaceum f. micans... 317,318
var. kirkbridei Cit 4L 4 51k 321, 348, 378, 388,
571, 574, 577, 635
уаг. pees (Schott) Стол 311, 316, 317,
318, , 460, 5, 571, 574, 578
бодна sensu Kunt 340
heleniae Croat ... 311, 335, 339, 345, 349, 350, 353,
355, 356, 357, 359, 360, 362, 370, 377, 378, 380,
5—467, 478, 570, 574, 578, 636, —€—
heterophyllum Poe
hoffmannii Schott -..... 316, 317, 319, 320, 332, 387
hoffmannii Schott sensu Engl. |... , 459, 469
holtonianum Schott 35, 336, 546
houlettiana En 337
ylaeae G. E: Bunting 335, еј
imbe Schot
immixtum Cra ED 311, 334, 339, 343, 345, n
350, 351, E 353, 355, 357,360, 362, 364, 365,
378, 379, 84, 407, 467—469, 481, 551, 570,
574, 578, 638
impolitum Schott 316, 507
inaequilaterum + AA 315, 317, 328, 369
inconcinnum Schott 364, 468
jacquinii Schott 316, 317, 318, 319, 320, 332, 335,
337, 338, 341, 343, 345, 349, 357, 363, 364, 365,
369, 370, 376, 378, 379, 388, 389, 413, 459, 460,
469-471, 571, bies 578, 637, 639, hey
320, 391
jamapanum (5. de age жааса шыны
јејепзе ___ Cro LB 312, 3 3, 339, ssh 357, 366,
370, 376, sm. 471-40) e 571, 574, 578, 639
jenmanii K. Kra 330, 335
jodavisianum G. $ Bunting .... 318, 319, 320, 321,
331, 339, 340, 347, 349, 350, 353, 357, 359, 360,
361, 362, 363, 378, 379, 386, pai 472-475, 495,
542, 571, 574, 578, 640, 641,
juninense Engl. 328
arsed Schot
knappiae Croat .. З 333, 339, 342, 349, 355, 357,
359, 365, 376, 390, 411, 475, 571, 574, 578, 639,
krebsii Schott
lacerum (Jacq.) Schott
laciniosum Schott
lancigerum Standl. € L. O. Williams
latilobum Schott...
latipes K. Koch & Augustin
latisagittium Matuda 319, 320, 489
lazorii Croat ...... 311,322, 331, 339, 347, 352, 353,
360, 387, 472, 476-477, 495, 571, 574, 578, 583,
27 229
315, 337
315
319, 516, 518
336
2, 64.
"a Mayo БИ Bars -< < — 344
chlerian „венци: 329, 498, 504
lehmannii Engl.
[етй cerdo E Grayum ..... 311, 321, 333, 339, 342,
702
Annals of the
Missouri Botanical Garden
349, 352, 354, 355, 457, 369, 383, 384, 466,
477-479 9, 484, 493, 571, 574, 578, 645, 646
unting
ligulatum Schott ___ 316, 317, 318, 320, 334, 339,
345, 349, 350, 351, 355, 357, 359, 360, 362, 397,
401, 422, 468, 479-481, 500, 559
var. ligulatum ...... 354, 357, 378, 379, 380, 384,
, 481, 481-482, 484, 500, 571, 574, 578,
var. heraclioanum Croat 311, 334, 349, 353,
57, 364, 367, 384, 480, 481, 482, 482-483
500, 559, 571, 574, 578, 648
var. oratum Croat 311, 334, 349, 357, 360, 383,
2, 483-484, 571, 574, 578, 647,
limnestis стене & Daghlian
lindenii Schot
327,
боце sie -- 911/2322; | 339, 349, 350, s
360, 362, 370, 386, 387, 439, 484-486, 524,
78, 646, 649
longilaminatum Schott < эт е у 327
sti kami. o Hm anc ones А27
чы ar -e PEE A We Maran ОЕ 334
longist Пит, K KARE uu җи и 338
madronense Croat __ 1, 312, 336, 338, 351, 352,
354, res 376, 382, MES 486-487, 545, 571, 574,
78, 64
maguirei G. s MERE к a 540
malesevichiae Croat _ 311, 331, 338, 342, 345, 350,
351, 357, 360, 378, 380, 385, 487—489, 555, 571,
574, ie 580, 585, 586, 650
таи K Ke NAS
mela. anochrysum d & André
melinonii Brongn. ex E NE зс о
mexicanum Engl. ___ 317, 318, 319, 320, 333, "330,
342, 345, 352, 354, 357, 358, 359, 360, 362, 363,
370, 378, io 383, 388, 389, 397, 489-490, 571,
574, 578, 6
micans K. Koh "e sh 317, 318, 329, 459, 460
micranthum Poepp. ex Schot
microphyllum K. Koc
wont Standl. & L. O. Williams __ 319, 335,
349, 351, 352, 353, 354, 355, 357, 358, 369,
ЗҮ 383, 418, 490, 538, 571, 574, 578, 652
miduhoi Маша 319, 320, Mia 460
1
19, 320, or 392
311, 334, 339, 352, 353, 365, 370,
, 384, 492-493, 571, 574, 578, 6
P o а 3
simi "e PUN 311, 333, 339, 348, 353, 355,
365, 367, 376, 371, 385, 493, 571, 574, 578, 653
obtusilobum чч
ornatum Schot
oxycardium Schott __ 315, 316, 317, 318, 329, 459,
460, 464.
осурган SORELLE E oi sS 461
Jechyeaul K. Күанве 8S un а 327
334
rause 317, 318, 319, 320, 330,
331, 339, 348, 351, 352, 353, 355, 357, 358, 360,
362, 365, 376, 386, 387, 473, 477, 493-495, 526,
571, 574, 578, 654
entume Weis Kumbh o ies 321, 336
pas n Kk Kause iu dixic eoe 336
ianum а К. ERMME а iu Bul 321
pedatum (Hook) Kunth sullo 337
рее 6. S, Bunting мз des 327
phlebodes С. S; Bunting ic он 327
pleasant End. LIS ety) sea 331, 553
pimichinense С. S. Bunting _____________ 3
isi uia (Jacq.) Schot RE qat tror 315; 328,:337
e Croat ... 311, 331, 339, 347, 348, 350, 353,
354, 357, 376, 387, 495—496, 571, 574, 578, 655,
656
ринен Engl. 2. 317, 318, 320, 335, 377, 461
platypetiolatum Madison 319, 320, 333, 339, 350,
352, = 355, 357, 378, 379, 388, 496, 498, 571,
56
peoneiam Standl. & L. O. Williams __ 319, 449
polytomum Schott _ ‚ 318, 507
444
311,322, 884: 339, "348,
351,352, 355, 358, 359, 369, 378, 380, 384, 405,
433, 434, 483, 498-501, 552, 571, 574, 578, 588,
654, 657, 658
pitas ко ads 319, 320, 510
(ues A дк 327, 328, 361
pterotum K. Koch ¢ & Augustin _ 316, 317, 318, 319,
320, 329, 331, 338, 345, 349, 352, 357, 359, 360,
361, 363, 376, 379, 385, 441, 501-503, 571, 574,
578, 657, 6
pulchrum С. M. Barroso _
purpureoviride Eng 2327318, 330: 838: 345, 347,
dos. 357, 369, 370, 376, 378, 389, 459, 503-504,
538, 571, 574, 578, 659
Lo TH 319320; 331339, "350,
352, 353, 355, 357, 358, 359, 370, 376, 379, 387,
392 90, 517, 572, 574, 578, 660,
quitense е Eng
us 316, 317, 318, 319, 320, 321,
337, 338, 342, 349, 352, 354, 355, 357, 360, 361,
363, 364, 367, 370, 374, 376, 377, 379, 436, 506,
507, 537, 557, 580,
UE 349, 378, 382, 507, 507—510,
572, 574, 578, 662, 663
var. pseudoradiatum (Matuda) Croat ...... 311, 319,
320, 337, 357, 371, 382, 436, 507, 510, 537,
reichenbachianum Ѕећой Issn 327
remifolium R. E 327
robustum Schat 2 ево Пра пат EE 328
řobustum Sodiro ined. с ос 432
foraimae К. Kame су. 524 0 ла 328
roseospathum Croat .. 311, 328, 331, 341, 347, 349,
‚ 353, 355, 360, 366, 370, 384, 511-512, 666
var. rosespathum .. 342, 512, 513, 572, 574, 578,
665
TT LLB Ut
Volume 84, Number 3
1997
var. angustilaminatum Croat 311, 312, 359, 512,
3, 572, 574, ie 667
ponia (Engl. & K. Krause) Croat & Grayum
6, 338,345, 348, 351, 352, 357, 358, 359,
360, e 370, 379, е 513-516, 572, 574, 578,
580, 582, 663, 664.
cl 516-521, 537, 545, 550, 552, 553, 565, 572,
78, 666, 667, 668
кшн SM ii dun 316, 339, 401
КОНИ e Lu
sanguineum Regel з es 317, 318, 320, 516, 518
sc MRNA o ә.
a Croat € Grayum __ 311, 322, 331, 339,
7, 349, 350, 353, 355, 356, 358, 360, 378, 380,
HA 521—523, 572, 574, 578, 669, 670
scandens K. Koch & Sello - SEEN 315: 316, 317, 318,
329, 377, 459, 460, 4
f. micans (Schott) G. S. Aid
subsp. oxycardium (Schott) G. ©. Bunting ....
var. cuspidatum (К. Koch & Bouché) Engl. .. 460
schottianum Н. Wendl. ex Schott ___ 317.318, 320:
330, 331, 339, 345, 347, 348, 350, 351, 352, 353,
356, 357, 358, 359, 360, 361, 385, 386, 387, 396,
435, 485, 486, 523-525, 524, 542, 572, 574, 578,
580, 584, 671, 672
ерле боро o a ао 315, 317
«Деши K: Koch e о 372. 548
ерен Hook, Ба ка ата Dal, 530
мен (Пор. Род у
.) Don
smithii Engl. us 317, 318, 320, 334, 335, 339, 345,
350, 351, 353, 359, 360, 361, 364, 368, 369, 370,
379, 389, € 925-528, 561, 572, 575, 578, 670,
672, 673, 6
sodiroanum = рол UM - 539
sodiroi hort.
sousae Croat |... 211, 312, 333. 339; 352: 357. 359,
364, о 376, 377, 389, 409, 528-529, 572, 575,
578,6
speciosum cibi R4 rr 372
sphalerum Schott = ______ 442, 468, 492
splitgerberi м 329
squamicaule Croat & Grayum .. 311, 331, 338, 342,
345, 347, 350, 352, 353, 354, 355, 357, 358, 365,
366, 378, 380, 385, 529-531, 555, DIR ISO,
673, 675
squamipetiolatum Croat ___ 811. 33L po 345, 350,
352, 354, 357, 365, 378, 380, 383, 389, 447, 455,
488, € 555, 572, 575, 578, е, 676
standleyi Стаушт _____ 370
steyermarkii С. s. Bunting 827
straminicaule Croat _ 311 322, 335, 339, 344, 345,
351, 353, 355, 358, 359, 364, 386, 390, 394, 395,
410, 526, 532-534, 561, 572, 575, 578, 581, 673,
677
strictum С. S. Bunting __ 319, 321, 330, 339, 347,
353, 355, 350. 360, 370, 378, 379, 385, 456, 457,
534-536, 542, 572, ad die 678, 679
subhastatum Engl. € K. K
subincisum Schott
Croat
Philodendron Subgenus Philodendron
703
353, 360, 365, 377, 388, 536-537, 572, 575, 578,
678, 680
subovatum Schott ________ 317, 318, 332, 334, 391
sulcicaule Croat & Grayum ___ 11, 321, 335, 339,
343, 345, 352, 353, 354, 355, 357, 362, 388, 389,
537-539, 572, 575, 578, 679,
surinamense (Miq. ex Schott) Engl. Зи ние Mis SS 8 368
П A Ss ПИ 317
carpa за E E 316, 516
A со алашы шыт
tenue K. Koch & Augustin ___ 315, 316, 317, 318,
320, 330, 331, 339, 348, 349, 350, 351, 352, 857,
360, vn 379, 388, 539-541, 572, 575, 578, 680,
681,
tenuipes En er E RN
thalassicum Croat & Grayum __ 311, 321, 322, 330,
339, 349, 350, 351, 354, 355, 385, 386, 456, 524,
, 940, 541-543, 572, 575, 578, 681, 682, 683
— Schott I Ta uu. 329
ie NINE a iU dE T d Li T
tripartitum (Jacq.) Schott .. 315, 316, 317, 318, ne.
320, 326, 336, 338, 345, 350, 352, 355, 359, 361,
362, 364, 367, 368, 378, 379, 382, 398, 399, 423,
Bi 514, 543-549, 572, 575, 578, 683, 684, 685
e Wicuspiaatum Eng. — . .. — 543
irc BENE E 319, 320, 397
tla 320
untin 0, 516
tysonit Croat __ 311, 322, 333, 339, 348, 352, 353,
356, 357, 359, 361, 370, dl 388, 402, 0 419,
549-551, 572, 575, 578
ubigantupense Croat 311, 3 2, 334, 339, a 855,
359, 376, 377, 383, 551, + 575, 578, 688
uleanum Engl.
urbanianum K. a. MESES 329
gren Cos 312, 334, 339, > 357,
7, 384, 551-552 P 575, 519, 68
МИ
v ense Croat ___ 1, 312, 319, 320, 333, 339,
ony 352, 355, 857; Pi 389, 552—553, 572, 575,
9, 688
моин L. Mathieu ex Schott... 316, 317, 318,
320, 331, 338, 345, 350, 353, 357, 358, 365, 377,
378, 379, 385, 387, 447, 488, 530, 531, 553—556,
912, 919, is 689, 691
victoriae G. S. Bunting 336, 337
viride En 334
wallisii Regel ex Engl. 336
warszewiczii К. Koch & Bouché ____ 316, 317, 320,
321, 325, 337, 338, 342, 345, 347, 348, 349, 350,
351, 352, 353, 357, 359, 364, 365, 366, 367, 369,
370, 379, 382, 408, 436, 507, 556—558, 572, 515,
579, 583, 690, 691, 692, 697
weberbaueri Engl.
ot 1 1
334, 338, 339, 345, 351, 355, 358, 360, 363, 369,
wilburii Croat & Grayum _ 311, 321, 322, 335, 339,
, 362, 369, 370, 389, 377, 388, 475, 492,
504, 526, 559-561, 562
var. wilburii .. 319, 321, 357, 377, 561-563, 572,
515, 519, 692, 693, 694
var. longipedunculatum Croat € Grayum ___ 311,
321, 352, 353, 358, 366, 389, 525, 534, 561,
, 563-564, 572, 575, 579, 694, 695
williamsii Hook. uo EES ESE EIS УМЕ ПОЗВА 372
704 Annals of the
Missouri Botanical Garden
илаш Eel. A АН 334 Synandrospalliz.Empl. SS оа as йды 323
wurdackii G. S Bu üng о d. 334 Syngonium (Engl.) Стон +! 314, 315, 343, 370, 565
num Croat 311, 333, 339, 350, 353, 356, 357, armi, yee (Standl. & L. O. Williams) Croat __ 319
360, 364, 388, 390, 564—565, 572, 575, 579, 696
Philodnidrum Schot LAE Ss 315, 380, 576 haran (Standl. & L. O. Williams) Croat __ 319,
Piptpaha NB A 322
NE diia A онь 314 ње ics pi а ЕВЕ 513
go. P A E ta. УР ЗОЯ 31 Taccarum Brongn. ex Schot E ы A 323
Rhodospatha Pap. Аа A E 9207359 Трой Rak os 20100 © s emis 380
auromatum Kunth i BL 37 grandifolius-(Jacq.) Raf... — 12. и 380
омањи Zoll. & AAA 324 Thaumatophyllum Schott ZC REE 380
Sue Soho o ЕН 323 spracepnum Soho NC EO a O 380
Spathica OOK A е AN А NE 32 Bphonodonim Schott ccoo cic 322, 360
Spathiphyllum Se h O о 315, 324, 353 B cens SOME. uL LOO umo SE.
Sphincterostigma [unranked] _______________ 32 5 (К. Koch & qus К. Koch коло 374
Stenospermation Schott. inpoi 2 i used 353 ovt jn B eeu EAR 314, 360
Volume 84, Number 3, pp. 311-704 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN
was published on October 24, 1997.
- Geographical Distribution and Endemism
Petioles
Blades
Blade Shape __
Posterior Lobes
Posterior Ribs
Lesser Order Veins
Morphology of Reproductive Structures
Inflorescences
Peduncles
Spathes
Spathe Color
Spadix
Male Flowers
Pollen
Female Flowers
Placentation
Ovary Locule Number |
Ovules per Locule
Style and igne Morphology
Style Type
Friis and 1 Seeds
Pollination Biology
Breeding Behavior |
Phenology
A. Flowering i in Dry Season and Wet ‘Semon
B. Flowering only in Wet Season
C. Flowering only in Dry Season
D. Flowering АП Year
E. Flowering Bimodally
Cytology
Taxonomic Treatment
. Generic Description
+ Mey 40 Bue E Philodendron |
Subgeneric Descript
Keys to Species, of "Philodendron Subg. Philodendron
| Spec à
Excluded Ne Kasa | cd
- Literature Cited | )
Appendix 1; Geographic Distribution of Central American Philodendron Subg. Philo- +
dendron
Appendix 2: Technical Data on Pistils -.
_ Appendix 3: Composition of Philodendron Subg. Philodendron in (rs America
про, 4: Зарвон Patterns of Central American Philodendron — — — —
Figures |
Index.
СОМТЕМТ5
А orion of Philodendron Fes Philodendron A for Mexico and Central
merica B. Croat . Sis
- Materials and Methods _ A | БАА) ELT БУ
— Acknowledgments |... n EE PS ANS Maat A I г
.ı History of the Genus Philodendron "S ab ыл жогарЫ
Heinrich Wilhelm Schott .___. | и Баија ~
_ Adolf Engler ____ eae
Кип Krause _
Modern Won 22222.
Collecting History _.
-Fossil History РА
Intrageneric TSS EAS
Infrageneric Relationships _______
Key to Subgenera of Philodendron __ |
- Relationships Within P. Subg. Philodendron — A
. Discussion of Subgeneric Classification - EAN А
М А Р ` Baursia · әсе - а on
2521 9 eek. Philopsami iius Si
САС P Sect. Philodendron —
LE Subsect. Macrolonchium 28. 2.P CR тате не (328); de
| P. Subsect. Platypodium (329); 4. P. Subsect. Psoropodium (329); 5. _
> Subsect. Solenosterigma (329); 6. P. Dubseet: Philodendron ет
7. P. Subsect. Achyropodium TE. ы
4, P. Sect. Calostigma _ =
AR барве. Macrobelium. (332) 2. R Зарнен, С СЕ 3 P i
у ах ~
› Sect. "Schizophyllun - Set TRNA ie
› Sect. Polytomium => ped ime E
› Sect. азаа =a reas x
9. P Sect: Camptog
Key to Sections and узебез ог e Р Sube. Phen. A LE
Morphology of Vegetative Structures iria moran ES. Ta |
Anatomy 2 чый СЫ
`. Vegetative Anatomy by R. € R. с. I
Vascular ' Anator -
Annals
` of the
_ Missouri
. Botanical
Garden |
1997 Y
Number 4
Volume 84, Number 4
. Fall 1997
Annals of the
Missouri Botanical Garden
The Annals, published quarterly, contains papers, primarily in systematic botany,
contributed from the Missouri Botanical Garden, St. Louis. Papers originating out-
side the Garden will also be accepted. All manuscripts are reviewed by qualified,
independent reviewers. Authors should write the Managing Editor for information
concerning arrangements for publishing in the ANNALS. Instructions to Authors are
printed in the back of the last issue of each volume.
Editorial Committee
Michael H. Grayum
Editor, —— |
Missouri Botanical Garden
Amy Scheuler McPherson
Managing Editor, |
Missouri Botanical Garden
Diana Gunter
Editorial Assistant,
Missouri Botanical Garden
Vicki Couture
Secretary |
Ihsan A. Al-Shehbaz
Missouri Botanical Garden
For subscription information contact ANNALS
OF THE MISSOURI GARDEN, % Allen Marketing |
& Management, Р.О. Box 1897, Lawrence, KS
66044-8897. Subscription price is $120 per
volume U.S., $130 Canada & Mexico, $155 all
other countries. Four issues per volume. The
journal Novon is included in the subscription
price of the ANNALS, — х dive.
amcpherGadmin.mobot.org (editorial queries)
http://www.mobot.org l
© Missouri Botanical Garden 1997
The mission of the Missouri Botanical Garden is to discover and share knowledge about plants and
their environment, in order to preserve and enrich life. oe ^s b arit.
Gerrit Davidse
Missouri Botanical Garden
Roy E. Gereau
Missouri Botanical Garden
Peter Goldblatt
Missouri Botanical Garden
Gordon McPherson
Missouri Botanical Garden
P. Mick Richardson
Missouri Botanical Garden
Henk van der Werff
Missouri Botanical Garden.
. The ANNALS OF THE MISSOURI BOTANICAL
GARDEN (ISSN 0026-6493) is published quar-
terly by the Missouri Botanical Garden, 2345
Tower Grove Avenue, St. Louis, MO 63110. Pe-
riodicals postage paid at St. Louis, МО and ad-
. ditional mailing offices. POSTMASTER: Send ad-
dress changes to ANNALS OF THE MISSOURI:
BOTANICAL GARDEN, % Allen Marketing &
Management, P.O. Box 1897, Lawrence, KS
66044-8897. |... | | |
AC aan m
Ф This paper meets ће requirements of ANSI/
Volume 84
Annals
of the
Missouri
Botanical
Garden
Y
A REVISION OF STYRAX
(STYRACACEAE) FOR
WESTERN TEXAS, MEXICO,
AND MESOAMERICA!
Peter W. Fritsch?
ABSTRACT
Despite a recent taxonomic treatment of Styrax from North America and the Caribbean, the taxonomy of this genus
has remained poorly understood. pa
morphological characteristics of o
phasized vegetative characters for circumscription, whereas the p
evision comprises 19 species and 24 taxa in on area amet nearly ice | 7ши
arnatus,
features of the androecium. This r
w species
as тё species аз е previously. It includes six new species (5.
uxtlensis), two new subspecie
, 5. steyermarkii, S.
effort to clarify the ми of Styrax
"2500 s specimens from
from western Texas through Mesoamerica,
erbaria were cxpunined, The е previous treatment em-
includes
i S. inc
. nicar gue. nsis, S.
s (5. platan ifolius « dien. 5. nicarague nsis
subsp.
ellipsoidalis) three new combinations A ات stellatus, S. pats anifolius r texanus, A is nifolius
two ne
The genus Styrax comprises about 120 species
of trees and shrubs distributed in eastern and
southeastern Asia, the New World, and the Medi-
terranean region. It is a member of the family Styr-
acaceae (12 genera, about 150 species), which,
along with the Ebenaceae, Lissocarpaceae, Sapo-
taceae, and, according to some authors, the Sym-
rie comprise the order Ebenales (Cronquist,
1981; Dahlgren, 1980; Takhtajan, 1980, 1987;
. austr romexican
rescens var.
us, 5. lur latus). Los.
Thorne, 1992). A phylogenetic analysis of the
Ebenales based on chloroplast DNA rbcL sequenc-
es (Morton et al., 1997) tentatively indicates that
Styracaceae are diphyletic, with Styrax sister to
Clethra (of the monotypic Clethraceae). Styrax has
essentially a Laurasian Tertiary relict distribution
(as defined in Tiffney, 1985) except for a large neo-
tropical component and a single species that ex-
tends east of Wallace’s Line to New Guinea and
d imens available: A, AAU, B, BH, BR, CAS,
! I thank the curators of the following herbaria, who kindly made imi X
Paul S. Man s, Stephen A. Spongberg,
for justam. This re was supporte
ent of Botany, Duke University,
of Жыш California Albany of Sciences, Golde
HCIB, IBUG, INB, K, LL, ME
dead uiis K. Fritsch for help with the collection database, Porter P. Low
and Robert L. Wilbur for helpful comments o
d by a grant from the A. W.
Durham, North Carolina 27708-0339, U.
n Gate Park, San Francisco, California 94118-4599, U.S.
U, MICH, MO, MSC, NY, POM, эө ө,
п the manuscript, ca Lisa dn
Mellon ш ен and Duke Univ
S.A. Current ea pines
А.
ANN. MISSOURI Bor. Garp. 84: 705-761. 1997.
706
Annals of the
Missouri Botanical Garden
several islands of the South Pacific (Perkins, 1907;
van Steenis, 1932; Fritsch, 1995). The genus is
characterized by a campanulate, white or pink, usu-
ally 5-parted corolla with petals connate at the
base, usually twice the number of stamens as pet-
als, a 3-locular, superior ovary, a linear style, and
a 1-3-seeded drupe, nut, or 3-valved capsule with
a persistent calyx. Leaf arrangement is alternate,
and the vesture consists of scales or stellate hairs.
INFRAGENERIC CLASSIFICATION
Styrax is currently divided into section Foveo-
laria (Ruiz & Pav.) Perkins, comprising taxa with
3- to 5-ovulate gynoecia (two species, Cuba and
Peru), and section Styrax, with 16- to 24-ovulate
gynoecia (all the remaining species; Perkins, 1907).
Perkins (1907) maintained two series within section
Styrax: series Imbricatae Giirke (about 30 species)
and series Valvatae Giirke (about 90 species). As
the names suggest, the species of series Imbricatae
possess imbricate corolla aestivation, and those of
series Valvatae possess valvate corolla aestivation.
However, some species of section Valvatae are vari-
able for aestivation type—sometimes even within
the same individual (Perkins, 1907). For this rea-
son van Steenis (1932) suggested that the two series
are not “natural” (i.e., monophyletic) but offered no
alternative classification. A recent study of the in-
ternal transcribed spacer regions of nuclear ribo-
somal DNA (Fritsch, 1995, 1996a) has supported
the non-monophyly of the two series, but because
of limited sampling in this study, the development
of a revised classification at this time is premature
Ongoing morphological studies of Styrax
(Fritsch, unpublished data) suggest that characters
in addition to corolla aestivation are important in
the delimitation of infrageneric categories. The 60
or so neotropical evergreen Styrax species possess
a drupe, although the fruit is nearly always mistak-
enly described in the literature as dry, hard-walled,
and rugose because of the appearance of immature
fruits on herbarium specimens (the mature fruit is
rarely collected). The rest of the genus has dry
fruits: in the paleotropical evergreen taxa they are
thick-walled and either indehiscent or dehiscent
via three valves; in the deciduous taxa of the north-
ern subtropical and north-temperate zones they are
thin-walled and usually dehiscent via three valves
(rarely they are indehiscent). Corolla aestivation is
valvate in all neotropical and paleotropical ever-
green species, but valvate and/or imbricate in the
deciduous species (Fritsch, 1995). On the basis of
these characters, the delimitation of neotropical ev-
ergreen, paleotropical evergreen, and deciduous
groups within Styrax appears to be justified. Ad-
ditional phylogenetic work in the genus using both
morphological and molecular characters is needed
to lay the groundwork for a revised infrageneric
classification.
PRIOR WORK ом MESOAMERICAN STYRAX AND
PRESENT OBJECTIVES
The present taxonomic revision was undertaken
in conjunction with a treatment of Styracaceae in
the ongoing series Flora Mesoamericana. There are
two modern (20th century) treatments of Styrax in
Mesoamerica (the region east of the Isthmus of Te-
huantepec south through Panama) for use as ref-
erence works for the Flora Mesoamericana treat-
ment. The first (Perkins, 1907) provided a
treatment of the species of the region as part of a
worldwide monograph of the Styracaceae and is the
most recent worldwide treatment of Styrax to date.
Perkins (1907) recognized six species from the re-
gion: S. argenteus, S. conterminus, S. glabrescens,
S. polyanthus, S. polyneurus, and S. warscewiczit.
The second (Gonsoulin, 1974) comprised part of a
revision of North American, Central American, and
Caribbean Styrax. In this treatment, the number of
species in this region was reduced to three: S. ar-
genteus, S. conterminus, and S. glabrescens. As a
basis for this reduction, Gonsoulin (1974) argued
(with some justification) that Perkins was a notori-
ous splitter who disregarded continuous character
variation. Gonsoulin (1974) also placed several new
species that had been described subsequent to Per-
kins's (1907) monograph in synonymy under S. ar-
genteus. Gonsoulin's revision, having been based on
much more data than Perkins's, initially appeared
to provide an adequate context for undertaking the
Flora Mesoamericana treatment.
Bongo it became clear during early exami-
f Mesoamerican material that Gonsoulin
peo had made a number of problematic taxo-
nomic decisions, particularly concerning the cir-
cumscription of S. argenteus. This species was di-
vided into three loosely defined varieties that
encompassed all evergreen Styrax in Mexico and
Mesoamerica except S. conterminus, which pos-
sesses an unusual trichome type easily delimiting
it from the rest of the material examined by Gon-
soulin. He justified this circumscription by com-
paring it with the variable nature of the characters
used in Perkins's (1907) key to differentiate addi-
tional species, specifically pubescence on the lower
laminar surface, flower length, and calyx length. I
agree with Gonsoulin (1974) that these characters
by themselves provide poor support for species cir-
Volume 84, Number 4
Fritsch
Revision of Styrax
Figure 1.
cumscription in Styrax, and it is unfortunate that
Perkins provided only these characters in her key,
because she included a number of critical androe-
cial characters in her descriptions that at once dis-
tinguish S. argenteus from many of those taxa
placed in synonymy by Gonsoulin. Apparently Gon-
soulin surmised that, because she rarely included
them in her key, Perkins considered androecial
characters inconsequential. On this basis, he ap-
parently decided not to pursue the potential utility
of androecial characters for circumscription, for he
did not employ them in the key to species or de-
scribe them to any significant degree.
The questionable concepts of circumscription in
Gonsoulin (1974) were compounded by various no-
menclatural and bibliographic errors in the same
work, making the application of names to taxa I had
recognized in the Mesoamerican region uncertain.
Although Perkins’s (1907) treatment was careful, I
considered this work outdated and the species con-
cepts within it too narrow for reliable use. There-
fore, I decided to examine not only material from
Mesoamerica, but also all Styrax material from
western Texas and Mexico, as well as a limited
amount of South American material, to provide the
general context for a clear and unambiguous treat-
ment for Flora Mesoamericana.
GEOGRAPHIC DISTRIBUTION, ENDEMISM, AND
EcoLocy
Styrax exhibits a somewhat discontinuous distri-
bution in western Texas, Mexico, and Mesoamerica
(Fig. 1). The northern half of the range can be di-
vided into two parts. In the northeastern part of the
General distribution of Styrax in western Texas, Mexico, and Mesoamerica.
region members of the genus occur from the Ed-
wards Plateau and the Davis Mountains of Texas
south along the Sierra Madre Oriental to Oaxaca;
in the northwestern part they extend from the Cape
Region of Baja California Sur and southern Sinaloa
to Oaxaca along the western slope of the Sierra Ma-
dre Occidental, the southern slope of the trans-
Mexican volcanic belt, and the Sierra Madre del
Sur, bypassing the low elevations of the Rio Balsas
basin. Collections are also known from the Tuxtlas
Range in Veracruz. The southern half of the range
extends from the Sierra Madre and Central Plateau
of Chiapas through the mountainous regions of
Guatemala, northwestern El Salvador, and Hondu-
ras to a small portion of northern Nicaragua, and
then along the Pacific slope of the cordillera from
extreme southern Nicaragua to western Panama.
Isolated localities have been recorded in Belize,
central Nicaragua, and central and eastern Panama.
The 19 species of Styrax recognized in this treat-
ment are all endemic to the region of interest ex-
cept S. glabratus and S. peruvianus, both of which
also occur in South America. Only six species are
relatively widespread and fairly common: S. argen-
teus, S. glabratus, S. glabrescens, S. radians, S. ra-
mirezii, and S. warscewiczii. Nearly all the rest are
narrow endemics and are likely to be rare, vulner-
able, endangered, or extinct (terms sensu Lucas &
Synge, 1978). Two species are known with certainty
only from single collections (i.e., S. tuxtlensis, S.
panamensis). Styrax platanifolius has a relatively
large range but is uncommon; it comprises five sub-
species, all of which are rare. In contrast, S. mag-
nus occurs only from central Chiapas to southwest-
708
Annals of the
Missouri Botanical Garden
ern Guatemala yet has been collected many times;
thus it is probably common within its narrow range,
although it is unclear to what extent sampling ar-
tifact has affected the perceived abundance of this
and other species. Locating additional populations
of all narrow endemics, especially 8. gentryi, 8. in-
carnatus, S. panamensis, S. steyermarkii, and S. tux-
tlensis, will be invaluable for comprehensive docu-
mentation of morphological features, biogeographic
studies, and conservation.
Species of Styrax are found most often from 500
to 2500 m elevation. Several species (S. argenteus,
5. glabratus, 8. panamensis, 8. radians) occur ad-
ditionally or exclusively at elevations < 500 m;
others (S. austromexicanus, S. glabratus, S. rami-
rezii, S. warscewiczii) extend to 3000 m elevation or
more. According to herbarium specimen informa-
tion, the genus is most often associated with Quer-
cus and Pinus but can occur in a wide variety of
forested situations, such as tropical humid forests,
subtropical deciduous forests, seasonal evergreen
forests, temperate deciduous forests, coffee plan-
tations, and open savannas. Styrax platanifolius
subsp. youngiae grows in woodlands composed of
Acer, Arbutus, Cercis, Juniperus, and Quercus (Wendt
582), and also those composed of Pinus (pifion) and
Juniperus (Johnston et al. 1 1888). Many species
show a distinct preference for mesic microhabitats,
such as canyons, draws, and other riparian situa-
tions. However, individuals of various Styrax spe-
cles are also found in less protected situations,
such as pastures and wind-swept ledges. Within
primary forests, the tree species (i.e., all except S.
Jaliscanus and S. platanifolius) are usually under-
story or mid-canopy trees, seldom attaining a height
of > 20 m. They also occur in secondary forests
and other disturbed situations such as roadsides,
burned-over areas, plantations, and open fields.
Locating populations of Styrax species in the
field using herbarium label information can be dif-
ficult, primarily because they generally are local,
sporadic, and normally small (« 30 individuals).
urthermore, members of the genus lack easily ob-
servable diagnostic features that would otherwise
facilitate identification at a distance. The smooth or
cracked gray bark and alternate, simple leaves with
entire margins typical of the evergreen members of
the genus are similar to many associates, e.g., Oco-
tea of the Lauraceae.
The deciduous species of Styrax generally flower
in April and May and produce mature fruit by Sep-
tember; however, at low latitude they appear to pos-
sess a broader range of flowering and fruiting times.
In contrast, the evergreen species flower and fruit
at variable times throughout the year. The factors
controlling flowering times are unknown. The fruits
in the deciduous group (and likely in the evergreen
group) take several months to mature; in the ever-
green group, maturation from the green, relatively
hard phase of fruit development to the purplish
black, fleshy mature phase is not synchronous with-
in an infructescence, and change of single fruits to
the fleshy phase likely occurs quickly (within days).
Some neotropical species (S. argenteus, S. gla-
bratus, S. nicaraguensis, S. panamensis, and S. stey-
ermarkii in the present treatment) bear relatively
long and stiff, stellate hairs that form a dense mass
on the ventral side of the distinct portion of each
stamen filament (Figs. 11, 12). The arms of the
hairs are predominantly oriented along the longi-
tudinal axis of the flower. A high proportion of the
arms belonging to hairs nearest the distal end of
the filament point upward; these are usually the
longest in the hair mass (0.8-2 mm long). In the
same location on each filament, two longitudinally
oriented auricles can occur (S. argenteus, S. nicar-
aguensis, 5. peruvianus, and S. steyermarkii in the
present treatment; Figs. 12, 13). They are usually
obscured by the dense hair mass that surrounds
them; in S. peruvianus the trichome arms (in this
case, scales) are relatively short and the auricles
are larger and more prominent than those of the
other species in the treatment (Fig. 13). The dense
hairs and auricles occupy much of the space within
the stamen tube and partially cover the ovary. The
auriculate and densely pubescent conditions are
both probably derived within Styrax: the genus
Pamphilia Mart., likely the sister group to Styrax,
lacks auricles and pubescence. In the floral bud
stage, the auricles and/or pubescence occupy the
cavity between the basal end of the anther and the
apex of the ovary, and therefore may help to protect
the bud from small herbivores.
In all other species of Styrax treated here, the
ventral side of the distinct portion of the stamen
filament lacks both auricles and a mass of stiff-
armed stellate hairs, and the hairs nearest the distal
end of the filaments are not noticeably longer nor
do they have predominantly upward-pointing arms.
The pubescence that is present is instead short-
armed (to + 0.7 mm long) and situated mainly
along the filament margins (Fig. 10). The face of
the filament proper (as opposed to the margin) is
often glabrous proximally, becoming sparsely to
densely pubescent distally.
Little comparative work has been done on the
reproductive biology of Styrax species, especially
those within the region of study. The most detailed
analysis of breeding systems and pollination biol-
ogy in the genus has been conducted by Sugden
a
-——.Üao AA AA
ПР RRA dale was _ йй “ч
«шеле ч pies P... adii
Volume 84, Number 4
1997
Fritsch 711
Revision of Styrax
“thieving,” because the flower was not pollinated
by the insect during this process even though it
relinquished its reward.
It is not known whether the pollination syndrome
documented in S. redivivus is typical of other Styrax
species. Because evolution of floral morphology in
Styrax been conservative, the general syndrome
of bee and butterfly pollination combined with xenog-
amy might be expected to apply across much of the
genus. The flower-visiting fauna of S. redivivus resem-
bles that of S. americanus Lam. and S. grandifolius
Aiton, both from the southeastern United States, and
sucrose-dominant nectar is also produced by S. ar-
genteus (Sugden, 1986). "Morphological gynodioecy
has been documented for the genus Pamphilia (Miers,
1859) and has recently been observed (pers. obs.) in
the two species of Styrax comprising section Foveo-
laria (S. foveolaria Perkins and S. obtusifolius Gri-
seb.). These two Styrax species might be expected to
exhibit a pollination syndrome divergent from that of
most or all other species in the genus. Additional
study of the pollination and breeding systems of a
wide variety of Styrax species (including those of sect.
Foveolaria) and Pamphilia would provide further in-
sight into the evolution of reproductive systems in the
genus.
MATERIALS AND METHODS
Over 2500 herbarium specimens from 29 herbaria
were examined for this study. South American mate-
rial from BR, BM, CAS, F, GH, MICH, MO, NY, P,
US, and W, and Asian material from GH were also
examined to aid in understanding the taxonomic sig-
nificance of character variation in the Mesoamerican
members of the genus and to determine whether the
ognized here occur also in other regions. One
week of fieldwork in Texas (S. platanifolius subspp.
stellatus and texanus) and two weeks in southwestern
Mexico (S. jaliscanus and S. radians) supplemented
observations on the herbarium specimens.
All descriptions were derived from examination of
herbarium specimens except where noted. Flowering
and fruiting times, elevation ranges, habitats, distri-
butions, common names, and uses were derived from
label information of herbarium specimens. Descri
tions of leaves refer to those of the fertile branches;
leaves of sterile branches are consistently larger and
often possess more variation in hair quantity and
quality than those of fertile branches. Leaf and petiole
measurements were taken from the larger examples
on each herbarium sheet. All floral descriptions refer
to the anthesis stage only, except where noted. Calyx
dimensions are presented in terms of height (from the
end of the pedicel to the apical margin) Х width at
the apex, and thus include the short hypanthium.
Measurements of corolla length do not include the
hypanthium. In the descriptions, the phrase “free por-
tion of the stamen tube” refers to the section of the
androecium from where it becomes free from the co-
rolla to where the filaments become distinct (i.e., free
from each other). Fruit length was measured from the
base of the fruiting calyx to the tip of the fruit (the
calyx is persistent). Fruit measurements of the drupe-
bearing species were taken from the larger examples
on each fruiting branch of fully expanded but unripe
fruits (fully mature drupes are rarely present on her-
barium specimens); those of the dry-fruited species
were taken from the larger examples of mature fruits.
Most observations were made by eye or with the aid
of a dissecting microscope (maximum magnification
= 60X). Examination of trichome morphology was
“ринен мић ЗЕМ.
Seven infraspecific taxa аге tegel in the
treatment. The category “subspecies” has
been used in all cases in accordance with the con-
cepts of Hultén (1967) and Thorne (1978). These
authors use the subspecies category for infraspe-
cific taxa that are geographically as well as mor-
phologically distinct.
TAXONOMIC TREATMENT
The following taxonomic treatment of Styrax from
southern North America differs substantially from
that of Gonsoulin (1974). A comparison using only
Gonsoulin’s annotated material shows that nearly
three times as many species are recognized in the
present revision as the last (Appendix 1). The num-
ber of deciduous species is lower (three vs. five)
and the number of evergreen species is much high-
er (ten vs. two). Specimens representing six addi-
tional species were collected or accessioned sub-
sequent to publication of the last revision
(Appendix 1). The total number of species and taxa
circumscribed in the present revision is 19 and 24,
respectively, versus 7 and 11 in the last revision
(Appendix 1). Therefore, the genus is much more
diverse in Mexico and Mesoamerica than previous-
ly thought. Six new species and two new subspecies
are described. In addition, three new combinations
and two new species names result from this study.
Styrax L., Sp. pl. 444. 1753. TYPE: Styrax offi-
cinalis L. [*officinale"].
Strigilia Cav., Diss. 7: 358, t. 201. 1789. Tremanthus
rs., Syn. pl. 1: 467. 1805. TYPE: Strigilia race-
mosa Cav
Cyrta Lour., Fl. cochinch. Ed. 1: 278. 1790. TYPE: Cyrta
agrestis Lour.
712
Annals of the
Missouri Botanical Garden
Корени Ruiz & Pav., Fl. региу. ргодг. 57, 1. 9. 1794.
E: [not designat ed].
pees ae Fl. ludov. 56. 1817. TYPE: Adnaria odorata
R
Epigenia Vell., Fl. flumin. 183. 1829. TYPE: [not desig-
ated].
Ке Hayne, Getreue Darstell. Сем. 11, t. 24. 1830,
поп Boerh. ex Schaeff., 1760. TYPE: Benzoin offi-
nalis Hayne [= Styrax benzoin e |
Г көне Torr., Proc. Amer. Assoc. Adva
191. 1851, nom. rej. TYPE: баа redicina
Torr.
Evergreen or deciduous trees or shrubs; bark
smooth or cracked, gray; outer layer of older twigs
fibrous, dull brown or more often gray, inner layer
dull maroon; buds superposed, lepidote or stellate-
pubescent, naked. Vesture consisting of scales,
stalked or unstalked stellate hairs, and often sessile
or stalked glandular hairs; evergreen species usu-
ally with inconspicuous scattered stalked glandular
hairs on at least some of the vegetative parts: often
adaxially on petiole and base of laminar midrib,
abaxially on laminar surface, and on inflorescence
bracteoles. Leaves alternate, simple, pinnately
nerved, estipulate. Inflorescences bracteolate, ax-
illary or false-terminal racemes or panicles, some-
times corymbose in appearance, sometimes two or
more arising from the same node (or a single inflo-
rescence branched at the base), sometimes 1-2-
flowered, then usually axillary. Flowers actinomor-
phic, bisexual or rarely gynodioecious, fragrant,
with a short hypanthium adnate to the basal third
or less of the ovary wall; calyx gamosepalous be-
yond the hypanthium, campanulate or cupuliform,
teeth (2-)4-6 or absent; corolla gamopetalous be-
yond the hypanthium proximally, the petals distinct
distally, campanulate, the lobes 5(-10) and almost
always longer than the tube, valvate or imbricate in
bud, white, pink, or rarely yellow; stamens adnate
to the corolla tube beyond the hypanthium proxi-
mally, free distally, 10 when the number of corolla
lobes = 5, up to twice the number of the corolla
lobes when the number of lobes > 5, uniseriate but
appearing biseriate in bud (with five stamens in
each series), the inner series often exceeding the
outer at the distal end of the androecium, free por-
tion of the filaments often connate proximally and
distinct distally, sometimes completely distinct, the
ventral side of the distinct portion glabrous to stel-
late-pubescent or lepidote and sometimes with two
juxtaposed and longitudinally oriented auricles,
these often obscured by a dense mass of hairs,
many of which have arms predominantly oriented
along the longitudinal axis of the flower, the hairs
nearest the distal end of the filaments with arms
predominantly pointing upward; anthers introrse, 2-
locular, basifixed, longitudinally dehiscent, the an-
ther sacs glabrous to moderately stellate-pubescent;
pollen light or golden yellow; ovary superior,
3-locular at base, 1-locular toward apex; style fili-
form, slightly exceeding the stamens; stigma ter-
minal, obscure, capitate or + 3-lobed; ovules (3-
5)16–24, axile or basal, anatropous. Fruit a drupe,
a capsule dehiscent via 3 valves, or nut-like (dry
and indehiscent), globose or ellipsoid, 1(-3)-seed-
ed, with calyx persistent. Seed globose or ellipsoid,
brown, completely filling the fruit cavity, usually
smooth except for 3(-6) longitudinal grooves, with
a broad hilum; endosperm copious; embryo straight;
cotyledons flattened. About 120 species. U.S.A. to
Argentina, E Mediterranean, E and SE Asia.
Note. The generic name Styrax has been treated
by various authors as masculine, feminine, or neu-
ter. Nicolson and Steyskal (1976), after conducting
a thorough evaluation of the issue, advocated the
use of masculine as the gender that best fits the
wording of the botanical rules of nomenclature.
Thus, Styrax is treated as masculine here.
KEY TO SPECIES OF STYRAX IN TEXAS, MEXICO, AND MESOAMERICA
la. Corolla lobes imbricate in bud, — velan at anthesis; fruit dry, indehiscent or dehiscent via three
valves; PME deciduous (Styrax sect. Styrax mbricatae).
2a о 30 m tall, small pri not pua es stemmed from the base; lamina membranous; fruit nearly
д. ae rarely dehiscent via three valves, the larger fruits 10-17 mm lon 6. S. glabrescens
2b. Shrubs to 3 tall, multi- TET from the base; lamina chartaceous; fruit dehiscent by boe valves,
the larger fruits 7-11 mm
За. Longer petioles 4—5 mm ا leaf bases subnet or Founded, rarely | subcordate but then not
attenuate; secondary and tertiary laminar ve wall coarsely
and weakly Cone e striate, not ot transversely waked post-dehiscence M 8. 5. jaliscanus
ЗЬ. Longer petioles 6-20 mm long; зо bases on each plant cordate or truncate and slightly
attenuate; secondary an t ab tertiary lam ini inar veins evident but not ч abaxially; undehisced _
i fruit wall smooth, usually transversely wrinkled post-dehiscence ____-__------- 14. 5. nifolius
ser. Valvat
ruit
Corolla lobes valvate in bud, non- overlapping at anthesis; fruit a drupe; а evergreen (Styrax sect. Styrax
ae).
4a. Abaxial laminar surface (and calyx) with vesture of peltate or lacerate-margined scales; scales at mid-
m di
calyx 0.27–0.44 mm diam.
9a. Ventral side of the distinct portion of the stamen filaments not auriculate; leaves 2.8-3.9 times
— ——
m ea
ud TTR. —— 3 ~ ·
AAA AAA AAA SI, A A AS - A
т ee
Volume 84, Number 4 Fritsch 713
1997
Revision of Styrax
4b.
c
as long as wide; connectives not or only slightly prolonged beyond the non- — anther sacs
3. S. conterminus
5b. Ventral side of the distinct portion of the stamen filaments strongly auriculate; bein 2.0-3.1
times as long as wide; connectives distinctly prolonged beyond the tapered anther sacs ............
13. S. peruvianus
Abaxial — surface We a vesture of stellate MÀ or rarely radiate scales or nearly glabrous; scales
at mid-calyx 0. m diam., or else lackin
6a. pry sie disi obliqui 18. S. tuxtlensis
6b. Leaf bases symmetric or aii slightly oblique.
Та. Arms of the hairs nearest the distal end of the ventral side of the distinct portion of the
stamen filaments predominantly pointing upward, some of these up to (0.8-)1-2 mm long;
connectives distinctly prolonged beyond the tapered anther sacs
. Calyx vesture uniformly golden brown; lamina 1.5 times as long as wide .................
8b. pa vesture predominantly or completely grayish green, occasionally with rng
cattered yellow or orange-brown trichomes; lamina 2.1-3.8(-5.2) times as long as
vile
9a. Calyx vesture of stellate hairs.
10a. Larger upward-pointing hairs on "€ ventral side of the distinct portion of
stamen filaments with at least some arms 1-2 mm long; abaxial laminar
surface and outer calyx grayish P кик without scattered
orange-brown stellate hairs; larger drupes 13-18 mm long (excluding beak,
if present), ellipsoid or ovoid-ellipsoid: corolla lobes 1.7-2.3 mm wide,
widely spreading or recurv l. 5. argenteus
10b. Larger upward-pointing hairs on the ventral side of the distinct portion of
the stamen filaments with arms to 1 mm long; abaxial laminar surface and
outer calyx with scattered, orange-brown, stellate hairs in addition to gray-
green pubescence; larger drupes (15-)17-28 mm long, ovoid or obo-
huido corolla lobes 2.5—4 mm wide, recurved or reflexed 00000000000...
11. 5. nicaraguensis
b. Calyx vesture of radiate scales.
Па. Abaxial laminar surface glabrous or nearly so; calyx 4-7 mm long, in fruit
funnelform, 4-8 mm long; anthers 5-8 mm long; corolla lobes qon
ratus
llb. Abaxial laminar surface densely pubescent; calyx 2.5-3 mm long, i in o Бин
shallowly cupuliform, to 1.5 mm long; anthers 4.5 mm long; corolla lo
linear 17. S. steyermarkii
Tb. Arms of the hairs nearest the distal end of the ventral side of the distinct portion of the
tamen filaments not predominantly pointing upward, typically not exceeding 0.5(-0.7) mm
long; connectives not or only slight - or: Бона the non-tapered anther sacs.
12a. Calyx teeth 1-2 mm long, con 4. S. gentryi
=
@
[71
FS
EE
3
E
3
qu
3
$
©
H
а 3.0-4.8 mm long, 1.9-5.3 times as long as the distinct pim of the
filaments S. ramirezii
14b. Чыг on the outer distal third of the corolla lobes of ам scales
anthers 1.7-3.6 mm long, 1.1-2.2 times as long as the distinct portion of
he fi 19. S. warscewiczii
13b. Vesture just below mid-calyx lomi of stellate hairs, sometimes also wit
a few scale-like hairs scattered among n stellate ones.
15a. Abaxial surface of t nd tertiary veins obscured by the to-
mentum; * evenly асай, orange-brown hairs (in addition to whitis h
pubescence) present on the abaxial surface of most or all fully expanded
ves.
16a. Laminae chartaceous, lanceolate, the larger 7.7-9.7 ст long; fruit 9-
11 mm long; corolla 11-15-17) mm long, white ......... 9. S. lanceolatus
. Laminae thick-c 'еоив, clliptie, slightly oblanceolate, or ces
the larger 11-18 ст long; fruit 17-19 mm long (unknown in <
carnatus); corolla (10—)14—24 mm long, usually pink, sometimes
white.
17a. Larger laminae 3.5-5 cm wide, usually narrowly elliptic, some-
times slightly — — ovate, or a T
icels 11-15 mm long; calyx x 5-6 mm
1
=
СЧА АРИОН ustromexicanus
17b. Larger laminae 5.5-9.5 cm wide, ovate or E elliptic. lower
pedicels 3-9 mm long; calyx 3-3.5 X 2.54.5 mm .............
714
Annals of the
Missouri Botanical Garden
do sax O Ln al qeu Ju magnus
19a. Distinct portion of stamen filaments 2.5-3.5 mm long; abaxial
laminar surface usually soft to the touch; calyx not stellate-
villous; corolla white, thin, the petals connate up to the calyx
margin, the lobes moderately or widely spreading; stigma 0.6—
i 15
abaxial laminar surface slightly or moderately rough or rarely
soft to the touch, if soft then at least the lower third of the calyx
villous; corolla white or pink, thickened, the petals connate 1—
0.8 mm wide ... : . 15. S. radians
- Distinct portion of stamen filaments 0.6-2.0(-2.6) mm long;
mm
y
spreading; stigma 0.3-0.5 mm wide _________
1. Styrax argenteus C. Presl, Reliq. haenk. 2:
60. 1831-1835. Strigilia argentea (C. Presl)
Miers, Ann. Mag. Nat. Hist., ser. 3, 3: 282.
1859. TYPE: Mexico. Guerrero: “Ad portum
et urbem Acapulco" (protologue), T. Haenke
s.n. (or 148; Perkins, 1907) (holotype, PR not
seen; isotypes, F, MO, W, photo of W in A, F,
MICH). Figures 2, 12, 14-16.
Styrax myristicifolius Perkins, Bot. Jahrb. Syst. 31: 481.
1902. TYPE: Guatemala. Santa Rosa: Las Vi as,
Sep. 1894, Heyde & Lux 6182 (lectotype, designated
by Standley & Williams (1967), F; isolectotypes,
GH, MO, NY, US).
Styrax polyanthus Perkins, Bot. Jahrb. Syst. 31: 479,
902. TYPE: Guatemala. Guatemala: prope Guate-
mala, February 1890, J. Donnell Smith 2266 (lec-
totype, designated by D’Arcy (1979), US; isolecto-
types, F, GH).
Evergreen tree to 20(-30) m. Young twigs grayish
yellow-, yellowish brown-, light brown-, or orange-
brown-stellate-pubescent; outer layer of older twigs
not exfoliating. Petiole 10-25 mm long. Lamina
chartaceous, 6.5-18 x 2.4-7.5 cm, 2.1–3.7 times
long as wide, ovate, lanceolate, narrowly or
broadly elliptic, or rarely slightly oblanceolate, sec-
ondary veins 9-11; apex acute to acuminate; base
cuneate to rounded, often slightly attenuate, sym-
metric or slightly oblique; adaxially glabrous except
along the major veins; abaxially covered with min-
ute grayish green stellate hairs and often also larg-
er, soft, grayish green or yellowish brown stellate
hairs, the minute hairs usually absent from the pri-
mary through tertiary veins, thus the vein surface
easily visible; margin entire. Inflorescences axill
or false-terminal, racemose or rarely paniculate, the
racemes or panicle branches 3-14 cm long, 3-22-
flowered. Lower pedicels 2-1 1(-15) mm long. Са-
lyx 2.5-5.0 x (3.524—6(-7) mm, cupuliform or
d the calyx margin distally, the lobes slightly
1 16.
S. ramirezii
broadly cupuliform, grayish green-stellate-pubes-
cent; calyx teeth 5, to 0.5 mm long, deltoid, calyx
margin truncate or rarely slightly concave between
the teeth, inner margin often glandular. Corolla
white, 10-16 mm long, petals usually connate up
to the calyx margin, rarely to 2 mm distally beyond;
corolla lobes 5, valvate in bud, 8-14 x 1.7-2.3
mm, non-overlapping, widely spreading or re-
curved, thin, linear-deltoid, vesture on the outer
distal third of the lobes consisting of stellate hairs.
Free portion of stamen tube 0.7-1 mm long; dis-
tinct portion of filament 2-2.5 mm long, of equal
width throughout, ventrally with small auricles
bearing a dense mass of white stellate, stiff hairs
with at least some arms 1-2 mm long, the hairs
nearest the distal end of the filament with arms
predominantly pointing upward, also pubescent be-
low the auricles and at the sinus between adjacent
filaments; anthers (5-)6-8 mm long, the connec-
tives distinctly prolonged beyond the tapered an-
ther sacs. Ovary densely stellate-hirsute; style gla-
brous, or pubescent at base only; stigma 0.3-0.5
mm wide. Drupe 13-18 X 8-14 mm (wider when
2-seeded; does not include occasional beak), ellip-
soid or ovoid-ellipsoid; wall irregularly and coarse-
ly rugose on herbarium specimens. Seed coat gla-
brous
Near Acapulco, Guerrero, Mexico, to central
Panama; most common on the Pacific slope of the
cordillera of Central America; one collection from
Cayo District, Belize; 100-1700 m; tropical humid
forests, seasonal evergreen forests with Pinus and
Quercus, tropical deciduous forests, pastures, open
scrub, coffee plantations; found often along water-
ways with other riparian vegetation, and in moist
ravines, but also on dry limestone ridges, at forest
edges, disturbed slopes, and with various types of
Volume 84, Number 4
1997
Fritsch 715
Revision of Styrax
| Bahia de Campeche
Pa | аҹ бб.
Caribbean
|
|
|
|
|
|
Figure 14. Distribution of Styrax argenteus.
ae old growth. Flowering: August-March; fruit-
tober—May.
names. Alamo (Honduras, Erazo 63,
Pereg 1 15947. von Hagen & von Hagen 1253;
Nicaragua, “7-1” s.n.), alomo (Honduras, Molina &
Molina 34417), chucam-ai (Chiapas, 1919, Becerra
s.n.), duraznillo (Guatemala, Steyermark 51100),
estorac (El Salvador, Standley 20147), estoraco
(Costa Rica), estoraque (El Salvador, Calderón
1404 and 1405, Standley 20405; Guatemala, Mo-
rales 1208, Steyermark 50801), estoraque común
(Guatemala, Aguilar 186 and 214), naranjo (Gua-
temala, Standley 82830), palo blanco (Honduras,
Erazo 63, Molina 20321), roble (Guatemala, Stey-
ermark 30624), tepe aguacate (Guatemala, Steyer-
mark 50699). Standley (1924) has listed additional
common names for this species: bracino (Costa
Rica), capulín (Oaxaca), chilacuate (Guerrero, Mi-
choacán), estorac (El Salvador), hoja de jabón (Oa-
xaca), resina, resino (Costa Rica, Nicaragua), ruín
(Sinaloa), and sahumerio (Panama), but because his
circumscription of S. argenteus differs from mine,
the = of these names in this revision is
. In Standley (1938) the names listed
i B lire: resina, tubtis; Costa Rica) are more
likely to apply to S. argenteus as conceptualized in
Standley (1924).
Uses. Fruits are sold in local markets (Chiapas,
Bauml et al. 531); flowers are boiled with lime
leaves and the water used as a perfume (Oaxaca,
Dec. 1961, MacDougall s.n.); bark is used to poison
fish (El Salvador, Standley 20405). Standley (1924)
reported that the gum of S. argenteus is burned as
incense in Costa Rican churches, but it is not clear
whether the gum used is from S. argenteus as cir-
cumscribed in the present revision.
Selected specimens examined. BELIZE. Cayo: Río
Frío Caves road, Augustine, Meave & Howe 1162 (MO).
COSTA RICA. Alajuela: Santiago de San Ramón-Finca
Barranca, Gómez et al. 20777 (CAS, CR, DUKE, MO).
Cartago: Turrialba, Córdoba 754 (CR, DUKE). Guana-
caste: Parque Rincón de la Vieja, ha Santa Marfa,
Herrera 854 Ke F, MO, TEX). He 1
li GH). Puntarenas: Monte
W). San José: m N
17390 ) (F. MO, US. EL SALVADOR. Ahuachapán: Si-
erra de Apaneca, Standley 20147 (GH, NY, US). San Sal-
vador: Comasagua, Calderón 1404 (GH, NY, US). a7:
Ana: vicinity of UE Ana, Standley 20405 (GH,
US). GUATEMALA. Baja Verapaz: 10 mi. S of “КТАН,
Harmon & Dwyer 3056 (GH, МО, NY). Chimaltenango:
along road from Chimaltenango to San Martin Jilotepeque,
Standley 57958 (F, NY). Chiquimula: along Tepotun Riv-
er, vicinity of Esquipulas, Molina & Molina 25328 (F,
„ЮР : Sierra de las Minas, | Jan. 1908,
Kellerman s.n. (F). Eseuintla: below Las Lajas, Standley
64767 (A, F). Guatemala: Guatemala City, Harmon 3461
(DS, MICH, MO, N Huehuete : Aguacatán,
Skutch 1935 (A, F, yj dace El Aguilar, just E of San
Sebastián, Standley 82830 (F, MO, em Jutiapa: Que-
brada above E Sandi; 77656 ;
lar 758 (F). Santa Rosa: Уојсап ње N of Chi-
quimulilla, Seeders 33138 (A, F). Sololá: Finca Belén,
716 Annals of the
Missouri Botanical Garden
gures 15-24. 15, 16. Styrax argenteus. не Four adjacent stamens, Palacios 34 (CAS). —16. Fruit, Skutch
1 935 a 17-20. Styrax nicaraguensis s ubsp. nicaraguensis. —17. Flowering branch. —18. Flower. —19. Four adjacent
stamens. 17-19, Webster et al. 12496 (LL). —20. Fruit, Atwood s.n. 8 June 1975 (MO). —21. Styrax nicaraguensis
subsp. ellipsoidalis, fruit; Dryer 1420 (F). 22-24. St tyrax steyermarkii. —22. Flower. —23. Four adjacent stamens,
Lundell & Contreras 19214 (LL). —24. Fruit. 22, 24. Lundell & Contreras 21219 (LL).
Volume 84, Number 4
1997
Fritsch 717
Revision of Styrax
Lake Amatitlán, White 5260 (F, LL, MICH, US). 5
tepéquez: southern lower slopes of Volcán Zunil, т
ermark 35418 (F). Zacapa: between La Reforma and Те-
culután, Steyermark 42075 (F, NY). HONDURAS.
Choluteca: vicinity о soe Marcos de Colón, Standley
Y). El Pardleo near
5 Danli, Williams & Molina 10491 (DS,
F, GH, LL, MICH, " Morazan: near Cuesta Grande,
Williams & Molina 13270 (BH, F, GH, LL, MEXU, MO,
egucigalpa: Mont. de la Flor, von Hagen & von
Hagen 1253 Œ NY. MEXICO. Chiapas: Intersection of
H 90 and road to Tuxtla Gutiérrez airport, Bauml et
al. EF С, US); 13 km М of Arriaga along Hwy. 195,
Breedlove 28487 (DS, DUKE, F, LL, MEXU, MICH, MO,
NY, RSA); El Zapotal, al SE de Tuxtla Gutiérrez, Palacios
34 (CAS). Guerrero: La Providencia, Mar. 1926, Ferris
s.n. (DS). Oaxaca: between Trapiche Santa Ana and T.
del Lazo, Conzatti 4558 (NY, US); San Bartolo Yautepec,
Dec. 1961, MagDougall s.n. (MEXU). NICARAGUA.
Boaco: Cerro Mombachito, Aranda et al. 40 (MO, NY).
Chontales: ca. 2.8 km above (N of) Cuapa, Stevens 22709
(MO). Estelí: Paso de León, 3.5 km al NW de Estelí,
Moreno 10854 (DUKE, MEXU, MO, NY). Madriz: Cerro
El Fraile, Moreno 23512 (MEXU, MO). PANAMA. Coclé:
vicinity of Olá, Pittier 5076 (GH, NY, US).
Styrax argenteus is one of the most common spe-
cies of Styrax in the area circumscribed by this
revision. It is distinguished from sympatric species
by its evergreen leaves, grayish green stellate pu-
up to the calyx margin, and slightly auriculate sta-
men filaments bearing long-armed and relatively
stiff stellate hairs ventrally; the hairs nearest the
distal end of the filament possess the longest arms,
which predominantly point upward. It is a relatively
low-elevation species, unlike most other Styrax
species in the region. Generally this species does
not extend lower than 400 m, but it does occur at
or below 100 m in Guanacaste Province, Costa Rica
(OTS Research Area A, Frankie 239a) and Coclé
Province, Panama (Pittier 5076).
The differences between S. argenteus and S. ni-
caraguensis are subtle. Styrax nicaraguensis has or-
ange-brown hairs on the calyx and the lower lam-
inar surface, a larger drupe, a highly restricted
range, and a different habitat. Sterile specimens of
S. argenteus are difficult to distinguish from the
equally common S. warscewiczii: S. argenteus tends
to have larger leaves that are fulvescent when
young, and often has longer, relatively loose and
soft hairs abaxially. However, some sterile speci-
mens cannot be determined, particularly those from
saplings or stump sprouts, or from undocumented
elevations and/or habitats.
My treatment of S. argenteus differs from those
of Perkins (1907) and Gonsoulin (1974). I agree
with Gonsoulin (1974) that S. argenteus was un-
necessarily divided into several species by Perkins
. Styrax myristicifolius and S. polyanthus
both fall within the range of variation encompassed
by my concept of S. argenteus [Perkins later (1907)
placed S. myristicifolius in synonymy under $. ar-
genteus]. However, I strongly disagree with Gon-
soulin’s (1974) revised concept of S. argenteus. This
concept includes not only the clearly distinct spe-
cies $. magnus, S. ramirezii, and S. warscewiczii,
all of which were described prior to Gonsoulin's
revision, but also the species now recognized as S.
austromexicanus, S. radians, and S. steyermarkii.
Gonsoulin annotated specimens of the latter three
species as S. argenteus despite examining material
with diagnostic characters. He also annotated the
type of S. panamensis as S. argenteus but failed to
include this name in the treatment.
To account for the high amount of variation pres-
ent in his circumscription, Gonsoulin (1974) rec-
ognized two varieties of Styrax argenteus in addition
to the typical variety: variety hintonii and variety
ramirezii (— var. micranthus). The three varieties
were distinguished by the type and amount of pu-
bescence on the lower laminar surface and the ca-
lyx, but essentially exhibited no ecological or geo-
graphic distinctions. Although calyx hair types are
critical characters in the delimitation of Styrax spe-
cies, the hairs on the lower laminar surface tend to
be more variable and unreliable. When Gonsoulin
attempted to reconcile the relatively constant na-
ture of the calyx pubescence with the variable na-
ture of the leaf pubescence, the result was the rec-
ognition of a large number of intermediates
throughout the ranges of the varieties. For example,
S. argenteus var. ramirezii is distinguished from va-
riety argenteus by a thin versus thick tomentum on
the lower laminar surface. However, S. argenteus in
my view is polymorphic for this character in that
individuals always possess the thin tomentum, but
in addition often have an additional layer of longer
hairs. The majority of specimens examined by Gon-
soulin that are assigned here to S. argenteus were
annotated by him as variety argenteus, primarily
because S. argenteus usually possesses the longer
hairs. However, a considerable portion were anno-
tated as variety ramirezii (those with the long pu-
bescence absent), or intermediates between this
and the typical variety. When the hairs are unusu-
ally long, specimens of S. argenteus were annotated
by Gonsoulin as variety hintonii or as a variety hin-
tonii intermediate (it is never stated what the sec-
ond taxon is in cases of purported intermediacy).
The high number of intermediates assigned to va-
riety argenteus and variety ramirezii by Gonsoulin
~
oud
718
Annals of the
Missouri Botanical Garden
Figure 25. Distribution of Styrax austromexicanus, S. radians and S. tuxtlensis.
is accounted for to a large extent by the variable
amount of the longer hairs present in individuals
of S. argenteus. In the present treatment S. argen-
teus vars. ramirezii and hintonii are synonyms of S.
ramirezii.
The collection date of the type of S. argenteus is
probably October-December 1791 (McKelvey,
1955).
"
Styrax austromexicanus P. W. Fritsch, nom.
et stat. nov. Based on: Styrax argenteus var.
grandiflorus E. Carranza, Acta Bot. Mex. 36:
15. 1996, non S. grandiflorus Griff., 1854.
TYPE: Mexico. Oaxaca: Mpio. de Juxtlahuaca,
3 km S of the road to San Martín Peras on the
road to Coicoyán de Las Flores, 27 Sep. 1993,
J. L. Panero et al. 3475 (holotype, MEXU not
seen; isotypes, ENCB not seen, IEB not seen,
MSC). Figures 25-27.
Evergreen tree to 17 m. Young twigs predomi-
nantly and densely orange-brown-stellate-pubes-
cent or -villous, with intermixed scattered short-
armed white stellate hairs; outer layer of older twigs
not exfoliating. Petiole 13-21 mm long. Lamina
thick-chartaceous, 11-18 X 3.5-5 cm, usually nar-
rowly elliptic, sometimes slightly oblanceolate,
slightly ovate, or elliptic, secondary veins about 13;
apex acute to abruptly acuminate; base cuneate to
subrounded, symmetric or slightly oblique; adaxi-
ally glabrous except along the major veins or rarely
sparsely stellate-pubescent; abaxially covered with
minute whitish stellate hairs, larger whitish hairs
and evenly scattered orange-brown hairs, the three
types occurring on the secondary and tertiary veins,
thus obscuring the vein surface; margin entire. In-
florescences axillary or false-terminal, racemose or
paniculate, racemes or panicle branches 3.5-6.5
cm long, 2-11-flowered. Lower pedicels 11-15 mm
long. Calyx 4-5 X 5-6 mm, conical or broadly cu-
puliform, light grayish green-stellate-pubescent,
with a variable amount of larger orange-brown stel-
late hairs; calyx teeth 5, to 0.5 mm long, deltoid or
more often indistinct from the calyx margin, rarely
absent, calyx margin truncate or more often slightly
concave between the teeth, inner margin rarely
glandular. Corolla pink or white, 14-23 mm long,
petals connate 2-5 mm beyond the calyx margin
distally; corolla lobes 5, valvate in bud, 8-17 X
2.5-3.5 mm, non-overlapping, erect or spreading,
thickened, linear-deltoid, vesture on the outer dis-
tal third of the lobes rE of stellate hairs.
Free portion of stamen tube 3.5-6 mm long; dis-
tinct portion of filament 1. 5-2 mm long, of equal
throughout, ventrally not auriculate, sparsely
tdeo mar йыш at the somewhat incurved
filament margins, glabrous or nearly so on the face
proximally, increasing in pubescence distally, the
hairs with arms to 0.5 mm long; anthers 4.5-5.2
Volume 84, Number 4 Fritsch 719
Revision of Styrax
ESTE
ES
=v =. E E PH
"А5
Se а
т -
P ин
res 26-31. 26, 27. Styrax austromexicanus. — 26. Pedicel and flower, Hinton et al. 14750 (Е) —27. Fruit,
L).
Fi
Reedowski 18529 (TEX). 28-30. Styrax incarnatus, Lundell & Contreras 19618 (LL). —28. Flowering branch. —29.
Pedicel and flower. —30. Four adjacent stamens. —31. Styrax lanceolatus, pedicel and flower; Johnston 7408 (TEX).
720
Annals of the
Missouri Botanical Garden
mm long, the connectives not or only slightly pro-
longed beyond the non-tapered anther sacs. Ovary
densely stellate-hirsute; style pubescent from base
to 45-90% of the total length; stigma 0.3-0.5 mm
wide. Drupe 17-19 X 10-12 mm, obovoid, slightly
beaked by the base of the persistent style; wall ir-
regularly and coarsely rugose on herbarium speci-
mens. Seed coat sparsely white-stellate-pubescent.
Guerrero and western Oaxaca, Mexico, in the Si-
erra Madre del Sur; bosque mesofilo de montaña,
Quercus-Pinus-Abies forest; 2100—3000 m; found in
rocky limestone areas. Flowering: June, September,
October; eg February, April, June, September,
November.
це specimens examined.
MEXICO.
16 km SW of Filo de Caballo, Breedlove & Alme
64929 (CAS); W of Puerto El Gallo alo ong road to Toro
о
Е
al, 2642 (МО); Mpio. де Chilpancingo, entre los
la Aguililla y Plan de Potrerillos, Valdez 616 (MEXU).
ахаса: E de Xatu Yahta, а! W Coicoyán, Avila 742
(CAS); km 18 on the road to Саћада de Lobos, Gorge of
the Río Ratón, Panero 5254 (MSC).
Styrax austromexicanus is a rare endemic to the
cloud forests of the Sierra Madre del Sur in Guer-
rero and Oaxaca, Mexico. Except for S. incarnatus,
it has the largest flowers of any evergreen Styrax
species in the region. It can be distinguished from
sympatric species by its stellate pubescence
throughout, narrowly elliptic evergreen leaves, mix-
ture of white and orange-brown hairs on the lower
laminar surface and calyx, large, usually pink flow-
ers with a long corolla tube, and stamen filaments
that lack both auricles and stiff, long-armed, up-
ward-pointing hairs ventrally. In addition, the sec-
ondary and tertiary veins of the lower laminar sur-
face are obscured by the tomentum, distinguishing
this species from most specimens of S. ramirezii.
This species appears to be allied to S. incarnatus
of Guatemala and Honduras, from which it differs
principally by its narrowly elliptic leaves.
As the first known specimen of this species was
not collected until 1939 (Hinton 14750), Perkins
did not see this species. Gonsoulin (1974) annotat-
ed both collections he examined as Styrax argen-
teus var. argenteus, and Carranza (1996) also con-
sidered this species to be a variety of S. argenteus
(var. grandiflorus). However, in contrast to S. ar-
genteus as circumscribed here, S. austromexicanus
possesses white or pink (vs. consistently white)
flowers, non-tapered anther sacs, non-prolonged
anther connectives, and stamen filaments that lack
both auricles and stiff, long-armed, upward-point-
ing hairs ventrally. I consider the sum of these
character differences to justify recognition of this
taxon at the species level.
3. Styrax conterminus Donn. Sm., Bot. Gaz. 18:
5. 1893. TYPE: Guatemala. Quiché: San Mi-
guel Uspantán, Apr. 1892, Heyde & Lux 2915
(holotype, US; isotypes, B destroyed, GH,
HRCB not seen, M not seen, MO, NY). Figures
7, 32-35.
Evergreen tree to 18 m. Young twigs lepidote,
the scales brown mixed with olive-green; outer
layer of older twigs not exfoliating. Petiole 13-
25 mm long. Lamina thick-chartaceous, 8.5-16.5
X 2.7-5.0 cm, 2.8-3.9 times as long as wide,
narrowly elliptic or narrowly oblong, secondary
veins 11-13; apex acuminate; base cuneate to
slightly attenuate, symmetric or slightly oblique;
adaxially glabrous or sparsely lepidote; abaxially
covered (surface and veins) with olive-green pel-
tate or lacerate-margined scales, and more scat-
tered orange-brown scales; margin entire. Inflo-
rescences axillary or false-terminal, racemose or
paniculate, racemes or panicle branches 3-5.5
cm long, 4—9-flowered. Lower pedicels 8-12 mm
long. Calyx 3-4 X 4-5.5 mm, conical, olive-
green-, light brown-, or orange-brown-lepidote,
scales at mid-calyx 0.33-0.42 mm diam., peltate
or lacerate-margined; calyx teeth 5, to 0.7 mm
long, deltoid; calyx margin slightly concave or
rarely truncate between the teeth, inner margin
rarely glandular. Corolla white or pink, 10-13
mm long, petals connate 2-4 mm beyond the са-
lyx margin distally; corolla lobes 5, valvate in
bud, 7-10 X 3 mm, non-overlapping, erect or
spreading, thickened, linear-deltoid, vesture on
the outer distal third of the lobes consisting of
peltate or lacerate-margined scales. Free portion
of stamen tube 2-3 mm long; distinct portion of
filament 1-2 mm long, of equal width throughout,
ventrally not auriculate, sparsely radiate-lepidote
at the somewhat incurved filament margins, gla-
brous on the face, the scales with arms to 0.4 mm
long; anthers 3.5 mm long, the connectives not
or only slightly prolonged beyond the non-ta-
pered anther sacs. Ovary lepidote; style glabrous
or pubescent at base only; stigma 0.4-0.5(0.8)
mm wide. Drupe 12-15 X 7-9 mm, ellipsoid;
wall irregularly and coarsely rugose on herbari-
um specimens. Seed coat glabrous.
n AAA. A
—— CE.
A A ee e e COR
Volume 84, Number 4
1997
Fritsch
Revision of Styrax
Figure 32. Distribution of Styrax conterminus and 5. magnus.
Southern Chiapas to western Honduras and
northern El Salvador; + 1500-2800 m; Pinus-
Quercus-Ostrya forest, broad-leaved cloud forest.
Flowering: April-June, December; fruiting: Janu-
ary, February, June, August.
Common names. Copalillo (El Salvador, 4 June
1976, Reyna s.n. p.p.).
Additional specimens examined. EL SALVADOR.
Santa Ana: Cerro Montecristo, Allen 7184 (F, GH, LL,
MICH, US), 4 June 1976, Reyna s.n. p.p. (CR). GUATE-
M nango: above Puente Alto,
45107 (451047) (F, GH, MEXU, MO, NY); above San
Juan Ixcoy, Sierra de los Cuchumatanes
49982 (F, US). VW" slopes of Volcán Gemelos, Stey-
ermark 43279 (A, F, NY). HONDURAS. Ocotepeque:
Alrededor de la Laguna Verde, 12 km al SE de Nueva
Ocotepeque, Dario 59 ара MEXICO. Chiapas: on
ridge NE of Cerro Boquerón rom El Rosario to
Niquivil, Breedlove 68928 (CAS); пи near Motozin-
tla, Matuda 15507 (CAS, F, LL, NY); Cerro del Вод
Purpus 7422 (GH, NY).
Styrax conterminus is a rare endemic to northern
Mesoamerica. It is easily distinguished from other
species within its range by the possession of peltate
South American species, e.g., 5. cordatus (Ruiz &
Pav.) A. DC. [likely = S. ovatus (Ruiz & Pav.) A.
DC.] and S. leprosus Hook. € Arn. and is clearly
derived within the genus. These species possess
other putative derived characters, such as auricu-
late stamen filaments and tapered anther sacs,
whereas S. conterminus has non-auriculate stamen
filaments and non-tapered anther sacs (polarity as
determined from the outgroup taxa Pamphilia and
S. foveolaria). Many other South American species
e.g., S. guianensis A. DC., S. subargenteus Sleu-
mer) have stellate-pubescent calyces but tapered
anther sacs and/or auriculate stamen filaments.
This combination of characters in S. conterminus
suggests three possible evolutionary pathways for
this species: (1) derivation from a peltate-scaled an-
cestor followed by multiple losses of auriculate sta-
men filaments and tapered anther sacs; (2) deriva-
tion from a non-auriculate ancestor followed by
convergent evolution of peltate scales; (3) specia-
tion resulting from hybridization involving a non-
auriculate, stellate-pubescent taxon and a peltate-
scaled, auriculate taxon, whereby non-auriculate
anthers, non-tapered anther sacs, and peltate scales
were fixed in the hybrid species. Detailed phylo-
genetic analysis
characters would help to determine the likelihood
of each of these possibilities.
The protologue of Styrax conterminus does not
specify the holotype location. ли (1974) in-
722 Annals of the
Missouri Botanical Garden
Figures 33-39. 33-35. Styrax conterminus. —33. dr Sus Le adjacent stamens. 33, 34, Sharp 45107 i"
451047; “Po — 85. Fruit, Allen 7184 (MICH). 36-39 39. Styr us. —36. Flowering branch. —37. Flower. — 38.
Four adjacent stamens. 36-38, Croat 37190 (MO). — 39. Fr vit "TIN INBio 16 (MO).
|
Volume 84, Number 4
1997
Fritsch 723
Revision of Styrax
Figure 40. Distribution of Styrax gentryi and S. jaliscanus.
dicated that the holotype of S. conterminus was de-
posited at MO. However, the label heading of the
type collection states “Ex plantis guatemalensibus,
quas edidit John Donnell Smith,” and John Donnell
Smith donated his herbarium to US. If the collec-
tion originated from Donnell Smith’s herbarium, as
seems likely, then the holotype is best considered
to be at US rather than MO.
4. Styrax gentryi P. W. Fritsch, sp. nov. TYPE:
Mexico. Sinaloa: Ocurahui, Sierra Surotato, 1—
10 Sep. 1941, H. S. Gentry 6244 (holotype,
DS; isotypes, GH, MICH, MO, NY). Figures
40-44.
Arbor r sempervirens; lamina subtus alba-stellata-pubes-
us 1-2 mm longis, contiguis; corolla
lobis in aestivatione valvatis, x anthesi apertis; filamen-
a stamin rtibus distinctis ad margines ventrales pilis
stellatis aequiliter dispersis instructa, aliter fere glabra
ventraliter praeter ad apices, brachis pilorum plus min-
usve radiatis ex axibus centralibus; connectivum antherae
loculos non-contractos non vel leviter superans.
Evergreen tree to 17 m. Young twigs grayish
green-stellate-pubescent mixed with scattered or-
ange-brown stellate hairs; outer layer of older twigs
not exfoliating. Petiole 11-18 mm long. Lamina
chartaceous, 13-18 X 6-8 cm, ovate, broadly el-
liptic, or slightly oblanceolate, secondary veins 16–
20; apex acuminate to abruptly acuminate; base cu-
neate to rounded, symmetric or slightly oblique;
adaxially glabrous except along the major veins;
abaxially covered with minute greenish white stel-
late hairs and more scattered larger greenish white
stellate hairs, both types almost always covering the
secondary and tertiary veins, thus obscuring the
vein surface; margin entire. Inflorescences axillary
or false-terminal, racemose or paniculate, the ra-
cemes or panicle branches 3—6.5 cm long, 2-7-
flowered. Lower pedicels 6 mm long. Calyx 6-8 X
5-7 mm, campanulate, whitish green-stellate-pu-
bescent; calyx teeth 5, 1-2 mm long, deltoid, con-
tiguous, inner margin often glandular. Corolla of
unknown color, 17-20 mm long, petals usually con-
nate 3 mm beyond the calyx margin distally; corolla
lobes 5, presumably valvate in bud, 15 X 2 mm,
non-overlapping, spreading or recurved, thickened,
linear, vesture on the outer distal third of the lobes
consisting of stellate hairs. Free portion of stamen
tube 4 mm long; distinct portion of filament 1.0—
1.4 mm long, of equal width throughout, ventrally
not auriculate, white-stellate-pubescent at the
somewhat incurved filament margins, glabrous or
nearly so on the face, the hairs with arms to 0.4
mm long; anthers 6 mm long, the connectives not
or only slightly prolonged beyond the non-tapered
anther sacs. Ovary densely stellate-hirsute; style
glabrous or pubescent at base only; stigma 0.3 mm
wide. Largest drupe observed (only immature fruits
available for study) 10 X 7 mm, ellipsoid; wall ir-
724 Annals of the
Missouri Botanical Garden
Figures 41-46. 41—44. Styrax gentryi. —41. Fruiting branch, Gentry 6244 Miis MO) and Breedlove 15588
Tire cnt F). —42. Flower. —43. Four adjacent stamens. 42, 43, Do mínguez 28 (HCIB). —44. dm Gentry
6244 (MO). 45, 46. Styrax tuxtlensis. —45. Leaf, LaFrankie 1306 (CAS). —46. Fruit, LaFrankie 1306 (MO
A AE AAA O ——— — ID и
Volume 84, Number 4
1997
Fritsch 725
Revision of Styrax
regularly and coarsely rugose on herbarium speci-
mens. Seed coat observed immature, glabrous.
Sierra de La Laguna, Baja California Sur and
Sierra Surotato, Sinaloa, Mexico; 1600-2200 m;
Quercus-Pinus forest, mixed dominants forest;
found in riparian situations, such as arroyos, steep
moist ravines, and deep rocky canyon bottoms.
Flowering: October, probably also in March, Au-
gust; fruiting: March, October
Common name. Aguacatillo (Baja California Sur,
Domínguez 28).
Additional specimens examined. MEXICO. Baja Cal-
ifornia Sur: Sierra de la Laguna, Arroyo de los Encinos
Blancos, Domínguez 28 (“R.D.C. ») (HCIB); Sierra de la
ar Los Orno
sisse 15588 (CAS, F, NY); Sierra Surotato, Mpio. de
Badiraguato, Ocurahui valley, ge ove & Kawahara
16893 (CAS); Sierra Surotato, 5 mi. NW of Los Ornos,
Breedlove & Thorne 18269 (CAS, RSA); Sierra Surotato,
mi. N of Los Ornos, Breedlove & rne 18414 (CAS);
Sierra Surotato, 5 mi. NE of La Ciénega, Breedlove &
Thorne 18584 (CAS, RSA).
Styrax gentryi is a rare species with a disjunct
distribution in the Sierra de la Laguna of Baja Cal-
ifornia’s Cape Region and the Sierra Surotato in
northwestern Sinaloa, Mexico. It is particularly ap-
propriate that the species be named after Howard
Scott Gentry (1903-1993), considering that he was
able to collect this plant in both disjunct locali-
ties—from Baja California Sur in 1939 (the first
collection of the species) and Sinaloa in 1941.
This species is easily distinguished from all oth-
er species of evergreen Styrax by the prominent
calyx teeth, evident even in fruit. This is one fea-
ture that probably led Gonsoulin (1974) to include
Gentry 6244 and Gentry 4420 within the deciduous
capsular-fruited species S. jaliscanus. These collec-
tions were made just after anthesis, so no flowers
or fruits are available for examination. However, a
third collection (Breedlove 15588) with the same
calyces and leaves as the Gentry collections is an-
notated as S. argenteus var. argenteus by Gonsoulin.
This collection has immature fruit that establishes
its affinity with the evergreen, drupe-bearing spe-
cies rather than the deciduous, capsule-bearing
species. A relatively recent flowering collection
(Dominguez 28) further confirms the affinity of this
species with the evergreen taxa by virtue of its non-
overlapping corolla lobes.
is species’s disjunct distribution seems not to
be strictly paralleled in other flowering plants, al-
though distribution patterns of at least two species
are similar: Populus monticola Brandegee and Ilex
tolucana Hemsl. both occur in the Sierra de Ја La-
guna of the Cape Region of Baja California Sur and
in the same habitat as Styrax gentryi. However, on
the mainland, P. monticola occurs in Sonora, and
I. tolucana is disjunct from Sonora to Veracruz and
Oaxaca (L. Lenz, pers. comm.). Additional collec-
tions contributing to the still poorly known flora of
Sinaloa may reveal more taxa that are disjunct from
the Sierra Surotato and the Sierra de la Laguna.
The Cape Region first split from northern Jalisco
in the mid-Miocene (13-14 Mya; Smith, 1991) and
existed as an island until the establishment of the
Isthmus of La Paz in the Pleistocene (López Ramos,
1983; Fenby & Gastil, 1991). Because the two ar-
eas circumscribed by the range of S. gentryi were
never directly connected, explaining this disjunc-
tion by vicariance would require a prior mainland
distribution that extended at least partly into Nay-
arit and Jalisco. Although the narrow range of this
species in the two disjunct areas suggests relictual
status, S. gentryi is presumably bird-dispersed and
the current mainland distribution does not extend
south of northern Sinaloa. Thus, dispersal across
the Gulf of California remains a viable explanation.
Genetic analysis of disjunct populations of S. gen-
tryi and additional information regarding its distri-
bution and ecology would help to clarify this bio-
geographic issue.
5. Styrax glabratus Schott, in Spreng., Syst. v
4(2): 406. 1827. Strigilia glabrata (Schon)
3,
А ys
807 (holotype, ?BP; isotypes, F, US 09). Figure
Styrax не ee Pohl [“S. erymophyllum"], Pl. bras.
scr. 2: 57. 1830. TYPE: Brazil. Rio de Ja-
neiro: Schot 4184 (holotype, W; isotypes, BR(2), F,
GH, W(3)).
Epigenia integerrima Vell., Fl. flumin. 183. 1829. TYPE:
Icon. 4, t. RS in Vell, FI. flumin., UT ле
Styrax a 66. 1844 [“S. psi-
lophyllum"]. Pu peilophylla T DC.) Miers,
pos Mag. Nat. Hist., ser. 3, 3: 283. 1859. TYPE:
French Guiana. Cayenne, Martin s.n. (holotype, P;
pe. US).
Styrax d ен" in Lindl., Уер. kingd.
, t. 402 p.p. 1853 E leiophyllum"]. а
leiophylla (Miers) Miers, Ann. Mag. Nat. Hist., ser.
3, 3: 283. 1859. TYPE: Brazil. Rio io Janeiro: Val-
ença, John 1345 (holotype, BM not seen; photo o
olotype,
Styrax lauraceus Perkins, Bot. Jahrb. Syst. 31: 478. 1902.
PE: Venezuela. Distrito Federal: Galipan, Karsten
E (holotype, B destroyed; photo of holotype, NY(2);
sotype
sibs squamulosus M. F. on Acta Amazon. 1: 23. 1971
[^S. squamulosa”]. TY Brazil. Amazonas: Ma-
naus, Reserva Florestal Ducke, 1390 Levantamento,
726
Annals of the
Missouri Botanical Garden
2 | deo
e 47. Distribution of Styrax glabratus, S. incarnatus, S. nicaraguensis subsp. nicaraguensis, S. nicaraguensis
Figu
diii, ellipsoidalis, S. peruvianus, and S. steyermarkii.
14 Mar. 1971, W. Rodrigues & D. Coélho 8342 (ћо-
lotype, INPA not seen; isotype, NY).
Evergreen tree to 20 m. Young twigs sparsely to
densely greenish gray-stellate-pubescent or radiate-
lepidote; outer layer of older twigs not exfoliating.
Petiole 5-16 mm long. Lamina subchartaceous, 9—
19 X 2.56 cm, 2.2-3.8(-5.2) times as long as
wide, elliptic or oblanceolate, secondary veins 6—
9; apex acute to acuminate; base cuneate, symmet-
ric or slightly oblique; adaxially glabrous or sparse-
ly pubescent on the midvein at the base; abaxially
glabrous except for the midvein at the base, or rare-
ly sparsely stellate-pubescent or radiate-lepidote,
occasionally with domatia in the axils of the sec-
ondary veins, the hairs white to grayish green; mar-
gin entire. Inflorescences axillary or false-terminal,
racemose or rarely paniculate, the racemes or pan-
icle branches 2—7 cm long, E ey Lower
pedicels 2-13 mm long. Calyx 4–
cupuliform, usually grayish ie Wee.
(always in Costa Rica), sometimes with a few or-
ange-brown radiate scales toward the base, rarely
bes Brazil) stellate pubescent; scales at mid-calyx
diam.; calyx teeth 5 or often absent,
to 05 mm long, linear-deltoid, the calyx margin
truncate between the teeth, inner margin eglandu-
lar. Corolla white to pale pink, 13-20 mm long,
petals connate 1-4 mm beyond the calyx margin
distally; corolla lobes 5, valvate in bud, 8-13 х
2.5-3 mm, non-overlapping, reflexed, slightly
thickened, linear-deltoid, vesture on the outer dis-
tal third of the lobes consisting of radiate scales or
stellate hairs. Free portion of stamen tube 2-2.5
mm long; distinct portion of filament 1.5-2 mm
long, somewhat incurved, of equal width through-
out, ventrally not auriculate, each margin with a
cluster or longitudinal line of stiff white stellate
hairs, these sometimes also occurring on the ventral
face, the hairs with arms to 1.8 mm long, the hairs
nearest the distal end of the filament with arms
predominantly directed upward; anthers 5-8 mm
long, the connectives distinctly prolonged beyond
the tapered anther sacs. Ovary densely short-stel-
late-pubescent; style glabrous; stigma 0.3-0.4 mm
wide. Drupe 13-18 X 7-9 mm, ellipsoid; wall ir-
regularly and coarsely rugose on herbarium speci-
mens; fruiting calyx funnelform, 4—8 mm long. Seed
coat glabrous.
Osa Peninsula, SW Costa Rica; also Colombia,
Venezuela, Suriname, French Guiana, and Brazil;
400 m in Costa Rica. Flowering: October (buds) in
Costa Rica.
Specimens examined. COSTA RICA. Puntarenas: Can-
tón de Golfito Jiménez, Río Piro camino a Cerro Osa, Her-
rera 4534 (CR, INB).
In North America, Styrax glabratus has thus far
been documented only from the Osa Peninsula in
Costa Rica; the major portion of its range is in
South America, from Colombia to southeastern Bra-
zil. The existence on the Osa Peninsula of a species
otherwise found only in South America is not dis-
cordant with the general composition of the Osa
flora, which comprises many endemics and species
otherwise not known in Costa Rica and has strong
affinities to the flora of the Chocó region in Colom-
Volume 84, Number 4
1997
Fritsch 727
Revision of Styrax
poe Pasce | a es
| i | | | | | |
| | | | | |
ч Gulf | | |
| of | | | |
‚| Mexico | | |
| | | | gs |
nw | | [ |
| | | |
| | | |
| | ? | |
| Caribbean
| h^ | |
| | 6: | Sea
, Га | |
| | 1 + (2 |
id | Ч |> co E. 4 2 mus if |
| | | .
©
©
e
r£.
9 2
3 Ф
ce
wT
э
Ф
a
E
-
z
Ф
En
33
> =
Wi
MANY
=
* a
— 59. Flower. 58, 59, Breedlove 68742 (CAS).
McVaugh & Koelz 541 (MICH). —62. Four
(CAS). 61-63. Styrax radians. —61. Flower,
Tenorio 15605 (MO).
Figures 58-63. 58-60. Styrax magnus. —58. Flowering branch.
adjacent stamens. —63. Fruit. 62, 63,
—60. Fruit, Breedlove 46223
736
Annals of the
Missouri Botanical Garden
stellate-hirsute; pies "— or —— at base
only; stigma 0.5 wide. D 15-20 x 9-12
mm, ellipsoid; эй ин баг coarsely rugose
on herbarium specimens. Seed coat glabrous.
Uncommon in the central and southern moun-
tains in Chiapas, Mexico, extending into south-cen-
tral Guatemala along the Pacific slope; 1700-2700
m; montane rainforests and dry forests, evergreen
and mixed forests, Magnolia-Podocarpus and Quer-
cus-Drimys forests; found on dry ridges, on volcanic
slopes, in sandy soil. Flowering: May, August; fruit-
ing: September—January.
Additional specimens examined. GUATEMALA. Que-
zaltenango: above Mujuliá, Standley 85559 (F). San
cos: Barranco Eminencia, above San Rafael Pie de
la Cuesta, Standley 68618 (F); Río Vega, near San Rafael,
Steyermark 36234 (F). MEXICO. Chiapas: near summit
of Huitepec, Alexander 1214 (MICH, NY); 45-50 km NE
of Huixtla, Breedlove & Thorne 31045 (DS, MEXU, МО);
SW side of Cerro Mozotal, Breedlove & Thorne 31106 (DS,
MEXU, MO); Selva Negra, 10 km above Каубп Mezcala-
pa, Власа & ip 32394 (B, DS, DUKE, F, LL,
E Y, RSA); near the summit of Muk'ta
vits (Cerro Монако). Г вето 39600 (DS, МО, RSA);
near the summit of Muk’ta vits aed Huitepec), Breedlove
slo
41446 (DS, MO, TEX, US); E
(MO); Pueblo Nuevo a Simojovel, Miranda 9170 (US); La
Yerbabuena, al NW de Pueblo Nuevo Solistahuacán, Pa-
lacios 161 (CAS); Cerro Huitepec al W de San Cristóbal,
Ton 7963 (CAS, F, MO); Santa Cruz en San Filipe, Ton &
Concepción 9734 (CAS, GH, MICH, MO, NY, TEX)
Styrax magnus is endemic to the broad summit
of the Chiapas Plateau's wet eastern escarpment
and the volcanic slopes of the Sierra Madre in
southern Chiapas and southwestern Guatemala, al-
though the species seems to be relatively common
within its narrowly restricted geographic range. The
large, usually obovate leaves readily distinguish it
from other sympatric species of Styrax. Styrax ar-
genteus occurs at lower elevations and often has
leaves nearly as large as those of S. magnus, but
they are not obovate. Styrax incarnatus has leaves
of similar size and could conceivably be found
within the range of S. magnus. However, its leaves
are ovate or broadly elliptic and thick-chartaceous,
with the secondary and tertiary veins completely
covered by pubescence abaxially; the leaves of S.
magnus are chartaceous, with the abaxial surfaces
of the veins visible through the pubescence.
Gonsoulin (1974) subsumed Styrax magnus un-
der his S. argenteus var. hintonii. I have maintained
5. magnus as a species and assigned most of the
remaining specimens annotated by Gonsoulin as S.
argenteus var. hintonii to the stellate form of S. ra-
mirezii. There are two consistently reliable char-
acters that distinguish S. magnus from S. ramirezii:
the consistently larger mature drupe of S. magnus,
and the rather subtle leaf-shape difference between
the two species. Two other features of S. magnus
(branched, many-flowered inflorescences, more
widely spreading corolla lobes) also delimit this
species from S. ramirezii, but are not always pres-
ent. The individuals of S. ramirezii that have been
collected within the range of S. magnus are all the
large-scaled form of the species and are easily dis-
tinguished from S. magnus by the radiate-lepidote
calyx.
11. Styrax nicaraguensis P. W. Fritsch, sp. nov.
PE: —— Matagalpa: Cordillera de
Darién, 28-31 m E of Sebaco, 21 July
1962, G. L. Webster et al. 12496 (holotype,
MO; isotypes, DUKE, F, GH, LL, U). Figures
17-21, 47
sempervirens; lamina 2.1-3.0plo longior quam
densis cinerascentibus-viridibus-stellatis et saepe dis
sis ferrugineis-stellatis instructus; corolla lobis in aesti-
vatione valvatis, 2.5—4.0 mm latis, sub anthesi apertis,
recurvis vel reflexis, linearibus-deltoideis; filamenta stam-
inum partibus distinctis ad margines ventrales auriculatis,
auriculis pilis albis-stellatis dense instructis, brachis pi-
mm longae, ovoideae vel sign dea
Evergreen tree to 25 m. Young twigs orange-,
yellow-, or gray-brown-stellate-pubescent; outer
layer of older twigs not exfoliating. Petiole 11-16
mm long. Lamina chartaceous, 8.5-14.5 X 3.4-5
cm, 2.1-3.0 times as long as wide, elliptic or ovate-
oblong, secondary veins 8-11; apex narrowly acute
to acuminate; base cuneate to rounded, symmetric
or slightly oblique; adaxially glabrous except along
the major veins; abaxially densely covered with
short-armed, grayish green stellate hairs and scat-
tered larger orange-brown stellate hairs, the latter
particularly prevalent near and on the veins, the
secondary and tertiary veins glabrous or pubescent;
margin entire. Inflorescences axillary or false-ter-
minal, racemose or paniculate, racemes or panicle
branches 2.3-7 cm long, 2-10-flowered. Lower
pedicels 5-9 mm long. Calyx 3.5-5 X 46 mm,
cupuliform, grayish en n often
with larger orange-brown stellate hairs especially
toward the base; calyx teeth 5, to 0.3 mm long,
EA OS УНА
Volume 84, Number 4
1997
Fritsch 737
Revision of Styrax
deltoid, calyx margin truncate between the teeth,
inner margin often glandular. Corolla white, 13-15
mm long, petals not or only slightly connate beyond
the calyx re pcd corolla lobes 5, valvate
in bud, 9-11 mm, non-overlapping, re-
curved or ас thin, linear-deltoid, vesture оп
the outer distal third of the lobes consisting pre-
dominantly of radiate scales, best seen in the ma-
ture bud stage. Free portion of stamen tube 1.5 mm
long; distinct portion of filament 2 mm long, of
equal width throughout, ventrally with auricles
bearing a dense mass of stiff white stellate hairs
with arms to 1.0 mm long, also pubescent below
the auricles and at the sinus between adjacent fil-
aments, the hairs nearest the distal end of the fil-
ament with arms predominantly pointing upward;
anthers 5.8-7.8 mm long, the connectives distinctly
prolonged beyond the tapered anther sacs. Ovary
densely stellate-hirsute; style glabrous or pubescent
at base only; stigma 0.3 mm wide. Drupe (15-)17-
28 X (10—11–14(–15 if 2-seeded) mm, ovoid ог
obovoid; wall irregularly and coarsely rugose on
herbarium specimens. Seed coat glabrous.
Northwestern Nicaragua and northwestern Costa
Rica; 1100-1550 m
Styrax nicaraguensis is a rare endemic with a
disjunct distribution in the cloud forests of north-
western Nicaragua and northwestern Costa Rica.
This disjunction reflects a paucity of high moun-
tainous regions with cloud forest habitat between
the two areas where it is known to exist. Enough
time has apparently elapsed during separation for
the disjuncts to have differentiated morphological-
ly; hence, two subspecies have been designated
ere.
This species is probably the closest relative of 5.
argenteus, from which it can be distinguished by
the presence of orange-brown hairs on the calyx
and lower laminar surface, and the shorter stiff,
long-armed, upward-pointing stellate hairs on the
stamen filaments ventrally. ough S. argenteus
does reach up to 1500 m elevation in many parts
of its range, it does not occur in cloud forest habitat
where S. nicaraguensis oc
Gonsoulin (1974) identified the only collection
of this species seen by him at the time of his re-
vision (Williams 23955) as Styrax argenteus var. ar-
genteus (the annotation label on one of the dupli-
cates states “close to var. ramirezii”). Because these
specimens are either sterile or possess inflores-
cences with only pedicels and calyces, it is under-
standable that Gonsoulin overlooked the subtle
vegetative differences between this species and the
others in Central America. However, the collection
Gonsoulin saw does present unusual features with
respect to the rest of the material available to him,
and it is surprising that no mention was made of
this in his revision.
KEY TO THE SUBSPECIES OF STYRAX NICARAGUENSIS
la. dirum, өп: + rotate; ure ovoid, deeply ru-
n herbarium specim
ST lla. in iii subsp. nicaraguensis
lb. Fruiting calyx cupuliform; e oiim shal-
lowly rugose on herbarium spec
11b. 5, пагани e ellipsoidalis
lla. Styrax nicaraguensis subsp. nicaraguen-
sis. Figures 17-20, 47.
Fruiting calyx + rotate; drupe 17-21 X 13-14
(-15 if 2-seeded) mm, ovoid, deeply rugose on her-
barium specimens.
Rare in northwestern Nicaragua; 1300-1550 m;
cloud forests. Flowering: June—July; fruiting: June.
Additional specimens examined. NICARAGUA. Es-
telí: Laguna de Miraflores, Atwood A316 (MEXU, MO,
а Jinotega: те па de Miraflores, 8 June 1975, At-
wood s.n. (MO); km 146 carretera Мамин,
^s 16366 (MO). along Hwy. 3, с of
Fundadora entrance, Stevens & Henrich 20401 (МО). Ма-
tagalpa: Sta. María de Ostuma, Williams et al. 23955 (Е,
NY, US, W
llb. Styrax nicaraguensis subsp. ellipsoidalis
P. W. Fritsch, subsp. nov. TYPE: Costa Rica.
Puntarenas: Cordillera de Tilarán, Monteverde
Comunidad, 6 June 1977, V. J. Dryer 1420
(holotype, F; isotypes, CR, MO). Figures 21,
47
Calyx fructifer cupuliformis; drupa (15-)19-28 X
(10-)11-13 mm, obovoidea, rugosa tenuiter in sicco.
па calyx cupuliform; drupe (15-)19-28 X
0-11-13 mm, te shallowly rugose on her-
e speci
dr sin dn Rica in the Cordillera de
Guanacaste and the Cordillera Tilarán on the Pa-
cific slope; 1100-1500 m; forest edges. Flowering:
July, November; fruiting: April—June.
a
Additional specimens examined. COSTA RICA.
uanacaste: Cerro Cacao, Chávez 540 (CR, INB); Parque
Nacional aint mien Mengo, Cerro Cacao II IN-
Bio 164 (CR, M nas: Rio аа. Bello 932
(CR, INB); e uires rie 1420 (CR, F), Dryer 1575
CR, MO), Fuentes 192 (INB, MO), Haber 230 (ААЏ, CR,
P in Mata's pasture, Monteverde, Hartshorn 1894 (CR,
MO).
—
The II INBio 164 collection has smaller leaves
and fruit (15 X 10 mm) than other collections, but
its other features leave no doubt as to its identity.
738
Annals of the
Missouri Botanical Garden
| 100 km.
at a е
cw i |
de с
ae '
=
3
| 83W
Figure 64. Distribution of Styrax panamensis; see text
12. Styrax panamensis Standl., Contr. U.S. Natl.
Herb. 18: 121. 1916. TYPE: Panama. Сојбп:
Loma de la Gloria, near Fató (Nombre de
Dios), July and Aug. 1911, H. Pittier 4242 (ћо-
lotype, US-679343; isotypes, GH, NY, US).
Figures 11, 64.
Evergreen tree to 10 m. Young twigs golden
brown-stellate-pubescent; outer layer of older twigs
not exfoliating. Petiole 6-16 mm long. Lamina
chartaceous, 11.5-12 x 8-8.2 cm, 1.5 times as
long as wide, broadly obovate, secondary veins 6–
8; base cuneate, symmetric or slightly oblique; apex
acuminate or emarginate through abortive tip de-
velopment; adaxially glabrous except along the ma-
jor veins; abaxially covered with minute white stel-
late hairs and scattered larger, grayish white,
somewhat stiff stellate hairs, the latter occurring
most frequently on the veins, the minute hairs ab-
sent from the primary through tertiary veins thus
the vein surface easily visible; margin entire. Spec-
imens in bud only. Inflorescences racemose, axil-
lary or false-terminal, 3-4.5 cm long. Lower pedi-
cels 5 mm long. Calyx 6 X 5 mm, narrowly
cupuliform, uniformly golden brown-stellate-pubes-
cent; calyx teeth 5, minute, deltoid, the calyx mar-
gin truncate or slightly concave between the teeth,
inner margin apparently eglandular. Corolla in bud
11 mm long; corolla lobes 5, valvate in bud, 7-8
X 2 mm, linear-deltoid, vesture of the outer distal
third of the lobes consisting of stellate hairs. Free
portion of stamen tube 0.8 mm long; distinct por-
tion of filament 0.8 mm long, slightly flexuous, of
equal width throughout, ventrally not auriculate,
densely covered across the face with tawny, undu-
lating to somewhat stiff stellate hairs with arms to
0.8 mm long, the hairs nearest the distal end of the
filament with arms predominantly pointing upward;
anthers 5 mm long, the connectives distinctly pro-
longed beyond the tapered anther sacs. Ov.
short-stellate-pubescent; style glabrous. Fruit un-
nown.
for explanation of dubious record.
Endemic to western Panama; 10—104 m; forests.
Flowering (in bud only): July, August.
tyrax panamensis is known with certainty only
from the type collection. Its occurrence in low-el-
evation tropical forest habitat is unusual with re-
spect to the other Styrax species treated in this
revision. Fruiting specimens from 1550 to 1650 m
elevation in the Serranía del Darién [Gentry & Mori
13797 and 14073 (MO)] might belong to this spe-
cies, but they have smaller leaves with long-acu-
minate apices. А fruiting specimen from 150 m el-
evation in Mpio. de Hidalgotitlán, Veracruz, Mexico
(17%15'45"N, 94?29'30"W, Wendt 3576), also may
be this species. More material, particularly good
flowering and fruiting material of plants from near
Pittiers collection site, as well as material at an-
thesis from the Darién region and the locality at
Veracruz will be required to better understand Sty-
rax in these areas.
Although only flower buds are available on the
type, careful analysis of internal floral morphology
suggests that 5. panamensis is allied to other low-
land South American species (e.g., S. guianensis)
that lack auricles but possess a dense mass of hair
with relatively long arms (7 0.7 mm) on the ventral
side of the stamen filaments.
Perkins apparently did not see the Pittier collec-
tion. Gonsoulin (1974) annotated it as Styrax ar-
genteus var. ramirezii (— S. argenteus var. micran-
thus; see comments under $. ramirezii).
13. Styrax peruvianus Zahlbr., Ann. K. K. Na-
turhist. Hofmus. 7: 4. 1892. TYPE: Peru. Ca-
jamarca: Tambillo, Jelski 16 (holotype, W; is-
otypes, B destroyed, photo in F, GH, MICH,
MO, NY). Figures 13, 36–39, 47.
Evergreen tree to 10 m. Young twigs lepidote,
the scales brown mixed with olive-green; outer lay-
er of older twigs not exfoliating. Petiole (7-)10-15
mm long. Lamina thick-chartaceous, 6.5-12 *
2.2-5 ст, 2.0-3.1 times as long as wide, lance-
Моште 84, Митбег 4
1997
Fritsch 739
Revision of Styrax
elliptic or oblanceolate, secondary veins 7-9; apex
abruptly short-acuminate to rarely acuminate; base
cuneate to slightly attenuate, symmetric or slightly
oblique; adaxially glabrous or sparsely lepidote;
abaxially covered (surface and veins) with olive-
green peltate or lacerate-margined scales, and more
scattered orange-brown scales; margin entire. Inflo-
rescences racemose, axillary or false-terminal, 3.5—
8.5 cm long, 3-9-flowered. Lower pedicels 8-14
mm long. Calyx 3-3.5 X 4.5 mm, cupuliform, ol-
ive-green lepidote with scattered light orange-
brown scales, scales 0.2 mm diam., peltate
or lacerate-margined; calyx teeth 5, to 0.5 mm long,
deltoid; calyx margin slightly concave or rarely
truncate between the teeth, inner margin often
glandular. Corolla pink, 10-16 mm long, petals
connate 1-3 mm beyond the calyx margin distally;
corolla lobes 5, valvate in bud, 7-10 X 1.8-3.2
mm, non-overlapping, spreading to recurved, line-
ar-deltoid, thickened, vesture on the outer distal
third of the lobes consisting of peltate or lacerate-
margined scales. Free portion of stamen tube 1-1.7
mm long; distinct portion of filament 1.5-2.0 mm
long, of equal width throughout; ventrally with
prominent auricles bearing cales or stellate
hairs with arms 0.1–0.3 mm long, glabrous else-
where; anthers 4—5.5 mm long, the connectives dis-
tinctly prolonged beyond the tapered anther sacs.
vary lepidote; style glabrous or pubescent at base
only; stigma 0.3-0.4 mm wide. Drupe 11-20 X 5-
10 mm, ellipsoid; wall irregularly and coarsely ru-
gose on herbarium specimens. Seed coat glabrous.
Northwestern Costa Rica to western Panama;
also Colombia, Ecuador, and Peru; 1600-2100 m;
lower montane rainforests, mixed forests; found on
windswept ridges and in forest understory. Flow-
ering: July-August; fruiting: January, June, Octo-
ber.
Additional specimens examined. COSTA RICA. Ala-
juela: Monteverde Reserve, El Valle Trail near continen-
tal divide, Haber ex Bello 3173 (MO). Guanacaste:
RE элечи Guanacaste, Fila vernis I INBio 16
(CR, F, MO). Puntarenas: Monteve eserve, Senderos
Chomogo and Pantanoso, Haber & pe ام 9318 (CR,
INB). PANAMA. Chiriquí: Cerro Colorado, along mining
road 31.6 km beyond bridge over Río San Félix, Croat
37190 (MO, NY); (стр Pate Macho, de Nevers & Mc-
Pherson 6848 (CAS,
Styrax peruvianus is easily distinguished from
other sympatric species by the peltate or lacerate-
margined scales on the leaves, branches, and inflo-
rescences. The only other species in the region with
this feature is S. conterminus, which has narrowly
elliptic leaves, non-auriculate | stamen filaments,
and non-tapered anther sacs.
This species has been only rarely collected
throughout its range. The description given here is
based on the specimens cited above and the follow-
ing: L. Ellemann 66559 (LOJA, QCA; Ecuador), С.
Lozano C. 2537 (COL; Colombia), R. Sandino-Par-
do 11 (COL; Colombia), and B. Wallnéfer 18-1588
(W; Peru). Styrax peruvianus is considered to be-
long to a lineage comprising several species in
South America with peltate scales, stamen fila-
ments with auricles and/or dense pubescence with
long-armed hairs (> 0.8 mm) ventrally, and рго-
longed anther connectives, including S. cordatus
and S. leprosus.
14, Styrax platanifolius Engelm. ex Torr.,
Smithsonian Contr. Bot. 6: 4. 1854 [“S. platin-
ifolium”]. TYPE: U.S.A. Texas: Comal Co.,
near New Braunfels, 1851, Lindheimer s.n.,
(holotype, MO; isotype, GH). Figures 55, 56,
65, 66.
Deciduous shrub multi-stemmed from the base, to
3 m. Young twigs glabrous to densely white-stellate-
pubescent, sometimes somewhat glaucous, often to-
ward the base with scattered orange-brown or dark
brown stalked stellate hairs; outer distal layer of older
twigs often exfoliating into long strips. Petiole 6-20
mm long. мера - ert on P" twig usually alter-
nate, someti . Lamina chartaceous,
4.5-9. 0(-12) > x пун ue cm, depressed-orbicular
or very broadly ovate, sometimes slightly condupli-
cate, as evidenced by longitudinally folded laminae
on herbarium specimens, secondary veins 5-6; apex
rounded or obtuse, rarely acute, sometimes broadly
cuspidate; at least some leaf bases on each leaf cor-
date or truncate, others rounded, usually slightly at-
tenuate, symmetric or slightly oblique; adaxially
glabrous or pubescent; abaxially glabrous or thickly
covered with white stellate hairs, with scattered, or-
ange-brown or dark brown stalked stellate hairs es-
pecially prevalent on veins and the lowest pair of
leaves on each shoot, the stellate portion often decid-
uous from the base, the secondary and tertiary veins
evident but not prominent; margin usually egland
wg entire or apically with 2-several lobes or coarse
th. Inflorescences false-terminal, sometimes con-
ao and thus appearing umbellate, 2-5 cm long,
1-7-flowered. Lower pedicels 4-9 mm long, becoming
wider from base to apex, 1.3-2.3 times as long as the
calyx. Calyx 3—5(-6) X 4.5-5.5 mm, campanulate,
glabrous or densely white-stellate-pubescent, margin
and/or teeth at least sparsely glandular; calyx teeth
0—7, irregularly distributed, to 1 mm long, narrow-
deltoid or linear, the calyx margin truncate between
the teeth. Corolla white, 12-21 mm long, petals con-
740
Annals of the
Missouri Botanical Garden
New
Mexico
32 №
e
[X]
z
xas 3
^ e A
Coahuila ® е
©
5
100 km o |
S. platanifolius
e subsp. platanifolius
O subsp. stellatus
a subsp. texanus
e 65. пере of Styrax platanifolius subsp. platanifolius, S. platanifolius subsp. stellatus, and S. platan-
ifolius. sinn: texan
nate up to the calyx margin qeu corolla lobes 5(6),
imbricate in bud, 11-18 mm, overlapping,
spreading, thin, elliptic, vesture on the outer distal
third consisting of stellate hairs. Free portion of the
stamen tube 1—6 mm long; distinct portion of filament
2-9 mm long, of equal width throughout, ventrally not
auriculate and sparsely white-stellate-pubescent, be-
coming glabrous toward the distal end of the filament,
ма - with arms to 0.15 mm long; anthers 3—5.5
ng, the connectives not or only slightly pro-
said d the non-tapered anther sacs. Ovary
densely short-stellate-pubescent; style pubescent at
base only to nearly throughout; stigma 0.3-0.7 mm
wide. Capsule 7-10 X 7-11 mm (broader when 2—
3-seeded), globose, 3-valve-dehiscent, the wall hard,
— smooth but usually transversely
wrinkled post-d Seeds 6—9.5 mm long; seed
coat glabrous.
| peers
West-central Texas on the Edwards Plateau es-
carpment and in the Davis Mountains, to Coahuila
and Tamaulipas, Mexico, where it exists along the
@ S. platanifolius subsp. mollis
€ S. platanifolius subsp. youngiae
Figure 66. Distribution of Styrax platanifolius subsp. mollis and S. platanifolius subsp. youngiae.
Volume 84, Number 4 Fritsch 741
1997 Revision of Styrax
Table 1. Characters differentiating Styrax platanifolius from S. redivivus (modified from Fritsch, 1996b).
Character S. platanifolius S. redivivus
larger Bremse capsules 7-10 mm long 11-15 mm long
NER 6—9.5 mm long 9.5-12 mm long
caps м риђезсепсе canescent tawny or fulvous
seré pen length (ratio) 3–2.3 5-1.4
calyx
leaf margins
3-5 (6) X 4.5-5.5 mm
often coarsely lobed or irregularly undulate
4-7 X 5-7 mm
never lobed
northeastern escarpment of the Sierra Madre Ori-
ental, and the Chihuahuan desert mountains; 200—
2000 m; dry wooded bottomlands, rocky stream
banks and ledges, wooded canyons, izotal vegeta-
tion; found on limestone, and according to Cory
(1943), igneous soil in the Davis Mountains.
After пи M of S. platanifolius (Torrey,
1853), no new Styrax taxa from Texas were pub-
lished ael. Cory (1943) described two new species
(S. texanus and S. youngiae) and one new variety
(S. platanifolius var. stellatus) from the Edwards
Plateau and the Davis Mountains. Gonsoulin (1974)
agreed completely with Cory's assessment of the ge-
nus in Texas and incorporated material collected in
northern Coahuila and not seen by Cory into S.
youngiae.
Data from morphology (Fritsch, 1996b), isozymes
(Fritsch, 1996a), and DNA sequences (Fritsch,
1995) strongly suggest that these taxa are more
closely related to each other than to other taxa of
tyrax. However, in my opinion the morphological
variation within and among these taxa warrants the
recognition of only one species of Styrax from Tex-
as. Cory (1943) and Gonsoulin (1974) presented a
summary table of differences among S. platanifol-
ius, S. texanus, and S. youngiae. Most of the char-
acters in this summary (i.e., upper leaf surface col-
ог, calyx color, presence of calyx glands, vein
reticulation on lower leaf surface) are not taxonom-
ically reliable upon close inspection of the collec-
tions. The remainder involve differences in the
quantity of pubescence on the vegetative parts,
pedicels, and calyces that I consider to be taxo-
nomically inconsequential within many other spe-
cles of Styrax, such as S. glabrescens, S. grandifol-
ius, and S. redivivus. To place this variation in
perspective, there are reliable characters separating
members of this group from the closely related spe-
cles S. redivivus, endemic to California (Fritsch,
1995, 1996b; Table 1)
Additional evidence for the recognition of only a
single species of Styrax from Texas comes from iso-
zyme analysis (Fritsch, 1996a). Pairwise genetic
identities (J) estimated from isozyme allelic varia-
tion between populations of S. platanifolius subspp.
texanus and stellatus range from 0.986 to 0.991,
which is within the 0.90-1.00 range expected
among infraspecific taxa (Crawford, 1983). In con-
trast, pairwise estimates of 1 between populations
of S. redivivus from California and all taxa from
Texas ee is 0.446, well below the reported
average among congeneric species (0.67; Gottlieb,
1977; NEES! 1983, 1989). Because only four
populations within 5. platanifolius were sampled, it
would be desirable to obtain more isozyme data
from additional populations, = pna for subspe-
cies platanifolius, youngiae mollis. In this re-
gard, one individual Ы of subspecies mollis
(= S. youngiae in Fritsch, 1996a) exhibited several
alleles not shared by any individuals of subspecies
stellatus or texanus (= “S. platanifolius stellatus”
and “S. texanus” in Fritsch, 1996a, respectively).
Although the variation present is not considered
sufficient to confer species status to Stryrax texanus
and S. youngiae, an argument can be made for re-
taining these taxa as subspecies within S. platani-
folius. Variation in pubescence quantity in other
New World species of Styrax is either not at all or
only poorly associated with geography, ecology, or
life history (S. glabrescens, S. grandifolius), or at
most is distributed along latitudinal clines (S. amer-
icanus, S. redivivus), and I do not recognize infra-
specific taxa within these species. In contrast, tri-
chome morphology or abundance within
platanifolius is distinctly regional and facilitates
the delimitation of nearly or completely allopatric
taxa. Therefore, I have recognized five subspecies
within S. platanifolius based on minor but distinc-
tive differences. Trichome characters are those most
reliable for the delimitation of these subspecies;
characters of secondary importance include leaf
form, jap gland density, and surface features of
the st
свиње (1974) incorrectly attributed the de-
scription of S. platanifolius to “Boston J. Nat. Hist.
6: 146-147, 1854.” The article to which I presume
he was referring (Gray, 1850) is the second of a
three-part account of F. Lindheimer's collections in
742
Annals of the
Missouri Botanical Garden
western Texas. Because it covers the collections
from the years 1845-1848, this article would not
be expected to list the 1851 S. platanifolius collec-
tion in any case. The other two parts to the series
(Engelmann & Gray, 1845; Blankinship, 1907) also
do not cite the collection.
KEY TO THE SUBSPECIES OF STYRAX PLATANIFOLIUS
la. Abaxial laminar surface glabrous or visible
through the pubescence
2a. Leaves, pedicels, ind calyces without white
stellate hairs
a. S. platanifolius subsp. cogi Sede
2b. Leaves, pedicels and calyces with wh
stellate
a. кыта of еа |
abaxial ente iab — About
0 long, often exceeding 1 mm;
teeth to 0.3 mm long, the calyx margin
sparsely glandular
4b. S. platanifolius subsp. mollis
w
c
ds
Ф
Ф
=
5
e
е.
Ф
@
б
=
Ф
2
а
==
w
E
®
Е
E
~
©
5
еф
a
o
n-
dular 14c. S. platanifolius subsp. stellatus
lb. Abaxial laminar surface тр covered and
obscured by the pubesce
4a. Young twigs faintly uitis abaxial lami-
iar ponis from base to ca. 15-35% of
de total le
. S. platanifolius a. texanus
4b. Young twigs кайа ر а
зсепсе;
axial laminar surface stellate-pubescent;
style pubescent from base to ca
the total length :
14e. 5. platanifolius subsp. youngiae
14a. Styrax platanifolius subsp. platanifolius.
Figure 65
Young twigs glabrous except for scattered or-
ange-brown or dark brown stalked stellate hairs to-
ward the base. Lamina glabrous on both sides ex-
cept for scattered — or dark brown
stalked stellate hairs abaxially some leaves,
margin often irregularly ee re 3-lobed,
or coarsely toothed, less often entire. Pedicel and
calyx glabrous. At least some calyx teeth to 1 mm
long, the teeth and calyx margin densely glandular.
Style pubescent from base to 15-35% of the total
length
Rare, occurring in west-central Texas on the Ed-
wards Plateau escarpment; 200-700 m; dry wooded
bottomlands, rocky banks and ledges. Flowering:
— fruiting: July-September.
mon name. Sycamore-leaf snow-bell (Gon-
nc 1974).
Additional specimens examined. UNITED STATES.
exas: County u
mer 1897, Stanfield s.n. (NY); Bull Cr., Mr. J. rd's
реч 14 Арг. 1915, Young s.n. (TEX). "Эви Бе Little
Blanco River near intersection of Hwy. 281 and 473, Dorr
& Nixon 1910 (NY); bluff overlooking Little Blanco River,
5 of Twin Sisters, Gonsoulin 1289 (NY); Little Blanco Riv-
r and Hwy. 66-281, 24 Aug. 1947, Parks s.n. (LL); in-
tersection of Hwy. 281 and 473 on N bank of Little Blanco
River, 29 Apr. 1972, Welborn & Sanstrup s.n. (TEX). Bur-
WSW of = 16 Јипе 1950,
well Co.: 4.5 mi. W of Delhi on TX 713, Солин
1287 NY). Edwards Co.: ravine near W Nueces River,
ca. 19 mi. SW of Rocksprings, Gonsoulin 1306 (NY). Hays
Co.: near Lake Travis, 9.3 mi. WNW of Cedar Creek, Gon-
soulin 1288 (NY). Kendall Co.: near Edge Falls along Cur-
ry Creek, Correll 29152 (GH, LL); below Edge Falls, ca.
5 mi. S of Kendalia, Correll & Smith 29572 (GH, LL);
Edge Falls, Correll & Correll 37024 (LL). Kerr Co.: near
1 Hunt, Gonsoulin 1305
1 0907 (A); Big Saline Creek, Reverchón 1551 (GH). Kin-
ney Co.: on banks of Nueces River, Gonsoulin 1307 (NY).
Llano Co.: Enchanted Mountain, 26 June 1932, C. Е. A.
s.n. (TEX); Enchanted Rock, Jermy s.n. (NY), Jermy 105
(MO), 12 June 1930, Whitehouse s.n. (GH, NY), 25 Apr.
s.n.
Спис - 1206 (МУ). Travis Co.: cultivated at 1405 Rabb
d. in Austin, Correll & Correll 37003 (LL); Austin, Apr.
1913, Huppetz s.n. (TEX); Travis Peak, Austin, 3 May
1930, Prowse s.n. (TEX), 3 May 1930, Whitehouse s.n.
(GH); near Austin, Wright s.n. (GH).
This is the most widespread of the three subspe-
cies of Styrax platanifolius endemic to Texas, but
it is extremely rare throughout its range. Popula-
tions apparently were larger and more common in
the early 1900s, but the subspecies has since de-
clined dramatically (Cory, 1943; J. Poole, Texas
Parks and Wildlife Department, Austin, Texas,
pers. comm.; pers. obs.). Many populations have
been extirpated, and those that remain often consist
of only a single individual or a few plants. Docu-
mentation and monitoring of remaining populations,
assessment of genetic variation among all remain-
ing individuals, and management plans are desper-
ately needed to ensure the survival of members of
this subspecies.
14b. Styrax platanifolius subsp. mollis P. W.
Fritsch, subsp. nov. TYPE: Mexico. Coahuila:
Mpio. de Muzquiz, Rancho Agua Dulci, E
slope of the Sierra de San Manuel, 28 June
1936, Е L. Wynd & С. H. Mueller 340 (holo-
type, MO; isotypes, A, MICH, NY). Figure 66.
—
Volume 84, Number 4
1997
Fritsch
Revision of Styrax
743
Lamina alba-stellata-pubescens utrinque, subtus ad
tactum mollis, brachis longissimis pilorum stellatorum
typice 0.8 mm longis, saepe excedentibus 1 mm longis,
pagina abaxiali visibili infra tomento; pedicelli albi-stel-
lati-pubescentes; calyx alba-stellata-pubescens, margine
inferme glandifero, dentibus usque ad 0.3 mm longis.
Young twigs densely white-stellate-pubescent in
addition to scattered orange-brown or dark brown
stalked stellate hairs toward the base. Lamina
above with scattered stellate pubescence, the hairs
most often simple, sometimes stellate and then to
about 4-armed, below moderately densely white-
stellate-villous in addition to scattered orange-
brown or dark brown stalked stellate hairs on some
leaves, soft to the touch, the surface visible through
the pubescence, the longest arms of each stellate
hair typically about 0.8 mm and often exceeding 1
mm long, margin often usually entire, less often
weakly undulate. Pedicel and calyx with white stel-
late hairs of variable length. Calyx teeth to 0.3 mm
long, the calyx margin sparsely glandular. Style pu-
bescent from base to 30-75% of the total length.
Rare, Coahuila and Tamaulipas, Mexico, along
the eastern escarpment of the northern extension of
the Sierra Madre Oriental and Chihuahuan desert
mountains; 1200-1400 m; wooded canyons. Flow-
ering: April.
Additional s ipe examined. MEXICO. Nuevo
León: Monterrey, Chipinque, Peterson 1243 (TEX); Mon-
terrey, La Silla Trail, Smith M149 (TEX).
I have also seen individuals of this subspecies
cultivated at the Berkeley Botanic Garden, Berke-
ley, California (accession number 91.1305), origi-
nally collected by J. Fairey (Yucca Do Nursery,
Waller, Texas) from Purificación, Tamaulipas, Mex-
ico.
Styrax platanifolius subsp. mollis is a rare en-
demic to Coahuila and Tamaulipas, Mexico. It is
the southernmost of the five subspecies recognized
in 5. platanifolius, having no apparent range over-
lap with the nearest subspecies to the north (subsp.
youngiae), from which it is distinguished by the soft
pubescence that does not completely cover the low-
er laminar surface. The distinctness of this subspe-
cies from subspecies youngiae is highlighted by
Gonsoulin's (1974) ambiguity regarding its identi-
fication. He cited the only specimen available to
x of this subspecies (Wynd & Mueller 340) as 5.
ungiae, but annotated it as S. platanifolius var.
cim probably because subspecies stellatus re-
sembles subspecies mollis more closely than does
subspecies youngiae.
14c. Styrax platanifolius subsp. stellatus (Cory)
P. W. Fritsch, comb. et stat. nov. Basionym:
Styrax platanifolius var. stellatus Cory, Madro-
fio 7: 111. 1943 [*S. platanifolius var. stella-
ta^]. TYPE: U.S.A. Texas: Bandera Co., Sa-
binal Canyon, 16 June 1940, V. L. Cory 34765
(holotype, A; isotype, NY). Figure 65.
Young twigs sparsely white-stellate-pubescent in
addition to scattered orange-brown or dark brown
stalked stellate hairs toward the base. Lamina
above with scattered stellate pubescence, the hairs
with 6—14 arms, below with scattered white stellate
pubescence in addition to scattered orange-brown
or dark brown stalked stellate hairs on some leaves,
somewhat rough to the touch, the surface visible
through the pubescence, the longest arms of each
stellate hair typically about 0.4 mm long, rarely if
ever reaching 1 mm, margin often irregularly un-
dulate, weakly 3-lobed, or coarsely toothed, less of-
ten entire. Pedicel and calyx with white stellate
hairs of variable length. At least some calyx teeth
to 1 mm long, the teeth and calyx margin densely
glandular. Style pubescent from base to 50—7046 of
the total length.
Rare in west-central Texas on the Edwards Pla-
teau escarpment; 500—700 m; dry wooded bottom-
lands, rocky banks and ledges. Flowering: April-
Ed lign July-September.
name. Hairy sycamore-leaf snow-bell
биш 1974).
Additional specimens examined. UNITED STATES.
Texas: Bandera Co: 5 1 omi. N of anser 15 June
11528 (GH, MO, NY, POM); 2 mi. N of Vanderpool, Parks
1009 (A); Vanderpool, Parks 1942 (TEX), Parks 40988
(A); Lost Maples im National Area, along the Sabinal
i n 1205 эч Sabinal Canyon State
274 q Sabinal тубе са. 7 т
Арг. 1972, Welborn n. (TEX); Hwy. 407 (4702) көзе
Utopia - Тагрјеу, 5 bank of Hondo pes 9 Apr. 1972,
Welborn s.n. (TEX). Kendall Co.: ca. 6.5 mi. NW of Boer-
ne, on заћи of Cibolo Creek, 24 Sep. 1983, Brecken-
n. (A
Co eh Palmer 11474 (GH, MO, NY); Cibolo, 15 June
1913, Schattenberg s.n. (A). Real Co.: along TX 337, 3
mi. W of Leakey, Miller et al. 5145 (MO). Uvalde Со.:
Garner State Park, SE boundary of park along Frio River,
744
Annals of the
Missouri Botanical Garden
Riskind 1691 (LL); Garner State Park, southern portion of
park, 9 Apr. 1972, Welborn s.n. (TEX).
The range of Styrax platanifolius subsp. stellatus
is contiguous with the southern boundary of sub-
species platanifolius, occurring at the southern
edge of the Edwards Plateau. It is rare throughout
its range, with the exception of Sabinal Canyon
State Park, where apparently populations are com-
mon and often composed of many individuals (Ris-
kind 1682).
Gonsoulin (1974) incorrectly cited the holotype
at TEX and an isotype at Е
14d. Styrax platanifolius subsp. texanus (Cory)
P. W. Fritsch, comb. et stat. nov. Basionym:
Styrax texanus Cory, Madrofio 7: 112. 1943
[“S. texana”]. TYPE: U.S.A. Texas: Edwards
Co., Polecat Creek, 4 July 1941, V. L. Cory
34940 (holotype, GH). Figure 65.
Young twigs faintly glaucous, glabrous except for
scattered orange-brown or dark brown stalked stel-
late hairs toward the base. Lamina above glabrous,
below thinly and evenly white-stellate-pubescent
except for scattered orange-brown or dark brown
stalked stellate hairs on some leaves, the surface
completely covered and obscured by the pubes-
cence, margin usually entire or weakly undulate,
rarely lobed or coarsely toothed. Pedicel and calyx
thinly and evenly white-stellate-pubescent. At least
some calyx teeth to 1 mm long, the teeth and calyx
margin densely glandular. Style pubescent from
base to 15-35% of the total length.
Rare, occurring in west-central Texas on the Ed-
wards Plateau escarpment; 500-700 m; steep lime-
stone cliffs. Flowering: April-May; fruiting: July—
September.
Common name. Texas snow-bell (Gonsoulin
1974; Cox, 1987).
парене specimens examined. UNITED STATES.
Texas: Edwards Co.: Cedar Creek, canyon walls above
nnd springs, 25 Sep. 1983, Breckenridge s.n. (TEX);
epe Creek, 4 July 1940 or 1941, Cory s.n. (GH, RSA),
r. 1942, Cory s.n. (GH, POM), Cory 34937 (TEX),
Cory 34938 (POM), Са 34939 (ТЕХ); Cedar Creek, 1.25
ті. above Roberts Greek, و 37767 (TEX), Cory 37769
8763 (TEX), Cory 38764 (A);
at Creek, Cory (TEX); Polecat Canyon, Cory
38034 (NY) Little Hackberry Canyon, Cory 42953 (NY);
Little Hackberry Creek, 14.5 mi. E of Rocksprings, Cory
42956 (TEX); Ра ries Cory 49179 (LL); cliff above
= Creek, Barksdale, Dorr & Nixon 1918
Cedar Creek fi oa ре of — Johnston
et 9 7428 8 (T — Jessup Ran n Little Hackberry
Creek, Keeney 145 е Yu Расне Gael. Keeney 1457
(MO). Real Co.: Nueces Canyon, 4.5 mi. N of Hackbe
Cory 42659 (F), Cory 42661 (TEX), Cory 42664 (NY),
Cory 42666 (TEX); Nueces River, 4.25 mi. above Hack-
wt Cory 42668 (A), Cory 42669 (POM); Nueces River,
1/3 mi. above Hackberry, Cory 42673 (A), Cory 42677
o Cory 42678 (F); 9 mi. ENE of Vance, W side of
Bullhead Creek, Johnston 7304 (TEX)
Styrax platanifolius subsp. texanus has the nar-
rowest distribution of the five S. platanifolius sub-
species, with documented localities in Edwards and
Real Counties, Texas, and one population reported
in Del Rio County, Texas. It is probably also the
rarest of the subspecies, with only nine populations
known, four of which consist of more than two in-
dividuals (J. Poole, Texas Parks and Wildlife De-
partment, Austin, Texas, pers. comm.). Although
seed production and the germination rate of 5. pla-
tanifolius subsp. texanus are relatively high, grazing
by both native and exotic herbivores has prevented
regeneration within populations (Cox, 1987). Re-
covery efforts by the Texas Parks and Wildlife De-
partment are under way to help ensure the survival
of this taxon.
Gonsoulin (1974) incorrectly cited the type as
“Cory 19 Apr 1942” with isotypes at GH and NY.
14e. Styrax platanifolius subsp. youngiae
(Cory) P. W. Fritsch, comb. et stat. nov. Bas-
ionym: Styrax youngiae Cory, Madroño 7: 113.
1943 [“5. Youngae”]. TYPE: U.S.A. Texas:
Limpia, canyon (on MO specimen only), Davis
Mountains, 12 May 1914, M. S. Young s.n. (ho-
lotype, TEX; M. S. Young 23 isotype, MO).
Figures 55, 56, 66.
Young twigs densely white-stellate-pubescent in
addition to scattered orange-brown or dark brown
stalked stellate hairs toward the base. Lamina
above with scattered stellate pubescence, the hairs
with 6-14 arms, below with a thick layer of white
stellate pubescence except for scattered orange-
brown or dark brown stalked stellate hairs on some
leaves, the surface completely covered and ob-
scured by the pubescence, the hairs of variable
length, margin entire, undulate, or coarsely toothed.
Pedicel and calyx with a thick layer of white stel-
late pubescence, the hairs of variable length. Calyx
teeth to 0.6 mm long, usually shorter, the teeth and
calyx margin sparsely glandular. Style pubescent
from base to 60-80% of the total length.
Rare, Davis Mts., Texas, and eastern slopes of
Chihuahuan desert mountains in northern Coahui-
la; 000 m; izotal vegetation (Yucca, Nolina,
Dasylirion associates), pifion/juniper woodland, in
canyons; found on limestone; according to Cory
(1943), found on igneous substrates in the Davis
Mts., Texas, but this needs confirmation. Flowering:
April-May; fruiting: July-September.
Volume 84, Number 4
1997
Fritsch 745
Revision of Styrax
Common name. Young’s snowbell (Gonsoulin,
1974).
ELA specimens examined. MEXICO. Coahuila:
. E of Boquillas in Sierra del Carmen, Henrickson
s 499 (TEX), Henrickson 11561 (TEX); 17 за N of Кап-
cho, El Jardín, Johnston et al. 11888 (CAS, F, LL, MO,
of Rancho Lagunita, one 9 а 1915 (ТЕХ);
Mpio. de Villa Acuña, Serranías del Burro, Rancho El
Bonito, Riskind 2079 (TEX): Sita del Carmen, 1/2 mi.
(TEX); Sierra del Carmen, in fork o:
Cañón del Diablo, Wendt 582 (MEXU, TEX).
Styrax platanifolius subsp. ers occurs in
the Davis Mountains, Texas, U.S.A., and the Sierra
del Carmen and Макон Ri н regions
in the Chihuahuan Desert, Coahuila, Mexico. The
type locality in the Davis Mountains (“Limpia; can-
yon”) has not been rediscovered. The Davis range
consists primarily of igneous substrates, so Cory
(1943) presumed the type to have come from an
igneous rather than a limestone substrate. This fact
and the higher elevation at which this subspecies
occurs were considered to be features that distin-
guish 5. youngiae from S. platanifolius and 5. tex-
anus. However, all subsequent collections of S. pla-
tanifolius subsp. youngiae that indicate substrate
have come from limestone, placing in doubt Cory’s
(1943) assumption of an igneous substrate for the
type. The elevation difference is still a distinguish-
ing feature, although it is not as wide as Cory re-
ported: Cory described S. platanifolius (= S. pla-
tanifolius subspp. platanifolius and stellatus) and S.
texanus (= S. platanifolius subsp. texanus) as oc-
curring below 600 m, and S. youngiae (= S. pla-
tanifolius subsp. youngiae) as occurring above
1200 m; current estimates of elevation ranges
(based on data from herbarium labels) are 200-700
m for subspecies platanifolius, stellatus, and tex-
anus combined, and 900-2000 m for subspecies
youngiae.
The holotype is without number, but the isotype
(Young 23) is clearly part of the same collection as
the holotype, as indicated by the identical dates,
general locality, and stage of leaf development.
15. Styrax radians P. W. Fritsch, sp. nov. TYPE:
Mexico. Jalisco: (Mpio. de) San Sebastián, Ha-
cienda del Cura, Sierra Madre, 2 Jan. 1927,
Y. Mexía 1351 (holotype, CAS; isotypes, A.
DS, F, GH, MICH, MO, NY, UNC not seen,
US). Figures 25, 61-63.
or sempervirens; lamina ovata-oblonga vel elliptica,
йш lanceolata vel suborbicularis, subtus alba-stel-
lata-pubescens, ad tactum mollis, venis secondariis et ter-
tiariis evidentibus; calyx stellata-pubescens; corolla alba,
is
moderate vel late; filamenta staminum partibus distinctis
.9 mm longis et ad margines ventrales pilis stellatis
ја а dispersis instructa, aliter fere glabra ventraliter
praeter ad apices, brachis pilorum plus minusve radiatis
ex axibus centralibus; connectivum antherae loculos non-
contractos non vel leviter superans; stigma 0.6-0.8 mm
lata.; drupae longiores 9-13 X 5-7 mm.
Evergreen tree to 12(-20) m. Young twigs grayish
white-stellate-pubescent, often mixed with tawny-
or orange-brown stellate hairs; outer layer of older
twigs not exfoliating. Petiole 9-17 mm long. Lam-
ina chartaceous, (8.5-)11-15(-24) х 4-9(-11.5)
cm, broadly ovate-oblong or elliptic, sometimes lan-
ceolate or suborbicular, secondary veins 9—15; apex
acute to rounded, rarely broadly cuspidate; base
subrounded to rounded, rarely truncate, subcorda-
te, or cuneate, often slightly attenuate, symmetric
or Amd oblique; adaxially glabrous except along
e major veins; abaxially covered with minute
urs stellate hairs and also larger, woolly, whitish
hairs rendering the surface soft to the touch, the
minute hairs absent from the primary through ter-
tiary veins, thus rendering the vein surface easily
visible, the soft larger hairs rarely absent; margin
entire. Inflorescences axillary or false-terminal,
racemose or especially the larger usually panicu-
late, the racemes or panicle branches 2-6 cm long,
2—14-flowered. Lower pedicels 3-8 mm long. Calyx
2.5-5 X 2.5-5 mm, cupuliform or conical-cupuli-
form, grayish green-stellate-pubescent, rarely also
with scattered orange-brown stellate hairs; calyx
teeth 5, to 0.4 mm long or absent, deltoid, the calyx
margin truncate or sometimes concave between the
teeth, inner margin often glandular. Corolla white,
5-15 mm long, petals connate up to the calyx mar-
gin distally; corolla lobes 5, valvate in bud, 4-14
X 1-3 mm, non-overlapping, moderately or widely
spreading, thin, linear-deltoid, vesture on the outer
distal third of the lobes consisting of stellate hairs.
Free portion of stamen tube 0—4 mm long; distinct
portion of filament 2.5-3.5 mm long, of equal width
throughout, ventrally not auriculate, + densely
white-stellate-pubescent at the incurved filament
margins, glabrous or nearly so on the face proxi-
mally, increasing in pubescence distally, the hairs
with arms to 0.7 mm long; anthers 2-6 mm long,
the connectives not or only slightly prolonged be-
yond the non-tapered anther sacs. Ovary densely
stellate-hirsute; style pubescent from base to 40—
85% of the total length; stigma 0.6–0.8 mm wide.
Drupe 9-13 X 5-7 (wider when 2-seeded) mm (not
including occasional beak), ovoid or narrowly ellip-
746
Annals of the
Missouri Botanical Garden
soid; wall irregularly and coarsely rugose on her-
arium specimens. Seed coat glabrous.
Most common in Nayarit and extreme western
Jalisco, but also in southern Sinaloa, southern Mex-
ico, northern Guerrero, and Oaxaca, Mexico; 400—
1400 m; Quercus savannas, Quercus-Pinus forests,
ecotones between subtropical deciduous forest and
tropical subdeciduous forest, tropical subdeciduous
forests, tropical deciduous forests; found often in
valley bottomlands or barrancas. Flowering: No-
ан fruiting: February—July.
Com names. Aguacatillo (Oaxaca, Mac-
ЫРА Hi 56), azagar (Jalisco, Vázquez & Guzmán
4164), azajar (Jalisco, Cárdenas et al. 79, Robles et
al. 813), cortapico (Jalisco, Mexía 1351), levadura
(Nayarit, Gentry & Gilly 10863), mamuyo (Jalisco,
Cuevas & Rosales 1876), zajar (Jalisco, Santana et
al. 4776)
Uses. Wood is used for ox yokes, plows, and other
purposes where durable, tough wood is desired (Jal-
isco, Mexía 1351).
Additional specimens examined. MEXICO. Guerre-
ro: Tlalchapa, Aguirre 51 (MEXU). Jalisco: са. 35
km E of Cabo Corrientes, Anderson & Anderson 6109
(MICH); El Tuito, 11. 5 qa al NW del pueblo por el cam-
ino a Ixtlahuahey, Ayala 678 (F, MO, NY); 14 km W of
Villa de Purificación, Bullock 2024 (CAS); Cuautitlán, 2
km NE of El Durazno, Cárdenas et al. 79 (WIS); Cabo
Corrientes, 15-78 km SW of Puerto Vallarta, Cochrane et
al. 11942 (WIS); new dirt road from San Sebastián W of
La Estancia, Cochrane et al. 12047 (WIS); Mpio. де Pur-
ificación, El Ocotillo, 10-12 km al NE de Purificación,
Cuevas et al. 709 (WIS); 4-6 km al NE de Casimiro Cas-
tillo, Cuevas et al. 2391 (WIS); 4-5 km al W de Casimiro
Castillo, De Niz et al. 23 (WIS); gorge of the Río Horcones,
27 km by road S from Puerto Vallarta, Dieterle 4023
(MICH); center of El Durazno, Iltis et al. 389 (WIS); 2-7
km ENE of El Durazno, Iltis et al. 459 (WIS); Ayotitlán,
Judziewicz et al. 5212 (WIS); 7.6 km al E de la carretera
Puerto Vallarta, Lott 2540 (CAS, MICH, MO); ca. 4 km al
N
itas, Межа 1861 (CAS, US); Cuautitlán, 29 km al SW
de Telcruz, Nieves 411 (WIS); E de Telcruz,
Nieves 546 (WIS); Cerro Huehuentón, FS 814 (NY);
along hwy. from Puerto Vallarta S to Chamela at Río Hor-
cones, Plowman 14547 (F); Mpio. de Cuautitlán, Río Ay-
otitlán entre Ayotitlán y Chancol, Ramírez & Alcocer 477
(WIS); 6.5-7.5 km al NE de Cuautitlán, Robles et al. 813
(WIS); 11 km al NE de auge Santana & Guzmán
tana et al. 4776 (WIS); Mpio. de d - E cam-
ino de Talpa a Cuale, 4 km después Vázque.
1195 (WIS); 1-2 km al ` del Tecolote, El ade o Cas-
imiro Castillo, Vázquez & Guzmán 4164 (WIS). México:
Mpio. de Temascaltepec, Luvianos, Hinton 5309 (A, F,
RSA), Hinton et al. 6193 (CAS, NY, RSA, US); Mpio. de
Temascaltepec, Villaneda, Hinton et al. 7410 (A, MO, U,
US); 3 mi. W of Luvianos, Moran 10161 (DS, RSA). Nay-
arit: Camichín de Juaja, Calzada 17840 (RSA); 4 km al
NE del poblado de Sta. María del Oro, Calzada et al.
18805 (MICH); Hwy. 28 between Tepic and Jalcatlán at
m 14-16, Croat 45262 (CAS, MO); 3 km al SW del Izote,
camino al Cuarenteño, Flores & Ruenes 1907 (F, MO); 11
km al У de Jalcocotán, Flores & Ramírez 2498 (MICH,
MO); 11 km al NE del Izote, camino a Palapita, Flores-
Franco et al. 3344 (MICH); 8 to 10 mi. W of Tepic, Gentry
& Gilly 10485 (LL, MICH); 15 mi. 5 of Tepic along hwy.
to Compostela, Gentry & Gilly 10811 (LL, MICH); 15 mi.
S of kem along road to Compostela, Gentry & Gilly 10863
(LL, ; mountains 9 mi. N of Compostela, McVaugh
& cea 541 ; ca. 10 road-mi. E of Jalcocotán on
road to Tepic, McVaugh 12100 (МИ) 3 km NW a! Es
Ocotillo, Mg 23515 (MICH); along dirt road to С
entefio, 2-10 km from El Izote, Miller & Téllez 3140 (CAS,
MO); ca. 3 mi. E of Jalcocotán, Norris 14959 (MICH);
Rancho de la Siempreviva, Ortega 60 (A, US); 2 a 10 km
de Mazatan, Ramírez & Flores 894 (MICH, MO); La Es-
condida, 14 km al NE de Tepic, Téllez 10282 (MO); km
9.5 de la terraceria al Спагетећо, Téllez et al. 10412
ecd MO); between bs and Ixtlan del Río, ну Коу.
(MICH, MO, RSA); 68 km al МЕ de San Pedro Ixzcatan,
Tenorio et al. 16929 (MICH, MO); 3 km al S del Izote,
Tenorio et al. 16984 (MICH, MO); Volcán Sanganguey,
Tepic, Walker 73H07 (NY). Oaxaca: Mpio. de San Jerón-
imo Coatlán, 38.6 km al SW de San Jerónimo Coatlán,
Campos 3467 (MEXU); Arroyo Limón, San Vincente, Put-
la, MacDougall H155 (F, NY); San Vincente, Putla,
MacDougall H156 (F, pto vicinity of Concordia, Pochu-
tla, Makrinus 551 (05); С afetal Concordia (Cerro ee
Reko 3642 (US m
Agua Colgada, San Ignacio, Montes
714 (US); San Juan, “шыл: 4118 (05); бака PR Ortega
5082 (BH, US); La Copetona, Arriba de Guadalupe de
Los Reyes, Tenorio 8364 (RSA, TEX); San Juan 14 km al
SE de San Ignacio, Tenorio 8441 (RSA).
Styrax radians is a relatively common and wide-
spread species, occurring at mid elevations along
the western escarpment of the Sierra Madre Occi-
dental and the southern slopes of the trans-Mexican
volcanic belt in Mexico. It appears to be most com-
mon in the states of Jalisco and Nayarit. The spe-
cies occurs in three disjunct areas: Sinaloa to
southern Jalisco, northern Guerrero and the south-
ern portion of the state of México, and southern
Oaxaca. The disjunction between western Mexico
and eastern Mexico is likely to be real rather than
an artifact of inadequate collecting because nu-
merous collections of Styrax, especially S. ramire-
zii, have been made in the intervening areas. Plants
occurring in the eastern part of the range have
leaves that are a little larger on average than those
Volume 84, Number 4
1997
Fritsch 747
Revision of Styrax
in the west, but they are otherwise essentially in-
distinguishable. Some populations in western Mex-
ico have extremely small flowers, but this is only
an extreme form of a completely continuous char-
acter and cannot be used for the designation of in-
fraspecific taxa.
Perkins apparently did not see any collections of
this species. Gonsoulin (1974) considered the col-
lections placed here in S. radians to fall within the
range of morphological variation exhibited by S. а ar-
pered anther sacs, non-prolonged connectives, and
stamen filaments that lack both auricles and stiff,
long-armed, upward-pointing hairs ventrally. Fur-
thermore, the ranges of the species overlap only in
Oaxaca. Styrax radians can be distinguished from
other sympatric species of Styrax by its evergreen
habit, drupaceous fruit, stellate pubescence
throughout, and corolla lobes that are united only
to the calyx margin. Pubescence on the lower lam-
inar surface, consisting of a short, dense tomentum
and a longer tomentum soft to the touch, is usually
a good character distinguishing sterile specimens
of S. radians from S. ramirezii; in S. ramirezti, the
lower laminar surface is usually at least somewhat
rough to the touch. However, the soft tomentum is
not always present in S. radians, and some speci-
mens of the stellate form of S. ramirezii approach
S. radians in this respect. In these cases, the lower
elevation at which S. radians occurs (400—1400 m,
usually below 1000 m) as compared to that of S.
ramirezii (770-3100 m, usually above 1500 m) can
aid in identification.
16. Styrax ramirezii Greenm., Proc. Amer.
Acad. Arts 34: . 1899. Styrax argenteus
var. ramirezii (Greenm.) Gonsoulin, Sida 5:
227. 1974. TYPE: Mexico. Morelos: Mountain
canyons above Cuernavaca, 15 May 1898, C.
G. Pringle 6848 (holotype, GH; isotypes, A,
BH, BR, F, MEXU, MO, NY, PH not seen,
POM, US, W). Figures 3, 4, 8, 57, 61—10.
Styrax micranthus Perkins, Bot. Jahrb. Syst. 31
1902. Styrax ramirezii var. micranthus (Perkins) Per-
kins, in Engl., Pflanzenr. IV. 241 (Heft 30): 54. 1907.
Styrax argenteus var. micranthus (Perkins) D'Arcy,
i 1 . Gard
x Temascaltepec, Ehrenberg 498 (ho-
lotype, B ama isotype,
Styrax chacal Perkins, Repert. Spec. Nov. Regni Veg.
> 1906. TYPE: Mexico. Oaxaca: Rincón, May
H. Galeotti 1687 (holotype, B destroyed; photo
of holotype, F, GH, US; isotypes, BR (2), G-DEL not
seen, NY, US,
Styrax Анар Perkins, Repert. Spec. Nov. Regni Veg.
2: 1906. Styrax ramirezii var. orizabensis (Per-
kins) ge in Engl., Pflanzenr. IV. 241 (Heft Ys
54. 1907. TYPE: Mexico. Veracruz (probably) o
Puebla: "Orale 1856, Botteri s.n. or 100 6
(holotype, B destroyed: isotypes, G-DC not seen, NY,
Styrax hinsonit Bullock, Bull. Misc. Inform. Kew 1936: 9.
1936 [“5. Hintoni”]. Styrax argenteus var. hintonii
(Bullock) Gonsoulin, Sida 5: 226. 1974. TYPE: Mex-
ico. México: Dto. de Temascaltepec, Calera, 19 Apr.
1933, G. B. Hinton 3798 (holotype, K; isotypes, F,
GH, MO, US)
Evergreen tree to 20(-25-27) m. Young twigs
short- to long-stellate-pubescent, or radiate-lepi-
dote, the hairs r e-brown, or dark
brown
outer layer of older twigs not exfoliating. Petiole 5—
25 mm long. Lamina chartaceous, (10.1—)11. ада
X 2.9-9.5(-10.5) ст, 1.64.4 times as long
wide, narrowly or broadly ovate-oblong, tities or
sometimes ovate, secondary veins 10-15;
rounded to acuminate, sometimes ien
through abortive tip development; base cuneate to
rounded, symmetric or slightly oblique; adaxially
glabrous except along the major veins; abaxially
covered with greenish white stellate hairs of vari-
e length or sometimes radiate scales, rarely
nearly glabrous, the surface slightly to moderately
rough or rarely soft to the touch, if soft then at least
the lower third of the calyx villous; veins glabrous
to densely tomentose; margin entire. Inflorescences
axillary or false-terminal, racemose or sometimes
paniculate, the racemes or panicle branches 2.5—
12 cm long, (2-)5-17-flowered. Lower pedicels 4—
11 mm long. Calyx (1.5-)3-6 X (3-)3.5-6.5 mm,
cupuliform, grayish green- or brown-radiate-lepi-
dote, or short- to long-stellate-pubescent; scales at
mid-calyx (when present) 0.05-0.17 mm diam.; ca-
lyx teeth 5, to 0.5 mm long, deltoid, the calyx mar-
gin slightly or distinctly concave between the teeth,
inner margin often glandular. Corolla white or pink,
8-)11-14 mm long, petals connate 1-3 mm be-
yond the calyx margin distally; corolla lobes 5, val-
vate in bud, 6-10 X 2-3 mm, non-overlapping,
slightly spreading, thickened, linear-deltoid, ves-
ture on the outer distal third of the lobes consisting
of stellate hairs. Free portion of stamen tube 1-3
mm long; distinct portion of filament 0.6-2.0(-2.6)
mm long, of equal width throughout, ventrally not
auriculate, white-stellate-pubescent at the some-
what incurved filament margins, glabrous or nearly
so on the face proximally, increasing in pubescence
distally, the hairs with arms to mm long; anthers
0—4.8 mm long, 1.9-5.3 times as long as the dis-
tinct portion of the filaments, the connectives not
or only slightly prolonged beyond the non-tapered
anther sacs. Ovary densely stellate-hirsute; style
~
з
748 Annals of
Missouri Aidan Garden
>
Е Fe
A
р и ры
ИТАР
ч;
t
it
РИК MU —
Figures 67-73. 67-70. Styrax ramirezii. —67. Flower, Pringle j i
. gle 9000 шр» —68. Four adjacent stamens, Hinton
et al. 7605 (TEX). —69. Leaf of small-scaled form, Prin, gle 9000 (MO). — "s 0. Leaf of Melius bat Koc nchez
еч e 71-73. Styrax warscewiczii, Davidse 24632 (MO). —11. Flowering nd —72. Flower. — 73. Four adjacent
Volume 84, Number 4
1997
Fritsch 749
Revision of Styrax
glabrous or pubescent at base only; stigma 0.3-0.5
mm wide. Drupe (7~)9-15 X 5-9 mm, ellipsoid;
wall irregularly and coarsely rugose on herbarium
specimens. Seed coat glabrous.
Southern Sinaloa and Durango south to western
Jalisco and northern Colima, east to northern Mo-
relos and Oaxaca, and from the southern portion of
the state of México south to central Guerrero; 770—
00 m; bosque mesofilo de montafia, Quercus-
Abies forests, Quercus-Pinus forests, tropical semi-
evergreen cloud forests, seasonally dry tropical
subdeciduous forests; found often along waterways,
sides of arroyos, humid canyons, deep riverbeds,
disturbed situations, on granitic rock, rich soil, or
sandy clay. Flowering: various times throughout
year; fruiting: various times throughout year.
Common names. Azajar (Jalisco, Niz et al. 290),
chilaquate (Morelos; Standley, 1924), escaramuza
(Jalisco, Mexía 1504), garrapata (Michoacán, Sharp
45499), jaboncillo colorado (Michoacán, 17 Dec.
1971, Rees s.n.), mamullo (Jalisco, Jardel et al.
214), mamuyo (Jalisco, Cuevas 1251, 1294, and
1366), memelita (Estado de México, Hinton et al.
6145), palo de casa (Jalisco, Cevallos et al. 72, Mu-
ñoz & Vázquez 3
Additional specimens examined. Stellate form (select-
ICO. Guerrero: Yesceros, Hinton
Arroyo Santa Gertrudis, Méxia T 504. (A, CAS, DS, F, GH,
MICH, MO, NY, US). México: Calera, Temascaltepec,
Hinton 7605 (A, DS, GH, LL, MICH, MO, RSA, TEX,
US). Michoacán: Pontezuelas, 20 km al E de Morelia,
Rzedowski 18371 (DS, MICH, NY, TEX, US). Nayarit:
base of Cerro Sanganguey, ca. 20 km SE of Tepic, Miller
3114 (MO, TEX).
Large-scaled form: MEXICO. Chiapas: Paraje Vo'bits,
Laughlin 408 (DS, LL, US); 2 km camino Cabecera =
acantan a pobablado Salinas, Zinacantan queda al W
pueblo, Ton 9329 (CAS, TEX, WIS). Oaxaca: Dto.
Ixtlán, 6 km NNE of La Luz, "Boyle 3841 (MO); 20 жы
NE of Тео Ап del Camino, Breedlove 59921 (CAS, NY);
Mpio. de San Jerónimo Coatlán, 36 km al SE de Cruz de
Honduras, Campos 4740 (CAS, MEXU); Dto. de Cuica-
Чап, паана "санне 2338 (Е); La Loma, Cuya-
450 (F); 13 mi. E of Тео ап, Croat
48197 (MO); Ма de Comaltepec, 5 Comaltepec, Her-
ndndez 322 (MEXU); Tonaguia, E 596B (F); Cam-
ino de Coyula a E Smith 347 (GH); Sierra Ma-
zateca, Plan de Guadalupe, Solheim 1397 (NY, WIS); 30
km al NE de Теош Ап, Tenorio 14351 (MEXU, MO, RSA,
TEX).
е form (selected оя МЕХІСО. Со-
lima: Mpio. de Minatitlan, Cerrito Вгейоѕо, Santana 2695
: along Майды Date о Hwy.
. Dur : the :3-
15 km toward El Salto pal Sinaloa, McVaugh 23503
). : 4
9059 (B, CAS, F, GH, LL, MO, NY, U,
Rancho El ште, 1.5 km adelante del бе шы, Flores
2680 (MEXU, TEX). México: 8 km al W de Cuernavaca,
Koch 7929 (CAS, F, MO, NY). Michoacán: San Felipe
Zitácuaro, Hinton 11831 (F, GH, LL, MO, NY, US, W).
orelos: Sierra de Tepoxtlan, Pringle 8023 (A, B, BH,
BR, F, GH, MEXU, MO, NY, POM, US, W). Sinaloa: 1
mi. S of El Palmito on Hwy. 40, 1 mi. W of hwy., Sanders
4405 (RSA, TEX).
Styrax ramirezii is one of the most common and
widespread species treated in this revision. It is
endemic to Mexico, occurring at relatively high el-
evations on the western slopes of the Sierra Madre
Occidental from Sinaloa to Jalisco, the southern
slopes of the trans-Mexican volcanic belt from Jal-
isco to Morelos, the Sierra Madre del Sur, the east-
ern escarpment of the Sierra Madre in Oaxaca, and
rarely in the mountains of Chiapas. The distribution
of this species shows a striking pattern along the
trans-Mexican volcanic belt, essentially occurring
only within a narrow band north of the Rfo Balsas
asin.
This species exhibits a great deal of morpholog-
ical variation across its range. The indumentum on
the calyx can be of long or short stellate hairs,
small or large radiate scales, or a mixture of these
types. The pubescence on the lower laminar surface
can consist of a thin to thick layer of long and/or
short stellate hairs or radiate scales, or it may be
nearly absent. The veins on the lower laminar sur-
face are usually visible through any pubescence,
but in most specimens from Oaxaca and Chiapas
they are not. Scattered orange-brown stellate hairs
or radiate scales sometimes occur on the calyx
Leaf and flower size varies a great deal throughout
the range of the species.
Differences between S. ramirezii and other spe-
cies of Styrax are addressed in the discussions un-
der S. argenteus, S. austromexicanus, S. magnus, S.
radians, and S. warscewiczit.
The extensive morphological variation exhibited
by this species is often not random, either in terms
of the correlations of characters or geographically,
and this complexity is likely responsible for the
contrasting treatments of S. ramirezii by Perkins
) and Gonsoulin (1974). Perkins (1907) de-
fined S. ramirezii relatively narrowly, although her
concept of this species was still broad enough to
maintain three varieties within it (vars. micranthus
and orizabensis, as well as the typical variety). She
considered S. cyathocalyx and S. warscewiczii to be
distinct species. Gonsoulin (1974), who had access
to more material, considered the variation in most
evergreen Styrax from Mexico as inconsequential
or highly intergrading, and thus circumscribed 5.
argenteus var. ramirezii to include seven previously
recognized taxa: 5. cyathocalyx, 5. micranthus, $.
orizabensis, S. polyanthus, S. polyneurus, S. rami-
rezii, and S. warscewiczii. Gonsoulin (1974) consid-
750
Annals of the
Missouri Botanical Garden
Table 2. Mean and standard error (SE) of lamina length/width ratio (= 1/w) and elevation for four groups within
Styrax ramirezii recognized on the basis of calyx trichome differences. Measurements were obtained
shape was estimated by dividing length by width (average of two largest leaves per: collection, one
imens. Lamina
from herbarium
spec
duplicate sampled per collection; sample list available from the author upon request). N — sample s
l/w Elevation (m)
Group N Mean SE N Mean SE
stellate form 43 2.19 0.06 39 1903 51
stellate/small-scaled
intermediates 10 2.49 0.12 9 1977 107
small-scaled form 79 2.82 0.04 64 2193 122
large-scaled form 12 2.82 0.11 7 2289 40
ered S. hintonii, a species described subsequent to
Perkins’s published research on the Styracaceae, to
be a variety of S. argenteus, and treated S. magnus
and 5. vulcanicola, also described after Perkins’s
publications, as synonyms of this variety.
The present revision follows Perkins’s (1907)
concept of Styrax ramirezii more closely than that
of Gonsoulin (1974). The species status of both S.
ramirezii and S. warscewiczii is recognized, as in
Perkins (1907). However, in contrast with Perkins
(1907), S. cyathocalyx, S. ramirezii var. micranthus,
and 5. ramirezii var. orizabensis are reduced to syn-
onymy under 5. ramirezii. The present treatment is
concordant with Gonsoulin (1974) in that S. cyath-
ocalyx, S. micranthus, and S. orizabensis are treated
as synonyms, but differs in that S. ramirezii and S.
warscewiczii are treated as species, and S. polyan-
thus and S. polyneurus are placed in synonymy un-
der S. argenteus and S. warscewiczii, respectively.
Furthermore, S. argenteus var. hintonii is placed
here in synonymy under S. ramirezii, and S. mag-
nus is treated as a distinct species; S. vulcanicola
is treated as a synonym of the latter name
The present circumscription of S. ramirezii com-
prises three fairly distinctive infraspecific entities.
One of these has scales at mid-calyx that are 0.15—
0.17 mm diam. This “large-scaled” form occurs
only in Oaxaca and Chiapas. ен form has
scales at mid-calyx that are 0.0. mm diam.
This “small-scaled” form is sf aera occurring
from Sinaloa to northern Morelos and the Sierra
Madre del Sur. In a third form the calyx is stellate-
villous on at least the lower third. This “stellate”
form is also widespread, but occurs entirely within
the range of the small-scaled form, from Nayarit to
the southern portion of the state of México and the
Sierra Madre del Sur. Specimens exhibiting calyx-
trichome morphology intermediate between that of
the small-scaled and stellate forms comprise about
10% of the collections of S. ramirezii; most often
they are from western Mexico and the state of Méx-
ico. Twig pubescence of the three forms generally
corresponds to the calyx pubescence: the twigs are
stellate-villous in the stellate form, and radiate-lep-
idote or rarely short-stellate-pubescent in the
small-scaled and large-scaled forms. Also, most
specimens of the large-scaled form possess a thick,
dense layer of stellate hairs or radiate scales,
whereas the other two forms have a thin layer of
stellate hairs or radiate scales. In the stellate form
this pubescence is often mixed with variable
amounts of larger, white, stellate and somewhat stiff
hairs. The small-scaled form is especially prevalent
in northern Morelos, where the stellate form does
not occur, and also extends farther north in western
Mexico than the stellate form. The large-scaled
form is apparently allopatric with respect to the oth-
er two forms. It is somewhat disjunctly distributed
in the mountains of Oaxaca and a small area west
of San Cristóbal de Las Casas. In this region S.
ramirezii is difficult to distinguish from S. warscew-
iczii if flowers are not available.
The stellate form of S. ramirezii in general pos-
sesses more broadly elliptic leaves and occurs at
lower elevations than small-scaled and large-
scaled forms (Table 2). Analysis of variance tests
showed significant differences among means of the
three groups (level = 0.05) for the length:width ra-
tio and elevation (P < 0.001 for both). The small-
scaled and large-scaled forms did not differ signif-
icantly for length:width ratio or elevation in a post
hoc comparison. To assess the distinctness of these
forms, canonical discriminant analysis was рег-
formed with the computer program SYSTAT (Systat,
Inc.). In this procedure, the proportion of individ-
uals classified correctly by the discriminant func-
tion reflects its ability to distinguish among groups
(see, e.g., Murrell, 1994). Mahalanobis distances
(Sneath & Sokal, 1973) among the groups were sig-
0.001] and the three groups were ide
rectly “%, (68.8%), Y, (55.6%), and 2%, (71.8%) of
Volume 84, Number 4
1997
Fritsch 751
Revision of Styrax
the time, respectively, on the basis of the discrim-
inant function. Thus, the discriminant function
showed that calyx trichome type tends to be asso-
ciated with leaf shape and elevation differences.
Despite this general correspondence, however,
formal recognition of the forms of S. ramirezii at the
infraspecific level is not warranted, mainly because
reliable key characters are lacking and there is a
relatively high proportion of small-scaled and stel-
late intermediate forms scattered throughout the
range of the stellate form. Furthermore, in Jalisco
and Nayarit intermediates seem to be of a different
nature from those in patty areas in that they possess
short-stellate hairs narrowly elliptic leaves
(e.g., Miller 3114, Téllez 10139); other intermedi-
ates seem to have the calyx and leaf characters
combined more or less independently (especially
collections near Nanchititla, in the state of México,
e.g., Bartholomew 2914). More detailed studies fo-
cusing on associations between morphological and
molecular variation, as well as differences in ele-
vation and habitat, would shed further light on the
ecology and population genetics of these forms.
I have placed Styrax cyathocalyx in synonymy
under 5. ramirezii with some hesitation. Perkins
(1907) maintained the two species on the basis of
differences in the size of the flowers: 1 cm long in
S. cyathocalyx and 1.5-2 cm long in S. ramirezii.
Styrax cyathocalyx was based solely on Galeotti
1687. The duplicates that I have seen of this col-
lection possess floral buds and no flowers at anthe-
sis; presumably Perkins had mature flowers avail-
able to her, because she described the corolla as 1
cm long, whereas the longest buds I could measure
were only 0.6 cm long. If Perkins’s observation is
correct, the corollas are unusually short for S. ra-
mirezii. The calyx (2 mm long) is also short, es-
pecially relative to the unusually long pedicels (14
mm). Perkins (1906) also described the style as
denticulate, but expressed uncertainty as to wheth-
er this was an artifact or real. In the specimens I
examined, the style margins are entire, i.e., in no
way do they differ from those of other specimens of
Styrax. A fruiting specimen from the same area in
Oaxaca (Liebmann 596B, Tonaguia, Oaxaca) shows
the same calyx and pedicel characters, and the fruit
is unusually small (7 X 5 mm) for S. ramirezii. It
is therefore possible that these collections represent
a distinct variety of 5. ramirezii, or even a distinct
species. More collections of evergreen Styrax from
Oaxaca, especially in flower, are needed before a
conclusive taxonomic decision regarding the status
of S. cyathocalyx can be reached. It would be es-
pecially helpful to rediscover populations at sites
where Galeotti and Liebmann collected.
Perkins (1902) listed only Ehrenberg 498 in the
protologue of S. micranthus; the location of original
material was not precisely indicated. From the pro-
tologue of 5. ramirezii var. micranthus in Perkins
(1907) it can be inferred that this collection was
deposited at B and/or W. Because no material of
Ehrenberg 498 occurs at W, however, it is reason-
able to assume that the holotype was at B, and has
been destroyed. The location of the original mate-
rial of 15 species described in the publication that
included S. cyathocalyx and S. orizabensis (Perkins,
1906) was indicated only vaguely as existing partly
at B, BR, G-BOIS, G-DC, P, and/or W. I have as-
sumed that the holotypes of these two species
names were located at B, because that is where
Perkins conducted her work; they have in all like-
lihood been destroyed. See the discussion under $.
glabrescens for information concerning the collec-
tion date of Galeotti 1687.
When Gonsoulin (1974) made the combination
Styrax argenteus var. ramirezii, he listed S. micran-
thus Perkins as a synonym. The latter name is the
basis of S. ramirezii var. micranthus (Perkins) Per-
kins (1907), making Gonsoulin's varietal name su-
perfluous. D'Arcy (1979) corrected Gonsoulin's er-
ror by making the combination S. argenteus var.
micranthus.
The protologue of Styrax ramirezi Greenm.
(Greenman, 1899) does not specify the holotype lo-
cation. The NY duplicate of Pringle 6848 is cited
as the holotype of S. ramirezii in Gonsoulin (1974).
However, Greenman worked at GH until 1899,
studied at the University of Berlin from 1899 to
1901, and returned to GH in 1902 (Woodson,
1951). On this basis, I consider it best to regard
the GH specimen as the holotype. Gonsoulin (1974)
also cited the MO duplicate of Hinton 3798 as the
holotype of S. hintonii. However, Bullock worked
at K, so the specimen there should be regarded as
the holotype.
17. Styrax steyermarkii P. W. Fritsch, sp. nov.
TYPE: Guatemala. Baja Verapaz: Unión Bar-
rios, E of km 154, 16 Apr. 1975, C. L. Lundell
& E. Contreras 19214 (holotype, CAS; iso-
types, LL, MEXU, MO, NY). Figures 6, 22-
24, 47.
Arbor sempervirens; lamina 2.1—3.0plo longior quam
latior, subtus pilis densis bir i instructa; calyx radia-
пеи dotus, squamis densis cinerascentibus-viridis et
pe dispersis ferrugineis ad medicalycem 0.130
mm кайшы. с. corolla lobis in aestivatione valvatis,
pilis albis-stellatis dense instructis, brachis pilorum usque
1.5 mm longis, illis pilorum apicem versus pro parte
752
Annals of the
Missouri Botanical Garden
maxima ordinatis apicem floralibus versus; antherae 4.5
mm longae; connectivum antherae loculos contractos man-
ifeste superans
Evergreen tree to 20 m. Young twigs grayish
green- to dark orange-brown-radiate-lepidote; outer
layer of older twigs not exfoliating. Petiole 9-20
mm long. Lamina chartaceous, 11-16 X 3.5-7.5
cm, 2.1-3.0 times as long as wide, elliptic, second-
ary veins 6-8; apex acute to abruptly short-acu-
minate; base cuneate, rarely subrounded, symmet-
ric or slightly oblique; adaxially glabrous except
ong the major veins; abaxially covered with min-
ute grayish green stellate hairs and sparsely scat-
tered, larger, grayish green or orange-brown stellate
hairs or radiate scales, the latter particularly prev-
alent near and on the veins; margin entire. Inflo-
rescences axillary or false-terminal, racemose or
paniculate, the racemes or panicle branches 2.54
cm long, (1-)4—7-flowered. Lower pedicels 5-8 mm
long. Calyx 2.5-3 X 4 mm, conical-cupuliform, in
fruit shallowly cupuliform, grayish green-radiate-
lepidote, sometimes with scattered orange-brown
radiate scales toward the base; scales at mid-calyx
0.13–0.25 mm diam.; calyx teeth 5, to 0.3 mm long,
linear, the calyx margin slightly concave between
the teeth, inner margin often glandular. Corolla
white, 12-15 mm long, petals connate пр to the
calyx margin distally; corolla lobes 5, valvate in
bud, 10-12 X 2 mm, non-overlapping, spreading,
slightly thickened, linear, vesture on the outer dis-
tal third of the lobes consisting of radiate scales.
Free portion of stamen tube 1 mm long; distinct
portion of filament 2-3 mm long, of equal width
throughout, ventrally with auricles bearing a dense
mass of stiff white stellate hairs with arms to 1.5
mm long, the hairs nearest the distal end of the
filament with arms predominantly pointing upward,
also pubescent below the auricles and at the sinus
between adjacent filaments; anthers 4.5 mm long,
the connectives distinctly prolonged beyond the ta-
pered anther sacs. Ovary densely short-stellate-pu-
bescent; style glabrous or pubescent at base only;
stigma 0.3 mm wide. Drupe 16-18 X 9-11 mm,
ellipsoid; wall irregularly and coarsely rugose on
herbarium specimens; fruiting calyx shallow, to 1.5
mm long. Seed coat glabrous.
In the mountains of Guatemala, departments of
Baja Verapaz, Huehuetenango, and Suchitepéquez,
and in the Cordillera Talamanca in Costa Rica;
1500-2600?) m; “in high forest on top of hill"
(Lundell 19214), riverbanks. Flowering: March,
June, July; fruiting: April.
Additional specimens examined. COSTA RICA. Pun-
tarenas: Buenos Aires, Ujarrás, Quebrada Dorora, Her-
тета 5866 (CAS, CR). GUATEMALA. Baja Verapaz:
Nifio Perdido, Lundell & Contreras 21219 (LL). Huehue-
tenango: Cerro Huitz, between Mimanhuitz and Yulhuitz,
Steyermark 48666 (F); Maxbal, ca. 17 mi. N of Barillas,
Sie tl 48848 (F). Suchitepéquez: Volcán Santa Cla-
a, Holdridge 2299 (US).
Styrax steyermarkii is rare in the high mountains
of Guatemala and Costa Rica. It is named after Jul-
ian А. Steyermark (1909-1988), prolific collector
of the Guatemalan flora, as well as that of Missouri
and of Venezuela, who first found the species in
1942.
This species is similar to Styrax glabratus by
virtue of its radiate-lepidote calyx and stiff, long-
armed, upward-pointing hairs on the stamen fila-
ments ventrally, but differs most obviously from this
species in its densely pubescent, as opposed to
nearly glabrous, lower laminar surface. Gonsoulin
(1974) mistook S. steyermarkii for S. argenteus var.
ramirezii, most likely because the lepidote calyx
and general appearance are very similar to that of
S. warscewiczii, which he had placed in synonymy
under variety ramirezii. However, the flower buds
on Steyermark 48666 clearly exhibit the stiff, long-
armed, upward-pointing hairs on the stamen fila-
ments ventrally, which are absent in S. warscewiczii,
and the leaves are larger.
The closest relatives of S. steyermarkii appear to
be several species from Andean South America.
These include S. buchtienii Sleumer, S. ferax J. F.
Macbr., S. heterotrichus Perkins, 5. leptactinosus
Cuatrec., S. pentlandianus Remy, S. socialis J. F.
Масђг., and S. subheterotrichus Herzog. The tax-
onomy and relationships of these species are un-
certain, but their types share with S. steyermarkii
the lepidote calyx and lower laminar surfaces that
possess either stellate hairs throughout or a mixture
of stellate and scale-like hairs. Styrax steyermarkii
can be distinguished from all these species by the
longer upward-pointing stellate hairs on the stamen
filaments ventrally, and by the possession of nar-
rower anther connectives. These species plus 5.
steyermarkii (= the S. steyermarkii group) appear
to be allied to the S. ovatus group, i.e., those spe-
cies with peltate or lacerate-margined scales cov-
ering the leaves, stems, and calyces (including 5.
conterminus, 8. cordatus, S. leprosus, S. ovatus, and
S. peruvianus). Some of the species in the 5. stey-
ermarkii group appear to be polymorphic for the
strongly peltate scales represented by the S. ovatus
group and the radiate-lepidote scales found within
most individuals of the S. steyermarkii group. А
clearer understanding of the taxonomy and phylog-
eny of these groups and their phylogenetic positions
Volume 84, Number 4
1997
Fritsch 753
Revision of Styrax
Bahia de Campeche
S
hl [ | |
| |
Caribbean
Pacific
¡Ocean
(90W
Figure 74. Distribution of Styrax warscewiczit.
within Styrax must await more detailed study of
Styrax in South America.
18. Styrax tuxtlensis P. W. Fritsch, sp. nov.
TYPE: Mexico. Veracruz: Los Tuxtlas Range,
Mpio. de Mecayapan, Volcán Santa Marta, 18
May 1986, J. V LaFrankie 1306 (holotype,
CAS; isotypes, GH, MO, TEX). Figures 25, 45,
46.
Arbor sempervirens; Hune lanci- ovata stellata-pubes-
cens, basi manifeste obli ignotae; calyx
stellatus-pubescens; fores ignoti; drupa angusta-ellipso-
idea, 15-18 х 8 mm
Evergreen tree to 10 m. Young twigs dark or-
ange-brown-stellate-pubescent; outer layer of older
twigs not exfoliating. Petiole 9-11 mm long. Lam-
ina chartaceous, 7.5-10 X 3.3-3.7 cm, lance-ovate
or lanceolate, secondary кеше ah apex, gradually
or abruptly short-acuminate; di y oblique
and slightly attenuate; € ИБ except
along the major veins; abaxially covered with olive-
gray stellate hairs and scattered orange-brown stel-
late hairs, the latter particularly prevalent near and
on the veins; margin entire. Inflorescence and flow-
ers unknown. Corolla presumably valvate in bud,
non-overlapping at anthesis. Infructescences axil-
ary or false-terminal, racemose, 3-5 cm long, with
about 3-6 peducles or peduncle scars. Peduncle
5-6 mm long. Fruiting calyx 2-3 mm high, shal-
lowly cupuliform, slightly concave between the 5
minute (0.25 mm long) teeth, the exterior surface
densely covered with appressed stellate or scale-
like hairs, inner calyx margin glandular. Drupe 15—
18 X 8 (if 1-seeded) or 12 (if 2-seeded) mm, nar-
row-ellipsoid; wall irregularly and coarsely rugose
on herbarium specimens. Seed coat glabrous.
Known only from the type locality on Уојсап
Santa Marta in the Tuxtlas range, Veracruz, Mexico;
800 m; steep volcanic slopes. Fruiting: May.
This species is known only from one fruiting col-
lection. It can be distinguished from all other spe-
cies treated here by the distinctly oblique leaf bas-
es. Styrax tuxtlensis is apparently geographically
isolated from all other Styrax species except S. gla-
brescens, from which it can be easily distinguished
by its evergreen habit and drupaceous fruit. More
collections of this unusual species, especially in
flower, are needed to better understand its taxono-
my and relationships.
19. Styrax warscewiezii Perkins, Bot. Jahrb.
Syst. 31: 480. 1902. TYPE: Costa Rica or Pan-
ama. “Veragua,” (1848?), Warscewicz 203 (ho-
lotype, B destroyed; photo of holotype, GH,
MICH, US; isotype, F). Figures 5, 9, 10, 71-
74.
Styrax polyneurus Perkins, in Donn. Sm., Bot. Gaz. 35: 5.
1903. TYPE: Costa Rica. [San José:] “In silvis ad
Copey" pee Feb. 1898 (protologue; “1896”?
U еп), Tonduz 11744 (holotype, В de-
stroyed; uiia CR, F, US).
Evergreen tree to 20(-30) m. Young twigs stel-
late-pubescent, the hairs usually orange-brown,
sometimes yellow-brown or dark brown, rarely
Annals of the
Missouri Botanical Garden
greenish gray mixed with brown hairs; outer layer
of older twigs not exfoliating. Petiole 5-20(-25) mm
long. Lamina chartaceous, 5.2-12.5(-16.0) X 1.6–
4.5(-5.4) cm, narrowly or broadly elliptic, second-
ary veins 9-12; apex sometimes subrounded or
acute, more often cuspidate to acuminate; base cu-
neate or less often subrounded, symmetric or slight-
ly oblique; adaxially glabrous except along the ma-
jor veins; abaxially thickly covered with greenish
white and often larger orange-brown stellate hairs,
the secondary and tertiary veins glabrous to densely
tomentose; margin entire. Inflorescences axillary or
false-terminal, racemose or paniculate, the racemes
or panicle branches 2-6 cm long, 2-21-flowered.
Lower pedicels 2.5-8(-11) mm long. Calyx 2.5-4
X 3-5 mm, broadly conical or cupuliform, grayish
green-radiate-lepidote, often with larger, scattered
orange-brown scales especially toward the base;
0. :
teeth 5, to 0.5 mm long, deltoid, the calyx margin
concave between the teeth, inner margin often glan-
dular. Corolla usually pink, sometimes white, some-
times white with pink lobes, 9-17 mm long, petals
connate (0—)2—6 mm beyond the calyx margin dis-
ul corolla lobes 5, valvate in bud, 6-13 x (1-)
5-2.5(-2.9) mm, non-overlapping, spreading or
€ erect, thickened, linear-deltoid, vesture on
the outer distal third of the lobes consisting of ra-
diate scales. Free portion of the stamen tube 1—3
mm long; distinct portion of filament (1.0—)1.3-2.5
mm long, of equal width throughout, ventrally not
auriculate, white-stellate-pubescent at the some-
what incurved filament margins, glabrous or nearly
so on the face proximally, increasing in pubescence
distally, the hairs with radiate arms to 0.3 mm long;
anthers 1.7—3.6 mm long, 1.1–2.2 times as long as
the distinct portion of the filaments, the connectiv
not or only slightly prolonged beyond the non-ta-
pered anther sacs. Ovary densely а
style glabrous or pubescent at base only; stigm
0.3-0.4 mm wide. Drupe 7-12(-15) X 4-168)
mm, ellipsoid; wall irregularly and coarsely rugose
on herbarium specimens. Seed coat glabrous.
Far western Panama north to western Chiapas,
Mexico, disjunct to Guerrero in the Sierra Madre
del Sur. One specimen collected from San Luis Po-
tosí in the Sierra Madre Oriental, Mexico, is of un-
certain status but resembles S. warscewiczii; most
common from Chiapas to Guatemala, and in the
Costa Rican cordillera; 1100—3200 m; cloud for-
ests, Quercus-Pinus forests (Guerrero, Chiapas),
lower montane wet forests, montane rainforests, dry
montane forests, pastures, savannas; found in pri-
mary or secondary growth, in valleys and along
streams, in the understory to unshaded, open situ-
ations, and roadsides, on rock outcrops, lava flows,
or heavy clay loam. Flowering: apparently through-
out the year; fruiting: apparently throughout the
year.
Common names. Azulillo (Costa Rica, Madriz 9),
el comün (El Salvador, Allen 6882), estoraque
т 5а 58420), езїогасе (ин
Steyermark 32827), estorach (Guatemala, La
115), oreja - уепадо (El Salvador, 4 June ges
Reyna s.n. p.p.), sapuyulillo de montafia (Guate-
mala, Steyermark 33315). Standley (1938) has list-
ed under S. polyneurus and S. warscewiczii the Cos-
ta Rican common names aguacate ascá, ascá
laurel, quiquicirrí, and resina, all of which likely
apply to the concept of S. warscewiczii presented
here.
Uses. Firewood (Guatemala, Steyermark 33315).
cted s specimens examined. COSTA RICA. "ena
d
ana
Ре of Volcán Barba, Hatheway 1258 (CR, DS, DUKE,
F, GH, US). Limón: Valle de Silencio, Davidse et al.
pb (CAS, CR, DUKE, MO, NY). Puntarenas: Cantón
de Buenos Aires, Ujarrás, El Carmen, Cerro Betsú, /. Cha-
cón 494 (CR, F, MO); Sitio Cotón (Cotonsito), along the
road to Sitio Coto Brus, Davidse 24632 (CR, MO). San
José: overlooking the village of Boquete and about 20 km
N of San Isidro del General, Wilbur 21135 (AAU, BR,
CAS, CR, DUKE, F, LL, MICH, MO, NY, TEX, US). EL
SALVADOR. Chalatenango: Los Esesmiles, E of La Pal-
ma, Allen & van Severen 7314 (F, GH, LL, NY, US). Santa
Ana: Cerro Verde, Allen 6853 (F, US); northern slopes of
Volcán Santa Ana, Allen 6882 (F, US); Cerro Montecristo,
4 June 1976, Reyna s.n. p.p. (CR). GUATEMALA. Alta
Verapaz: Río Frío, entre Tactic y Santa Cruz, Molina &
Molina 12236 (F, NY, US). Baja Verapaz: Sierra de las
Minas E of Chilascó, Lamb 115 (F); Niño Perdido, Lundell
& Contreras 20452 (CAS, LL, MEXU, MO, NY). El
, MICH, U). Guatemala: upper slopes
of Volcán Pacaya, Harmon 4160 (MO, NY); Volcán de
mark 4844] (A, F). Jalapa: Volcán Jumay, N of Jalapa,
Steyermark 32340 (F, US); Monataña Miramundo between
rmark 32827 (A, F).
№,
hills of Finca Carmona, SE of Antigua, Standley 63
arcos: Volcán Tajumulco,
o, Steyermark 36712 (F). — Volcán Atitlán,
47397 (A tepéque
? ,
: Volcán Santa
arán, Mol 27
Yashse, Molina 6522 У (Е GH, US). La Paz: cn del
————— MÀ
Volume 84, Number 4
1997
Fritsch 755
Revision of Styrax
Huis, Molina 6460 (F, US). Morazán: San Juancito, Wil-
liams & Molina 13318 (F, GH, MO, US). tip cho: trail
between La Chorrera campsite and 1900 m camp on ridge,
Thomas et al. 474 (MO). MEXICO. Gane Mpio. de
DUKE, F, MEXU, MICH, NY); Barranca Honda near Sil-
tepec, Matuda 4135 (A, DS, LL, MEXU, MICH, NY, US).
Guerrero: Piedra Ancha, Puerto eo Hinton et al.
14742 (F, GH, LL, MO, NY, RSA, US, W). Oaxaca: Dto.
de Сшеа ап, La Loma азн Conzatti 3465 (US).
an Luis Potosí: Río de las Gallinas, Rascon, Purpus
5317 (CAS, F. GH, MO, NY, US). NICARAGUA. Jino-
tega: along Hwy. 3, ca. 1 km NW of La Fundadora en-
trance, Miller & Stevens 1402 (DUKE, MEXU, MO, NY).
atagalpa: Cerro El Picacho above Hotel Santa María
de Ostuma, Nee 27636 (DUKE, LL, MO, NY). PANAMA.
Bocas del Toro: Valle de Silencio, Antonio 1640 (MO).
Chiriquí: along road to Bambito, Allen 4851 (BR, F, GH,
MICH, MO, NY, U, US).
Styrax warscewiczii is а widespread, relatively
high-elevation species, occurring from Guerrero,
Mexico, to northern Panama. A collection of Styrax
from San Luis Potosí may be this species as well
(see below). In Mesoamerica, it is the species that
generally replaces the equally common S. argenteus
ve 2000 m. It is most common in Chiapas and
Central America; in Mexico west of Chiapas S. war-
scewiczii has been collected only once, in the Sierra
Madre del Sur, Guerrero.
Gonsoulin (1974) placed this species in synon-
ymy under 5. argenteus var. гатігеги. His annota-
tions of specimens referred to here as S. warscew-
six as "intermediates" of varieties argenteus or ra-
mirezii. Styrax warscewiczii is easily distinguished
from S. argenteus as circumscribed in the present
revision by the lack of both auricles and stiff, long-
armed, upward-pointing hairs on the stamen fila-
ments ventrally, as well as by the presence of ra-
diate scales rather than stellate hairs on the calyx.
Styrax warscewiczii and S. ramirezii are morpho-
logically similar in many respects. Nonetheless,
they exhibit enough differences throughout their
ranges to justify the recognition of these two entities
as species, although they are likely to be closest
relatives. They can be diagnostically differentiated
from each other by vesture on the outer distal third
of the corolla lobes, which consists of stellate hairs
in S. ramirezii and radiate scales in S. warscewiczii.
The two species can be further distinguished by the
other characters in couplet 18 of the key: the an-
thers are longer and the ratio of anther length to
the length of the distinct portion of the filaments is
higher in S. ramirezii. Besides the characters listed
in the key, S. warscewiczii differs from S. ramirezit
by a tendency toward smaller leaves, smaller flow-
ers, and shorter leaf internodes. However, these
characters exhibit some degree of overlap.
The two species are most similar in Oaxaca and
Chiapas, where the range of the large-scaled form
of S. ramirezii overlaps the range of S. warscewiczii.
There are a number of morphological patterns ex-
hibited by S. warscewiczii and S. ramirezii that sug-
gest bidirectional introgression between the two
species in this region. First, the sizes of the calyx
scales are equivalent in S. warscewiczii and the
large-scaled form of S. ramirezii, whereas in the
other forms of S. ramirezii the calyx pubescence
consists of smaller scales or stellate hairs. Second,
the large-scaled form of S. ramirezii exhibits a ten-
dency toward the anther length and anther-filament
ratio typical of S. warscewiczii, whereas in the other
two forms of S. ramirezii these two characters clear-
ly distinguish the species. Third, north of Honduras
and especially in Chiapas many individuals of S.
warscewiczii possess larger leaves and longer leaf
internodes than is typical for the species, and thus
the measurements of these characters approach or
overlap those typical for S. ramirezii.
these two species are in fact closest relatives,
it may be difficult to ascertain whether the patterns
of morphological variation displayed in the zone of
sympatry are a result of primary speciation or sec-
ondary contact. А phylogenetic estimate that re-
solves the sister-group status of these species would
make the pursuit of this issue practical. Further
investigation of this problem should focus on the
area of geographic overlap, especially where the
two species apparently grow together in the area
west of San Cristóbal de Las Casas in Chiapas.
Several other characters are variable across the
range of S. warscewiczii. The southernmost popu-
lations (Costa Rica, Panama) tend to possess a
higher percentage of brown versus grayish green
radiate scales than those farther north. In Panama
the radiate scales have somewhat longer arms (e.g.,
Schmalzel 1608). Character variation in some col-
lections of S. warscewiczii (i.e., Haber 357, Haber
2917, Haber 5541) in extreme northern Puntarenas
Province, Costa Rica (Monteverde and San Luis),
suggests hybridization with 5. argenteus. These col-
lections have a mixture of grayish green radiate
scales and nearly stellate hairs on the calyx. They
were found at relatively low elevation and may grow
with typical S. argenteus, which is known from the
general area where these collections were made.
The flowers do not deviate from the range of vari-
ation within S. warscewiczii as defined here. Addi-
tional study of individuals in this area is needed to
establish whether these collections are hybrids or
an unusual form of S. warscewiczii.
756
Annals of the
Missouri Botanical Garden
Two specimens from San José Province, Costa
Rica (San Gerardo de Dota), here assigned to S.
warscewiczii, have nearly glabrous abaxial leaf sur-
faces (Kappelle MK118 and MK1530). These are
the only specimens of this species that show this
type of variation. The specimens are sterile. It is
possible that they are hybrids between S. warscew-
iczii and S. glabrescens, the latter of which grows
in the same area (cf. Kappelle MK23), but this is
unlikely given the relatively great evolutionary dis-
tance between the two species. It is also possible
that they represent an undescribed taxon
A single collection of Styrax Бана to the
Mise evergreen group has been made in the
state of San Luis Potosí, Mexico € ca
Brandegee referred this collection to S. p urus
(= S. warscewiczii); Standley (1924) vi that it
might represent an undescribed species. The leaves
of this collection are narrowly elliptic with the apex
broadly acute or obtuse, and the leaf internodes are
unusually short. In these respects it matches no
other Styrax collection precisely. It is not a variant
of S. lanceolatus, which occurs both north and
south of the locality at which Purpus made his col-
lection, because that species has a stellate-pubes-
cent calyx and distinctive lanceolate leaves. The
Purpus collection resembles S. warscewiczii in its
large radiate scales on the calyx and small leaves
(relative to S. ramirezii), so I have tentatively
placed it in S. warscewiczii. However, the fact that
this collection is so far removed geographically
from any other collection of S. warscewiczii makes
this placement uncertain. Until more material is
available from the area, especially in flower, it will
remain unclear whether Purpus 5317 represents (1)
an unusual form of S. warscewiczii, (2) a distinct
infraspecific taxon of S. warscewiczii, (3) an inter-
mediate between S. warscewiczii and S. ramirezii,
or (4) a new species.
EXCLUDED NAMES
Styrax diplotrichus Diels, Notizbl. Bot. Сам. Berlin-
Dahlem 13: 508. 1937. TYPE: Mexico. Guer-
rero: Sierra Madre del Sur, Gipfel des Telpit-
zahua, Jena 458 (holotype, B destroyed).
I have located no authentic material referable to
this name. If the collection cited by Diels does not
actually represent a new species, then on a geo-
graphical basis it could be a specimen of either S.
austromexicanus or S. ramirezii. The description in-
dicates that the leaves of the specimen are 7-9 cm
wide, thus excluding S. austromexicanus from con-
sideration. The description, although quite general,
is not inconsistent with the characters of S. rami-
rezii. However, there is sufficient uncertainty to pre-
clude the placement of this name in synonymy.
Styrax — — & Bonpl.) Miers, J. Linn.
oc., Bot. 1880. — Symplocos limon-
cillo die is re pl.
Literature Cited
Blankinship, J. W. 1907. Plantae ier idm те part III.
Rep. (Annual) Missouri Bot. Gard. 1 23.
Carranza, E. 1993. Una nueva о. A = Lyrax argen-
teus (Styracaceae) del NE de Querétaro (Mexico). Acta
Bot. Mex 5—40.
a nueva variedad de Styrax TET
(Sivratacesc} del estado de Oaxaca, Mexico. Acta Bot
Mex. 36: 15-19
Cory, V. L. 1943. The genus Styrax in central and western
Texas. Madroño 7: 110-115.
Cox, P. 1987. Chasing the wild Texas snowbells. Center
Pl. Conservation 2: 1,
Crawford, D. J. 1983. уеде and systematic infer-
ences from electrophoretic studies. /n S. O. Tanksley &
T. J. Orton (editors), Isozymes in Plant Genetics de
Breeding, part A, 257-287. Elsevier, Amsterdam
. Enzyme electrophoresis and plant аі
ансв. Рр; 146—164 in D. E. Soltis & Р. 5. Soltis (edi-
tors), Isozymes in Plant Biology. Dioscorides Press,
Portland.
Cronquist, A. 1981. An Integrated System of Classification
of Flowering Plants. Columbia Univ. Press, New Yor
. G. 1979. Flora ^ Panama. Styracaceae. Ann.
Missouri Bot. Gard. 66: 165-172.
Dahlgren, R. T. 1980. A bird: system of dee m
of the angiosperms. Bot. J. Linn. Soc. 80: 9
Engelmann, С. & A. Gray. 1845. Plantae ERR
Ап enumeration of F. Lindeimer' collection of Texan
plants, with remarks, and descriptions of new species,
etc. Proc. Boston Soc. Nat. Hist. 5: 210-264.
Fenby, S. S. & R. C. Gastil. 1991. Geologic tectonic map,
Gulf and Peninsular Province of the Californias, plate
10. In J. P. Dauphin & B. R. T. Simoneit (editors), The
Gulf and Peninsular Province of the Californias. Amer-
ican Association of Petroleum Geologists, Tulsa, Okla-
oma.
Fritsch, P. 1995. Systematic and biogeographic studies i
the genus Styrax. Unpublished Ph.D. Dissertation
Claremont ee School, Claremont, Californ
a. Isozyme analysis of an nn dis-
jiincts within Styrax (Styracaceae): Implications for the
Madrean-Tethyan hypothesis. Amer. J. Bot. 83: 342
996b. Population structuring and patterns of
morphological variation in Californian Styrax (Styraca-
ceae). Aliso 14: 205-218.
Gonsoulin, G. J. 1974. A revision of Styrax Siac
in North America, Central America, and the Caribbean
Sida 5: 191-2
Gottlieb, L. D. 1977. ia نوو сее and plant
systematics. Ann. Missouri Bot. Gard. 64: 161-180.
Gray, A. 1850. Plantae беткі, й І. Ап ас-
і е by Е Lindheimer
апа 1847—1848
new species, еіс. Proc. Воз
250.
ston Soc. Nat Hist. 6: 141-
Volume 84, Number 4
1997
Fritsch 757
Revision of Styrax
Grayum, M. H. & G. C. de Nevers. 1988. New and rare
understory palms from the Península de Osa, Costa
Rica, and adjacent regions. Principes 32: 101-114.
Сібейлай, . M. . Some new species, extended rang-
es, and newly noted Bere among the Mexican
phanerogams. Proc. cad. Arts 34: 568-578
A
Hartshorn, G. S. 1983. Pisces introduction: Pp. 118-15
in D. H. Janzen (editor), Costa Rican Natural History.
Univ. Chicago Press, Chicago-London.
Hinton, J. & J. Rzedowski. 1972. George B. Hinton, col-
ector Pe -— in southwestern Mexico. J. Arnold Ar-
bor. 5 –181.
Hultén, E pens Бх оп the Flora of Alaska апа
Yukon. Ark. Bot., n.s. 7: 1-147.
López Ramos, E. 1983. Geologia de México, ed. За, vo
3. le n.; Secretaría de educación, Pública No. 6402/19],
Mexi :
Lucas, G. рч H. Synge. 1978. The IUCN Red Data Book.
Unwin Brothers, The Gresham Press, Old Woking, Sur-
rey.
McKelvey, S. D. 1955. Botanical Exploration of the Trans-
Mississippi West. Arnold Arboretum, Jamaica Plain,
Massachusetts.
McVaugh, R. 1972. Botanical exploration in Nueva Gali-
cia, Mexico from 1790 to Ei e present time. Contr. Univ.
inane? Herb. 9: 205-357.
1978. Galeotti’s bonito orks in Mexico: The
вена of his collections and a brief itinerary. Contr.
Univ. арена: Herb. 11:
Miers, J. 1859. On the natural order Styraceae, as distin-
guished fon the Symplocaceae. Ann. Mag. Nat. Hist.,
ser. 3, 3: 394—404.
Morton, C. M., M. W. Chase, K. A. Kron & S. M. Swensen.
1997. A Ras evaluation of the monophyly of the
order org iid upon rbcL sequence data. Syst.
Bot. 21: 567-5
Murrell, Z. E. ns Dwarf dogwoods: Intermediacy and
the morphological er, Syst. Bot. 19: 539-556.
Nicolson, D. H. & G. C. Steyskal. 1976. The masculine
gender of the generic name Linnaeus (Styraca-
ceae). Taxon 25: 581-587
Pacheco, L. 1983. Styracaceae. Flora de Veracruz fasc.
32: 1-11. Instituto ¿a nal de Ме tee une sobre
Recursos Bióticos, Xalapa, Veracruz, Mexico.
Perkins, J. 1902. — zur Kenntnis ia Styracaceae.
Bot. Jahrb. Syst. 31: 478—488.
1906. Styracaceae io t novae. Repert.
st Ми. кеч Мег. 2: 1
7. Styracaceae. = A Enger (editor), Das
Estes IV. 241 (Heft 30). 1-11
Smith, J. T. 1991. Cenozoic marine маса and paleo-
geography of the Gulf of California. Pp. 637—666 in
‚ Dauphin & B. R. T. Simoneit (editors), The Gulf nd
WEB Province of the Californias. American As-
sociation of Petroleum Geologists, Tulsa, Oklahoma.
Sneath, H. A. & R. R. Sokal. 1973. Numerical Taxonomy.
W. H. Freeman, San Francisco.
Stafleu, м = & R. S. Cowan. 1988. a. literature,
» 7: W-Z. Regnum Veg. 116: 1-653.
знање P = 1924. Styracaceae. Trees and shrubs of
Mexico. Contr. U.S. Natl. Herb. 23: 1129-1130.
. 1938. Styracaceae. In Flora of Costa Rica. Publ.
Field Mus. Nat. Hist., Bot. Ser. 18: 913-915.
— —— & L. O. Williams. 1967. —— In Flora of
ну, aca Bot. 24(3): 2
Steenis, C. G. G. J. van. 1932. The снови ни of Neth-
erlands India. Bull. Jard. Bot. Buitenzorg sér. 3, 12:
-212.
Sugden, E. A. 1986. Anthecology and pollinator efficacy
га Styrax officinale subsp. redivivum (Styracaceae).
mer. J. Bot. 73: 919-930.
Yat A. 1980. Outline of the classification of flow-
ering 0 (Magnoliophyta). Bot. Rev. (Lancaster) 46:
ga hg Magnoliophytorum. Nauka, Len-
in od: (Я Russi
Poach i 978. ‘New subspecific арта ни for
southern Califa plas. Aliso 9: 1
ae An updated rr nie ^ the flowering
plants: Aliso 13: 365-389.
Tiffney, B. H. 1985. The Eocene north Atlantic land
bridge: Its importance in Tertiary and modern phyto-
С е ће погіћегп teen е J. Arnold Arbor.
Torrey, J: sans On the Е californica. Smith-
r. 1951. idi More om 2 Sanne
1951). Ann: cd Bot. Gard. 38: 95-100
Numerical List of Styrax Species
argenteus C. Pres
wn
jaliscanus S. Watson
lanceolatus P. W. Fritsch
magnus Lundell
nicaraguensis P. W. Fritsch subsp. nicaraguensis
nicaraguensis subsp. ellipsoidalis P. W. Fritsch
12. S. panamensis Standl
To © ооо: слу фә оз
dH doe.
Y
3.
14a. s Paene Реде ex Torr. subsp. platanifol-
14b. 5, сны subsp. mollis Р. W. Frit
14c. S. platanifolius subsp. stellatus (Cory) РА W. Fritsch
14d. S. platanifolius subsp. texanus (Co ry) P. W. dx
14e. S. Vetus Prat subsp. youngiae (Cory) Р.
sch
radians P. W. er
amirezii Gre
Dii P E Fritsch
tuxtlensis P. W. Fritsch
warscewiczii Perkins
pat
ла:
RIDE
Index to Exsiccatae. Specimens are listed alphabetically
by collector. Numbers in parentheses correspond to I
on the label is
the d e qeda to belong to collections with-
out number when followin
(i.e., А and Sharp collections) ! кае duplicate of
collections where transcription errors apparently occurred.
Abbots R. 0. 113 (16); : 223 (16). Aguilar, J. I. 186 (1);
214 (1); 456 (1); 758 (1); 1335 = 9). Aguilar, R. 1126 (6);
1159 (19). Aguirre B., C. 100-51 (15). Alexander, E. J.
1214 (10). Allen, P. H. 4851 ay 6853 (19); 6882 (19);
7184 (3); 7314 (19). et F. 2400 (19); 3044 (19);
3291 (19); 3761 (19); 5642 (6). Anderson, W. R. 6109
(15). Andrie 513 (6). Sena T. 1640 (19). еы М.
758
Annals of the
Missouri Botanical Garden
40 (1). —Ó M. 363 ). Arriaga, R. 176 A
Arsene, B. G. 2841 (16); E аб, 8472 (16). Atwood
J. A316 (Па). Avendafio R., 5. 238 (6); 258 ae 2 1 6;
353 (6); 793 (6). ^ A. de 742 (2). Ayala, M
6). Barkley, F. A. 7440 (16). Barthol-
16). Bauml, J. 531 (1). Bello, E. 932
qua; vem Ln 2158 (6); cM (6); 5310 (6). Berlandier,
Boege, W. 1765 (16). Bossé, G. 8222 (1); 8335
eri 1006 (16). Boutin, F. 2878 (4). Boyle, B.
1 (16). Brave 2015 (19). Breedlove, D. E. 6366 (1);
6439 (1); 7619 (1); 8917 (6); 10188 (6); 14085 (1); 15588
(4); 16893 (4); 18269 (4); 18414 (4); 18584 (4); 20577
(1); 20635 (1); 21827 (19); 21943 (1); 22841 (19); 23352
(1); 24991 (6); 25152 (1); 25292 (19); 27646 (19); 28186
(6); 28487 (1); 29307 (1); 29696 (1); 30187 (6); 30255
(1); 30989 (19); 31045 (10); 31106 (10); 31368 (19);
3
=
2394 (10); 32773 (1); 33804 (1); 34377 (6); 39600 (10);
ју 0% A ۹ О е
`
41611 (19); 42421 (1); 45097 (8); 45657 (16); 46223 (10);
46979 (1); 48026 (1); 49750 (10); 50020 (19); 50206 (1);
1); 51000 (1); 52664 (6); 52994 (19); 55749 (19);
57053 (10); 58137 (6); 59921 (16); 61642 (8); 61786 (16);
64233 (16); 64437 (16); 64929 (2); 64955 (16); 65176
(2); 67582 (6); 68742 (10); 68828 (6); 68928 (3); 69019
(6); 69083 (19). Brenes, A. М. 180 (6); 298 (6); 345 [258]
(1); 3683 (1); 3772 (19); 3971 (19); 3986 (6); 4462 (19);
5403 (19); 5506 (19); 6079 (6); 6380 (19); 6410 (1); 6464
а); 6134 (б); p 6; 11458 (19); 13599 (6); 17008 (1).
ock, S. Н. 2024 (15). Burger, W. 7390 (19); 7707 (19);
7713 (19; 12004 (6); 12064 (19); 12153 (6); 12400 (6);
1247
Cabr r E. 1404 (1); 1405 (1); 1988 (1); 3805 (10).
6
» 17816 (8); 17822 (8);
eim
7(1
5 0) 4740 (б)
SS
X
Cárdenas, A. L. 79 (15). Сабин; м ye 19);
(6). Carranza, E., 4033 (9); 4465 (9); 4519 00 Conil
C., G. 1573 (6); 2203 (6). Cedillo T., R. 6 ; “4
(1); 2356 (6). Cevallos, J. 72 (16). под: \ю 09
Chacón, I. A. 1562 (19); 1577 (1). Chavarría, M. M.
(19). Chávez, С. 521 (19); 540 (Па). Cházaro В., M. Е
(6); 1511 (6); 4675 (6); 5485 (8). Cházaro, М. 2685 (6);
2691 (6); 3759 (6); 6993 (8). Chinchilla, M. 183 (6).
Chrysler, М. A. 5590 (19). Cochrane, Т. 5. ое
11942 (15); 12047 (15); 12546 (16). tetas E. 4915
(19); 4924 (19); 5102 Mim 11029 (19). Conzatti, C. 2338
(16); 2450 Ce 3465 e
3 (140)
о
—
È
sr
~
(14d); 34938
; 38934 (14d);
42659 (14d); 2664 ; 42666 (14d);
42668 (14d); 42669 (14d); 42673 (14d); 42677 (14d);
42678 (14d); 42953 (14d); 42956 a 49179 (14d);
49453 (14c); 49454 (14c). Cowan, C. P. 4612 (19). Croat,
34830 (19); 34981 (19); 37190 аз; 45262. (15);
48197 (16); 64211 (6). Cruz С., R. 534 (16). Cuevas G.,
R. 709 (15); 1251 (16); 1294 (16); 1366 (16); 1876 (15);
1927 (16); 2213 (16); 2391 (15); 2423 ee 2864 (16);
2890 e 4032 = 6). eai G. 461
9992 (19). aea M. 59 (3);
72 (6); "119 (6); 142 aged Davidse, G. 1511 (19); 1654 (19);
1656 (6); 10164 (19); 10216 (19); 24017 (19); 24409 (19);
24511 (19); 24632 (19); 26146 (19); 28381 (6); 28554
iris 28606 (19); 28668 а 34298 (6). Davidson, М. Е.
6 (19). Delgado 5., A. 19). de Nevers, G. 6018
d 9); 6848 (13). De Niz E ps 23 (15); 290 (16). Díaz B.,
H. 2106 n 2265 (16); 3656 es —€ Ate Díaz Lane;
C. L. 3592 (16). Dieterle, bee 3 (15). Diggs, G.
3804 d Domínguez С., о dion J. 574 (6).
. 1910 (14a); vers (14d). Dryer, V. J. 1261 (6);
1420 ids 1575 ря — J. D. 454 (1); 7006 (19).
wen d J. E. 716 tene B. P-290 (1). Eh-
renberg 498 (16). Enric G. M. 60 (16). Erazo, M. 63
063 (16); 1783 (16). Espinoza, R.
128 (1); 169 (1; 234 0 (1); 1254 (19). Eugenio, J.
388 (1). Evans, R. 1726
Fairey, J. کا وپ (9); T39M-3.5h-031992
(6). Fernández, А. 807 (6); 1444 (19). Ferrera B., A. 176
(1). Flores C., J. 315 (16). Flores F., G. 793 (16); 1907
(15); 2498 (15). Flores M., A. 2680 (16). Flores-Franco,
G. 3344 (15). Folsom, J. : 6047 (19). Fonseca J., R. M.
163 (2). Frankie, G. W. 239 (1). Fuentes O., M. 56 (16);
90 (16). Fuentes, Z. 192 (11a).
Galeotti, H. 1687 (16); 2851 (6); 2852 (6). Gamboa, B.
71 (19). García, A. R. 1902 (19). Garwood, N. 334 (19);
761 (1); 1404 (6). Gentry, A. 13797 (12?); 14073 (12?).
Gentry, H. S. 4420 (4); 5164 (15); 6244 (4); 10485 (15);
10811 (15); 10863 (15). Ghiesbreght, A. B. 620 (1); 811
(19). Gilly, C. L. 64 (6). Glassman, S. F. 1956 (19). Gliess-
man, S. R. CH-54 (19). Gold 129 (16). Goldman, E. A.
872 (1). Gómez, L. D. 20777 (1). Gómez-Laurito, J. 8012
(6); 9681 (19); 11345 (6); 11383 (19); 11445 (6). Gómez-
Pompa, A. 793 (6). Gonsoulin, G. 1287 (14a); 1288 (14a);
1289 (14a); 1305 (14a); 1306 (14a); 1307 (14a); 1308
(14a). González, J. 108 (1); 340 d 627 (19). González
L., M. 78 (16). González Q., L. 3347 (16). González-Es-
pinosa, M. 1148 (19); 1279 Aor 1659 (19). González-
Ману 4053 (1). Grayum, Sd bos 2 ym Сен E
5 (8). Guerrero 322 (8). "aes J. J. 649 (16).
а С. 124 (6). rt M. 239 (16). pit M.,
R. 2516 (16).
aber, W. 230 (11b); 357 (19); 363 (6); 404 (1); 2917
(19); 3173 (13); 3533 (1); 4330 (6); 5541 (19); 6116 (1);
7353 (6); 7526 (19); 9318 ыл 10631 (6); 11118 (19);
11381 (1). Наепке, T. s.n. . Hagen, С. von
1253 (1). Halbinger, C. 220 (16). Наши, В. 5734 (19);
6729 (19); 13812 (6); 15271 (6); 19068 (19); 20076 (6).
Bese B. 1479 (16). H 3056 (1); 3461 (1);
60 (19). Hartshorn, G. S. 1075 (19) 1805 (1); 1894
ен 2119 (6). Hartweg 489 (6). Н . 1258
ise Hazlett, D. 18-22 Nov. 1974 (6) 667 (1). Heller, С.
234 (6). Helmrich, Н. 1690 (6). Henrickson, J. 11499
(14е); 11561 (14е). Непгу, М. С. 6557 (14а); 6560 (14с).
Hernandez, G., L. 322 (16). Herrera, С. 297 (1); 472 (6);
ridge, L. R. 2299 (17); 2338 (6). Инеш, M., M. 233 (16
Iltis, H. H. 337 (16); 389 (15); 459 (15); 2516 (16). |
INBio 16 (13). II InBio 164 (11b).
Janzen, D. H. 12108 (1). vaa E. 214 (16). Jermy, G.
05 (14a). Jiménez M., A. 345 (1); 1285 (1); 1974 (19);
2305 (19); 3511 (1); 3653 (19); 3983 (19). Jiménez, О.
= 2 (0 Jiménez, Q. wes 2 6 аз (6); 749 (1). терете
99 (19). Јоһпѕоп, Ј. С. 5 4c). Johnston, М. С. 7304
dm 7408 (9); 7428 (ad. am (14e); 11925 ле;
12845 (9). Juárez, L. С. 62 (6). Judziewicz, E. J. 4
m7
~
Volume 84, Number 4
1997
Fritsch 759
Revision of Styrax
(16); 4899 (16); m ee 5038 (16); 5098 (16); 5212
eo Juzepezuk, 5. 1
Kappelle, МК: a MK74 (19; МК118 (6);
MK119 (19); MK on (19); MK417 (19); MK483 (19);
MK505 (6); MK861 (6); MK892 (19); MK1088 (19); MK
116 (19); оу dii 2800 (19); 2875 (19). Keeney,
( та quer W. A. 5664
643 (16); 7929 (16).
19
~
"x
М «ы
Bg
8 5
Ф
л
Pc
14600 (6); 15533 (6); 26612 (6). Linden, J. J. suppl.
а - (6); 358 (19). Little, E. L. 6043 (19); 9932 D
1). "€ E. 7555 (1). López R., R. M. 503 (6).
tm H., F. G. 43 (16). Lott, E. J. 2540 (15). Lundell, C.
L. 12631 (6); 19214 (17); 19450 (7); 19618 (7); 20452
(19); 20933 (7); се А 21172 (6); 21219 (17). Гуоп-
net, Е. 2106 (16); 1
MacDaniels, L. " e (6). иерин T. H155 (15);
H156 (15). Machuca N., J. A. 6636 (16). Madriz V., A. 9
(19); 67 (1). Magallanes, A. S. 1534 (15); 3432 (15). Ma-
gafia, R. H. 5607 (6); 5785 (6); 6109 (6). Makrinus, E
551 (15). Mancera O., A. МО–315 (16). Marquez, W. 141
(6); 847 (6); 882 (6). Martin, G. J. 501 (6). Martin J., S.
1208 (1). Martinez A., M. 55 (16). Martinez, J. L. 1300
(6). Martínez S., E. 327 (16); 429 (16); 2857 (16); 3908
(16); 4906 (2). Martinez-Ico, M. 160 (19). Matuda, E. 467
(19); 568 (19); 2668 (19); 2982 (10); 3930 (19); 4135 (19);
4179 (6); 4335 (6); 5102 (6); 5215 (1); 15410 [5410] (6);
15507 (3); 15922 (6); 15995 [5995] (6); 16265 (19);
17816 (6); 28364 (16); 30405 (16); 30566 (16); 3
(16); 30858 Сх 31979 (16). Маућеја, М. Н. 1632 (15).
McCaffrey, D McPherson, G. =; A Pee i
R. 541 (15); 104 (16); 11764 (16); 1 see
(8); 12100 (15); 13369 (8); 13914 (16); 14279 (16); 13025
23515 (15); 23593 (16); 25492 (15); 2602
(15). Meave, J. 1162 (1). Medina G., C. 175 e
ano, F. G. (6); 5032 (16); 5044 (16); 5176 (16);
5191 (16); 6201 (19); 6706 (2); 6785 (16); 6821 (16);
oza, А. ud 3802 FR
Miller J. S. 140 (1
3114 (16); 3140 (15); 5145 (146). Miranda, F. 161 (16);
1214 (16); 1580 (16); 3066 (16); 3460 (6); 5295 (1); 5927
(1; 7020 (6); 7055 (19); 7232 (19); 9170 (10); 9173 (6);
9199 (6). Molina R., А. 627 (19); 6460 (19); 6522 (19);
7233 (6); 7524 (1); 12236 (19); 17086 (19); 20321 (1);
21341 (1); 21505 (19); 24404 (6); 24561 (1); 25328 (1);
27644 (19); 34417 (1). Montes, M. 714 d P. Mos oore, H.
E. 3231 (6); 6225 (6). Mora, C. E. 3, p.p. (19). Mora, С.
229 (19). Moraga, M. 263 (6). ан, Е 111 (19);
4266 (6). Moran, К. 10161 (15). Moreno G., 5. 38 Bint
176 (16). Moreno, P. Р. 480 (19); 1883 (1); 10854 (1);
16366 (11a); 17875 (1); 21064 (1); 21279 (1); 22001 (1);
22569 (1); 23512 (1). Mori, S. 7359 (19); 7416 (19). Mo-
rones G., A, 209 (16). Motte, M. us M m Mufioz, E. 4
(19); 36 ad. Museo Nacional 1
Narave, H ‚4 o 55 (6); i Noe, M. 24719 (6);
26290 (6); 27636 (19). Nees s.n. [18857] (1). Negrete А.,
М. 45 (16). Neill D. 2313 (19). ben E. E. 3750 (6);
6886 (16). день Lit њој мај o Miem ta G. 50 (8);
411 (15); 546 (15). Norris, D. . Núñez, J.
С. 5. 4739 (16); 4785 (16): evs Sg ЫН (16); 6328
(16
Өр, Р. А. 523 (1). Orozco, J. М. 329 (1). Orsted, А
S; nag 0 11194 (19). Ortega, J. G. 60 (15); 4118 (15);
n E., E. 34 (1); v^ (1); 161 (10). — R.,
M. 1 (6). Palmet, C. W. 97 (6). Palmer, E. J. 9 y
10237 (14c); 10907 (14a); wen (14c); 11528 ies Pa-
nero, J. 3475 (2); 5254 (2). Paray, L. 681 (6). Parks, H.
B. 1009 (14c); 1942 (14c); 40988 (14c). Pérez J., L. A.
814 (15). Pérez, S. 17 Sep. 1987 (8). Peterson, C. D. 1243
(14b). Pichardo A., V. 14 (16); 28 (16); 111 (16); 212 (16).
Pittier, H. 606 (1); 1583, p.p. (1); 1583, p.p. (19); 2140
(16); 11012 (8); 13104 (6). Proctor. E
25299 (19); 25514 (19); 31886 (19). Purpus, C. A. s.n.
ay 1925 “with 10057” (1); 46 (1); 82 (19); 157 (19);
1925 (6); 5317 (19); 7422 (3); 9280 (6); 10082 (19);
10088 (6); 10521 (6); 10531 (1); 10545 (6); 10555 (19);
10612 (6); 13001 (1); 14301 (1
Quirós, M. 180 (6).
Ramírez D., R. E (8); 477 (15). a T 77 о
Ramírez R., К. 89 isi Ramírez-Marcial, N. 504 (6);
553 (19). Ramos, P. X. 86 (16). Reko, B. У 3642 (15);
4136 А Reverchon, J. 1551 (14а). бене ЗА ы
(1); 463 (1). Кеуез, Ј.
(16). Riskind, D. H. 1682 (140); 1691 (14), 1915 hides
2079 әз Rivera, С. 198 (19); 749 (1); 1851 (6). Robles,
obl odríguez, J. V. 2004 (1);
(16). Rubio, H. 1583 (6);
(16); 17211 (6); 18362 (16); 18371 09; 18529 (2); 19771
(16); ve eg (9: 30304 (16); 397
Salaza 375 (8). 5 T — A. ive 5 (16). Sandoval
din: M., 8 (16); 2695 ~ 3303 (15);
4536 (16); 4776 (15). ud (6). es V., A. 1437
(15). Schmalzel, R. J. 1608 (19); 1756 ( (19). Schmitz 505
(16). Schwabe, W. 224 (1); 77566 (16). =» п, 5. wd );
258 (6); 1056 (6). Sessé, M. = 6). Sharp, А. J. 3460
(6); 4673 (6); 4676 (6); 4677 (19); e (6); 45107
[451047] (3); -— ee 45382 (6); 45499 (16); 45559
(6); ru dr 54 (6); 46173 (6); 46209 (6). Skutch,
A. F. 1935 ou th, A. A113 (19); P1971 (19); P2623
(19); 65 ci 176 (19); tj iv; 4165 (19); 4232 (19);
10079 (19). Smith, C. 3 (19); 4512 (6). Smith, J.
517 (140); 678 bie Sith, 1 D. 2266 (1). Smith, J. F.
47 (16). Smith, R. F. M149 (14b).
ча ЧУ
do
© бф;
get
pe
>
@
_ 3
un
——
o
©
=!
и
©
>
3
=
– E
=
>
oo не gr M. eges а Standley, P. C. 1674 (1);
н (1); 6344 (1); 8019 (19); 12351
(1); 14121 ip 15170 d» 15875 (1); 15947 (1); 20147
1); 20405 (1); 32549 (19); 33500 (6); 34672 (19); 41410
(19); 42216 (19); 42254 (19); 42345 (19); 42656 (19);
51137 (6); 51175 (6); 51309 (6); 57958 (1); 58420 (19);
62879 (1); 63714 (19); 63786 (19); 64767 (1); 67865 (6);
68618 (10); 70316 (6); 77656 (1); 80536 (19); 80605 (19);
); 80684
al
80659 (19); 81378 (1); 82830 (1); 82938 (1);
83018 (1); 83697 (6); 85559 (10); 91569 (19); 91577 (19);
92302 (19); 92400 (19); 92574 (19); 92631 (19). Stern,
W. L. 1993 (19); 2045 (19). Stevens, W. D. 86 (19); 6687
(б); 13694 (19); 15494 (1); 20401 (11а); 22180 (19);
760 Annals of the
Missouri Botanical Garden
09 (1). Steyermark, J. A. 30624 (1); 30672 (1); 30728 M., A. 4529 (16). Vela G., L. 2050 (16). Velazquez H., A.
D 31150 (1); 32340 (19); 32827 (19); 33138 (1); 33315 13 (8). Velzen, H. van 2765 (19). Ventura A., F. 784 (6);
(19); — 9); 35418 (1); 36234 (10) 36712 (19); 866 (6); 1195 (6); 3408 (6); 3511 (6); 5211 (6); 8071 (6);
10); 36807 (6); 37378 (6); 37989 (19); 42075 (1); 8116 (6); 12945 (6); 20283 (6); 20303 (6). Vergas, C. 285
pee (1); 43199 (19); 43279 (3); 43410 (19); 43593 (19); (6). Villarreal, L. М. 4585 (8). Villaseñor, J. L. 183 (16).
43937 ed. 46750 си 46759 (19); 47397 (19); 47924 — Vincelli, P. C. 424 (19). Voorhies, B. 3-11 (1); 5-3 (1);
(1); 48441 (19); 48666 (17); 48848 (17); 49738 (6); кен 16-9 (1).
(3); 50429 (1); 50699 (1); 50776 (1); 50801 (1); 51100 Walker, 5. 72.021 (16); 73HO7 (15). Warscewicz, J. 40
(1). Stone, D. E. 3286 (6). Sullivan, J. К. 543 (6). (1); 203 (19). Wawra 1011 (6). Weaver, К. Е. 1745 (6);
Tapia, L. 80 (6). Taylor, C. M. 3210 (19). Taylor, J. 2956 1754 (6). Webster, С. L. 12230 (19); 12496 (11а). Wendt,
(6); 11848 (19); 17390 (1). Téllez V., O. 10139 (16); Т. 148b (14e); 582 (14е); 3576 (127). Werff, H. van der
10282 (15); 10412 (15). Templeton, В. С. 7000 (15). Ten- 6301 (19). Wetter, М. A. 2007 (16); 2057 (16). yr G.
orio L., P. 2642 (2); 8364 (15); 8441 (15); 14351 (16); 70 (19). White, S. S. 5260 (1). Wiemann, M. C. 32 (19).
15605 (15); 16929 (15); 16984 (15). Tenorio, P. 5707 (1). Wilbur, R. L. 1766 (16); 8821 (19); 16166 (19); d
Terry, M. E. 1338 ed 1348 (19). Thomas, H. 26 (7); 474 (19); 18026 (1); 21135 (19); 22584 (19); 23494 (19);
(19); 496 (6). Ton, A. S. 404 (1); 3431 (1); 6494 (1); 7814 27228 (19); 29475 (19). Williams, L. O. 10491 (1); 11279
(6); 7963 (10); 9329 (16); 9734 (10). Tonduz, A. 459 (19); (1); 12136 (1); 12314 (1); 12648 (19); 12709 (1); 13270
7079 (1); 7447 (1); 7667 (6); 9816 (1); 11744 (19); 17924 (1); 13318 (19); 13581 (19); 13709 (19); 14028 (1); 16067
(1). Toriz A., С. 464 (6); 475 (6). Torres C., R. 2686 (6); (1); 16131 (19); 16260 (19); 17455 (19); 20001 (19)
4982 (6). тоа, К. 13321 (16). Tróchez, L. 17 (1). Tyson, 23955 (1 +" eie (19); 25087 (1); 28194 (19). Wood
E. L. 826 (19); 827 (19). son, R. E. yer 919 (19). Woronow, G. 99b (16
Ugent, D. 1635 (16). Umaña, С. 71 (19); 144 (19. Wynd, F. L. dr
Utley, J. 4557 (19). Yuncker, T. G. 5589 (1); 6175 (6).
Valdez, G. L. 616 (2). Valerio, Man. 1378 (1); 1396 Zamora У., N. 1213 (6). Zavala C., F. 295 (6). Zola В.,
(19). Valerio, Mar. 145 (19); 174 (19). Vásquez S., J. 2174 М. С. 526 (6); 612 (6). Zúñiga, R. 169 (6); 575 (6); 607
(16). Vázquez, А. 1195 (15); 4164 (15); 4625 (6). Vázquez (1).
Appendix 1. Comparison of Gonsoulin's (1974) treatment of Styrax from western Texas, Mexico, and Mesoamerica
to the present revision. Synonyms are indented. Specimens of the seven taxa at the end of the right column were not
available at the time of Gonsoulin's revision. (D) = deciduous.
Gonsoulin, 1974 Present revision
S. argenteus var. argenteus S. argenteus
S. panamensis S. panamensis
S. radians, s,
S. austromexicanus, nom. et stat. nov.
S. argenteus var. hintonii S. ramirezii
. magnus S. magnus
S. argenteus var. ramirezii S. ramirezii
S. orizabensis S. ramirezii
S. warscewiczii S. warscewiczii
8. Ld sp. nov.
5. conterminus S. conterminus
S. glabrescens var. glabrescens (D) D. delicia (D)
S. glabrescens var. pilosus (D) S. glabrescens var. pilosus
S. jaliscanus (D)
. jaliscanus (D)
. gentryi, sp. nov.
platanifolius var. platanifolius (D)
platanifolius var. stellatus (D)
texanus (D)
youngiae (D)
S
8
5. platanifolius subsp. platanifolius (D)
S. platanifolius subsp. stellatus, comb. et stat. nov. (D)
S. platanifolius subsp. texanus, comb. et stat. nov. (D)
5. platanifolius subsp. youngiae, comb. et stat. nov. (D)
5. platanifolius subsp. mollis, subsp. nov. (D)
S. glabratus
5
5
5
5
5
5
tn to үл л
. lanceolatus, nom. et stat. nov.
nov.
nicaraguensis subsp. nicaraguensis, sp. nov
. nicaraguensis subsp. ellipsoidalis, subsp. nov.
. peruvianus
. tuxtlensis, sp. nor.
Total: 7 species, 11 taxa Total: 19 species, 24 taxa
отини ии иь а иии иии еи
Volume 84, Number 4 Fritsch 761
1997 Revision of Styrax
Appendix 2. Index to scientific names in taxonomic part. Numbers in parentheses correspond to taxon numbers in
the text. Synonyms and excluded names are italicized.
nicaraguensis P. У. Fritsch (11) ................... 736
Adnaria Raf. 712 subsp. ellipsoidalis P. W. Fritsch (110); 737
Benzoin Hayne 712 officinalis var. jaliscanus (S. Watson) Perkins
Cyrta Lour. 711
Darlingtonia Torr. 712 orizabensis Perkins (16) 747
Epigenia Vell. 712 panamensis Standl. (12) 738
integerrima Vell. (5) 725 peruvianus Zahlbr. (13) 738
Foveolaria Ruiz & Pav. 112 pilosus (Perkins) Standl. (6) 727
Styrax L. 711 platanifolius Engelm. ex Torr. (14) ................ 739
argenteus C. Presl (1) 714 subsp. mollis P. W. Fritsch (14b) ............... 742
var. grandiflorus: E. Carranza (2) ................ 718 subsp. stellatus (Cory) P. W. Fritsch (14c) ..... 743
var. hintonii (Bullock) Candi (боја ево: 747 subsp. texanus (Согу) Р. W. Fritsch (14d) ...... 744
var. micranthus (Perkins) D'Arcy (16) .......... 747 subsp. youngiae (Cory) P. W. Fritsch (14e) ..... 744
var. parvifolius El Carranza (d) ainia dide 3 var. stellatus Cory (14c) 743
var. ramirezii (Greenm.) Gonsoulin (16) ........ 747 polyanthus Perkins (1) 714
austromexicanus P. W. Fritsch (2) ................. 718 polyneurus Perkins (19) 199
conterminus Donn. Sm. (3) 720 psilophyllus A. DC. (5) 725
cyathocalyx Perkins (16) 741 radians Р. W. Fritsch (15) 745
diplotrichus Diels 756 ramirezii Greenm. (16 741
вала Pohl (5) 725 var. micranthus (Perkins) Perkins (16) .......... 747
gentryi P. W. Fritsch (4) 123 var. orizabensis (Perkins) Perkins (00) 2252. 741
glabratus Schott (5) 725 squamulosus M. F. Silva (5) 726
glabrescens Benth. (6) 725 steyermarkii Р. W. Fritsch (17) 751
var. pilosus Perkins (6) 727 иан Согу (144) 744
malensis Donn. Sm. (6) 727 uxtlensis P W. Fritsch (18) 753
hintonii Bullock (16) 741 built Lundell (6) 721
incarnatus P. W. Fritsch (7) 730 vulcanicola Standl. & Steyerm. (10) .............. 734
jaliscanus S. Watson (8) 731 warscewiczii Perkins (19) 753
lanceolatus Р. W. Fritsch (9) 733 youngiae Cory (14e) 744
lauraceus Perkins (5) 725 Strigilia Cav. 711
leiophyllus Miers n" argentea (C. Presl) Miers (1) 714
limoncillo (Humb. €: Bonpl.) Miers ............... 156 glabrata (Schott) Miers (5) 725
magnus RU (10) 734 leiophylla (Miers) Miers (5) 725
micranthus Perkins (16) 747 psilophylla (A. DC.) Miers (5) 725
ا زر (1) 714 Tremanthus Pers. 711
SYSTEMATICS OF KALIMERIS
(ASTERACEAE: ASTEREAE)!
Hong-ya Gu? and Peter С. Hoch*
ABSTRACT
The genus Kalimeris (Asteraceae: basra- comprises a Re c group, with the exclusion of section Cordifol-
ium, characterized by laterally com
егегорарр, vealed
parisons and cytological studies. Its supposed i^: везне ‘with бо desa American genus
imeris are all closely relat
superficial. The species of
pappus bristles no longer than the length of the corolla
i i . It has
se a polyploid series ranging from diploid to
12-ploid with x — 9. Eight species and ten беча Ад of Кн v pas gasto ized in this treatment.
Kalimeris (Asteraceae: Astereae) is native in East
Asia, ranging from the Amur basin in
Asia at 50°N China, Korea, and Japan to
northern Indochina at 16°N, and westward to northern
Myanmar (Burma) at 97°E. Most species occur in Chi-
na. Although it is segregated from Aster by most Asian
botanists, its taxonomic status as well as its delimi-
tation have been controversial. In order to clarify the
taxonomic status of Kalimeris and its phylogenetic re-
lationships within Astereae, to establish its infrage-
neric delimitation and phylogeny, and to improve our
understanding of the biosystematically poorly known
јап Astereae, we undertook studies on the mor-
phology, cytology, and geography of Kalimeris and
some related genera. The results of the cytological
study and an RFLP analysis on rDNA of some Kal-
imeris species were published in separate papers (Gu,
1989; Gu & Tara, 1990; Gu et al., 1994). The present
treatment was based on results of those surveys, in
which we treat Kalimeris (excluding sect. Cordifolium)
as a monophyletic taxon of eight species, distinct from
Aster and other Asian Astereae.
TAXONOMIC HISTORY OF KALIMERIS
The name Kalimeris was first published as a sub-
genus of Aster by Cassini in 1822 (see Cassini in
& Dawson, 1975). The only species he de-
scribed in that subgenus was based on a single cul-
tivated plant that he illegitimately named Kalimeris
platycephala. In 1825, he elevated the subgenus to
generic level based on its distinctive short pappus
bristles, conical receptacles, and flat achenes.
Shortly thereafter, Blume (1826) published a
monotypic genus, Asteromoea, into which he trans-
ferred Aster indicus L. Subsequent treatments of
Astereae by other botanists have differed substan-
-— in the delimitation of Kalimeris and Aster-
"Neo (1832) expanded Cassini's concept of Kal-
imeris by transferring into it four species of Aster
with longer pappus bristles; these were treated in
the genus Heteropappus in most subsequent treat-
ments by other botanists. Nees maintained Aster-
omoea as a distinct monotypic genus. He altered
Kalimeris to Calimeris, a spelling that has been
! This "e$ was undertaken as part of the first author's doctoral dissertation research at Washington University, St.
topic and for his review of the manusc
T. Kawahara, J. poses M. Takahashi, H. Tobe, T. Yahara, D. С ya а . Li, C.
ul to the Missouri
hen i
Zhang. The first author's graduate studies were supported by the Ministry a Edasi d Chine and Weshington
L
University; to them she is very gratefu
We thank the curators of the following herbaria for providing access to and/or loaning plant specimens for this pm
M, E, G, GH, HIB, IBSC, ТЕР, K, KUN, KYO, LE, MAK, NAS, NY, P, PE, SYS, SZ, TI, TNS,
US, W,
2 College of Life Sciences, Peking University, Beijing, 100871, People's Republic of China.
3 Misso
ssouri Botanical Garden, P.O. Box 299, St.
Louis, Missouri 63166, U.S.A.
ANN. MISSOURI Bor. GARD. 84: 762-814. 1997.
Моште 84, Митбег 4
1997
би 4 Носћ 763
Systematics of Kalimeris
taken up by several botanists since then. De-
Candolle (1836) adopted Nees’s system and trans-
ferred two more species of Aster into Kalimeris.
Hoffmann (1890) also recognized Asteromoea as a
. separate genus, but treated Kalimeris sensu Nees
as a section of Aster (sect. Calimeris), a treatment
also followed by Nakai (1911, 1919) and Grau
(1977). Part of the reason for the consistent sep-
aration of Kalimeris from Asteromoea is that the
inclusion of the species with long pappus bristles
in Kalimeris by Nees obscured Cassini’s original
concept of the genus, making the single species
of Asteromoea with very short pappus bristles seem
more distinctive than it really is. Most Asian bot-
anists consider the types of Kalimeris and Aster-
omoea to be congeneric, and because Kalimeris
has priority, it has been adopted in the present
treatment.
The small North American genus Boltonia, de-
scribed by LHéritier (LHéritier € de Brutelle,
1788), shares several floral and achene characters
with Kalimeris. This led Bentham (1861, 1873) to
combine the two genera with the southern North
American Dichaetophora A. Gray to form a single
genus, Boltonia, and to designate each of the three
as a separate section. Dichaetophora is now gen-
erally recognized as a separate monotypic genus
(Shinners, 1946; Turner et al., 1961; Grau, 1977),
whereas the statuses of Boltonia and Kalimeris have
continued to be controversial. Debeaux (1877) ac-
cepted Bentham’s classification and described a
new species, Boltonia lautureana, collected in Chi-
na. Tamamshjan (1959) recognized this species as
the only Asian Boltonia and retained Kalimeris as
a separate genus. Only a few subsequent non-Asian
botanists have followed either Hoffmann’s (Grau,
1977) or Bentham’s (Grierson, 1964) treatments,
and only limited discussion on this subject is avail-
able in the literature (Grierson, 1964; Lauener,
1976; Koyama, 1983; Ito & Soejima, 1995).
Even among authors who recognize Kalimeris as
a distinct genus, there is considerable disagreement
about its delimitation and the number of taxa that
should be recognized. There is no recent treatment
that covers Kalimeris throughout its entire distri-
butional range, although several local taxonomic
treatments are available. In The Compositae of Chi-
na, Hu (1967) transferred 4 species of Aster into
imeris, treating a total of 11 species in the genus
in China. By contrast, in the most recent treatment
in Flora Reipublicae Popularis Sinicae (Ling et al.,
1985), 7 species were recognized. Kitamura
(1937b, 1957) recognized 6 species in Kalimeris in
is taxonomic treatment of Japanese Asteraceae
and a subsequent series of articles. However, in the
original Flora of Japan, Ohwi (1965) recognized
only 4 species. Kitamura (1937b) also established
2 sections: sect. Kalimeris and sect. Cordifolium,
which have been adopted by some other authors
(Ohwi, 1965; Ling et al., 1985). In Plantae Sinen-
ses, Handel-Mazzetti (1937-1938) used the generic
name Asteromoea and included 7 Chinese species
in it. Hu (1967) did a bibliographic study on the
genera and species of Asteraceae in China, and
transferred 3 more species from Aster into Kalimeris
to make a total of 11. Some taxa of Kalimeris sensu
Handel-Mazzetti and Hu were subsequently segre-
gated into Miyamayomena (Kitamura, 1937b,
1982).
In the treatment of Aster in the new Flora of
Japan, Ito and Soejima (1995) treated Kalimeris as
Aster sect. Asteromoea Makino, arguing that “recent
cladistical and molecular analyses reveal these spe-
cies [Kalimeris, Heteropappus, and Miyamayomena]
to be closely related to species of Aster s. str.,” also
noting that “further phylogenetic studies are need-
ed to reconstruct a complete system in Aster” (Ito
& Soejima, 1995).
TAXONOMIC DELIMITATION OF KALIMERIS
Generic delimitation in Astereae has long been
a source of disagreement among botanists. The
ways in which they treat the large and diverse ge-
nus Áster usually reflect their philosophy on generic
concepts, and although there are many variations,
in general there are two schools of thought. The first
approach maintains a very inclusive generic con-
cept of Aster, with subdivision of the genus into
several subgenera (Bentham, 1873; Hoffmann,
1890; Grau, 1977; Jones € Young, 1983). The sec-
ond approach is to segregate many distinctive small
genera from Aster, thus adopting a narrow generic
concept (see Cassini in King & Dawson, 1975;
Nees, 1832; DeCandolle, 1836; Tamamshjan, 1959;
Semple & Brouillet, 1980; Ling et al., 1985).
A number of small genera endemic to Asia have
been segregated out of Aster, including Heteropap-
pus, Kalimeris, Galatella, and Turczaninowia.
extreme case is the treatment of Astereae in the
Flora of URSS by Tamamshjan (1959), who adopted
a very narrow generic concept of Aster, segregating
two new genera (Kemulariella and Conyzanthus),
and accepting many small genera established by
Cassini, Nees, and Novopokrovski. Kitamura
(1937b) segregated another small genus, Gymnaster
(later changed to Miyamayomena [Kitamura,
1982]), in his major treatment of Japanese Astera-
ceae. In order to evaluate the status of Kalimeris,
a survey was made of 14 morphological characters
764 Annals of the
Missouri Botanical Garden
Table 1. Character comparison among several genera of Astereae.
Ray florets da Involucre
Taxa (no. of sp.)* Habit Fert. L (mm) L/W Lilobe) Shape Series
Aster sensu str. annual + 5.5-24 4.3-10 21—52 turbinate 2-4-8 subequal
(ca. 200) perennial campanulate imbricate
subshrub hemispheric
Heteropappus (12) biennial + 9.6-19.6 3.6-5.5 2.5-3.9 (long) hemispheric 2-3 subequal
(8) (short)
Kalimeris (8) perennial + 7-22 3.9-9.4 2.2-3 (long) сатрапшаіе 3—4 imbricate
3—5.1 (short) hemispheric
Aster miquelianus** ^ perennial + 87-112 5-7 T- hemispheric 2-3 subequal
Boltonia (4)*** perennial + 47-13 6.3-8.1 3.66 campanulate 3—4 imbricate
Miyamayomena perennial + 8-195 41-73 1.7-3.7 turbinate 2—1 subequal
(5)**** campanulate imbricate
emispheric
Asterothamnus (4) subshrub + 85-10 464 45-5 turbinate 34 imbricate
hemispheric
Galatella (22) perennial = E522 4.7-7.7 1.9-3.4 turbinat 34. imbricate
with woody campanulate
base hemispheric
Linosyris (3) subshrub X X X 3.2-9.7 turbinate several imbricate
Abbreviations: А: pappus as long as the disc corolla; B: pappus % as long as the disc corolla; C: pappus И— e as
long as the disc corolla; D: tw
o awnlike and fragile bristles; E: pappus
corolla: fert.: fertile; L: length; Ў longer lobe; short: shorter lobes; W: width; X: absent; +: yes;
s two or more series, and longer than the
—: no; +: more or
* Unless noted O the number of species is based on Kitamura (1937b), Jones (1980), Ling et al. (1985), and
Trae jan (195
* A.
idi was treated as a species in Kalimeris sect. Cordifolium by Kitamu
*** The number of species was estimated from the treatments of Cronquist (1980) = Anderson (1987).
986).
**** Based on У. L. Chen (1
of Aster sensu stricto, six Eurasian genera segre-
gated from Aster, and Boltonia; the results are listed
in el,
Asterothamnus, Galatella, and Linosyris are char-
acterized by their woody habit, unevenly com-
pressed achenes, and multi-seriate pappi. These
genera are distributed in northwestern and central
Asia and are very distinct from the other groups
listed in Table 1. Although Boltonia is similar to
Kalimeris in floral morphology, detailed analyses
indicate that the similarity is rather superficial.
Both genera have compressed achenes, but the
achenes of Boltonia are very flat with thin and al-
most transparent wings (Fig. 1), whereas the
achenes of Kalimeris are only laterally compressed
and have thickened margins (Fig. 2). The “shared”
short pappi have been used to support the argument
that these two genera should be combined, but on
careful examination the pappus bristles of the two
found on each side at the top of the achenes (Fig.
1). Kalimeris has subequal pappus bristles or, if
unequal, several longer bristles are interspersed
with shorter ones (Fig. 2). The texture of the pappus
is also different in Boltonia and Kalimeris: the for-
mer has fragile pappi similar to those of Erigeron,
whereas the latter has robust pappi like those of
Aster and Heteropappus. Boltonia also has entire,
glabrous leaves and narrow phyllaries with often
colored midribs, whereas Kalimeris has lobed or
incised, variously hairy leaves and broader phyl-
laries with green midribs. In summary, detailed
comparison of the achenes and pappi in Boltonia
and Kalimeris does not support a close relationship
between them; most of the purported “similarities”
are non-homologous and may reflect pides
evolution, such as that documented in m
of Asteraceae by Cronquist (1955) and ا д
(1977).
Kalimeris was subdivided into two sections by
Kitamura (1937b): sects. Cordifolium and Kalimeris
(as Asteromoea), which have been adopted by other
Asian botanists (e.g., Ling et al., 1985). The only
characteristic shared by these two sections is short
pappus bristles. They differ in many other funda-
Volume 84, Number 4 Gu 4 Hoch 765
1997 Systematics of Kalimeris
Table 1. Extended.
i i Ray (achene) Disc (achene)
t
E зен No. of No. of И
Shape Surface Shape ribs Shape ribs Indument Ray Disc
ooth cylindrical 6-8 cylindrical 6-8 glabrous A A
convex alveolar triangular 3-4 compressed 2-3
flat alveolar very compressed 2(-3) very compressed 2 hairy В-С A
= conv glandular
convex alveolar triangular 3(-4) compressed 2(-3) hairy C C
subconical glandular
flat alveolar cylindrical 6-7 cylindrical 4—6 glabrous C C
conical smooth + triangular 3 very flat 2-3 glabrous D D
loosely hairy
flat smooth cylindrical 5-6 cylindrical 4-5 glabrous A X
+ convex alveolar + triangular 3 = compressed 2-3 densely hairy
convex alveolar triangular 3 triangular 3 hairy E E
conical
flat alveolar compressed abaxially 1-2 compressed abaxially 1-2 densely hairy E E
convex scaled
flat convex | scaled X 0 compressed abaxially 1-2 hairy E E
mental characters; for example, species in section
Cordifolium have cordiform leaves with long peti-
oles, 2-3 series of subequal phyllaries, and cylin-
drical achenes with 5-7 ribs (Fig. 3, Table 2),
whereas species in section Kalimeris have oblong
and sessile leaves, 3—4 series of imbricated phyl-
laries, and laterally compressed achenes with 2-3
ribs. Cytological and anatomical evidence also
shows that these two sections are very different (Gu,
1987, 1989), and that retention of section Cordi-
folium in Kalimeris would make the genus diphy-
letic. These observations are also supported by ev-
idence from molecular studies on the Japanese taxa
of Kalimeris and some related Aster species (Nish-
ino & Morita, 1994; Ito et al., 1994, 1995). Con-
sequently, section Cordifolium is excluded from
Kalimeris in this treatment and left as part of Aster.
The relationships of this group remain to be deter-
mined.
Although Miyamayomena has only five species
(Chen, 1986), it is as variable morphologically as
the large genus Aster, as shown in Table 1. This
group may in fact be an artificial assemblage, but
further study clearly is needed.
Aster, even when circumscribed in the narrower
sense used here, is a very speciose and morpho-
logically diverse genus. Three main subgeneric
classifications based only on external morphology
are available for Eurasian Aster from regional flora
treatments (Kitamura, 1937b; Tamamshjan, 1959;
Ling et al., 1985). For convenience in the following
discussion, I have adopted Ling’s system of Aster,
in which the genus is subdivided into 3 sections,
Aster, Orthomeris A. Gray, and Alpigenia Benth.,
and 27 series.
Kalimeris is most similar to Heteropappus and As-
ter (especially sect. Orthomeris ser. Ageratoides Ki-
tamura). The main differences among these three
groups are habitat specialization, habit, leaf and
phyllary morphology, and floral and achene mor-
phology (Table 2). Phyllary and achene morphology
are the most useful characters for distinguishing the
three genera in the field. Heteropappus is charac-
terized by its two series of subequal herbaceous
phyllaries and dimorphic pappi in the ray and disc
achenes: shorter pappi on the ray achenes and lon-
ger pappi on the disc achenes (Fig. 4). The unequal
corolla lobes have also been used to distinguish
Heteropappus from the other two genera (Grierson,
1964), although this character state is also found
in several species of Aster and Kalimeris. Aster and
Kalimeris are similar in having 3—4 series of im-
bricated phyllaries, but differ in that Aster has ei-
ther laterally compressed achenes with pappus
bristles of the same length as the disc corollas (Fig.
5) or cylindrical achenes with shorter pappus bris-
tles or none, whereas Kalimeris has laterally com-
pressed achenes with short pappus bristles no lon-
ger than the length of the corolla tube.
As mentioned earlier, convergent evolution is
766 Annals of the
Missouri Botanical Garden
res 1-5. Morphology of achenes. —1. Boltonia diffusa; A. Front view. B. Side view. —2. Kalimeris indica; А.
Ray асћепе. В, С. Disc achene of front view and side view, respectively. —3. Aster miquelianus. —4. Heteropappus
hispidus; A. Ray achene. B, C. Disc achene with front view and side view, respectively. —5. Aster ageratoides. A. Front
view. B. Side view
|
Volume 84, Number 4
1997
Gu & Hoch 767
Systematics of Kalimeris
Table 2. Character comparison among Aster ser. Ageratoides, Heteropappus, and Kalimeris.
Character Aster ser. Ageratoides Heteropappus Kalimeris
Habit perennial biennial perennial
Habitat montane forest or dis- dry, sandy, or saline soil disturbed area
turbed area
Roo us tap fibrous; tap-root in one species
радиом stem absent or rhizome absent rhizome; absent in one species
Leaf venation 3 basal veins 1 midvein 1 midvein
Phyllaries 3-5 series, imbricate 2 series, subequal 3—4 series, imbricate
Fibers in cross section of continuous separate continuous
Receptacle + convex or flat flat convex or subconical
Ratio: long/short corolla lobes 1-1.7 1.5-2.4 1-1.8
Achene morphology = compressed very compressed compressed
Ratio: achene length/thick-
ness 2-3.1 6-7.7 3-4.2
Number of ribs on ray
achene 3-4 2(-3) 3(4)
Number of ribs on disc
chene 24 2 2-3
Ratio of = ray pappus/
disc p.
hire penmi of disc pappus same as disc corolla
1 (rarely less than 1)
shorter than disc corolla
less than 1 (rarely 1)
same as disc corolla
common in Asteraceae. In very few cases can a
single character be used to delimit a genus. For
example, short pappi have evolved many times in
different phylogenetic lines in Astereae. Several
have either been included in Kalimeris (Hu, 1967)
or (А. tongolensis Franch.) proposed for inclusion
(Jones & Smogor, 1983), but overall these species
share very few other characters with Kalimeris an
should not be included therein.
Although Kalimeris is treated as a distinct genus
here, the ranking of this morphologically discrete
taxon at the generic level is provisional, to the de-
gree that it ultimately must be consistent with a
comprehensive classification of the Astereae.
MATERIALS AND METHODS
Conventional morphological data were derived
from more than 3500 herbarium specimens of As-
tereae, 2000 of Kalimeris and 1500 of other mem-
bers of the tribe from Asia (Aster, Asterothamnus,
Doellingeria, Galatella, Heteropappus, c
Miyamayomena, and Turczaninowia) and North
America (Boltonia). Numbered EN are ref-
erenced in a collectors’ index. Morphological mea-
surements on this material were supplemented by
observations on living plants in the field and in
cultivation. Study of the morphology of rosettes was
made almost entirely on living plants in the green-
house. At least ten specimens of each species, from
different populations, were measured for morpho-
logical characters. The morphological terminolo
is based mainly on Lawrence (1951). Cauline
leaves were measured at the middle portion of the
stem; if the leaves in that portion were withered,
the leaves below the lowest branch were measured.
Floral features were measured on fully opened flow-
ers, as indicated in Figure 6.
We have adopted the "morphological species
concept" (Davis & Heywood, 1963; Stuessy, 1990)
in Kalimeris as the most practical approach in this
variable group. Our species are definable by mor-
phology, with sharp discontinuities between spe-
cies; where possible, each is defined by one or more
autapomorphies. For taxa whose morphological dis-
continuity is correlated with ploidy levels and/or
with allopatry, but where substantial intergradation
may occur in areas of overlap, we have used the
category of subspecies (Raven & Raven, 1976).
MoRPHOLOGY AND TAXONOMIC CHARACTERS
The eight species of Kalimeris are distinguished
by combinations of characters, and many of the fea-
tures differ only quantitatively. In the following text,
a. description of general morphological characters
is provided, with emphasis on some characters use-
ful in classification.
768 Annals of the
Missouri Botanical Garden
3-
a
! :
н 4r silo»
سم b—
па دو اا یه
c
Џ
rona чн.
— y لب
Џ Џ
D E
e6. Diagram of floral parts measured.
(b) oft e 20 Length of disc
(a) and width (b) of sce of ray floret; length
appendage (c). —F. Lengt
the margin of achene (c); length of pappus
HABIT AND LIFE HISTORY
All species of Kalimeris are perennial herbs. Ro-
settes grow out from winterized underground stems
(rhizomes) or adventitious buds in early spring, and
stems shoot out from the rosettes in late spring.
Prior to flowering, branches begin to develop from
the uppermost lateral buds, and the lower branches
develop subsequently from the lateral buds below
the first branch on the main stem. Generally the
flowering period begins in late May and ends in
early November. The species in the northern part
of the range, e.g., K. incisa and K. pinnatifida, usu-
ally bloom from late June to October, while south-
F G
—A. Length (a) and width (b) of involucre. —B. Length (a) and width
floret (a) and corolla tube (b); length (c) and width (d) of corolla lobe. —D. Length
c) of corolla
h of anther (a) and anther е жагу... (b). —G. Length (a) and width (b) of achene; width
s (d).
tube. —E. Length of style (a), style branch (b), and ate
ern species, such as K. indica and K. shimadai,
bloom for a longer period from May to November.
Achenes begin to mature in most areas by late Sep-
tember. The main stem dies back in winter and the
plants perennate from underground stems and ad-
ventitious buds.
STEMS AND BRANCHES
All species have erect and herbaceous stems 10-
200 cm tall. The stems are usually ridged and
greenish, sometimes purplish in K. indica, and cov-
ered with ascending, nonglandular, uniseriate-mul-
ticellular hairs interspersed with biseriate glandu-
Volume 84, Number 4
1997
Gu & Hoch
Systematics of Kalimeris
769
Figure 7. Perennating structures and cross section of a receptacle. —A. Stoloniferous rhizome (K.
yomena). —B. Condensed rhizome (K. lautureana subsp
—D. Cross section of a receptacle showing alveolar Pieds
lar hairs. The hairs are deciduous or restricted to
the ridges on the lower stems and become denser
upward. The vigorous growth of branches begins
when the main stem is terminated by a capitulum.
The angles between the branches and the main
stem are usually less than 45°, with the exception
of K. incisa subsp. macrodon and K. indica subsp.
collina, in which the angles are greater than 45*.
Tertiary branches are formed when the secondary
branches are terminated by a capitulum.
ROOT SYSTEM AND UNDERGROUND PERENNATING
STRUCTURES
All species have a fibrous root system, with the
exception of K. i folia, which has a tap-root
system. Stoloniferous rhizomes, i.e., underground
yomena
po tureana). —C. Adventitious buds (arrows, K. integrifolia).
stems with long internodes, are formed prolifically
in K. indica, K. pinnatifida, and K. yomena during
the flowering season and are characterized by in-
ternodes 5-10 mm long, with whitish scales at the
nodes (Fig. 7A). Condensed rhizomes, usually with
thicker stems and shorter internodes, are found in
K. incisa, K. mongolica, K. lautureana, and K. shi-
madai. They sometimes have almost indistinguish-
ably short internodes, and usually form big root-
stocks (Fig. 7B). Kalimeris integrifolia is
characterized by formation during the flowering
season of adventitious buds from the lateral roots
or near the top of the main root (Fig. 7C). These
rhizomes and adventitious buds either develop into
non-reproductive shoots in the same season, or re-
main dormant underground and grow out the fol-
lowing spring.
770 Annals of the
Missouri Botanical Garden
LEAVES
Leaf morphology varies greatly both among and
within species in this genus. During the rosette
stage, leaves are usually narrowly oblanceolate or
linear in Kalimeris integrifolia, and obovate, broad-
ly oblanceolate, or spathulate in other species, none
of which can be distinguished by this feature. The
margin is ciliate, serrate, crenate, or sometimes
deeply lobed on most species, but always entire in
K. integrifolia. The leaf apex is acute or obtuse or,
in upper cauline leaves, acuminate. The leaf base
is attenuate into a short petiole 5-20 mm long. The
indumentum is similar to that of the cauline leaves
of the same individual in most of the taxa, except
that K. lautureana and K. shimadai have longer and
denser nonglandular hairs on the rosette leaves
an on the cauline leaves. The rosette leaves grad-
ually wither as the stems mature. The cauline
leaves are obovate, oblanceolate, oblong, or linear,
and sessile. The leaves below the lowest branches
gradually wither after anthesis.
INFLORESCENCE
As in most genera of Astereae, the inflorescence
in Kalimeris is a radiate capitulum. The capitula
always terminate the stems and branches, and are
cymosely arranged. The corollas of the fertile pis-
tillate ray florets are whitish, light purple, or lilac
(but never yellow), and consist of a tube and an
oblanceolate ligule 5-28 mm long with three shal-
low distal notches. In the ray florets, the androe-
cium is absent or rarely wi nonfunctioning
anthers (or usually just remnant filaments) in the
tube; the gynoecium comprises an inferior ovary
with three vascular bundles and a single style with
two branches at the top. The lower portions of the
branches are stigmatic in function, and the style
always exceeds the corolla tube at receptivity.
The disc florets are bisexual, fertile, and always
yellow. The corolla throat is campanulate, abruptly
narrowed at the base into a tube, and 5-lobed dis-
tally. The lobes are triangular or broadly lanceolate
and usually %—% the length of the throat. The an-
droecium is syngenesious, i.e., the five anthers are
united laterally and their filaments are distinct. The
filaments are glabrous and adnate to the corolla
tube alternate with the lobes. The anthers are trun-
cate distally with triangular appendages. The gy-
noecium is the same as that of the ray florets except
that the ovary has two lateral vascular bundles
PEDUNCLE
The peduncles in Kalimeris are 0.3-18 cm long.
They are usually covered densely with the same
kind of nonglandular hairs as on the stems, and in
K. indica, K. integrifolia, and K. shimadai are in-
terspersed with glandular hairs. There are 1-13
bracts along the peduncles. The number of bracts
varies both among and within species. The bracts
on the lower peduncles are similar to the leaves on
the branches, and those on the upper peduncles are
similar to the phyllaries of the outermost whorl of
the involucre.
INVOLUCRE AND PHYLLARIES
The involucre is hemispheric or campanulate
and composed of 3—4 whorls of imbricated re
ies. Generally, it is 4.5—13 mm wide and 3.5-9
long, although smaller involucres are found in Kal.
imeris integrifolia, and larger ones in K. mongolica.
The size of the phyllaries increases inward. The
smallest phyllaries, those in the first whorl, are tri-
angular, lanceolate, or oblanceolate, and usually
herbaceous. The phyllaries in the middle whorl(s)
are oblong, obovate, or spathulate, and have mem-
branous margins. Some individuals of Kalimeris
mongolica have very broadly spathulate middle
phyllaries with broad, membranous margins. The
texture of the phyllaries varies from herbaceous (K.
incisa) to almost chartaceous (K. integrifolia). In
other species, the phyllaries are herbaceous on the
upper green portion and chartaceous on the lower
portion. The longest phyllaries, those of the inner
whorl, are oblong, oblanceolate, spathulate, or
sometimes linearly oblong, or linearly oblanceolate.
Phyllaries in all whorls have ciliate and sometimes
purplish margins.
RECEPTACLE
All species in Kalimeris have convex or subcon-
ical and alveolate receptacles 1.1-2.8 mm wide and
0.5-2.2 mm high. The alveolate appearance is
caused by the outgrowth of the epidermal cells and
a few layers of cells beneath them around the at-
tachment of achenes (Fig. 7D).
INDUMENTUM
The nonglandular trichomes on the leaves and
stems comprise multicellular and uniseriate hairs.
Both the type and the density of these hairs have
been used in distinguishing species within the ge-
“nus. For the convenience of this @iscussion, they
are termed thin or thick ascending hairs and ap-
pressed hairs. The thin ascending hairs are com-
posed of 4-6 long rectangular cells slightly swollen
at the top and bottom, and without a prominent
base (Fig. 8A); the thick ascending hairs are com-
Volume 84, Number 4
1997
Gu & Hoch 771
Systematics of Kalimeris
E F G
XR
Led ГВ
8
н |
Types of trichomes. —A. Thin ascending nonglandular hair (К. shimadai). —B. Thick ascending поп-
gure 8.
А аи hair d. ins коле indica). —C. Prostrate nonglandular
Ме Biseriate ораи шаг hair (К. indica su
ate, two-cell-tipped glandular hair (K. indica subsp. indica). —H.
hair (K. indica i
tipped анти pei (K. inte,
hair (K. integrifolia). —D. 4-celled nonglandular
bsp. indica). —F. Biseriate, one-cell-
ia). —G
А two-cell-tipped, навь айне чеч (К. pinnatifida). —1. Uniseriate glandular hair (К. yomena subsp.
na).
posed of 2—4 short rectangular cells and a promi-
nent base of several epidermal cells (Fig. 8B); and
the appressed hairs are composed of 3-5 rectan-
gular cells without prominent bases (Fig. 80).
Thick ascending hairs are scattered on the leaves
of Kalimeris indica subsp. indica, K. incisa, K. pin-
natifida, K. mongolica, and K. yomena and densely
cover the leaves of K. indica subsp. collina. Kali-
meris indica subsp. stenolepis and K. shimadai are
characterized by thin ascending hairs densely cov-
ering the leaves. The leaves of K. integrifolia are
densely covered with appressed hairs.
The nonglandular hairs on the achenes are 4-
celled, biseriate, and ascending (Fig. 8D) and are
found in all species except Kalimeris integrifolia.
In K. indica, the basal cells of these hairs are usu-
ally brownish. The nonglandular hairs on the co-
rolla are biseriate and multicellular (Fig. 8E).
There are two kinds of stipitate glandular hairs
on leaves. One type has several flat cells at the base
in two series and one large flat secretory cell at the
tip (Fig. 8F), and the other type has several cells
in two series at the base and two swollen secretory
cells at the tip (Fig. 8G). Both types are found on
the leaves of Kalimeris indica, K. integrifolia, K.
innatifida, and K. shimadai. Two somewhat differ-
ent kinds of glandular hairs are found on the corolla
of ray and disc florets of all species in the genus.
One type is a biseriate hair with a long stalk and
two secretory cells at the tip (Fig. 8H), and the
other is a uniseriate hair with one large secretory
cell at the tip (Fig. 8I).
PAPPI
Pappus morphology has been used as one of the
main characters in previous infrageneric classifi-
cations of Aster sensu lato. The pappus bristles of
Kalimeris are composed of elongated, lignified
cells; they are usually 2-12 cells thick at the base,
and multicellular in length. They tend to unite at
the base in K. integrifolia, K. lautureana, and K.
shimadai, but are separated in the rest of the spe-
cies. The pappus bristles range from 0 to 1.5 mm
long, with K. incisa and K. mongolica having par-
ticularly long bristles (0.6-1.5 mm long); тү
individuals are often found in К. indica subsp. in-
dica, and rarely in K. shimadai and K. o
Some individuals of K. incisa, K. integrifolia, K.
mongolica, and K. shimadai have ray pappi shorter
than disc pappi.
772 Annals of the
Missouri Botanical Garden
ACHENES abundantly from the Amur Basin to Korea and cen-
The achenes are laterally compressed, obovate
or broadly obovate. The disc achenes have two lat-
eral ribs (vascular bundles) or occasionally three.
The ray achenes in all species of Kalimeris are
3(4)-ribbed, with 2(—3) ribs abaxially and 1 adax-
ially (e.g., Fig. 3), and are isosceles-triangular in
cross section. The achenes are purple or dark pur-
ple in K. indica, K. pinnatifida, K. shimadai, and
K. yomena, and gray-green, straw-colored, or
brownish in K. incisa, K. integrifolia, K. lautureana,
and K. mongolica. The achenes have a whitish mar-
gin 0.1-0.4 mm wide. Glandular hairs are found at
the top of the achenes commonly in K. indica, K.
integrifolia, K. pinnatifida, and K. yomena, and oc-
casionally in some individuals of K. incisa, K. lau-
tureana, K. mongolica, and K. shimadai
CYTOLOGY
The basic chromosome number of Kalimeris is x
= 9. All eight species recognized in this treatment
were examined cytologically (Gu, 1989; Gu & Tara,
1990). Two tetraploid, a hexaploid, and three 12-
ploid taxa were counted for the first time in this
genus.
GEOGRAPHIC DISTRIBUTION
Kalimeris is native to East Asia, ranging from the
Amur Basin of northeastern China and adjacent
Russia, Korea, and Japan, through central and
southern China to northern Vietnam, Thailand, and
Burma. Most species have more limited ranges
within this region.
Kalimeris incisa and K. mongolica have the
northernmost distribution, occurring in the humid
temperate region along the Amur and Ussuri Rivers
and reaching Blagoveshchensk in Russia, the
northeastern limit of the genus. Both species also
occur in Korea, K. incisa from the Chinese border
to Cheju Island of South Korea and through the
western plains regions. Kalimeris mongolica is here
correctly attributed to Korea for the first time; prior
to this, it was misidentified as K. lautureana (e.g.,
in the Flora of Korea; Nakai, 1911). Kalimeris lau-
tureana is endemic to China and has a narrower
geographic distribution than usually has been at-
tributed to it, due to taxonomic confusion with K.
mongolica. Kalimeris shimadai is also endemic to
China and occurs abundantly on the Loess Plateau
and scattered throughout subtropical central and
southern China, extending to northern Taiwan.
Kalimeris integrifolia and K. indica are the two
most widely distributed species. The former occurs
tral China, with its southern limit in Jiangsu and
Anhui provinces, and its southwestern limit in Si-
chuan and Yunnan provinces. It is sympatric with
several species: K. incisa and K. mongolica in the
northern part, and K. indica and K. shimadai in the
southern part. Kalimeris indica occurs in central
and South China and northern Taiwan, and extends
to the Ryukyu Islands and southern Kyushu of Ja-
pan, and to southern Korea. It reaches its southern
limit in the plains regions of northern Vietnam and
Burma, and its northern limit at the northern base
of the Qinling Mountains and the southern edge of
the Loess Plateau in the Weihe and Fenhe Valleys,
where the original warm temperate forest-steppe
vegetation has been mostly removed by cultivation.
t is recorded from India, Indonesia (Java, Suma-
tra), and Malaysia (Malacca), where it may have
been cultivated originally for its edible rosettes,
medicinal uses, or ornamental purposes (Miquel,
1856), but it escaped from cultivation and has be-
come naturalized in these areas. The recent collec-
tion from Oahu, Hawaii Islands, also certainly re-
flects an introduction, perhaps through commercial
horticulture or by accident.
Japan has two endemic species, Kalimeris pin-
natifida and K. yomena. The former occurs in
southern Hokkaido, extending south to the main
island of Honshu, with its southern limit in north-
ern Kinki District (Kyoto, Nara, and Mie). Kalimeris
yomena is distributed from Niigata of northern Chu-
bu District to Shikoku, Kyushu, and the Ryukyu
Islands.
ALTITUDINAL DISTRIBUTION
All of the species of Kalimeris are found at ele-
vations below 900 m, but some species can also
grow as high as 2600 or 3100 m (K. lautureana or
K. indica, respectively) in the mountains and pla-
teaus of central and southwestern China.
HABITAT
All of the species of Kalimeris grow in relatively
disturbed habitats. They are basically heliophytes
and are usually associated with human activities.
They grow in sunny and dry places along roadsides,
at the edges of crop fields, or on abandoned lands.
Sometimes they are found in relatively mesic areas
on the floor of open forests. For example, K. incisa
and K. mongolica grow inside the temperate sec-
ondary-growth forests in the northern part of the
range of the genus, and at low elevations in mon-
tane regions in meadows and at the margins of
mixed coniferous and broad-leaved forests. Kali-
Volume 84, Number 4
1997
u & Hoch
Systematics of Kalimeris
meris indica subsp. stenolepis grows along margins
of subtropical mixed deciduous and evergreen
broad-leaved forests in the southern part of the
range. Kalimeris indica has the broadest ecological
range in the genus, occurring from cool, humid
temperate to warm subtropical regions at margins
of cliffs, in poor, often disturbed soils, at margins
or inside of open forests and thickets, or near crop
fields. Kalimeris integrifolia is usually found in rel-
atively disturbed areas, and especially when it is
sympatric with other Kalimeris species, it always
occurs in more disturbed habitats.
SYSTEMATICS OF KALIMERIS
Kalimeris (Cass.) Cass., in F. Cuvier, Dict. sci. nat.
37: 464, 491. 1825. Aster subg. Kalimeris
Cass., in F. Cuvier, Dict. sci. nat. 24: 324.
1822. Kalimeris sect. Kalimeris Ling, in Y.
Ling et al., Fl. reipubl. popularis sin. 74: 99.
1985. TYPE: Kalimeris platycephala Cass. ex
Nees, nom. illeg. [= K. incisa (Fisch.) DC.].
Asteromoea Blume, Bijdr. 907. 1826. Boltonia sect. Aster-
omoea (Blume) Benth. in Benth. € Hook. f., Gen.
pl. 2: 269. 1873. Aster sect. Asteromoea (Blume) Ma-
kino, Bot. Mag. (Tokyo) 12: 60. 1898. Kalimeris sect.
Asteromoea (Blume) Kitamura, Mem. Coll. Sci. Kyoto
Imp. Univ., Ser. B, Biol. 13: 305. 1937. TYPE: As-
teromoea indica (L.) Blume [= K. indica (L.) Sch.
Hisusua гђе, Prodr. 6: Hes isutsua can-
is (Lour.) DC. [= K "petis à) ewe Bip.].
Маш) о жељу Bull. Acad. Int. Géogr. Bot. 12: 31. 1903.
YPE: Martinia polymorpha Vaniot [= K. indica (L.)
Sch. Bip.].
Perennial erect herbs. Root system fibrous or
tap-root; perennating by condensed or stoloniferous
rhizomes and adventitious buds from main or lat-
eral roots. Stem forming from rosettes prior to flow-
ering, (10-)20-200 cm tall, cymosely branched,
ridged, covered with ascending or appressed, uni-
seriate nonglandular hairs, these denser along the
prominent ridge lines, sometimes interspersed with
biseriate or uniseriate, glandular hairs; the lower
part usually glabrous and not ridged, or rarely with
scattered nonglandular hairs after anthesis; branch-
es up to 48 cm long, the angle between the branch-
es and the stem usually less than 45° or sometimes
greater than 45°. Rosette leaves oblong, oblanceo-
late, or broadly obovate, less often narrowly oblan-
ceolate, (3.5-)4.5-11 х (1.2-)2.5-4.4 cm, glabrous
to densely covered with nonglandular hairs, the
apex acute or obtuse, the margin ciliate, remotely
crenate, serrate to deeply serrate, or sometimes en-
tire, the base attenuate or reduced abruptly to pet-
iole 0.5-2 cm long, withering after the stem has
formed; cauline leaves alternate, linear, narrowly
oblong, or oblanceolate to obovate, the margins cil-
iate, serrate to deeply pinnate-lobed, or less often
entire; gradually reduced in size upward, those on
the lower stem usually withering after anthesis, the
upper ones (0.7-)3.5-11(-15) X (0.2-)0.5-2.5(-
3.5) em, glabrous to densely covered with thin or
thick multicellular, nonglandular hairs and inter-
spersed with biseriate, 1—2-cell-tipped glandular
hairs, usually denser on the abaxial surface or
sometimes restricted to the veins, the apex acumi-
nate, acute, or obtuse, the base attenuate or cune-
ate, sessile. Capitula (5—)10-120(-200), terminat-
ing the stems or branches, loosely cymosely
arranged; peduncle (0.3-)2.5-12(-18) cm long,
often densely covered with ascending, thin, multi-
cellular, nonglandular hairs and interspersed with
biseriate, glandular hairs; bracts 1-10(-13) along
the peduncle, the lower ones leaf-like, narrowly ob-
long, oblanceolate, or lanceolate, entire, 6.1-16(-
30) X 0.7-3.2(-8) mm, the upper ones phyllary-
like, linear to oblanceolate, 1.2—4.5(-8) х 0.2-
1.4(-2.5) mm; involucre hemispherical or campan-
ulate, 4.5-13 mm broad, 3.5—6.5(—9) mm high. In-
volucral phyllaries in 3-4 loosely or tightly
imbricated whorls, gradually increasing in size in-
ward, the distal part of phyllaries usually herba-
ceous and green, the proximal part chartaceous and
whitish, the abaxial surface glabrous to densely
covered with nonglandular hairs and sometimes in-
terspersed with biseriate, glandular hairs, the ad-
axial surface usually glabrous or rarely with scat-
tered, thin, nonglandular hairs, the margins ciliate
and membranous, sometimes purplish; phyllaries of
the outermost whorl lanceolate or obovate, (1.5—)
5-5(-7.2) X 0.5-1.8(-2.8) mm, the apex acute,
obtuse, or sometimes acuminate (K. integrifolia);
those of the middle whorl(s) oblong, obovate, or
spathulate, 2.3—6.4(-8.1) х (0.7-)0.9-3(-3.5) mm,
the apex acute, obtuse, or acuminate; those of the
inner whorl(s) oblong, obovate, or spathulate, (3—)
4-7(8) X (0.6—)0.8-2.5(-3.2) mm, the apex acute,
obtuse or acuminate. Receptacle convex or occa-
sionally subconical, alveolate, 1.1-2.8 mm wide,
0.5-2(-2.2) mm high. Ray florets pistillate, con-
fined to the periphery of the capitulum, 10-30;
corollas whitish to light purple, ligules narrowly ob-
long, oblanceolate, or less often elliptical, (5.2—)
6.6-22(-28) X 1.1-4.5(-5.2) mm, the apex obtuse
with three very small notches, the tube 0.7-2.6 mm
long, covered with biseriate nonglandular and glan-
dular hairs; style exceeding the corolla tube, 1.3—
3.4(—4.5) mm long, glabrous, the two style branches
linear, (0.4—)0.6—1.1(-1.5) mm long. Disc florets bi-
sexual, yellow, (30-)45-115; corollas (2.1—)2.5—5(-
774
Annals of the
Missouri Botanical Garden
5.8) mm long, the throat campanulate, with 5
broadly lanceolate or triangular, equal, subequal,
or unequal lobes i (0.4—0.7-1.8(-2.2) х 0.4-
et ича
0.6-2 mm long, 4—% as wide as the throat, covered
with biseriate glandular hairs; androecium synge-
nesious, with five anthers 0.7-1.8(-2.2) mm long,
each with a triangular or broadly lanceolate ap-
pendage 0.2-0.5 mm long; style ibid,
the two branches linear, (0.4—)0.6—1.2(-1 -6) mm
inal stigmatic li
, whitish to
Ne bristles (0-)0.1-0.8(-
1) mm long; disc bristles (0—)0.1-0.8(-1.5) mm
long. Achenes straw-colored to dark purple, or
greenish, obovoid to broadly obovoid, compressed
laterally, loosely covered with biseriate, 4-celled,
hairs and interspersed with biseriate,
glandular hairs apically, or with only glandular
hairs at the top; the margins light-colored, ciliate,
thickened, 0.1-0.4 mm wide; ray achenes 3(—4)-
ribbed, 1.5-3.2 X 0.6-1.5(-2.2) mm; disc achenes
x Gib 1.5-3.2(-3.5) X (0.7-)1.1-2.1(-2.9)
number: x = 9, with aneuploids
ан Had els
Phenology. Flowering period from late May to
November.
Typification of this generic name is problematic.
When Cassini published Aster subg. Kalimeris in
1822, he ascribed to it only one species using the
illegitimate combination Kalimeris platycephala. In
1825, he elevated this subgenus to a genus, Kali-
meris (Cass. in F. Cuvier) Cass. in F. Cuvier, but no
species was transferred (see King & Dawson,
1975). Unfortunately, no specimens that were pos-
sibly used by Cassini have been located. Kalimeris
platycephala Cass. in F. Cuvier ex Nees (1832) was
validly published but with synonymy including ear-
lier species names, e.g., Grindelia incisa (Fisch.)
Spreng., Aster tataricus L. f. (as A. tartaricus), A.
incisus Fisch., and A. sibiricus Hort. plurr. It is clear
that K. platycephala was an illegitimate name then
because all other specific names included in the
synonymy were earlier, validly published names.
The correct name Nees should have used for that
species was K. tatarica, A. tataricus L. f.
(1782) is the oldest name (Article 11.4, Interna-
omenclature, Greuter et
originally described as K. platycephala and thus are
in conflict with it, and they both are treated as spe-
cies of Aster by contemporary botanists (Kitamura,
1937b; Tamamshjan, EE Ling et al., 1985). Be-
ified in this treat-
designation of the type of the genus Kalimeris is
followed here.
KEY TO THE SPECIES OF KALIMERIS
la. Disc pappus bristles more than 0.6 mm long.
lar hairs
Та. Pappus bristles 0-0.3 mm
long, or, if 0.4 mm, leaves
densely covered with
dular and glandular hairs
adaxially; widely жиы
ndi
Tb. Pappus bristles he mm
te leaves with scattered
hairs adaxially;
native in Japan ....... 5. K. yomena
6b. Disc pappus bristles 0.4-0.6 mm
re rarely wanting, the bases unit-
rhizomes condensed.
thin "pet, nonglandular ——
hairs _ _ 6. K. shimadai
8b. тањи i entire to veut lobed,
glabrous or sparsely covered
with thick, ascending, non-
glandular hairs _ 7. K. lautureana
1997
би 4 Носћ 775
Systematics of Kalimeris
1. Kalimeris incisa (Fisch.) DC., Prodr. 5: 258.
1836 ["Calimeris"]. Aster incisus Fisch., Mém.
Soc. Imp. Naturalistes Moscou 3: 76. 1812.
Grindelia incisa (Fisch.) Spreng., Syst. veg. 3:
incisa (Fisch.) Koidz., Bot. Mi. (Tokyo) 37:
56. 1923. TYPE: Russia.“Aster incisa, Sibi-
ria,” “Fisch. = Calimeris i. DC." (lectotype,
here designated, LE).
Kalimeris plat ees, Gen. sp. Aster. 226.
1832 rey TYPE: Bonis € Magers
пеогуре,
types, BM, ds. K, Ws
Root fibrous; rhizomes condensed, forming root-
stocks up to 3 cm diam. Stem 30—120 cm tall, cov-
ered with ascending, uniseriate, nonglandular
hairs. Cauline leaves narrowly oblong to obovate,
serrate, crenate, or deeply pinnate-lobed, rarely en-
tire, (2.6-)3.6-8(-10) X 0.7-2.4(-3.4) cm, glabrous
or with scattered ing nonglandular hairs
adaxially, sparsely to evenly pubescent abaxially
(mostly along the veins). Capitula 10—55; pedun-
cles (1.8—)4—14(-18) cm long; bracts 1-9 along the
peduncle, the lower ones narrowly to linearly ob-
long, 8-26 X 1—3(-6) mm, the upper ones linear,
(2.1-)3.4-8 х 0.3-1.4(-2.5) mm; involucre hemi-
spherical, 7-11 mm broad, 5-7.5 mm high. Phyl-
laries in 3-4 whorls, imbricate, covered with short
hairs or glabrous, the upper 424 part of the phyl-
laries or sometimes the entire phyllaries herba-
ceous and green, the lower part chartaceous, the
margins ciliate and membranous, sometimes pur-
plish; those of the outermost whorl narrowly #0188
or lanceolate, (2.5-)3.5-5(-5.2) X (0.7-)0.9-1.4
mm, apex acute, those of the middle whorl(s) ob-
lanceolate, (4~)4.2-5.5(-6.4) X 1.6(-2) mm, the
apex acute, those of the inner whorl oblanceolate
to oblong, (4.3-)4.7-6(-6.5) X 1.1-1.8(-2.2) mm,
the apex acute. Receptacles convex or subconical,
(1.1-)1.5-2.4 mm wide, (0.7-)1.2-2.1 mm high.
Ray florets (13-)16-27(-29); corollas pale lilac to
pale purple, ligules narrowly oblong or oblanceo-
late, (8.5-)11.5-20(-22) х (1.8-)2-2.8(-3.2) mm,
the tube (0.9-)1.1-1.6 mm long; style (1.6-)1.9-
2.8 mm long, the branches 0.7-1.2 mm long. Disc
florets (40-)58-115; corollas 3-4(4.2) mm long,
the lobes unequal, the longer one 0.9-1.6(-2) mm
long, the shorter ones 0.6—1.2(-1.5) mm long, both
0.3-0.6 mm wide; tube (0.6—)0.9—1.3(-1.5) mm
long and usually covered with biseriate, nonglan
dular or glandular hairs; anthers 0.9—1.4(-1.6) mm
1
long; style 2. 5-4.3 mm long, the branches 0.8-1.3
mm long, each with a triangular appendage 0.3-0.4
mm long. Ray and disc pappus bristles whitish to
light brown, sometimes unequal; ray bristles (0.2—)
0.5—0.8(—1) mm long; disc bristles (0.5-)0.6-1.1(-
1.5) mm long. Achenes greenish to straw-colored,
obovate to broadly obovate, covered with 4-celled,
biseriate, nonglandular hairs or these sometimes in-
terspersed with biseriate, glandular hairs at the top,
the margins 0.2-0.4 mm broad; ray achenes 3-
ribbed, 2.6-3.1 X 1.1-1.5 mm; disc achenes 2-
ribbed, 2.6-3.1 X 1.4-2 mm. Chromosome num-
ber: n = 9. Figure 9A—D.
mes ыс Flowering period: late June to Octo-
Р rom the two specimens Tamamshjan (1959) зе-
lected as the type of Kalimeris incisa, the one with
the annotated label “Aster incisa, Sibiria” and an
added note “Fisch. = Calimeris i. DC” is chosen
as the lectotype for K. incisa here. The type spec-
imen of K. platycephala was based on a cultivated
strain labeled as Aster sibiricus from the “Jardin du
Roi,” now the Muséum National d’Histoire Natu-
relle in Paris. No authentic specimens have been
located. In order to follow the typification of the
genus Kalimeris made by the first author, Cassini
in this case, a well-duplicated specimen collected
by F. Karo is selected as the neotype of K. platy-
cephala here (Greuter et al., 1994: Article 9.6 and
9.11).
This species has the most northeastern distri-
butional range in the genus and is divided into two
subspecies: subsp. incisa and subsp. macrodon.
KEY TO THE SUBSPECIES OF KALIMERIS INCISA
la. Branches usually diverging from the stem by less
than 45°; leaves oblanceolate to narrowly oblong,
serrate, or rarely entire; ray corolla Е mm
long a. subsp. incisa
1b. codo» usually diverging from the stem
more than 45*; leaves nad > elliptic, crenate;
ray corolla 8.6-15.5(-16) mm gs УВА
b. subsp. macrodon
la. Kalimeris incisa subsp. incisa
Aster incisus var. australis Kitag., Rep. Inst. Sci. чач
€— f 323. bre Kalimeris incisa var. a.
itag., Neo-lin. fl. Manshur. 652.
1979. TYPE: China. P Wui-hu-lin Railway Sta-
tion, 2 Sep. 1936, M. Kitagawa s.n. (holotype, ТТ).
Stems 40—120 cm tall; branches ascending at an
angle from the stem less than 45*. Cauline leaves
oblanceolate or narrowly oblong, serrate, less often
crenate, or rarely entire, 4.7-7(-10) X (0.7-)0.8(-
Annals of the
776
Missouri Botanical Garden
Моште 84, Митбег 4
1997
Gu & Hoch 777
Systematics of Kalimeris
à
F dem
у
ES
Figure 10. Geographical distribution of Kalimeris in-
cisa subsp. incisa (dots) and subsp. macrodon (star, indi-
cated by arrow).
2.2) cm, with scattered nonglandular hairs on both
sides or glabrous adaxially and loosely hairy abax-
ially. Capitula 10-55; peduncles (1.8-)4-10(-18)
cm long; bracts 3-9 along the peduncle, the lower
ones 8-17(-23) X 1-1.6(-6) mm, the upper ones
2.1-5.4(-8) X 0.3-1.4(-2.5) mm; involucre hemi-
spherical, 7-11 mm broad, 5-6.5 mm high. Phyl-
laries in 3 whorls, loosely imbricate; those of the
outermost whorl (3.1-)3.5-5.2 Xx (0.7-)0.9-1.3
mm, those of the middle whorl(s) 4.2—5.8(—6.4) X
1.1-1.6(-1.9) mm, those of the inner whorl 4.9-6(—
6.5) X 1-1.6(-1.8) mm. Receptacles convex or
subconical, (1.4—)1.8—2.3 mm wide, (0.8-)1.2-1.8
mm high. Ray florets (13-)16-27; corollas pale li-
lac, ligules 15.8-22 x (1.8-)2.1-3 mm, the tube
(0.8—)1—1.6 mm long; style (1.6-)1.9-2.8 mm long,
the branches 0.7-1.2 mm long. Disc florets (40—)
71-115; corollas 3—4(-4.2) mm long, the lobes un-
equal, the longer one 1-1.7(-2) mm long, the short-
er ones 0.5-1.1(-1.5) mm long, both 0.3-0.6 mm
wide, tube (0.6-)0.9-1.2(-1.4) mm long; anther
0.9-1.4(-1.6) mm long; style (2.1—)2.5—4(—4.3) mm
long, the branches 0.8-1.3 mm long. Ray and disc
pappus bristles equal or sometimes unequal; ray
bristles (0.2-)0.5-1 mm long; disc bristles 0.5—
1.2(-1.5) mm long. Achenes straw-colored, the
margins 0.2-0.4 mm wide; ray achenes 2.6-3.1 X
1.4—1.5 mm; disc achenes 2.6-3.1 X 1.4-2.2 mm.
Chromosome number: п = 9. Figure 9А-С.
Phenology. Flowering period: late June to Octo-
b
er.
Distribution (Fig. 10). Amur and Ussuri basins
(southern Far East region of Russia, possibly in
eastern Siberia but without accurate localities) to
the Daxing'anling Mountains in Heilongjiang and
to Nei Mongol Zizhiqu, and through Jilin and
Liaoning provinces, southward to Korea in the fol-
lowing provinces: Hambook, Hamnam, Pyong-
Book, Hwanghe, Gyong-gi, Seoul, Choongbook,
Choongnam, Gyong-nam, Boosan, and Junnam.
Most populations occur from sea level to about 900
m elevation. This taxon is fairly common and grows
in relatively mesic, semi-open secondary-growth
forests, in meadows, and along margins of mixed
coniferous and broad-leaved forests. It is rarely
found along dry roadsides, where the populations
are very small, sometimes comprising only one or
a few individuals.
Kalimeris incisa subsp. incisa is often sympatric
with К. integrifolia and К. mongolica, and all three
have been found growing side by side with very
slight habitat differentiation. Sometimes they can
be found mounted together under the same collec-
tion number (e.g., James s.n., May-Aug. 1886, K).
Usually, K. incisa subsp. incisa grows in relatively
mesic and protected areas, e.g., under trees,
shrubs, or among taller herbs, whereas K. integri-
folia and К. mongolica occur in drier and more
exposed places. It seems that K. incisa subsp. in-
cisa and K. integrifolia do not hybridize with each
other, because no intermediates have been found in
areas of overlap. Intermediates between K. incisa
subsp. incisa and К. mongolica (e.g., Kitamura s.n.,
15 Sep. 1939, KYO) are rarely found in the areas
where populations of the two taxa occur together.
This subspecies tends to grow more vigorously
in the northern part of its distributional range. In-
dividuals in the Amur basin are more robust than
those in the southern part of the range. Based on
this observation, Kitagawa (1936) gave varietal sta-
ee
Figure 9. Kalimeris incisa and K. pinnatifida. AC. К.
Achene. —D. Achene of K. incisa subsp. macrodon. E,
rhizomes.
incisa subsp. incisa. —A. Upper branch. —B. Root. —C.
F. K. pinnatifida. —E. Achene. —F. Root and elongated
778
Annals of the
Missouri Botanical Garden
tus to the southern populations, as Kalimeris incisa
var. australis. Kitamura also annotated herbarium
specimens from Tumenling, China, as “Kalimeris
tewakiana,” and noted on one of the labels that
the plants were smaller than typical K. incisa and
were cultivated at Kyoto University for further stud-
ies. It seems that the studies on these plants were
not completed because neither the name K. tate-
wakiana nor any discussion on this subject has
ever been published. There are many intermediates
between these two forms, suggesting a clinal pat-
tern, and because there are no other differences
correlated with size, these extreme forms are re-
garded as a single taxon in the present treatment.
The leaf morphology of this subspecies is very
variable. Some individuals have almost entire
leaves, whereas others have deeply pinnate-lobed
leaves. On Mt. Qianshan, Liaoning, China, individ-
uals with narrowly oblanceolate and broadly oblan-
ceolate leaves grow together in the same population
(H. Gu & W. Zhang 279, 280, MO). The length of
the pappus bristles tends to be reduced in popu-
lations at the southern part of the geographic range,
especially in South Korea. Generally, Korean plants
tend to have shorter pappus bristles and more slen-
der branches than those found on the continent.
Several specimens (e.g., Horsfield 788, K, BM)
from Java with Miquel’s annotation “Aster indicus B
horsfieldii” belong to K. incisa subsp. incisa, and
were most likely cultivated there. In any case, Aster
indicus В horsfieldii was never published. One
specimen, W. Souder 253 (BM) from “Rochester,
North America,” is either mislabeled (as noted on
the sheet) or was collected from cultivated plants,
since there are otherwise no collections known from
North America.
Additional specimens examined. RUSSIA. rudi Bla-
Karo 182 (BM, E, K, G, P). Primorski:
Saberkin 613 (NY), 770 (NY). Siberia
*Dahuria," Fischer 1 17 (О),
Turczaninow s.n. (GH); *Amur River," Max
).
3592 (ТІ); Togwon, Kitamura ѕ.п. (KYO
Kitamura ѕ.п. (KYO). Pyong-Book: Gangge, Mills 136
(ТІ), 145 (TD, 410 (TI); Yongsandong, Nakai 2785 (ТІ).
Hwanghe: Mt. Jangsoo, Koidzumi s.n. (KYO). Gyong-Gi:
Suwon, Sakata 498 (KYO). Seoul: Changduk Palace,
Okada s.n. (TI); Mt. Bookhan, Uchiyama s.n. (TI); Gwan-
grung, Lee et al. 6826 (TI), Nakai 13218 (TI); Mangwoori,
Seoul, Chung 9288 (TNS); Mt. Nam, Faurie 393 (KYO,
P), 403 (KYO), 404 (P), ded, e ‚ (TD; Seoul, Dann 54
(BM), Bodinier s.n. (E). Choongbook: rai City, Lee
et al. 6627 (TI). Ch : Gongjoo- ryong-
туо imura s.n. (KYO). Going не: Chinju. Uno
22976a (GH); Mt. Jili, Okamoto 16067 (KYO), 16068
(MAK). Boosan: Dongre, Kitamura s.n. (KYO). J m:
Kwangju, Mt. Kwang, wir 473 (TNS), 3107 > (KYO.
TNS). CHINA. “Manchuria”: Amur River, Mt. Burejae,
Radde 86 (P); “Chara,” ees 1507 (BM, P); Xing’an
сы Dzhalantun, Litvinov 2556 (NY), Karo 1507 (К, Р).
ngjiang: Acheng Xian, Yichuomao-shan, Baimaozi,
G. Wang 498 (IFP); Boli Xian, Hongqi Forestry Station,
Y. Zhang et al. 2117 (IFP); Mao’ershan Xian, Xingshi, 6.
Wang & Q. Li 26 (IFP); Mishan Xian, Hulin, Heilongjiang
Expedition Team 142 (IFP), 151 (IFP), 205 (IFP); Ning’an
Xian, Shihuan-zheng, Nanhutou, У Zhang et al. 1226
(IFP); Suiling Xian, aay Maguro s.n. (KYO); Sun-
wu Xian, Siji-dun i & С. Lin 1483 (IFP), 1499
(IFP); Yanxing Kian, fro mag pele to Xingdongcun, C. S.
Wang 128 (IFP), 172 (IFP). Nei Mongol: Bugt, Kitagawa
s.n. (TI); Ningcheng Xian, Cunjingou-gongshe, Songshul-
ing, 200 m, S. Li et al. 4655 (IFP); Solon Qi, Honggoli,
S. Liu et al. 8298 (IFP, PE); Xing’anling Mts., Ishidoya
s.n. (KYO). Jilin: Antu Xian, 285 m, S. Liu 3685 (IFP,
PE); Changbai-shan, Huapihe, 900 m, S. Liu et al. 1802
(IFP, PE); from Changbai-shan to TA James s.n.
G ; Fusong Xian, Donggang, m, S. Liu et al.
1282 (IFP, PE), 7363 (IFP, PE); pee Xian, Huancheng-
xiang, Qingshan, T. Wang 2465 (IFP), 2572a (IFP); from
Huapihe to Maton, гч ‚ Y. Zhang & S. Wang 657
(IFP); Hunchun Xian, nhua-xiang, Lanjiatangzi, T.
Wang et al. 2182 (IFP); Pm Shi, a bae Kitagawa
s.n. (ТІ); Xiaobai-shan, P. Fu et O (IFP); Taube
anto, ells, t Gus et al. 2259
)
han
(IFP); Jiutai Xian, Tumenling, Yamatsuta 99 (TNS), К.
);
900 т, J. Qian 850 (KUN), 955 а Tanghoko, James
s.n. (K); Wuihulin, Ma’an Shan, Kitagawa s.n. (TI);
Xing'anling Mts., Yamatsuta 87 (TNS), 93 (TNS); cor]
ngjiadian, T. Wang 1671 (IFP). Liaoning: Anshan
Qian- gud n, H. Gu & W. Zhang 276 ar т (МО), 280
(MO); Changbai-shan, Yuheng-shan, S. 760
Shenyang Shi, Donglin, H. Gu et al. 266a (MO), 270 (MO).
lb. Kalimeris incisa subsp. macrodon (Vaniot
& H. Lév.) H. Y. Gu, stat. et comb. nov. Aster
macrodon Vaniot & H. Lév., Bull. Acad. Int.
Géogr. Bot. 20: 141. 1909. TYPE: Korea. Che-
ju Island (Quelpart), 25 Sep. 1906, Faurie
1071 (holotype, E; isotypes, BM, KYO).
Stem 30—52 cm tall; branches ascending at an
angle from the stem greater than 45°. Cauline
leaves obovate to elliptical, crenate or less often
deeply serrate, 2.6-7.6 X 1-3.4 cm, glabrous or
with scattered nonglandular hairs adaxially. Capit-
ula (15-19-29; peduncles (2.8-)3.8-8.6 cm long,
bracts 2-5 along the peduncle, the lower ones 9-
12(-20) х 2.5-5.5 mm, the upper ones 4.7-7.5 X
1-1.8 mm; epe hemispherical, rarely cam-
panulate, (7-)8.5-10 mm broad, 5-6 mm hi
Phyllaries in 3-4 „ы imbricate; those of the
outermost whorl 3.2—4.5(—4.8) x 0.9-1.3 mm,
those of the middle whorl(s) (4-)4.5-6 X 1-2 mm,
those of the inner whorl 4.7-6.5 Х 1.6-2.2 mm.
Receptacles subconical, (1.1-)1.5-2.4 mm wide,
Volume 84, Number 4
1997
Gu & Hoch 779
Systematics of Kalimeris
(1.2-)1.6-2.1 mm high. Ray florets 19-29; corollas
pale lilac to pale purple, ligules 8.6-15.5(-16) х
2.2-3.2 mm, the corolla tube (1.1—)1.5 mm long;
style (1.5—)1.9—2.8 mm long, the branches 0.7-1.2
mm long. Disc florets (58-)68-112; corollas 3.34
mm long; the lobes unequal, the longer one 1.1-
1.8 mm long, the shorter ones 0.8-1.4 mm long,
both 0.5-0.6 mm wide, the tube 0.9-1.1 mm long;
anther 1-1.4 mm long; style 2.2-2.9 mm long, the
branches 0.9-1.3 mm long. Ray and disc pappus
bristles often brownish, unequal; ray bristles 0.2—
0.6 mm long; disc bristles 0.4-1 mm long. Achenes
straw-colored or greenish, the margins 0.1–0.3 mm
broad; ray achenes 2-2.8 X 1.3-1.5 mm; disc
achenes 2.2-3 X 1.5-1.9 mm. Chromosome num-
ber unknown. Figure 9D.
Phenology. Flowering period: July to October.
Distribution (Fig. 10). This subspecies is endem-
ic to Cheju Island, South Korea, the southern limit
of Kalimeris incisa, where it is sympatric with K.
indica subsp. indica. It occurs apparently from near
sea level to 800 m
This taxon may hybridize with Kalimeris indica
subsp. indica, as evidenced by some individuals of
K. incisa subsp. macrodon with shorter pappus bris-
tles.
Additional specimens examined. A. Cheju is-
: 800 m, Taquet 950 =: 951 ч е. 952 (С), 990
(Е, К), 991 (Е, TNS), 992 (TNS); “Hoabin,” 6236 (ТІ);
South Cheju, Nakai 6519 (TI); North Cheju, 6522 (TI).
2. Kalimeris pinnatifida (Maxim. ex Makino) Ki-
tam., Acta Phytotax. Geobot. 6: 50. 1937. Aster
быш var. pinnatifidus Maxim. ex Makino,
Bot. Mag. (Tokyo) 20: 41. 1906 [“pinnatifida”].
Aster pinnatifidus (Maxim. ex Makino) Makino,
in linuma, Somoku-Dzusetsu 4: 1106. 1912.
Aster incisus var. pinnatifidus (Maxim. ex Ma-
kino) Nakai, Bot. Mag. (Tokyo) 33: 215. 1919.
Asteromoea pinnatifida (Maxim. ex Makino)
Koidz., Bot. Mag. (Tokyo) 37: 56. 1923. TYPE:
Japan. Kanagawa: Yokohama, C. J. Maximow-
icz s.n. in 1862 (holotype, LE not seen; iso-
types, BM, GH, W).
Aster bp f. hortensis Makino, Bot. Mag. ee en
1912. Aster ну возе var. h
kino, in linuma, So EO ONRET 4:
lime nati ar. nsis Makino Kitam.,
Mem. Coll. Sci. Kyoto Imp Univ Ser. iol. 13:
309. 1937. TYPE: Ј Kochi: emo n, Sak-
awa-cho, 28 June 1887, T. Makino s.n. е
here тун aae MAK).
pinnati . albescens Makino, J. Jap. Bot. 7: 10.
1931. Kalimeris pinnatifida f. albescens (Makino) Ki-
tam., Mem. Coll. Sci. Kyoto Imp. Univ., Ser. B, Biol.
13: 309. 1937. TYPE: Japan. Iwate: Morioka, T. Ma-
kino s.n., s.d. (holotype, MAK).
Aster pinnatifidus f. caeruleus Makino, J. Jap. Bot. 7: 10.
1 f. caerulescens (Makino)
takawa-mura, S. Nemoto s.n., s.d. (holotype, МАК).
Root fibrous; rhizomes stoloniferous, 15-20 cm
long. Stem 56–130 cm tall. Cauline leaves oblong
to oblanceolate or less often broadly oblanceolate,
serrate to deeply pinnate-lobed, (3.4—)5—8(-10.5)
X (0.4—)1-2.5(-3.4) cm, glabrous or with scattered
nonglandular hairs adaxially, loosely or sometimes
densely covered with nonglandular hairs and with
scattered glandular hairs abaxially, lobes lanceolate
to triangular, 0.2—0.9 X 0.1-0.5 cm. Capitula 20-
88; peduncles (1-)2-8.5(-16) cm long; bracts 2—
6(-10) along the peduncle, the lower ones 10—25
X 1.5—5 mm, the upper ones 2.5-7.8 X 0.5-1.2
mm; involucre hemispherical or rarely campanu-
late, 7-10 mm broad, (3.5-)4.2-5.5(-6.5) mm high.
Phyllaries in 3—4 whorls, loosely imbricate; those
of the outermost whorl (2.1—)3—4.5(-5.1) X (0.6–)
0.9-1.4(-1.9) mm, those of the middle whorl(s) ob-
lanceolate, oblong, or less often spathulate, (3.2—)
3.5-5.6(-6.1) х 1-1.8(-2.2) mm, those of the inner
whorl oblanceolate, oblong, ог spathulate, 4–5.6 Х
1-1.8(-2) mm. Receptacles convex to subconical,
alveolate, 1.1-1.8(-2.2) X 0.8-1.3 mm. Ray florets
(10-)14-23; corollas pale lilac to whitish, ligules
10-17 X 1.6-3.5 mm, the tube 0.7-1.4 mm long,
style 1.5-2.1(-2.6) mm long, the branches 0.6-1.2
mm long. Disc florets (35-)46—71(-85); corollas
2.8-3.8(4) mm long, the lobes equal or subequal,
0.8-1.5 X 0.4-0.6 mm, the tube 0.7—1.2(–1.4) mm
long; anther 0.8-1.3(-2) mm long; style 2.2-3.3
mm long, the two style branches 0.8-1.2 mm long,
each with a triangular appendage 0.3-0.5 mm long.
Ray and disc pappus bristles similar, whitish, 0.2—
0.3 mm long. Achenes straw-colored to purplish,
obovate to broadly obovate, covered with biseriate
hairs and glandular hairs at the top, the margins
about 0.1-0.2 mm broad; ray achenes 3-ribbed, 2—
2.6 X 0.8-1.5 mm; disc achenes 2-ribbed, 2.1-3
X 1.2-2 mm. Chromosome number: n = 9. Figure
9E, F.
Phenology. Flowering period: June to October.
Distribution (Fig. 11). Endemic to Japan, where
it ranges from southern Hokkaido to northern Kinki
Districts. It is probably not very common in Hok-
kaido, because only a few specimens have come
from there. Kalimeris pinnatifida is common in
Honshu, usually occurring in disturbed and rela-
780
Annals of th
Missouri ои Garden
Geographical distribution of Kalimeris pin-
Figure 11.
natifida.
tively mesic habitats. This species ranges from sea
level to 1700 m elevation.
The epithet pinnatifida was first used by Maxi-
mowicz in an annotation as “Boltonia indica В pin-
natifida.” This name was not validated until Ma-
kino published the combination Aster indicus var.
innatifidus in 1906. Prior to that, Matsumura had
published Asteromoea indica var. pinnatifida, and
Boltonia indica var. pinnatifida in 1886 and 1895,
respectively, but did not provide a description or
diagnosis in either case. According to Article 32.1
of the International Code of Botanical Nomencla-
ture, ваге names were not validly published by
Matsum
меље is Kalimeris pinnatifida is most
similar to K. incisa, and, like it, also grows robustly
in the northern part of its distributional range.
However, its short pappus bristles, stoloniferous
rhizomes, equal corolla lobes, and glandular
achenes make it distinct from K. incisa (Fig. 9).
Populations of this species are generally not very
big; the only known exception is a large population
(about 200 m?) at Sakyo-ku, Kyoto, in an aban-
doned paddy field. This species has probably been
cultivated for a long time in Japan and other coun-
tries. Some selected strains have tubular disc florets
of the same length as the ray florets, whereas others
have long, tubular ray florets. Makino's Kalimeris
innatifida var. hortensis was based on such a cul-
tivated plant. Specimens with similar characteris-
tics have been collected from cultivation in Naga-
saki, Kumamoto, Tokyo, and Hyogo. Kalimeris
pinnatifida has also been introduced into the Unit-
ed States; a specimen from Durham County, North
Carolina (Wood s.n. in 10/1956, GH), clearly be-
longs here. Another collection from the Missouri
Botanical Garden (Kellogg s.n. in 6/1911, MO),
originally determined as Aster incisus, also repre-
sents K. pinnatifida.
Additional specimens examined. JAPAN. Hokkaido:
Asahikawa-shi, Ishikari, Arai s.n. (TNS). Aom
315 (MO); from Shirahama to Ashigesaki, H. Gu & Yahara
330 (MO), Kitamura s.n. (KYO); Kamikita-gun, Towadako-
cho, 2 km S of Yachionsen to Shirenhashi, H. Gu & Ya-
hara 326 M 326a (MO); Kazuno-shi, 5 km 5 of Towada
Lake, 380 m, H. Gu & Yahara 327 (MO); Tokera, 400 m,
H. Gu & Yahara 328 (MO); Kitatsugaru-gun, Tateoka-cho,
Hirataki-numa, Furuse 27 (KYO); Kuji-shi, Ohkawame,
Murata & Tabata 49 (KYO); Mutsu-shi, Mt. Osore, 300
m, along Futamatazawa, Murata & Koyama 41306 (KYO);
Natsudomari Peninsula, Asamushi, Natsudomarisaki, Koy-
ата 220 (KYO); не -gun, Pen کر cho, Hiratak-
imura, Furuse s.n. Shimokita-gun, Saimura,
Harada, Mori 53 (KYO, Кеа Shimokita у увече Hi-
gashidorimura, Furuse s.n. (MAK); from Imotomura to Ho-
robe, Furuse s.n. (A); from Tanabu to Mt. Osore, Koyama
775 (KYO); Tomada okt” Kawajuri, 7 km SW of Maita, H.
y m,
hara 325 (MO), Faurie 5159 (KYO), enr s.n. (TI).
Iwate: Asagi-shi, Morioka, Kikuchi s.n. (TNS); Higashii-
wai-gun . Tabashineyama, Kikuchi s.n.
Ichinoseki-shi, Toyoryu Shrine, Makino 28 (МАК); мае
п, Iwate Mts., from Daijigoku to Yashikidai, Koyama
1492 (KYO, TNS); Isawa-gun, Isawa-cho, SE of Mt. Yak-
eishidake, 300-500 m, Kurosaki 11365 (KYO); Mizusawa-
shi, Shoho-ji, Mt. Horai-san, Makino s.n. (MAK), Iw
s.n. (KYO, TNS); Morioka-shi, Makino s.n. (MAK),
тада s.n. (MAK); Noshiro-shi, Hi A Hemmi 355
(KYO, MAK). Akita: Akita-shi, Morbid 13 (TNS). Mi-
yagi: Mt. Fubou-san, Namariz m, Boufford &
Wood 19863 (KYO, MO); Kata Shichikasyukurcho.
Mt. Fubo-dake, 600 m, Murata 4456. ав gun,
Miyagi-cho, Mt. кыйт. гм cds 2458 (TD; ne
yoshi-gun, Shizugawa-cho, Naito & Otomo s.n. (KYO);
Sendai-shi, Dainohara, Ogura s.n. (TI); Kawauchi,
Ninomaru, Okazaki & Kanno ; Kozi, lishiba s.n.
(МО); Tokusayama, Kakuda, бара et al. s.n. (МО), Каи-
пе 6743 (P). Yamagata: Akumi-
; Higashitagawa-gun, Y:
Tsutsumishinden, Kato s.n. (MAK); Higashokitama-gun,
Takahata-cho, Yashiromura, Koidzumi s.n. (KYO); Yuza-
Volume 84, Number 4
997
Gu & Hoch
Systematics of Kalimeris
781
cho, Mt. Chokai, Kannonmori, 600 m, Ohashi s.n. (TI);
wa s.n. (INS);
Murayama-shi, Ohmaki, Takahashi 102 (МО); THEY ai
Inoe s.n. ; Yamatata-shi, Kabutoiwa, Yamadera,
arf 718022 (МО, TI); Kaminoyama, Beattie & Kurihara
11119 (GH); ад lan Narushima, Midorikawa 1368
d Fuk : Kit akata-shi, Makino 2 (MAK); Mina-
miaizu-gun, mi-cho, Asakusadake Mts., Yamasaki
s.n. (TD; Tajima- zs from Hariu to Akobara, 700 m, Mur-
ata 9908 (TI); Hinoemata-mura, Nasegawa s.n. (TI); Nish-
ishirakawa-gun, Shirakawakogen, 900 m, Koyama 407 (A,
KYO, NY); Yama-gun, Inowashiru-cho, Akaii et al. s.n.
(TD; from Niichyuraonsen to Hinosawa, Kurosawa s.n.
(TI); ii Do bone prn Ueno s.n. (TNS); T.
mura-gun, Mi Makino s.n. (MAK). Шакен
Hitachi-shi, мра, Sato ѕ.п. (KYO). Tochigi: Aso-
gun, Kuzuu-cho, Tagawa & Iwatsuki 1991 (KYO); Imai-
chi-shi, Fubasami, 300 m, Ohashi & Murata s.n. (KYO,
TI); Kanuma-shi, Itaga, 200 m, Ohashi et al. 1546a (TI);
Kuruizo-shi, Ikuyohashi, Hara s.n. (TI); Nasu- a cy Kuro-
"8 cho, Miyujintoge, 376 m, Hara s.n. aes amizo
Mts., Ungan-ji, Hara € Kurosawa s.n. (ТІ); Nasuno-gun,
авас сћо, Suzuki s s.n. (KYO); Nikko- sh, Bienes
imano, Мы ке
motsuga-gun, Kinagawa-mura, Биће s.n.
Numata-shi, ت Makino 149 (MAK): (cec ep
Џепо-тига, Fujidou, 7. m, Murata et al. 6257
(KYO); Tone-gun, Mt. Mitsumine-san, Tobe s.n. (TNS);
Usui-gun, Matsuida-cho, Mt. Myugisan, Hakuunzan, 700
m, Murata 27445 (KYO); from Usuitoge to Yokokawa, 550
m, H. Gu & Yahara 426 (MO). Chiba: Abiko-sh, Murata
& Togashi 39 (KYO, MAK); Awa-gun, Amatsuko minata-
n, from Takayanagi to Shinden, Kurihara et al. 160
(TD; Ызама, Yamamoto s.n. (ТІ); Kashiwa-shi, Ki-
takogana, Shirai s.n. (TNS). Saitama: Chichibu-shi, Mt.
Bukou, Moriya s.n. (TNS), Tagawa s.n. (KYO); Ohnomura,
Ohnotoge, 850 m, H. Gu & Yahara 427 (MO), 429 (MO),
430 (MO), 500 m, 428 (MO); Hannou-shi, Shirako, Ma-
K); Naguri River, Hannou-shi, Sugiyama s.n.
(TI). Tokyo: гава. shi, Mt. Mitakesan, 900 т, Murata
6075 (TI); Arakawa-ku, Dakan-yama, Makino s.n. (MAK);
Bunkyo-ku, Koishikawa Botanical Garden, Abe s.n. (TI
Edogawa-ku, Koiwamura, Makino s.n. (MAK); Fuchyu-shi,
Mizushima s.n. (MO, TI); Hachiouji-shi, Kaminukugitah-
en, Makino s.n. (MAK); Mt. Takaosan, Sato s.n. (ТІ); Ita-
bashi-ku, Kamiitabashi, Makino & Kitamura s.n. (KYO);
Itsukaichi-shi, Kobayashi 4294 (MAK); Mitaka-shi, Ino-
kami, Makino s.n. (MAK); Osawa, Elliott 144 (A); Nakano-
ku, Makino s.n. (MAK); Nelima-ku, Makino s.n. (MAK Pe
itama-gun, Hinohara-mura, Sengennoe, 500—
Fa prin od Motohachiwooj-mura, Shiroyam ~
Furus gakuzan, Mizushima 12032 (MAK,
ш ы а Makino s.n. (KYO); Shimura,
Okuyama s.n. (TNS); Suginami-ku, Maekawa 160 A32
(TD); Kugayama, Yamatsuta 4050 (TNS). Kanagawa: Ка-
makura-shi, from Yukinoshita to Juunisou, Kobayashi
3485 (MAK), Asai s.n. sere — from Atsugi-shi
to Yokohama-shi, Yamasaki s.n. (TI); Kawasaki-shi, No-
borito, SW of Tokyo, Me s.n. (KYO, MAK); “Kinu-
” Suzuki s.n. (A); Odawara-shi, Kankatai s.n. aca
E Makino s.n. gen ) Tsukui-gun, Tanza
Mts., Mt. Yake-yama, Yamasaki s.n. (TI); Yok Кората,
Hisanchi ѕ.п. (KYO); Zushi- dle Numakan, Momiyama 18
—
РҮ:
(MAK). Niigata: Arai-shi, Mt. Kazano, from Kamihira-
maru to Nagasawa, 500-700 m, Kurosaki 11432 (KYO);
Higashikanbara-gun, Kanose-cho, Nakai & Maruyama
s.n. (TNS); Kitakanbara-gun, Sasakami-mura, Deyuonsen,
Yatabe s.n. (ТІ); Kitauwonuma-gun, lrihirose-mura, from
.n.
gun, Muika-cho, Hakkatoge, Kanai 6481 (A, TI);
kanbara-gun, Yahiko-mura, Noda s.n. (TNS); cet E
Yamasaki 8986 (ТІ); Sado Island: Hamochi-cho, Ito s
oyama: Kaminikawa-gun, Ohyama-cho, вар
gai, Котон 12069 (КУ О); Naganiikawa-gun, Kamiichi-
cho, Hinoya to Ома, 2116 (KYO); Osawa, 2298 (KYO);
Toyama-gun, 17 km SW of зрана Kirino 386 (МО);
Uotsu-shi, Katakaigawa, Nagai s.n. (KYO); Tsubono, 300
m, Nagai s.n. (ТІ). Nagano: Chiisagata-gun, Sanada-cho,
Sugadaira, Kobayashi 13126 (KY О); Toubu-cho, from Nar-
ahara to Mt. Yunomaru, 1200 m, Murata 27540 (KYO);
Chino-shi, Kirigemine, Makino s.n. (MAK), Okamoto s.n.
(KYO); Hanishina-gun, Sakaki-cho, Makino s.n. (MAK);
lida-shi, Chiyo-mura, Yonegawa, Koidzumi 94955 (TNS);
liyama-shi, Mizuhokuni, Yokouchi s.n. (MAK); Kigashi-
chikuma-gun, “Katooka,” Kubota s.n. (KYO); Kamimino-
chi-gun, Shinano-cho, Lake Nojiriko, 700 m, Midorikawa
1028 (TT); Togakushi-mura, Koshimizuhara, 1250 m, 2002
(TD; Kiso-gun, Kisofukushima-cho, Mitakemura, Kawa-
saki s.n. (MAK); Mt. Togakushi, Nenbutsuike, Naruhashi
1856 (KYO); Kitaadzumi-gun, Kitoshiro-mura, Furuse
13944 (KYO); Kitasaku-gun, Karuizawa, Fox s.n. (BM);
Shinanooiwake, Karishuka, 1000 m, Kanai 419 (TI); To-
ge em ra s.n. (TNS); Minamisaku-gun, Kawakami-
mura Mikuniyama, Azusayama, Kitagawa 1535
(KYO), Nol 11818 (KYO); from Senjagahara to Mt.
Mikuniyama, 1700 m, Hotta 10181 (KYO); Koumi-cho,
> np e занар & Ohba 5382 (ТІ); Nishichiku-
aida- , Nishino, 1200 m, Nitta & Mabuchi
12012 |" ме a а Tadachi-mura, p
5386 (GH); V maU Bunkuitoge, 1250 m
7311 (KYO); from Kamazawa to Koshibuyu, 500-700 m,
Iwatsuki Е іт та 37 (KYO, MAK, TNS); Ooshika-mura,
from Ohgawara to ng eae Murata & Shimizu 1472
MO); Suwa-shi, M oriyasan, 1100-1650 m, Midori-
kawa 508 (TI), 5406 (TD, 540b (TI), 661 (TI); Ueda-shi,
Beshuonsen, ара 5361 (KYO); Yamakuchi-mura,
Aono, Okuhara s.n. (KYO). i i
Mt. Fuji, 12 200 m оа et al. 1162
Fujigawa-cho, Ivabuchi-eki Ohmura s.n. (KYO); E of
Kenashiyama, Fumoto m, Hino et al. 62 (KYO); Go-
teba-shi, “ath ean E of Mt. Fuji, 450 m, a
“~
1610 (KYO); Sunto-gun, Nagaizum mi-cho, Suiji Mt.
ae Fujimura & Fukui 29 (KYO); Зимно S of
Fuji, 900 m, Konta 11564 (KYO). Yamanashi: Hi-
ol gun, Sa ntomi-mura, Mt.
600-1700 m, Midorikawa 1729 (TI);
Hakushiyu-cho, Mt. Koma, Kitamura s.n. ;
Kitatsuru-gun, Џепоћага-сћо, Shiotsu, Okamoto s.n.
(KYO); Tanakami, Okamoto s.n. (KYO); абра d cm
Nanbu-cho, Kozori, 400—700 m, Tateishi et al. 4563 (TD;
Minamishuru-gun, Niahikashura-cho, Mt. азан ЫА
. (MAK, KYO); Nirasaki-shi, Mt.
m, Yamasaki s.n. (ТЇ); Nishiyatsushiro-gun,
Kamikuishiki-mura, Asagirikogen, Mt. — m, Mur-
ata et al. 33895 (A); Lake Motosu, Mt. Fuj
(A); Nanbu-cho, Kozori, 400-700 m “Tateishi et al. 4363
(TI). Aichi: Atsumi-gun, Fukue-cho, Жайына, Torii s.n.
(KYO); Kitashitara-gun, Inabu-cho, — To z i
(KYO); Ninamishitara-gun, Tsukude- , betwee
mogaya and Yuanami, 500 m, Murata "13232 (KYO): 4
782
Annals of the
Missouri Botanical Garden
Nokoenhan, Matsuoka s.n. (MAK); Sumikawa,
tara-cho,
890 m, Marai 14351 (KYO): Cushiyou-gun AES
cho, biet Shiota 689 (GH); Ohno-gun, Kuguno-cho,
o Mt. Funayama, Fulàioha 7518 (KYO,
MAK); Kashima-gun, Kahn cb Solido
an, m, Yamasaki s.n. (TI); Suzu-shi, Suzuzinzya, Mt.
— E Masamune 14176 mo Fukui: Mina-
mirokuro-shi, Karasawa, Kurosaki 11104
(KYO); Nanjo-gun, Imajo-cho, Shinbo, Kinometoge, Hi-
yoshi Shrine, Kanai 731271 (TNS); Ohno-shi, from Fukai
to Mt. Шипуата, Fukuoka 8265 (KYO); Mt. Kyogatake,
Murata & Shimizu 329 (KYO); Takefu-shi, Ohshio, 60—
160 m, Kurosake 8019 (KYO, TI); Tsuruga-shi, Ikenoko-
chi, 250 m, Kitamura & Murata 2079 (KYO). Shiga: Tak-
ashima-gun, Kawakami-mura, Hashimato 4670 (TNS);
Kuchiki-mura, Udoya, Inumaru, 300 m, Kurosaki 11893
(KYO), Ohba & Akiyama 1987 (TI). Kyoto: Atago-gun,
Kurama, Kitamura s.n. (KYO); Kitakuwada-gun, Miyama-
, Okamoto s.n. (KYO, TNS); Kyoto-shi, Sakyo-
wara, 40-60 m, H. Gu et al. 415 (MO); between
Kibune and Hanasetoge, Tagawa 106 (KYO); Nomi, N of
Hanase, 600 m, Murata 17491 (KYO, US); Kuta, 60-100
m, H. Gu et al. 414 (MO); between Kuta and Hirogawa,
Hiroe 16362 (KYO, ТІ); Sisigatani, Tagawa 2989 (MAK,
NY), Stowe s.n. (TI).
3. Kalimeris mongolica (Franch.) Kitam., Acta
Phytotax. Geobot. 6: 21. 1937. Aster mongoli-
cus Franch., Nouv. Arch. Mus. Hist. Nat. 6:
41. 1883. Aster lautureanus var. mongolicus
(Franch.) Kitag., Bot. Mag. (Tokyo) 48: 110.
1934. Asteromoea mongolica (Franch.) Kitam.,
Acta Phytotax. Geobot. 4: 21. 1937. Kalimeris
lautureana var. mongolica (Franch.) Kitag.,
Neo-lin. fl. manshur. 653. 1979. TYPE: China.
Hebei: Chengde (Mongolie: Géhol), July 1864,
A. David 1995 (lectotype, here designated, P;
isolectotype, P).
Kalimeris incisa var. holophylla Maxim., Prim. fl. amur.
146. 1856. Aster d var. holophyllus en
im.) F. H. Chen, Bull. F. m. Inst. Biol. 5
1934. TYPE: Amur: ког aximowicz s.n.
(holotype, LE not seen; isotypes, BM, G, K, P).
Aster associatus Kitag., Rep. Inst. Soc. Res. Manchoukuo
29. 1938. Kalimeris associata (Kitag.) Kitag.,
Neo-lin. fl. manshur. 652. 1979. TYPE: China. Hei-
longjiang: Ke-shan, 12 Sep. 1937, M. Kitagawa s.n.
(holotype, TI; isotype, TI).
Aster associatus var. stenolobus Kitag., Lin. fl. manshur.
5. 1939. Kalimeris associata f. pudore (Kitag.)
Lin. fl. manshur. 435. 1939. TYPE: China.
Jilin: n near Xinjing Xian, 17 Aug. 1938, M. Kitagawa
s.n. (holotype, TI).
Aster associatus n "beachslobéü Kitag., Lin. fl.
Kitag., Lin. fl. manshur. 439. 19 PE: Chi
т near Xinjing Xian, 17 Aug. 1938, M. Kitagawa
n. (holotype, TI).
Root fibrous; rhizomes condensed, forming root-
stocks up to 3 cm diam. Stem 50-122 cm tall. Cau-
line leaves narrowly oblong to oblanceolate, serrate
to deeply pinnate-lobed, (3.5—)5—8.5(-9.6) х 0.9-
2.1 cm, glabrous, less often loosely oet with
nonglandular hairs on both sides or only so abaxi-
ally, lobes lanceolate to triangular, 0.5-1.5 X 0.2-
0.4 cm. Capitula 12-45; peduncles 2.5-11 cm
long; bracts 2-7 along the peduncle, the lower ones
9-23 X 1.4—4 mm, the upper ones 4-8 X 0.8-1.3
mm; involucre campanulate or hemispherical, 6.8—
13 mm broad, 5.2-7(-8.5) mm high. Phyllaries in
3—4 whorls, imbricate, glabrous or loosely covered
with nonglandular hairs, chartaceous below, blunt,
sometimes purplish; those of the outermost whorl
lanceolate to obovate, (2.7-)3.5—6(-7.2) X 0.8-
1.8(-2.8) mm, those of the middle whorl(s) obovate
to spathulate, 4.2-7(8.1) X 1.5-3(-3.5) mm, those
of the inner whorl obovate to spathulate or few in-
nermost ones oblanceolate, 4.2-7(-8) X 1.1-2.8(-
3.2) mm. Receptacles convex, 1.6-2.7 mm wide,
0.7-1.6(-2) mm high. Ray florets (11-)14-25; co-
rollas ape lilac to lilac, ligules oblanceolate, 13.5—
25.5 X 2.7-4.8(-5.2) mm, the tube 1.1-1.8(-2.1)
mm long; je 2-2.8(3.4) mm long, the branches
(0.5-)0.7-1.4 mm long. Disc florets 38—86; corollas
3.5-4.9 mm long, the lobes unequal, the longer one
1.2-2.1 mm long, the shorter ones 0.8-1.7 mm
long, both (0.4—)0.5—0.7(—0.8) mm wide, the tube
0.8-1.6(-1.8) mm long; anther 1-1.6(-1.8) mm
long; style 3-4.2 mm long, the branches 0.9-1.5
mm long, each with a triangular appendage 0.4—0.5
mm long. Ray and disc pappus bristles brownish,
unequal; the ray bristles 0.3—0.5 mm long, rarely
up to 0.8-1 mm long; the disc bristles 0.5—1.2 mm
long, very rarely up to 1.5 mm. Achenes straw-
colored to brownish, broadly obovate, loosely cov-
ered with 4-celled nonglandular hairs, rarely scat-
tered with biseriate glandular hairs at the top, the
margins 0.3-0.6 mm broad; the ray achenes 2.8—
3.2 X 2-2.2 mm; disc achenes 3.2-3.5 x 2.3-2.9
mm. Chromosome number: n — 27, 54. Figure
12A-F
Phenology. Flowering period: June to October.
Distribution (Fig. 13). Reaches its northern limits
in the Amur basin in the Far East region of Russia
and in adjacent Heilongjiang province of China,
extending south and west in China to Jilin, Liaon-
ing, Nei Mongol, Hebei, and Beijing provinces; it
also extends southeastward in Korea to H
Hamnam, Pyongbook, Pyongyoung, Whanghai,
Kyonggi, and Seoul provinces. This distribution is
similar to that of K. incisa except that K. mongolica
extends further south into Laiyuan in Hebei Prov-
Volume 84, Number 4 Gu & Hoch 783
199 Systematics of Kalimeris
===, =; LB
х.
=
© IS A EAS SO ICO |
] ] ] ] — —C. Disc
i ¡meri lica and K. shimadai. A-F. K. mongolica. —A. Upper branch. —B. Root. —
e di rag viue tri led —E. Leaf. —F. Portion of the leaf showing indumentum. G-J. K. shimadai.
—G. Disc floret. —H. Phyllary at the middle layer. —I. Leaf. —J. Portion of leaf showing indumentum.
Annals of the
Missouri Botanical Garden
Figure 13. Geographical distribution of Kalimeris
mongolica.
ince, China. This species grows from near sea level
to 1200 m. In addition to growing in disturbed ar-
eas near human activity like most Kalimeris spe-
cies, К. mongolica is commonly found in semi-open
secondary forests and along edges of coniferous and
mixed broad-leaf forests in relatively mesic sites.
Two specimens were designated when this spe-
cies was published, i.e., David 1995 and 2189; the
former was chosen as lectotype because it has more
sheets. Kalimeris mongolica has been confused in
the herbarium not only with K. lautureana, but also
with K. pinnatifida and K. incisa. Handel-Mazzetti
identified some Chinese K. mongolica as K. pin-
natifida because of their pinnate-lobed leaves. In
fact, K. mongolica is relatively easy to distinguish
from these three species. It can be distinguished
from K. incisa by its bigger capitula and broad
phyllaries with the upper portion herbaceous and
the margins broad and membranous, and from K.
lautureana and K. pinnatifida by its thinner leaves,
longer pappi, and broader, thickened achene mar-
gins (Fig. 9).
is species is sympatric with Kalimeris incisa
subsp. incisa throughout most of its geographic
range. In the field, one can observe small differ-
ences in microhabitat, however, with K. mongolica
growing in more disturbed, open places close to
roads and paths, and K. incisa subsp. incisa grow-
ing in more sheltered and natural places.
Additional specimens examined. RUSSIA. Primor-
ski: Khanka Lake, Bohnhof 41 (E, G, NY, P), Rock 41
(NY); Ussurisk, "e s.n. (TI). Amur:
i gene s.n. (GH
Mori
ply Ha Tode Okuyama s.n. (TNS). КҮ аулы
Gang-ge, Mills 141 (TI). pa bem d so
oung, Smith s.n.
ailun Xian, Kitagawa s.n. (TI); Heli,
Jinfeng-cun, Tatewaki 31540 (KYO); Jinpohu, Diaoshuil-
ou, ae 10220 (IFP, PE); ig 10219 (IFP, PE);
(IFP, PE); Songhua-jiang River, Maximowicz s.n. (
Weiguo, T. N. Liu et al. 10275 (IFP); Xiaoxing'anling Mts.,
Tangwanghe River, Pare s.n. (KYO); Xunhe, Nanzhan,
Heda-tun, G. D. Ciu & C. 0. Lin 1528 (IFP); Zhaodong
Xian, from n to Dongxinsan- tun, G. Z. Wa
a
), pum s.n. (MAK
ang et al. 2367 (IFP, KUN), 2371 (IFP),
2373 (ТЕР), 2381 (IF P), 2386 (IFP), 2390 (IFP), 2644
(IFP); Daxing'anling Mts., Taidong, 160 m, Jilin Teachers’
College Sey 788 (PE); Bushs Xian, Dongshuai-wanzi,
G. D. . Lin 1668 (IFP, PE); Nanmenwai, Sato
= (IFP. PE), ји (KYO); Hunchun Xian, Chunhua-
ang, Sanjiat C. S. Wang et al. 1897 (IFP), 2138
(IFP). Jilin Shi шо shan, Y. L. Zhang et al. 945 (ТЕР,
PE); Xiaobai-shan, 190 m, P У Pu & У L. Chou 2180
(PE); Jiutai Xian, Тонар . Tatewaki s.n. (KYO), Tak-
enouchi s.n. (MAK); Tagan: Xin; 40 km SW of Tonghua,
P. Y. Fu & Z. H. Zhang 451 (IFP), 464 (IFP); Ji'an to
Toaghua, Takahashi 860 (TNS), Takahashi 547 (TNS);
Wangqing 2 Chunyang-xiang, Gaoli-cun, Р У Fu et
al. 959 (IFP, PE), C. S. Wang et al. 1846 (IFP, KUN),
Komarov 1507b (GH, NY, ТІ), 1510 (W). Liaoning: Dan-
dong Xian, vye ap Sato 1188 (IFP); Fenghuang-shan,
Yamatsuta 80 (ТІ), 277 (ТІ); Huanren Xian, C. S. Wan
20 (IFP, PE), 3830 (IFP, PE), 3855 (IFP, PE),
3868 (IFP, is 4231 (IFP); Hunhe, Yamatsuta 115 (TNS);
Jianchang Xian, Daheishan Forestry Station, C. S. Wang
et al. 3180 (IFP, KUN, PE); Jinzhou Shi, Hulu-dao
Wang 3451 (IFP, KUN), 3452 (IFP), 3453 (IFP), ' 3524
(IFP), W. Mo 3526 (IFP, PE); Jing Xian, Heshang-shan,
Y. Yabe s.n. (PE); Liaoyang Xian, Emei-zhuang, rosis
s.n. (KYO, MAK, TNS); Lushun Shi, Baiyu-shan, Y. X. Liu
& T. Wang 919 (IFP); Qian-shan, H. Gu & H. Zhang 279
(MO), 283 (MO), 284 (MO), 286 (MO), 288 (MO) Yamat-
suta 105 (TNS) Shenyang Shi, Beilin, Ki
(KYO, MAK, TI), Yabe s.n. (PE), Yamatsuta 84 (TNS
(TNS), 94 (TNS), 104 Seer? 114 (TNS), 290
Chang
et ba 2656 (IFP
Volume 84, Number 4
1997
Gu & Hoch 785
Systematics of Kalimeris
(IFP), Kitagawa s.n. (ТІ); Xiuyan Xian, Tangchi-xiang,
Toudao-dadui, Xiaozhezigou, W. Wang et al. 454a (IFP);
cil Xian, Namuqiantu-cun, T. Wang et al. 2721
(ЕР). Nei Mongol: Hulun Buir Meng, Ergun Youqi,
Shangkuli, Ibn s.n. (TD); " Ud Meng, e Qi,
Guangxingyuan-gongs m, Mengning Team 1150
(PE). Hebei: Декан ‘Sate 8870 (IFP, KYO, PE); Bei-
daihe, Cowdry 119 (K), Schnack 135d (W); Changli Xian,
be s.n. (PE); “Paita,” 1015
m, Licent 9 (W); Shanhaiguan, Jiao-shan, 1579 (BM, P,
W); Xinglong Xian, Wuling-shan, Hongmeisi, 650 m, P.
Y. Fu et al. 4568 (IFP, PE), Nakai et al. s.n. (TI); from
Chaihekou to Xinglongtang, Nakai et al. s.n. (TI); from
igou, PE Team 1 585 (PE); Gubeikou, Wuohu-shan, За
10287 (IFP, PE); Qinglongqiao, Капазто 4130 (KYO).
Kalimeris indica (L.) Sch.-Bip., in Zoll., Syst.
Verz. 125. 1854 [“Calimeris”]. Áster indicus L.,
Fl. Hongk. 174. 1861. TYPE: China: without
locality (holotype, LINN-997/42 not seen; pho-
tos, GH, NY; microfiche, MO).
Matricaria canton iensis Lour., Fl. омер. еа. 1:
1790. Hisutua cantoniensis (Lowe) D
Guangdong: Guangzhou; type not locate
Martinia polymorpha Vaniot, Bull. Ac ad. Int. Géogr. Bot.
12: 2 i Kalimeris indica f. (Va niot)
Kitam., J. Jap. Bot. 19: 340. 1943. Kalimeris indica
var. Бортас (Vaniot) Kitam. ex Y. Ling, in Y.
Ling et al., Fl. reipubl. popularis sin. 74: 102. 1985.
TYPE: China, Guizhou: Guiyang, 31 July 1897, E.
Bodinier 1739 (lectotype, designated by Lauener
(1976), E; isolectotypes, E, P (two sheets)).
e serrata Hook. & Arn., Bot. Beechey 65.
8. TYPE: Japan. akm Islands: type not locat-
M
Kalimeris indica f. epapposa J. Q. Fu, Bull. Bot. Res.,
Harbin 3: 111. 1983. TYPE: — Shaanxi: Kang
Xian, Qing-he Forestry Stat 1340 m, 18 Oct.
1963, Z. Y. Zhang 17324 redi) WUK).
Root fibrous; rhizomes stoloniferous, 0.5-18 cm
long. Stem green or purplish, (12-)20-150 cm tall.
Cauline leaves linearly ap ев оЫапсео-
late, ог obovate, entire to deeply pinnate- ог lan-
ceolate-lobed, (0.7-)2-7(-9) х (0. 3-0. 5-1.8(-2.5)
ст, glabrous, or densely covered with ascending
thin or thick nonglandular hairs, sometimes with
scattered biseriate glandular hairs. Capitula 8-120;
peduncles (0.3—)1–9(–13) cm long; bracts 1-8(-12)
along the peduncle, the lower ones 6.2-19(-27) X
1–4(–6) mm, the upper ones 1.2-6.3 X 0.2-0.9(-
1.3) mm; involucre hemispherical, rarely campan-
ulate, 4.5-10 mm broad, 3-6 mm high. Phyllaries
in 3-4 whorls, imbricate, usually obtuse at apex,
sometimes purplish; those of outermost whorl lan-
ceolate or obovate, 1.6-3.7(4) X (0.5—)0.8-1.1(—
1.4) mm, those of middle whorl(s) oblanceolate, ob-
ovate, or spathulate, 2.3—5.4 X 0.7-2 mm, those of
inner whorl oblong to obovate or spathulate, 2.5—
5.7 X 0.8-2 mm. Receptacles convex, 1.1-1.7(-
2.5) X 0.5-0.9-1.2) mm. Ray florets 10-26; co-
rollas pale lilac to pale purple; ligules narrowly ob-
long or oblanceolate to elliptical, glabrous or rarely
with scattered long biseriate glandular hairs near
the corolla tube, 5.2-16 X 1–3.2 mm, the tube 0.4—
2 mm long; style 1.3-2.6 mm long, the branches
0.4—1.3 mm long. Disc florets 40—105; corollas
2.2-4.2 mm long, the lobes subequal or unequal,
the longer one (0.8-)1-1.4(-1.7) mm long, к
shorter ones 0.6-1.2(-1.4) mm long, both 0.4–0
mm wide; the tube 0.6-1.2(-1.5) mm long; тш
0.8-1.6 mm long; style 1.7-3.8 mm long, the
branches (0.4—)0.5—1.1(-1.4) mm long, each with a
triangular appendage 0.2-0.4 mm long. Ray and
disc pappus bristles equal or unequal; ray bristles
(0-)0.1-0.3 mm long; disc bristles (0—)0.1—0.3(-
0.5) mm long. Achenes brownish to dark purple,
obovate to broadly obovate, covered with 4-celled
nonglandular hairs and with biseriate glandular
hairs at the top, the margins 0.1–0.3 mm broad; ray
achenes 3(-4)-ribbed, 1.5-2.6 X 0.6-1 mm; disc
achenes 2(-3)-ribbed, 1.5-2.9 X 0.7-1.4 mm.
Chromosome number: n = 18, 27. Figure 14.
Phenology. Flowering period: May to November.
This is a very variable and widely distributed
species. It ranges from central China to South Ko-
rea and Japan, and to northern Indochina. It is here
divided into three subspecies, as follows:
KEY TO THE SUBSPECIES OF KALIMERIS INDICA
la. Leaves glabrous or loosely covered with thick
nonglandular hairs 4a. subsp. indica
lb. Leaves densely to evenly covered with thin or
thick MPG hairs and with scattered glan-
dular hai
2a. слани leaves obovate to oblong, covered
with thick nonglandular hairs; phyllaries of
= whorl(s) %4—% as wide as long; rhi-
s 5-15 cm b. subsp. collina
. смање leaves ДЕ to oblong, covered
with thin nonglandular hairs; phyllaries of
middle whorl(s) %—% as wide as long; rhi-
zomes 0.5-1(-3.5) ст long ........................
de
N
=
786 Annals of the
Missouri Botanical Garden
Figure 14. Kalimeris indica. A-E. K. indica subsp. indica. —A. Upper branch. —B. Root. —C. Achene. —D. Leaf.
—E. Portion of leaf showing indumentum. F, G. K. indica subsp. collina. —F. Асћепе. —G. Leaf. —H. Portion of leaf
showing indumentum. I-K. K. indica subsp. stenolepis. —I. Achene. —J. Leaf. —K. Portion of leaf showing indumentum.
Volume 84, Number 4
1997
Gu 4 Hoch
Systematics of Kalimeris
4a. Kalimeris indica subsp. indica
Boltonia indica var. [*8"] rivularis Hance, Ann. Sci. Nat.,
Bot. sér. 5, 5: 219. 1866. TYPE: China. Fujian: Fu-
zhou, 1857, W. Gregory s.n. [sub Hance Herbarium
Number 1236] (lectotype, here designated, BM).
Kalimeris indica var. stenophylla Kitam., J. Jap. Bot. 19:
340. 1943. TYPE: China. Jiangsu: Nanjing, Zijin-
shan, 3 Sep. 1934, H. Migo 40 (lectotype, here des-
ignated, TI).
Kalimeris lancifolia J. Q. Fu, Bull. Bot. Res., Harbin 3:
112. 1983. TYPE: China. Shaanxi: Lueyang Xian,
700 m, Z. ¥ Zhang 17891 (holotype, WUK).
Rhizomes stoloniferous, 0.5—18 cm long. Stem
13—75 cm tall. Cauline leaves oblong, oblanceolate,
elliptical, or obovate, serrate to deeply pinnate-
lobed or less often entire, (1.7-)2.4—7.5(-9) х
(0.3—)0.7—1.8(-2.5) cm, glabrous or with scattered
thick nonglandular hairs and biseriate glandular
hairs abaxially, less often adaxially, lobes lanceo-
late, 0.9-2.1 X 0.2-0.4 cm. Сариша 8-30; pedun-
cles 1-10 cm long; bracts 2-7(-9) along the pe-
duncle, the lower ones 10.2-27 X 1-6 mm, the
upper ones 1.6—6.3 X 0.3-1.1 mm; involucre hemi-
spherical or campanulate, 4.5-10 mm broad, 3-6
mm high. Phyllaries imbricate, green, sometimes
purplish; those of the outermost whorl 1.6-3.4(4)
X 0.6-1.4 mm, those of the middle whorl(s) (2.3—)
2.8–4.5(–5.5) X 1-1.9 mm, those of the inner whorl
3—5(-5.5) х 0.9-1.8(-2) mm. Receptacles 1.3-1.7
mm wide, 0.6–0.9 mm high. Ray florets 10-58; co-
rollas lilac to pale purple, ligules 8-16 Х 1.5-2.8
mm, the tube 1-2 mm long; style 1.7-2.8 mm long,
the branches 0.5-1.1 mm long. Disc florets 40—
105; corollas 2.6-4.2 mm long, the lobes unequal,
the longer one 1-1.7 mm long, the shorter ones
0.6-1.2 mm long, both 0.4–0.6 mm wide, the tube
0.6-1.2(-1.5) mm long; anther 0.9-1.6 mm long;
style 2.4—3.8 mm long, the branches 0.5—1.1 mm
long. Ray and disc pappus bristles similar, (0—)
0.1-0.3 mm long. Achenes purplish, the margins
0.1-0.3 mm broad; ray achenes 2.1-2.6 X 0.6-1
mm; disc achenes 2.1-2.9 X 0.9-1.4 mm.
Chromosome number: n = 27. Figure 14A—E.
Phenology. Flowering period: June to early No-
vember.
Distribution (Fig. 15). Kalimeris indica subsp. in-
dica is one of the most widely distributed taxa in
Kalimeris, overlapping the ranges of the other two
subspecies of K. indica almost completely (it does
not occur on Hainan Island). In China, it extends
from Qingdao, Shandong, westward to the Qinling
Mountains, southwestward to the foot of the Qing-
zang Plateau, and occurs in all provinces of East
Figure 15. Geographical distribution of Kalimeris in-
dica subsp. indica.
and South China (Shanxi, Henan, Shaanxi, Ningxia,
Gansu, Anhui, Jiangsu, Zhejiang, Jiangxi, Fujian,
Taiwan, Guangdong, Guangxi, Hunan, Hubei, Si-
chuan, Guizhou, and Yunnan). It also occurs in
southern Korea, in Junnam province and extending
to Cheju Island. In Japan, it occurs on Kyushu (Na-
gasaki and southern Kagoshima prefectures), and
on the Ryukyu and Ogasawara Islands. In South-
east Asia, it is found in northern Vietnam (Dong
Dang, Langson, and Hung Hoa), northern Thailand
(Phitsanulok and Chiang Mai), northern Burma
(N’changyang), and Malaysia (Melaka). It has been
reported in Laos (Gagnepain, 1924), but no au-
thentic material has been seen. Collections from
Java are cultivated material (Miquel [1856] report-
ed this taxon in cultivation in Sumatra and Java),
and a recent collection from Oahu, Hawaii (Wagata
2457, K), may be a plant escaped from cultivation.
This subspecies occurs over a very broad altitudi-
nal range as well, from near sea level to 3900 m
in Yunnan, although it primarily occurs below 2000
m. Kalimeris indica subsp. indica is strongly as-
sociated with human activities. It grows primarily
along sunny, dry roadsides, margins of crop fields,
and semi-cultivated places near houses. It can also
be found in more stable natural environments, such
as margins of thickets or openings in forests, where
its populations are often large due to the extensive
formation of rhizomes.
Boltonia indica var. rivularis Hance was pub-
lished without a designated type specimen. There
are several specimens with the same Hance Her-
788
Annals of the
Missouri Botanical Garden
barium number, 1236, collected at different local-
ities. The only one annotated by Hance with that
name is the specimen chosen here as the lectotype.
The other Hance collections with this number are
from Guangdong Province, China (Nov. 1860, BM;
Dec. 1860, GH; 1860, W, two sheets).
Like that of other taxa in this genus, the leaf
morphology of this subspecies is very variable,
ranging from entire to deeply pinnate-lobed. Pop-
ulations found on islands, peninsulas, or along the
sea coasts tend to comprise short plants with broad,
crenate leaves, whereas plants from the continental
interior have various kinds of leaves. Individuals
with deeply pinnate-lobed leaves have been found
scattered throughout the geographic range. Several
specimens have been misidentified as K. pinnati-
fida or K. mongolica, mainly because of their deep-
ly pinnate-lobed leaves (e.g., I. B. Balfour s.n. in
1910, E). Such plants have been accorded taxo-
nomic recognition as K. indica var. polymorp
(Vaniot) Kitam. ex Y. Ling, but no other character
states are consistently associated with this leaf
character. Some plants from Yunnan, the western
limit of this subspecies, have narrowly oblong and
entire leaves and are less branched (Forrest 28457,
BM, M, and 30584, BM; Y Tsiang & H. Wang
, A; H. C. Wang 4093, PE); again, there are
no inia character states consistently associated
with these characters. Kalimeris lancifolia J. Q. Fu
was based on one specimen from Lueyang, Shaanxi,
which is very similar to the narrow-leaved individ-
uals from Yunnan. All these collections have been
assigned to K. indica subsp. indica in the present
treatment. Kitamura published a variety, Kalimeris
indica var. stenophylla, based on the individuals
with narrower leaves. Some of the specimens cited
by him clearly belong to K. indica subsp. stenolepis,
and some to K. indica subsp. indica.
The length of the pappus is another variable
character in K. indica subsp. indica, as is the case
in several other species of Kalimeris. Some popu-
lations from Yunnan have longer pappi or pappi
with united bases, conditions rarely found in pop-
ulations from other localities; but these character
states are not consistent even within populations.
For example, one collection from eastern Yunnan
(Forrest 22424) has duplicates with both longer
(about 0.4 mm, E) and shorter (about 0.2 mm, P)
pappi. These populations are distributed at the
margin of the range of the taxon, where extreme
variations are to be expected. Epappose individuals
also occur sporadically throughout the entire geo-
graphic range.
This taxon is sympatric with Kalimeris integri-
folia in Hubei and Jiangsu, the southern limit of
=
a
the latter taxon. It is also often sympatric with K.
shimadai. No intermediates have been observed in
the areas where their populations occur virtually
mixed together.
alimeris indica subsp. indica is sympatric with
subspecies stenolepis in central China, and with
subspecies collina in southern China, usually in
open areas or at lower elevations in montane
regions. In Wuchang, Hubei, and Lueyang,
Shaanxi, the senior author found nearly co-occur-
ring populations of subspecies indica (mostly in
more sunny, disturbed, and drier places) and sub-
species stenolepis (in relatively mesic and protected
areas). Based on herbarium information, subspecies
indica and collina are sympatric in South China.
The latter tends to grow in more mesic areas when
it is co-occurring with the former. These three sub-
species are distinguished mainly by indumentum,
branching pattern, and morphology of involucre and
pappi (Fig. 14).
е young rosettes of this subspecies are edible
as a salad (GH, pers. obs.) and leaves are used in
some areas as a folk medicine (Anonymous, 1990).
Because it is often cultivated for these purposes,
the wide range of this plant may be due in part to
cultivation and occasional naturalizing.
Additional specimens examined. CHINA. Shanxi: Heng-
u Xian, Tongshan-gongshe, Sanliyao, К. T. Fu 18643
(WUK). “ehas 900 m, Smith 6180 (W), Yecho-shan,
1 M, W); Jincheng Xian, S. Y. Bao € S. J.
Yan 2209 а ране Xian, Xiachuan-gongshe, Zhu-
wugou, 1700 m, K. T. Fu 18301 (WUK); Xiachuan, Yao-
shan, 1500 m, Licent 12526 (W). Shaanxi: Baocheng
Xian, Hedon adi ian, K. T. Fu 5513 (PE, WUK); Baoji Xian,
Hudian Railway Station, e m, 13859 (WUK); Hu bos
an ou, В. 2. Guo 48 (PE); Huaying Xian, Won
420 m, Huangtugaoyuan Team 1276 (WUK); Huang
Xian, Diantou-gongshe, Laohuzui, 1050 m, 487 (W
Huanglong-shan, Fengjiahe, K. T. Fu 3436 (WUK); ix
gao Xian, Silihe, from Diushan to Chapeng, 1800 m, P. Y.
Li 8439 i
WUK); Lueyang Xian, Baiquesi, C. L.
Fabi 561 (WUK), Guojiaba, K. T. Fu 5912 (PE, WUK),
from Jiangzheng to Maliutang, 760 m, C. L. Tang 399
(HIB, KUN, PE, WUK), from Wangiiatuo to Lueyang, Jia-
ling-jiang River, H. Gu & Q. Han 205 (MO), 219 Мој
from Lueyang to Wujiaying, 520 m, K. T. Fu 5965 (PE
WUK); Mei Xian, Taonggujiaokou, R. M. Zhang 679
(KUN, WUK); Mian Xian, Dingjun-shan, 900 m : Li
378 (WUK); Ningqiang ers Liejinba, de. $ 42 (KUN,
WUK), Simodi-gongshe, 950 m, J. Q. Xing 9660 (WUK),
760 m, T. N. Liu & C. Wang 107 (PE, wun, Pingli Xian,
Pingli- Sheng: 1150 m, Р У Li 1710 (WUK); Bre
Xian, Jiangxigou, 1000 m, S. B. He 547 (WUK), 400 m,
735 (WUK); Mt. Taibai- рн. Baiyunshagou, : P. Wei
1047 (WUK),
. Z. Guo 1839 pes WUK);
& P. C. Tsoong 3668 (PE), Yuan-
jia-zhuang, 1300 m, ۳ X Yang 2085 (WUK); Zhongnan-
Volume 84, Number 4
1997
Gu & Hoch
Systematics of Kalimeris
789
han, 1500 m, H. W. Kung 2845 (PE), Y. Y. Pai 1168
(PE); Siyang Xian, from Songshu Xian to Banjiuguan, P.
: Li 48 UK). Gansu: Hui Xian ji
EN PR Ay 2044 nn enm 900 m, 1
(WUK); —€— Xian, K nghe a
Ganyi Teams 1835 (PE), 1939 (PE): Wen Xian. Bikou-
gongshe, 750 m "t Zhang 13029 (WUK), 950 m
6573 (WUK), bio
gongshe, Dahai-dui, сет 1000 т, 6366 (У
2500 т, 6839 (ҰОК), 1800 т, 7094 (WUK), 3000 т,
7884 (ҰОК); үзе Xian, from Cike to goa on a
1660 m, 3541 (WUK), 1800 m, 3789 (WUK). Anhui:
Chaocheng Xian, Dong’an ng Field Station Team
3773 (PE); Shucheng Xian; Xiaotian, 4274 (PE). Henan:
Lushi Xian, Wulichuan to Hanhe, Dahegou, alt. 880 m,
J. Q. Fu 565 (HIB, WUK); Ye Xian, R. C. Ching 8679
(E, PE). Zhejiang: Tianmu-shan, T. Hong & P. K. Keng
в (KUN), Y W. Law 1220 (PE), елы i, 400 т, Zhe-
iang Expedition Team 28893 (PE), 29121 (PE), 29159
(PE): Taishun Xian, Wenping, 5. Y. Zhang 3617 (PE). Ji-
angxi: Duchang Xian, Shangshou-cun, Е Maekawa
119228 (TD); Gan Xian, Huangpodi, 630 m, Z. B. Yang &
J. Yao 1164 (PE); Jiujiang Xian, Allison 4 (GH), Sherrer
s.n. (К); Xiushui Xian, Moshagang, ed 300
S. (PE). Fujian: Gu-shan
—
`
(HIB, WUK), eti e qu 850 m, K. M. Liu 8995 (PE),
9039 (PE), 9048 angriwan, 340 m, K. R. Liu 393
(HIB, PE); Fengjie eins Shuangdian-xiang, Guanmen-
shan, 590 m, M. Y. Fang 24785 (HIB), 800 m, 24915
(HIB), Fuchikou, H. C. Chow 207 (NY); Hankou Shi, Mi-
chaelis 78 (W); Hefengcheng, Daping-nongchang, H. J. Li
2 (HIB, PE), Jigongdong, 310 m, H. J. Li 6493 (HIB,
PE) Erqu-qincaoping, 440 m 6530 (HIB, ves cd
(HIB, PE); Jiayu Xian, nogal 0:5
dian, Dashidong, 1400 т, 12738 (
tang, 1550 т, 11679 (SZ, WUK), 1610 т, 12619 (SZ,
WUK), Jingding, 1500-1700 т, 12820 (SZ, WUK), 13056
(SZ, WUK); Qixingshu, 980 m, 10725 (SZ, WUK), Wuy-
aling, 1020 m, 70901 (SZ, WUK), Zhufengxigou, 1300 m
13089 (SZ, WUK), Zixiaodian, 850 m, 10555 (SZ, WUK),
920 m, 10975 (SZ, WUK), 750 m, 11073 (SZ, WUK),
1020 m, 11273 (SZ, WUK); Laifeng Xian, Jiusi, Huangtu-
i H. J. Li 5307 (HIB, KUN, PE), 5757 (KUN,
pe 1584 (HIB, PE); Ningshan Xian, Shiga; 1120 m,
. W Zhang 1237 (WUK); Puqi Xian, Shengshan-gong-
0. 8. Wang 370 (HIB); Shijian Xian, Jinyanhe, Zuan-
dongyan, 290 m, L. У Dai & 2. H. Qian 425 (PE); Sui
Xian, Junchuan, Yao-shan, X. B. Chen 173 (HIB); Tong-
shan Xian, Jiugong-shan, Jiugong Forestry Station, L. Y
Dai et al. 2366 (IBSC); Wuhan Shi, Heyang, Shajiaping,
200 m, H. J. Li 8739 (PE), Hongshan, 200 m, C.
C. Chow 55 (A, G, NY), S. W. Teng 233 (W); Wuchang
Shi, Luojiashan, H. H. Chung 9129 (А, С, NY), 5. С. Sun
830 (NY), Wuhan University, Fukuoka c-245 (KYO), с-
258 (KYO, TI), Mo-shan, Fukuoka c-206 (TI), H. Gu $
D. Chen 122 (MO), X. 2. Long 241 (KYO, ТЇ), She-shan,
Migo 64 (KYO), Y. 2. Long 254 (KYO); Xianfeng Xian,
Disi-qu, Ganhegou, 1000 m, H. J. Li 9387 (HIB, WUK);
Xingshan Xian, G. X. Fu & Z. С. Zhang 259 (HIB, PE),
260 (HIB, PE), Fenglong-gongshe, Hongx
m, K. R. Liu 774 (HIB, PE), 1980 Sino-Amer. Bot. ~ -
pom 1105 (A, E, NY); Xuan'en Xian, Piandaoxi,
1100 m, H. J. Li 4767 (HIB, WUK), 59 (НІВ, а,
Yin an Xian, Hehua-gongshe, J. W. Wang & F. T. Wu
120 (HIB); Yichang Xian, прев ну 600 m, T. P.
Wang 11322 (HIB, PE, WUK), Sanyoudong, 200 m, Z.
Zheng 101 (HIB, PE), J. H. Zheng 96 (HIB, PE), Henry
835 (LE), 1343 (LE), 2099 (С, GH, TI), 2739a (K); Zhuxi
Xian, Dahui, 800 m, K. M. Liu 8574 (PE), Shuangzhu-
yuan, 850 m, K. M. Liu 8692 (PE), Henry 70 (GH), 1051
(E), 1100 m, H. J. Li 5486 (HIB, WUK), Wilson 1693 (K,
NY, P, W). Sichuan: Fengji Xian, Jiannong-xiang, Hon-
gyandong, 700 m, Z. R. Zhang 25884 (HIB, KUN, PE),
Dukouba, R. Z. Zhang 25652 (PE), Shuangdian-xiang, M.
Y. Fang 24783 (KUN, PE), 24785 (KUN, PE), Zhuyuan,
Gaoshi, 850 m, H. Е Chou 26593 (KUN, PE), 26886
(KUN, PE), S. X. Fu 110814 (HIB); Guangyuan Xian, P.
У Li Мыз (WUK); Jiange Xian, Lianhua-shan, 550 m, Т.
. Liu & C. Wang 342 (WUK); Wushan Жап; НН.
Chung 1702 (PE), Henry 7204 (A, GH, K). Guizhou:
Dejiang Xian, Yanmenkou, 1220 m, Z. F. Liu & T. P. Zhou
1781 (KUN).
4b. Kalimeris indica subsp. collina (Hance) H. Y.
Gu, stat. et comb. nov. Boltonia indica var. “a”
collina Hance, Ann. Sci. Nat., Bot. sér. 5, 5: 219.
1866. Kalimeris indica var. “a” collina (Hance)
Kitam., J. Jap. Bot. 19: 40. 1943. TYPE: China.
Guangdong: Guangzhou, Oct. 1859, Hance 5195
(lectotype, here designated, BM)
Rhizomes stoloniferous, 5-15 cm long. Stem green
or less often purple, 10-100 cm tall. Cauline leaves
obovate to oblong, serrate to lanceolate-lobed, 1.4—5.6
х 0.4-2 cm, densely to loosely covered with thick
nonglandular hairs on both sides and with biseriate
andular hairs abaxially. Capitula 15-50; peduncles
1.8-9(-12) cm long; bracts 1-9 along the peduncle,
the lower ones 6-9 X 1.1-3.7(-4.5) mm, the upper
0.2-0.7 mm; involucre hemispherical,
5-9 mm broad, 3—4.5 mm high. Phyllaries imbricate,
often densely covered with thick nonglandular and
glan airs, sometimes purplish; those of the out-
ermost whorl lanceolate ог obovate, 1.6-2.9 х 0.6–
1.4 mm, those of the middle whorl(s) obovate or spa-
thulate, 2.5—4.5 X 1-1.8 mm, those of the inner whorl
obovate to broadly oblanceolate or spathulate, 2.4—4.5
X 0.9-1.5 mm. Receptacles 1.1-1.7 mm wide, 0.5—
0.8 mm high. Ray florets 16-25; corollas whitish to
lilac, ligules 5.2-10(-13) X 1-2.6 mm, the tube
(0.4-)0.8-1.4(-1.8) mm long; style 1.5-2 mm long,
the branches 0.6-1.2 mm long. Disc florets (40—)70—
102; corollas 2.2-3.8 mm long, the lobes subequal,
the longer one 1-1.5 mm long, the shorter ones 0.8—
Annals of the
Missouri Botanical Garden
Figure 16. Geographical distribution of Kalimeris in-
dica subsp. stenolepis (stars) and K. indica subsp. collina
(dots).
1.4 mm long, both 0.4-0.6 mm wide, the tube 0.6-1
mm long; anther 0.8-1.4 mm long; style 2.2-3.3 mm
long, the branches (0.4—0.7—1 mm long. Ray and disc
pappus bristles similar, 0.1-0.3 mm long, or some-
times epappose. Achenes purplish, the margins about
0.1 mm broad; ray achenes 3-ribbed, 1.5-2 х 0.7-
0.9 mm; disc achenes 2-ribbed, 1.5-2 X 0.7-1 mm.
Chromosome number: n — 18. Figure 14F, G.
Phenology. Flowering period: late May to Novem-
be
г.
Distribution (Fig. 16). Kalimeris indica subsp.
collina is centered in South China, mainly in
Guangdong and Guangxi, just reaching northern Vi-
etnam. It barely overlaps at its northern limit the
range of K. indica subsp. stenolepis. It occurs in the
following provinces: Jiangxi, Fujian, Guangdong,
Hainan, Guangxi, Hunan, Guizhou, and Yunnan.
Its altitudinal range is 200-1700 m, but occurs pri-
marily at lower elevations. It commonly occurs in
disturbed habitats, usually along roadsides or at
edges of crop fields.
Although Hance did not indicate any specimen
when he published Boltonia indica var. collina, nor
in a subsequent article (Hance, 1873) discussing it,
the specimen collected by him (Hance Herb. No.
5195) from Guangzhou (Canton) with his annotation
“Boltonia indica a Benth. collina Hance” is here cho-
sen as the lectotype. Two other Hance collections from
the same locality with the same number were col-
lected on different dates, November 1859 (K, W) and
October 1863 (W), and lack his annotation.
This subspecies is characterized by its dense,
thick, nonglandular hairs and biseriate glandular
hairs on the leaves, especially abaxially, on the up-
per stems and branches, and by its densely dis-
posed peduncular bracts. Its capitula are smaller
than those of the other two subspecies, and its ray
florets are whitish rather than lilac to pale purple.
This taxon has sometimes been misidentified as K.
shimadai in the herbarium because of its robust-
ness and relatively dense hairs. But its smaller
heads, short and thick hairs, and elongated rhi-
zomes distinguish it from K. shimadai.
Additional specimens examined. CHINA. Jiangxi:
Anfu Xian, Wugong-shan, Wenjia, 320 m, J. S. Yue et
al. 2826 (KUN, PE), Kenzhichang, S. S. Lai 1797 (PE);
Dayu Xian, Guba, 500 m, M. X. Nie et al. 9650 (KUN);
Huichang Xian, Fucheng-xiang, Bankenshe, Q. M. Hu
3375 (IBSC, KUN, PE); Longnan Xian, Wuchi-shan,
Linwu-cun, S. K. Lau 4757 (A, BM, G); Nankang Xian,
from Fushi to Dashannao, 340 m, M. X. Nie et al. 9757
(KUN); Ningdu Xian, Q. M. Hu 5522 (PE); Ninggang
Xian, Dongshang-gongshe, 300 m, S. S. Lai et al. 5290
(KUN); Quannan Xian, Dongli-xiang, Xikenzhang, S. K.
Lau 4097 (A, G), from Lishuxia to Shangying, 330 m,
Jiangxi Teachers’ College Team 12037 (HIB); Shicheng
Xian, Kuangkou-xiang, Q. M. Hu 44617 (PE); Suichuan
Xian, Dafenghou, Linyang, 670 m, Q. S. Yue et al. 3865
(PE), from Nanjiangkou to Youduqiao, Q. Е Liu 30164
(PE); Xunwu Xian, Jianxi-gongshe, J. S. Yue et al. 1706
(PE). Fujian: Dehua Xian, Leifeng-xiang, K. M. Wu
60435 (IBSC); Fuzhou Shi, Lingpu-shan, 7. S. Ging 5233
(MO), 5399 (BM, MO), Wushi-shan, H. H. Chung 2402
(E, G), Carles 799 (E), H. H. Chung 3815 (PE); Hua’an
Xian, Jiao’an-gongshe, Futian, K. M. Wu 60165 (IBSC);
Liancheng Xian, Y Ling 3281 (PE), Guanxiang-shan,
3248 (PE); Minhou Xian, Gu-shan, H. H. Chung 2326
(A, МО, W), С. C. Tang & S. E. Ма 2906 (A), S. G.
Tang 5706 (BM, MO); Nanjing Xian, Dalingnan-shan, Q.
F. Guo 140036 (IBSC), Xiamen University Team 333
(PE); Sha Xian, Q. L. Chen 65 (PE), Fujian Botanical
Expedition Team 53428 (IBSC, WUK), P. C. Tsoong 26
(PE); Shaowu Xian, Guandun, H. N. Fan 9784 (A), Nan-
bao, 9728 (A), Nantian, 9718 (A), Tieyang, 9588 (A),
Xiaping, 9557 (A), 9610 (A), Xieshuken, 1250 m, 9654
(A). Zhangjia, 1170 m, 9623 (A); Yong'an Xian, Bai-
Ling 2965 (PE), Maoping, 2174 (PE).
Guangdong: Conghua Xian, from Lumuhe-shuitang to
Xiaoshui, L. Deng 8419 (KUN); Пари Xian, Fengxi For-
estry Station, 500 m, X. W. Wang et al. 192 (IBSC, PE),
Gucun-xiang, L. Deng 5081 (IBSC, PE), Shenpucheng-
zheng, Wenfushe, X. G. Li 202806 (IBSC, PE), Tonggu-
shan, W. T. Tsang 21594 (A, GH, IBSC, K, KYO, LE,
NY, PE); Guangzhou Shi, Sampson s.n. (BM), South Chi-
na Institute of Botany, H. H. Ye 462 (IBSC), Rechinger
22 ), Y Tsiang 3282 (NY); Hainan Island, Baisha
Xian, Yayi-xiang, Yaguanglu, H. D. Zhang 1929 (IBSC),
Dan Xian, Meiyang-cun, W. T. Tsang 776 (A, G, K, NY,
Volume 84, Number 4
1997
Gu & Hoch
Systematics of Kalimeris
PE); Huidong Xian, Pingshan-gongshe, Chenshuisha-
ngjiao, B hen et al. 221 (IBSC); Jiaoling Xian,
Changtan, X. H. Xu & H. Е Lin 153866 (IBSC), Nanli-
xiang, Sanxijiang, X. G. Li 202385 (IBSC); Lechang
Xian, Beixiang-qu, М. С. Dong 1472 (KUN), 5. Q. Chen
1472 (PE), Beixiangqu Forestry Station, H. Gu & 2. L
306 (MO), 307 (MO), 308 (MO), 309 (MO), 310 (MO),
314 (MO); Liannan Xian, Shi-shan, Sanjiang-zheng, 150
m, P. X. Tan 58868 (KUN, PE), Jinken-xiang, Huang-
dong, 300 m, 59479 (KUN, PE), Yao’an- xiang, Dapotou,
300 m, 59944 (KUN); Lingyun Xian, Yupi-xiang, X. Q.
Liu 28542 PE) Luoding Xian, “Kolan,” Y. Tsiang 1189
H, N ngmen, Nankun Forestry Station, G. С.
Zhang = a 81 (IBSC), > So
F. Wei 121999 (IBSC, KUN, PE); Maoming Xian, Dapo-
qu, Gexiang-xiang, L. -— 2062 (KUN): Mei € Jiay-
ing, Yingnashan, W. T. Tsang 21400 (GH, KYO, NY, PE);
Ruyuan Xian, Er-qu, Tixiashan, Z. Huang 42350 (IBSC,
SYS); Shantou Shi, Maclagan 124 (BM); Shixing Xian,
Luoba-xiang, Dongxing-she, Da'anken, 300 m, L. Deng
7027 (KUN, PE), from Luoba-xiang to Huangken, 200
m, 7108 (KUN, PE), Zhangdongshui Conservation Area,
Shixing Team 172 (IBSC); Wengyuan Xian, Diyi-qu,
Zhanyuan-xiang, Shangmiao-shan, X. Q. Liu 25137
(КОМ, PE), Huangzhuyi, Qingyun-shan, 5. K. Lan 664
(С, GH, NY), 2152 (GH, IBSC); Xinfeng Xian, Huangpo-
xiang, Lugushe, 650 m, L. Deng 7972 (KUN, PE); Xinx-
ing Xian, Tianjie-xiang, Sijialiao-kenshe, ¥. G. Liu 2538
(HIB, PE); Xinhui Xian, Gudou Forestry Station, Wus-
ongkou, Y. 2. Gao
~.
ang, Liantang, 1200 m, P.
60368 oe ip ut 1300 m, 60369 (IBSC, KUN);
Yangzhun Xian, . Z. Huang 37257 (IBSC, PE),
асель em e SYS) Zhaoqing, Dinghu-ca-
ochang, G. L. Shi spec (1850), Qixing-gongshe, So-
isk dung. Guangz isi Team 4956 (IBSC), Qix-
ingyan, G. L. Shi 12093 BSC) Levine 3446 (E, G, MO);
T. K. Ping 1884 (E, P i: Damiaoshan Xian,
Antai-qu, Sha litio ales: $: QA. Che
ене.
14484 (IBSC, KUN, PE), Pingshi-xiang, Jiuxianheng-
gang, 500-700 m, 16312 (IBSC, KUN, PE); Dayaoshan
Xian, 1700 m, C. — 40255 (A); Guilin Shi, ares
gyan, Putuo-shan, 200 m, Guangxi Team 3760 (PE),
Pangu-shan, Chaotian- fond Sanming-cun, W. d ^ uim
28036 (A), 220 m, Chow & Wan 79046 (KYO, NY); He
Xian, Babuzheng, 5. Q. Zhong А 60881 (КИМ), Gouzi-
tun, Y. K. Li 401235 (IBSC); Huaiji Xian, Dangshan, W.
T. Tsang 22759 (G, GH, W), Linwan . Lau 28542
p Tongthongeun, Touyueshan, W. E Tsang 23247 (G,
Zhong A
3
T
(KUN); e e e shan
S. 0. Chen 11800 (KUN, PE), Paizong-xiang, 360 m,
14007 (IBSC); Nanping, Nanpingdapuo L. 8. Ou 10
(IBSC); Shanglin Xian, E "тын Kimara s.n. (KYO
Tian'e Xian, — Z. Huan, pola (KUN, PE):
, W. T. Tsang
unan: n Xian, N : Sivndisi; H. Т.
Chang me (BSC); Xinhua Xian, Xikuang-shan, 500—
800 m, Handel-Mazzetti 2694 (A, E, W). Guizhou: Ап-
long Xian, Huali-gongshe, 900 m, Anshun Team 3221
PE, , 1700 m, 3947 (IBS K); Danzai
Xian, Xingren, Chenjiagou, 1300 m, an (IBSC, WUK);
Lugong- pe Niyu, m, 80 C); Dushan
Xian, 400 m i i
-shan, Qiannan Team 232
Meitan Xian, Yonghe-chang, 900 m, T. P. Zhou
Liu 2421 (KUN, PE); Weng’an Xian, Yonghe-gongshe,
Changlin-guan, liqu, Lipuo Team 2249 (KUN, PE). VI-
ETNAM. Tonkin: Langson, Bois 82 (P), Eberhardt 3284
(P), Lecomte & Finet. 93 (P).
4c. Kalimeris indica subsp. stenolepis (Hand.-
Mazz.) H. Y. Gu, stat. et comb. nov. Asteromoea
indica var. stenolepis Hand.-Mazz., Acta Horti
Gothob. 7: 225. 1938. Kalimeris indica var.
stenolepis (Hand.-Mazz.) Kitam., J. Jap. Bot.
19: 340. 1943. TYPE: China. Fujian: Yenping
Xian, 4 Aug. 1924, H. H. Chung 2895 (holo-
type, W; isotypes, BM, MO, PE).
Kalimeris indica f. gracilis J. Q. Fu, Bull. Bot. Res., Har-
bin 3: 112. 1983. зра China. Gansu: Wen Киш,
Bikou-eonethes T апђа, 900 т, 9 Зер. 1959, 7.
Y. Zhang 14791 пре WUK).
Rhizomes 0.5-1(3.5) cm long. Stem green, 45-
150 cm tall, densely covered with thin nonglan-
dular hairs and interspersed with biseriate glan-
dular hairs on the upper part, loosely so on the
lower part. Cauline leaves lanceolate to oblong, ser-
rate to deeply pinnate-lobed, 3-6 X 0.6-2 cm,
densely covered with thin nonglandular and biser-
iate glandular hairs abaxially, densely or loosely so
adaxially, lobes 0.9-2.3 X 0.3-0.5 cm. Capitula
40-100; peduncles 3-9.5 cm long; bracts 4-12
along the peduncle, the lower ones 6.2-19 X 1.1-
3.2 mm, the upper ones 1.6-3.6(-5) X 0.3-0.7 mm;
involucre hemispherical, 5-7.5(-10) mm broad, 3—
6 mm high. Phyllaries imbricate or sometimes
loosely so, covered with thin nonglandular and
glandular hairs; those of the outermost whorl lan-
ceolate to oblong, 1.9-3(-3.8) X 0.5-0.9(-1.1) mm,
those of the middle whorl(s) oblanceolate to obo-
vate, 2.4—5.7 X 0.7-1.4(-1.7) mm, those of the inner
whorl oblong, oblanceolate, or obovate, 3—5.7 X
0.8-1.4(-1.8) mm. Receptacles 1.2-2.5 mm wide,
0.5-1.2 mm high. Ray florets (12—)14—25; corollas
whitish to pale lilac, ligules narrowly oblong, 6-14
х 1.3-3 mm, the tube 1.1-2 mm long; style 1.5—
2.4 mm long, the branches 0.5-1(-1.3) mm long.
Disc florets 30-68; corollas 3-4 mm long, the lobes
unequal, the longer one 1-1.6 mm long, the shorter
ones 0.7-1.3 mm long, both 0.4-0.6 mm wide, the
tube 0.6-1.2 mm long; anther 1-1.5 mm long; style
2-3.5 mm long, the branches 0.6—1(-1.4) mm long.
Ray and disc pappus bristles subequal to unequal;
792
Annals of the
Missouri Botanical Garden
the ray bristles 0.1-0.3 mm long; the disc bristles
0.1-0.4(-0.5) mm long. Achenes brownish, the
margins 0.1-0.2 mm broad; ray achenes 3(—4)-
ribbed, 2-2.5 X 0.7-1 mm; disc achenes 2(-3)-
ribbed, 2-2.5 X 0.9-1.3 mm. Chromosome num-
ber: n = 18. Figure 141-K.
Phenology. Flowering period: June to late Octo-
T.
Distribution (Fig. 16). Kalimeris indica subsp.
stenolepis is endemic to China, where it is especial-
ly abundant in Hubei and surrounding central Chi-
na. It occurs in Shanxi, Henan, Shaanxi, Gansu,
Anhui, Zhejiang, northern Jiangxi, northern Fujian,
Hubei, Sichuan, and Guizhou. Its primary eleva-
tional range is m, but it occasionally
reaches 3000 m in Sichuan. It usually grows in
relatively shady and protected habitats.
The type collection is from Chaping, Fujian, at
the southern limit of the range, and as such does
not represent the taxon well, since it includes spec-
imens that are shorter and less vigorous than in-
dividuals from near the geographic center. Handel-
Mazzetti’s concept of this taxon was broader than
that in the present treatment. He included in it
some narrow-leaved plants that we treat as Kali-
meris indica subsp. indica, which has some char-
acters similar to this taxon but differs by its gla-
brous, thicker leaves and broader phyllaries. Some
specimens cited by him (e.g., Tang and Hsia 196
and Bois-Reymond 805a, W) doubtlessly belong to
subspecies indica.
Some plants from Jiujiang, Jiangxi (e.g., Allison
4, GH; Migo 46, KYO; Sherrer s.n. in 1873, K) have
less hairy leaves and relatively elongated rhizomes,
which are not typical for this subspecies, but other
characters indicate that they should be included in
subspecies stenolepis. The pappus bristles of this
taxon tend to be conspicuous and are often united
at the bases. Two collections from Hubei, H. C.
Chow 378 (NY) and Henry 2099 (GH, TI), have
pappi up to 1 mm long.
Additional specimens examined. JAPAN. N agasaki:
Nagasaki Shi, H. Ando 27 (MAK). Kagoshima: Ohshima
un, Amamiohshima, /to 510 (TNS); Tarumizu Shi, Shin-
oo, from Kami-shinmidoo to Horikiri, 50—500 m, Mur-
ata 11496 TD. Drs Ishigaki Shi, bhigakijima,
Iwasaki 2 (KYO), Yamasaki s.n. (ТІ); потап Shi, Furus
1589 (K); Kikai duc Miyagi 7277 (MAK), between Cu-
suku and Hyakunodai, Yoshinaga 523 (MAK); Nago Shi,
Harano, Коноје 18 (КУ О), Yaedake, 400 m, Yamazaki
;N
TNS), Y KYO);
Okinawa Shi, Nishihara, Tawada 3 (KYO); ERE ors
ma, Ohba 48 ( YO); Shimajiri = oO vst Koidzum
n. (KYO); Shuri, Hatusima 17309 (ТІ); Yoron hind.
uiui: Tagawa & Iwatsuki sais (KYO), Hatusima &
Sako 30887 (MAK). KOREA. Junn ^
s.n. (KYO); Namwon, ete 3 . Cheju Island: Mal-
laisan, Taquet 22 G, W), Taqet 9679 (TD, Nakai
6520 (TD. CHINA. e одеће Xian, Tong-pu rail-
way, T. Kanasiro 4757 (KYO, PE). Shandong: Qingdao
Shi, Zhongshan Park, H. Gu & У. Gao 292 (MO, PE), 293
(MO, PE), 293a (MO, PE), 295 (MO, PE). Henan: Lushi
unnam: Damyang
Huji-shan, 2. У. Zhang 189 (HIB, KUN, PE, WUK); Fop-
ing Xian, Gaojiaba, 1150 m, K. T. Fu 4945 (PE, WUK);
Langao Xian, Hengbin River, 1800 m, P Y Li 8153
(KUN, WUK), Tiefo-gongshe, 1800 m, 8512, (KUN,
WUK); Liuba Xian, Miaotaizi, 1600 m, K. T. Ки 6326 (PE,
МИК), Zibai-shan, 850 m, 6185 (PE, WUK); Ningqiang
Xian, Kuanchuan, 800 m, P. Y. Li 94 (WUK), Liejinba,
14 (KUN, WUK), Yangpingguan, Y. L. Qiao 108 (WUK),
750 m, T. N. Liu & C. Wang 50 (PE); Lueyang Xian, from
Luotuo-xiang to Lianghekou, 870 m, C. L. Tang 976
WUK), from Wangjiatou to Lueyang, 800 m, E Cu & E
e 206 ete 208 (MO); Yanglin-zheng, Weihe Riv
u & X. Hao 6 (MO), 7 (M зав 8 кне Pingli Xian,
de Pingli to Yaofu-shan, 1200 m 910
WUK), Yaofu-shan, 1200 m
~
(KUN, WUK); е Mts., Qinfeng-shan, 1500-1700 т,
Fenzel 136 (W); S ian, 750 m, T. P. Wang 16022
(РЕ, WUK); Pies Xian, Gangtie-gongshe, Huangguan
River, 1020 m, J. Q. Xing 7992 (WUK); Puhe-gongshe,
63
1100 m, X. X. Zhan et al. 1067 (WUK); Zhouzhi Xian,
Pai 1340 (PE); Ziyang Xian, Chengguan-gongshe, Moshi-
gou, P. Y. Li 7330 (КОМ, WUK); Liangmaya, 700 m
(KUN, WUK); Maoba, 700 m, 7056 (KUN, WUK); Shu-
an nghe- -gongshe, 1750 m, 7211 (KUN, WUK), 4791 (KUN,
WUK), 4933 (КОМ, WUK), 7194 (KUN, WUK). Ningxia:
Jinyuan Xian, Xisha, Ningxia Team 1-0228 (WUK). Gan-
su: Hucheng Xian, from Taihe-gongshe to Zu-zhuang,
1400 m, 2. У. Zhang 662 (WUK); Guyuan Xian, Wating,
1400 m, 7. P. Wang 17106 (PE, WUK); Kang Xian,
Changba, Shijiagou, Z. У Pug 16908 (WUK); one Da-
caochu uan to Xi
; Wen € Bikou-gongshe, 900
Z. Y. Zhang 14791 (WUK), 650 m, Y. Q. He 1102 (PE)
Dongjiaheba, 14897 (WUK). Anhui: Anqin Xian, Q.
Hu 3 (PE), Maekawa 741 (TI); Chu Xian, Langyia-shan
Huadong Field Station Team 3170 (KUN); Guangde Xiah,
m, Anhui Ex ловити pares 3300 (NAS); Huaining
Xian, Ganlusi, Maekawa 4 (TD; — 7 B
(TD; Linhu-xiang, 745 rn Maohuling, 8 B22 (TD;
Huang-shan, Chow 33 (A, K, KYO, MO, PE, US), from
Chuishilin to Tangkou, T. N. Liu & P. C. Tsoong 2170
dms around Tangkou, 2365 (PE); Jiuhua-shan, ~ m, С.
п € Y Y Lil 00-800 m, S. C. Sun 1239
(5, 1 259 (NY); Перуна; Huadong F ield nio Team
4795 (PE). Jiangsu: Baoying Xian, Kenzhichang, Nanzha,
Volume 84, Number 4
1997
Gu & Hoch
Systematics of Kalimeris
793
S. L. Liu et al. 18 (IBSC); Danyang Xian, Nanjing Forestry
College Team 7800-447 (IBSC); Jinjiang Xian, Zhulinsi,
Migo 37 (KYO); Nanjing si Wutai-shan, Merrill 1509
(MO); Xiaolingwei, Matsuda s.n. (TI); Suzhou Shi, ic
yin-shan, Migo 44 (KYO); bake A LEA
(GH); Shangfang-shan, Migo 43 (KYO); Yixing. Xian,
Longzhi-shan, W. Z. Fang et al. 102 (IBSC), C. N. Chen
8603 (W). Shanghai Shi: Minhang, Kimura s.n. (KYO);
Paoshan, Migo 36 (KYO), Zhelin, 41 (KYO); Shanghai
і NAS); Longquan ‘ge e
6828 (PE), 7107 (PE): Dd jte (PE
shan, 730 m, - & Wilson s.n. (GH); N
ber s.n. (E); Siming-shan, hia eit Y He 24938 (PE),
25882 (PE); ake 27448 (PE); Shouchang Xian, Shang-
cang, Zhejiang Team 27655 (PE); “Siakan, Fan-chiao,” R.
C. Ching 3629 (W); Shujiang d dee 35 a Tonglu
ian, Hengcun-zheng, 5. Chen 1829 (PE). Jiangxi: Dex-
= бар-а Mao-shan, M. X. E 5172 (PE); Guangchang
. Tangfan-xiang, Tianyao-she, Taishan, Q. M. Hu
5297 (IBSC, KUN); Tangwan-gongshe, J. 5. Yue et al.
2181 (KUN); Guixi Xian, Tianhua-shan, 800 m, S. S. Lai
& M. X. Nie 3809 (KUN, PE); Huchang Xian, Hedong,
Q. M. Hu 2966 (PE); Jinggangshan Xian, Jinggang-shan,
о
(5), Forbes 43 (BM), Schindler 308a (BM, E, K); Hanbok-
u, 1120 m, Migo 65 (K pied Tsiang 10770 (NY); Lian-
bie T. R. Liu s.n. (KYO); Louxiansi, K. J. Guan
74457 (PE), R. C. Ching pe (PE), 10260 (PE), 10462
(PE), S. Xiong 7109 (PE); Xinziyikou, Qianshusi, M. X.
Nie & S. L. Chen 7418 (KUN, PE); Jiujiang Shi, Lu-shan,
Migo 65 , 66 (TD, Bailey s.n. (GH), Carles s.n. (E).
Forbes 83 (BM), 927 (BM); Lichuan Xian, Wuyi Mts., De-
shengguan-nongchang, Maodian-fenchang, M. X. Nie et al.
3031 (PE); Lianhua Xian, Mi поарно pd
dadui, S. S. Lai 1651 (PE); Libei vow Libei B
7 n Xian, Wuyi. -shan,
* (PE); Xikeng, 751 m, M. X. Nie
152 (KU g Shi, T. N. Hsiung
539 cin E (GH), he (PE), X. X. pow. 10836 (PE);
Ningdu Xian, Fanxiangping, Xiapinglongkeng, Q. M. Hu
5522 (IBSC, KUN); Pingxiang Shi, from Xingdian to Ma-
tian, 420 m, Jiangxi Team 835 (PE); Xuande-qu, م m,
2419 (PE); Ruijin Xian, Lishupailiantang-she, Q. M. Ни
4223 (PE); Shangrao Xian, Wufu-shan, 700 m, M. X. Nie
& S. S. Lai 4681 (KUN, WUK); Shuichuan Xian, Dafen-
ks Linyang, 670 m, J. S. Yue et al. 3865 (KUN); Wangx-
ng Xian, Damao-shan, M. X. Nie & S. S. Lai 5172 (KUN);
1 1 Mts., Matoushan-xiang, 500 m, 3508
T. S. Ging 6657 (G, мо); northern Fuzhou, 5667 ri d
7143 (G, MO); sce a ок Migo s.n. (PE); Qushi-shan,
Nagasawa Xian, do [иб University
ijin
Team 50299 (KUN), 52402 (KUN). Taiwan: Danshui
Xian, Faurie 226 (BM, KYO), 1141 (BM. G, GH, KYO),
Nagasawa 523 (KYO, ТІ), Oldham 244 (BM, K, NY);
Jilong Shi, Ford 65 (K), Makino s.n. (TI); Taibei Shi, “Sek-
ten,” Keng & Kao 1276(1) (E), Shilin, Shike s.n. (TI),
Makino 169 (MAK), Asai s.n. (NY), Makino s.n. (TNS),
Tanaka 93 (BM, P); Taibei Xian, Shimada 5471b (KYO);
Yilan Xian, Dali, 5472b ме MM Va Congshi
ian, Huogangyuejin-gongshe, 650 m, L. Deng 8732
(KUN), 8736 (PE) Fengkai еқ Heishiding Natural
Conservation, B. H. Liang 12882 (IBSC); Fushan Xian, С.
Wenyon 1887 (K); Guangzhou Shi, Zhongshan University,
. Y Lau 20551 (IBSC), Levine 1081 (GH, MO), Sampson
319, (BM); Honam Island, Levine 272 (A, GH, МО), Mer-
rill 10021 (GH, NY); Huangpu Xian, Bei-qu, Huangpo-
yang Xian, Lingbei-gongshe, Lutian, Y. M. Cheng 170542
(IBSC); Leohaig Xian, Zhuangyuan-shan, Jinjiufeng, ЈУ. 7.
Tsang 20844 (K, on: MO, NY, PE, W); Ao
Callery s.n. (G). G i i
900 m, C. C. плина 10678 (KUN). Hunan: Wugang Зага
Yun-shan, Nantianmiao, 310 m, 1. Н. Liu С. Z.
16483 (PE); Xinhua Xian, Xikuang-shan, 5
Handel-Mazzetti 2694 (W). Hubei: Badong Xian, Si
gongshe, Baiqiao, 1000 m, Hubei Expedition Team 24197
(HIB); Shengnongjia, Niudongwan, Dongjiekou, G. X. Fu
& Z. S. Zhang 1145 (PE); пуна Xian, Taihe-gongshe,
Q. S. Wang 78 (HIB); Ensi Xian, Siqu-xiang, 1000 m, H.
J. Li 4791 (PE), Xinping-gongshe, Hejiadaping, 310 m,
8900 (HIB); Fengjie Xian, Zhuyuan, Gaozhi, 250 m, H.
F. Chou 26953 (HIB); Huangshi Shi, Bao'an, H. C. Chow
18412 (G, NY); Dayu, 348 (NY); Wuchang Shi, Luojia-
shan, S. C. Sun 772 (NY), 849 (NY); Mo-shan, H. Gu &
D. Chen 121 (MO), 127 (MO); Xianfeng Xian, Disi-qu,
000 m, H. J. Li 9387 (KUN); Shuishapi. ag-xiang, 1400
m, 9051 (HIB, PE); Xuan'en Xian, Piandaoxihou-shan,
1000 m, 4767 (HIB, PE); Yichang Xian, Dengcun,
m, H. Gu & D. Chen 134 (MO); Xiaoxita, 795 (MO),
200 (MO), Wilson 1693 (W), 1703 (K, NY, W); Zigui Xian,
Kongjiaqiao, 1300 m, T. P. Wang 11865 (PE, WUK);
Wuzhi-shan, 72098 (PE, WUK); Zhuxi Xian, Longdong-
he, 830 m, P. Y Li 11118 (KUN, WUK); Tangfonggou,
1500 m, 8797 (KUN, WUK), 9427 (KUN, WUK); from
Xiaohegou to Shizi-shan, 9976 (KUN, WUK), Henry 89
(GH), 216 (BM), 384 (GH), 5209 (GH). Sichuan: border
with Shaanxi Province, P. Y. Li 785 (WUK); Baoxing Xian,
Fumugou, 1100 m, K. J. Guan & W. T. Wang 2629 (PE);
Xikang, C. Pei 8083 (PE), T. P. Tsoong 39422 (KUN, PE);
Chengdu Shi, Ximen, W. P. Fang 12360 (A, BM, W), S.
S. Chien 5320 (W), 5948 (E), 500 m, PT 94 (P), T.
В ген 8282 (PE); Chengkou Xian, Qigan-shan, Erguai,
‚ T. L. Dai 102239 (PE), 102260 ets 102532
(PE), 600 m, 103190 (PE), 106939 (PE), 1300 m, 107099
(PE); Ebian Xian, S. L. Sun 982 (KUN), Z. X. Zhao 550
(KUN); Mt. Emei, C. Y. Chiao & C. S. Fan 208 (A), 2500
m, S. C. Sun & K. Chang 1170 (A), 2300 m, 1206 (A),
900 m, 1417 (А), 480 m, 1772 (A); Fuhusi, 510 m, K. J.
oon 1963 (PE), G. H. Yang 56939 (KUN); Heilongjiang,
, C. L. Li 1938 (CDIS); Leigongsi, 800 m, K. J.
Guan « ч al. 725 Fis Emei Xian, 5. С. Sun « K. Chang
7 (A); Hanyuan Xian, Xienline- -shan, 1200 m, 7. P.
Pb 887 (PE, sages 8910 (PE, WUK): Hechuan Xian,
450 m, W. P. Fan Wawu-shan,
Shuangdongxi, Z. ^ Yao 4042 ian
zi, 1850 m, Q. S. Zhao & S. S. Liu 5704 (IBCD); caes
Xian, Sanyu School, H. C. Chow 9008 (A); Ximen, 8940
(A); Luding Xian, Tianba-gongshe, 1470 m, K. J. Guan
W. T. Wang 1594 (PE); Nanchuan Xian, Jinfo-shan, Lan-
caoba, Longdong, G. L. Li 63217 (KUN), 1550 m, 63279
(PE); Sanjiaqianfoyan, 550 m, K. J. Guan et al. 854 (PE);
Xiaohenima-cun, J. H. Xiong & Z. Y. Chou 91840 (PE),
B
794
Annals of the
Missouri Botanical Garden
92444 (PE), 94009 (PE); Shizhu Xian, Muping-gongshe,
1100 m, У Chen 2968 (IBCD); Qiyaoshan Forestry Sta-
tion, 1710 m, 2137 (IBCD); Tianquan Xian, Chengxiang,
1090 m, D. У Peng 45389 (IBCD); Sihang-xiang, Xin-
. Hu & Z. He 12026 (PE); Xinchang-qu,
W. T. Wang 2264 (PE); Eryin-shan, 1900 m, X. M. Jiang
35676 (PE); Meine Xian, Zhongting to Fangjiatian, 800
m, P. Y. Li 6799 (WUK); Tianba-xiang 700 m, 6909 (KUN,
WUK); Dazhudong, 820 m, 4223 (WUK), 4420 (WUK),
5501 (WUK), 5560 (WUK), 5620 (WUK), 5963 (WUK),
and 6813 (WUK); Wuxi Xian, Gaoyuo-xianggongsuo, 1000
m, G. H. Yang 65084 (KUN), P. Y. Li 3308 (WUK); Zhao-
чё Xian, Shangsi, 900 т, 2. Р. Wei 3855 (WUK); SW
Sichuan, SE of Tia 3000 m, Forrest 22424 (E, W).
Guizhou: Anlong Xian, Longshan-gongshe, Qiaolingying-
qu, 1300 m, Guizhou Team 4107 (IBSC); Anshun Shi, An-
shun Team 1903 (KUN, PE); Biji Xian, Er-qu, Baohe-
xiang, 1300 m, P. H. Yu 489 (KUN); Ceheng Xian, Hu-
arong, 800 m, 2. Y. Cao 855 (PE); Rongbei, 1500 m, 2.
Y. Cao 394 (PE); Chishui Xian, Biji Team 1190 (KUN);
Dafang Xian, Baina-qu, Jiulong-shan, 1850 m, 983
(KUN); Dejiang Xian, Heba Forestry Station, 1050 m,
Qianbei Team 2697 (PE); Yanmenkou, 1781 (PE); Fenjin-
shan, Dahoyan, 900 m, Steward et al. 450 (E, GH, NY,
PE); Guiding Xian, 600 m, C. C. Chang 5577 (GH, NY,
E, W); Guiyang Shi, Войне 1913 (Е, Р), 2474 (К);
Guizhu Xian, Dangwu, Heishitou, 1150 т, Chuangian
Team 2076 (PE); Kaili Xian, Xumu-chang, Shanna Team
3723 (KUN, PE); Taojiang, Donglei-shan, 800 m, 2329
(KUN); Luke Xian, Jinjiagai, 1200 m, Qianbei Team 2336
(PE); Nayong Xian, Juren-qu, 1800 m, Biji Team 495
(PE); Pingba Xian, "med = m, Anshun Team 1759
(PE); Pingyuan-qu, Shawqi-xiang, 1000 m, 2. Е Liu & T.
P. Zhu 2697 (KUN); Pu' iin i Xian, Qingsh an-songshe, Pub-
exp Team 1376 (PE); Qing-
huanqian Team 1823 (PE);
Rongjiang Xian, Fruit Qiannan Team 3065 (KUN,
PE); Songtao Xian, Ganlong-qu, Hunq Z. F
Liu & T. P. Zhou 1439 (KUN, PE); Mu'erxi, 830 m, 1983
1
(KUN, PE); Xishui yh E -qu, Xinhua-zhuchang,
800 m, Biji Team 1564 (KU
gongshe, 1300 m, Gui
Fenghuang-shan, Chuangian Team 853 (PE); Liangfengya,
1100 m, Steward et al. 51 (A, LE, NY). Yunnan: Cheng-
jiang Xian, E E 1800 m, H. Wang 41433 (^,
KUN), 1 }
stry Station to Mugan-
Di ретту ибну Team 507 (KUN); Dali
Xian, Liujufeng, H. C. Wang 912 (PE), 4093 (PE); E of
Dali, 3300-3900 m, Forrest 4005 (BM, E), 3000-3300 m
CD); ree van
Guandu-qu, 1900 m, ‚ 77613 rh Heilongtan, 50787
(IBSC, PE); Huiliuwan, K. M. Feng 10290 (KUN, PE
Xishan, 1800 m, B. У Ош 77566 (BCD); Lufeng Xian,
1700 m = T Tsai 53573 (A, E, he PE); Lushui Xian,
“Maan-tsang, Shenn-mang-yean Wang
80971 (A, A. KUN, PE); Luanliang Xian, Eryuetianmalin-qu
17 iando
in; ngbei Team 772 (KUN), 773 (KUN);
Mengzi mg: “S of R m, He (Е,
K, LE, MO, NY); Mingjiulao-zhai, 1800 m, Y. У Hu & S.
K. Wen 580599 и Xichou, 1400 m, 580715 Mp
Ledug s.n. (P); Pingbian Xian, Maoyitou, 1250 m, T. W.
Wang & Y. Liu 82264 | (KUN, PE), 1300 m, 7. T. Tsai
60729 (A, KUN, PE), 61066 (A, KUN, PE), 400 m, 62482
(A, KUN, PE), 62827 (A, «с» PE); Tengchong Xian,
Siyuansishi-gongshe, Xian e, 1750 m Yi 60-
1238 (KUN); Y astro Xian, Howell 95 (E, K); Жер Xian,
m, T. У. Wang 6 A, KUN, PE); Lidiping, 3000
m, 67686 (A, PE, UK) Yanshan Xian, Tuguo-zhai, 1200
m, T. W. Wang & Y Liu we (KUN, WUK); Yanjin Xian,
Qinghe River, Pu’er-du, 500 m, Diandongbei Team
(KUN); Guanhe River, 892 (KUN); Yiliang Xian, 1800 m,
В. У Qiu 771226 (IBCD); Zhengxiong Xian, San-qu,
Guangduogian-laolin, 1950 m, P. H. Yu 992 (KUN, PE);
Ducloux 289 (P), 1585 (E), Forrest 3063 (BM, E), 8312
(E); northern Lijiang valley, 300 m, 13025 (BM, E), 28457
(BM, MO), 30584 (BM), Maire 50 (BM), 351 (BM), 454
(BM), 525 (BM), 2561 (NY), 2568 (NY), F. T. Wang 453
(KUN), 2247 (KUN). Hong Kong: Fortune 124 (BM,
mixed with Forbes 43 and 83), Hance s.n. (W). VIETNAM.
Dong Dang, Petelot 400 (P); Hung Hoa, Hanoi, s.c. 92 (P);
Kiendi, Prairie, Balansa 4498 (P); Langson, 840 (K, P),
Hautefenille 161 (P). THAILAND. Ban
medicine shop, Kerr s.n. (BM); no
1200-1685 m, Iwatsuki et al. T 9428 (KYO); Phitsanulok
in Phu Miang Mts., 1400 m, Shimizu et al. Т 11662 (KYO,
TNS). BURMA. N’Changyang, 25°50’ N, 97°48’ E, 1600
m, Kanlback 273 (BM); North Shau States: Lashio, 900
m, J. H. Lace 5841 (K). MALAYSIA. Malacca, D. Yuan
s.n. (С). JAVA. Cultivated: Blume s.n. т poit 895
(G, P), 899 (G); Beru, Zollinger s.n. (G). U ii
Oahu, Kaneohe, Keaahala Rd., Nagata dis a
5. Kalimeris yomena Veen Kitam., Kiku 40.
1940. Kalimeris incisa var. yomena Kitam.,
Mem. Coll. Sci. Kyoto Univ., Ser. B, Biol.
13: 311. 1937. TYPE: Japan. Kyoto: Yoshida,
N. Kinashi s.n. in 11/1921 (holotype, KYO;
isotypes, TI, TNS).
Root fibrous; rhizomes stoloniferous, 5-38 cm
long. Stem green or rarely purplish, 50-180 cm tall.
Cauline leaves narrowly oblong, oblanceolate, ellip-
tical, or obovate, serrate, crenate, or deeply pin-
nate-lobed, 3.5-8.2(-8.6) х 1.1-3(3.4) cm, gla-
brous or sometimes with scattered, thick,
nonglandular hairs adaxially and evenly or loosely
so and with scattered glandular hairs abaxially,
lobes 0.6—1.5 X 0.3–0.6 cm. Capitula 10—105; pe-
duncles 1.3-12(-16) cm long; bracts 2-13 along
the peduncle, the lower ones narrowly oblong, 7.8—
19(-30) X 1-4(-8) mm, the upper ones (0.8—)1.8—
9.5(-7) X 0.4-1.5 mm; involucre hemispherical or
campanulate, 4.7-10 mm broad, 3.5-7.5 mm high.
Phyllaries in 3-4 whorls, imbricate, glabrous or
loosely covered with nonglandular hairs, sometimes
purplish; those of the outermost whorl lanceolate to
obovate, 2.1—4 X 0.7—1.4(-1.7) mm, those of the
middle whorl(s) oblong, obovate or spathulate, 2.7—
5.5 X 1-2.2 mm; those of the inner whorl oblan-
ceolate, obovate or spathulate, 3.3-5.5 Х 1-1.7
mm. Receptacles convex to subconical, 1.1-2.2
—2.5) mm wide, 0.6-1.3(-1.5) mm high. Ray florets
14—22; corollas pale lilac to pale purple, ligules 6—
Volume 84, Number 4
1997
Gu & Hoch
Systematics of Kalimeris
795
18.4 X (1.3-)2.1-4 mm, the tube 1-2 mm long;
style 2-3.4 mm long, the branches 0.6-1.2 mm
long. Disc florets (33-)51-100; corollas 3.24.7
(-5.1) mm long, the lobes unequal, the longer one
1-2.1 mm long, the shorter ones 0.8-1.5 mm long,
both 0.4—0.6(-0.7) mm wide, the tube 0.8-1.4
(71.7) mm long; anther 0.8-1.5 mm long; style 2.7—
4.4 mm long, the branches 0.7-1.3 mm long, each
with a triangular or rarely broadly lanceolate ap-
pendage 0.3-0.5 mm long. Ray and disc pappus
bristles whitish or brownish, subequal or unequal;
ray bristles (0—)0.2—0.5 mm long; disc bristles 0.2—
0.6(-1) mm long. Achenes straw-colored, brownish,
or dark purple, obovate or broadly obovate, covered
with 4-celled biseriate nonglandular hairs and
topped by glandular hairs, the margins 0.1—0.3 mm
broad; ray achenes 3(4)-ribbed, 2.1-3 X 0.8-
1.6(-1.9) mm; disc achenes 2-ribbed, 2.2-3.5 X
0.8-2.3 mm. Chromosome number: 2n — 40, 41,
43, 45, 61, 62, 63, 66, 68, 69, 70, 71, 72, and 73.
Figure 17.
The nomenclature of Kalimeris yomena is com-
plicated. Kalimeris incisa var. yomena Kitam. and
K. yomena Kitam. were initially described only in
Japanese (Kitamura, 1937a), and so were invalidly
published (Article 36.1, ICBN, Greuter et al.,
1994). In a second article published the same year,
Compositae Japonicae, Kitamura (1937b) validated
the name K. incisa var. yomena by including a Latin
diagnosis. In an obscure book, again only in Jap-
anese, Kitamura (1940) published the combination
K. yomena (Kitam.) Kitam. for the first time, al-
though it is sometimes attributed to a later publi-
cation (Kitamura, 1957), where he published it in
Latin.
This species is endemic to Japan and unique cy-
tologically. Pentaploid, heptaploid, and octoploid
plants have been detected, and aneupolyploids are
also common (Gu, 1989; Gu & Tara, 1990). It com-
prises three subspecies, as follows:
KEY TO THE SUBSPECIES OF KALIMERIS YOMENA
la. Pappus more than 0.4 mm long; branches flexi-
ble 5b. subsp. angustifolia
lb. ا less than 0.4 mm long; branches 5
. Plants no taller than 80 cm, leaves ay
oblong or oblanceolate and covered with
thick nonglandular hairs on both sides .....
pak come 5c. subsp. dentata
2b. Plants up to 160 cm tall; if less than 80 cm
tall, leaves oblanceolate or obovate, glabrous
or loosely hairy on both sides -................-.-
______- ба. subsp. yomena
5a. Kalimeris yomena subsp. yomena
Kalimeris incisa |. tubulosa Kitam., Mem. Coll. Sci. Kyoto
m
24: 53. 1957. TYPE: Japan. Shimane (Izumo): Като,
30 Oct. 1927, C. Ishikura s.n. (holotype, KYO).
an govt pinnatifida f. leucantha Nakai, Bull. Natl. Sci.
Tokyo 33: 26. 1953. TYPE: Japan. reg o
ОМ сы Ады, 16 Oct. 1949, Т. Nakai & N. Ма
ата s.n. (holotype, TNS).
Kalimeris indica var. grandiflora Nakai, Bull. Natl. Sci.
Mus. Tokyo 33: 27. 1953. TYPE: Japan. Yamaguchi:
Yamaguchi Shi, Housenji, 14 Oct. 1949, T. Nakai &
N. Maruyama s.n. (holotype, TNS).
Rhizomes stoloniferous, 10-38 cm long. Stem
55—120(—160) cm tall; branches usually stiff. Cau-
line leaves oblong, oblanceolate, elliptical, or ob-
ovate, serrate, crenate, or lanceolate-lobed,
glabrous, or loosely covered with nonglandular
hairs, 3.5-8 X 1.2-3 cm. Capitula 10-90; pedun-
cles 1.3-12(-16) cm long; bracts 2-12 along the
peduncle, the lower ones 8-17(-30) Х 1-4(-8)
mm, the upper ones (0.8-)1.8—5.4 X 0.4-1 mm;
involucre 5.5-10 mm broad, 4.2—6.5(-7.5) mm
high. Phyllaries in 3—4 whorls, imbricate, some-
times purplish; those of the outermost whorl 2.1—4
X 0.7-1.4(-1.7) mm, those of the middle whorl(s)
2.7-5.5 X 1-2.2 mm, those of the inner whorl 3.3—
5.5 X 1-2.2 mm. Receptacles 1.1-2.1(-2.5) mm
wide, 0.7-1.2(-1.5) mm high. Ray florets (14—)16—
27; corollas lilac or pale purple, ligules 9.4—18.4
х 2.24 mm, the tube 1-2 mm long; style 2-3.1
mm long, the branches 0.6-1.2 mm long. Disc flo-
rets (33—)51—100; corollas 3.2-4.4(-4.7) mm long,
the lobes unequal, the longer one 1.2-2 mm long,
the shorter ones 0.9—1.3 mm long, both 0.4—0.6 mm
wide, the tube 0.8-1.4(-1.6) mm long; anther 1—
1.5 mm long; style 2.8-4 mm long, the branches
0.7-1.3 mm long. Ray and disc pappus bristles
subequal, 0.3-0.4(-0.5) mm long. Achenes brown-
ish or dark purple, the margins 0.1—0.3 mm broad;
ray achenes 3(4)-ribbed, 2.1-3 X 0.9-1.6 mm;
disc achenes 2-ribbed, 2.3-3 X 1.4-1.9 mm. Chro-
mosome number: 2n — 61, 62, 63, 66, 70, 71, 72.
Figure 17A-C.
Phenology. Flowering period: June to late Octo-
r.
Distribution (Fig. 18). Endemic to Japan. This
has the widest geographic range of any subspecies
of Kalimeris yomena. It occurs from Ajigata-mura,
Niigata in northern Chubu District, and Saitama
and Chiba prefectures in Kanto District south and
west to Shikoku and southern Kagoshima of Kyu-
shu. It also occurs on Iki, Fukue, and Yaku Islands,
796 Annals of the
Missouri Botanical Garden
URE
aS
INS
Volume 84, Number 4
1997
би 4 Носћ 797
Systematics of Kalimeris
Figure 18. Geographical distribution of Kalimeris yomena subsp. yomena.
and has been recorded in the Ryukyu Islands (Ha-
tusima, 1975). This taxon grows from sea level to
500 m elevation, usually in relatively sunny and
mesic areas frequently disturbed by human activi-
ties.
Individuals in areas heavily disturbed by human
activities often resprout from cut stems or rhizomes,
usually resulting in plants with shorter branches
and bigger leaves. One such specimen from Hyogo,
M. Togashi s.n., 25 Nov. 1965 (A, TI), is unusual
in having an almost creeping stem, bigger leaves
and achenes than usual, and later flowering time.
Kalimeris yomena subsp. yomena is very com-
mon and occurs sympatrically with subspecies den-
tata in Kanto and northern Kinki Districts. Some
populations of these two subspecies grow so close
together that duplicate specimens contain both
taxa. For example, K. Okamoto 690 from Tokyo has
been distributed to several herbaria; those depos-
ited at MO and NY are subspecies yomena, where-
as those at A, BM, K, KYO, TI, and TNS are sub-
species dentata. Subspecies yomena and
angustifolia often grow sympatrically, as well, and
intermediates commonly occur in the sympatric
zones
Additional specimens examined. JAPAN. Miyagi: Wa-
tari Gun, “Yukiana,” Saito s.n. (KYO). Chiba: Funabashi
Shi, Komuro-cho, Ohba 652000 (K); Matsudo Shi, Oku-
yama s.n. (TNS). Saitama: Minamisaitama Gun, Shi
ka-cho, Ohyamamura, Ehara 48 (MAK); Urawashi Shi, Ta-
jimagahara, Murata & Togashi 4 (KYO). Tokyo To: Hino
Shi, Tamagawahara, Makino s.n. (MAK); Katsushika-ku,
Mizumoto to Koaidame, Ohba 61101 (TT); Meguro-ku, Ko-
maba, Muramatsu s.n. (ТЇ); Mt. Osawa, Koshigaya, Ohwi
1002 (K); “Shiwasuda-machi,” Okamoto 6
Terasaki s.n. (K); Suginami-ku, Kugayama, Yamatsuta
4299 (TNS); Tama Gun, Mt. Mitake, Furusawa s.n. (TI);
Tama Shi, Sekido, Okuyama s.n. (TNS). Kanagawa: Yo-
kohama Shi, Kamishirane, Takahashi s.n. (MAK), Maxi-
mowicz s.n. (BM, G, GH, NY), Milne s.n. (BM); Zushi Shi,
Mt. Jinmuji, Makino s.n. (MAK); Mt. Shinmujisan, Makino
s.n. (MAK), Okuyama 17476 (TNS). Niigata: Kashiwazaki
Shi, Abe s.n. (MAK); Minamikan Gun, Shigeru 200 (KYO);
Nishikanbara Gun, Ajigata-mura, Kurihara 99 (MAK);
Takata Shi, Minamishiro, Junkan s.n. (KYO); Tsubame
Shi, Tsubakuro, Eguchi s.n. (TI). Toyama: Nakaniikawa
Gun, Kamiichi-cho, Oiwagawa, Kurosaki 2607 (TNS). Na-
gano: Kiso Gun, Yamaguchimura, Oda s.n. (KYO); Mat-
sumoto Shi, Koidzumi 27236 (TNS), 65094 (TNS). Shi-
zuoka: Ihara Gun, Fujigawa-cho, Аппазе River 150-350
m, Tateishi et al. 4496 ; Kamo Gun, Kawazu-cho, Mi-
neyama, 200 m, Inoue & Im 2252 (TI); Ogasa Gun, Ogasa-
yama, Sato s.n. (MAK). Aichi: Toyohashi Shi, Torii 2392
(KYO); Toyokawa Shi, Mikami, Torii s.n. (TNS). Gifu:
Gifu Shi, Shiota s.n. (TNS); Toki Shi, Tsumaki, Shiota
Es
Figure 17. Kalimeris уотепа. A-C.
K. yomena subsp. yomena. —A. Shoot. —B. Root. —C. Асћепе. —D. Achene
of K. yomena subsp. angustifolia. —E. Achene of K. yomena subsp. dentata.
798
Annals of the
Missouri Botanical Garden
8134 (GH). Ishikawa: a, Monzen-cho, Kita-
mura s.n. (KYO). Fukui: Nan un, Kono mura, Jindo,
150-250 m, Kurosaki 1 101 7 аю тани Сип, Eih-
eiji-cho, Daibutsujiyama, Murata KYO). Shiga:
Kamo Gun, Uchiike,
чэч ea 350 m, Murata 196
MO, NY, TI, TNS); Uda Gun, Haibara-cho, 10069 (TD.
yama: Naga Gun, Kokawa-cho, Kasukami, Mt.
Ryumon-zan, Ogawa s.n. (KYO). Osaka: Ibaraki ie
Beattie & Kurihara 11205 (GH); n Katsuragi, Kitamu.
s.n. (KYO). Kyoto: Ayabe Shi, from Ohbatake to don,
00 m, Persius et al. 4008 feris чина sin
Aracki s.n. (KYO); Каза Gun, Оое-сћо, Naigu, М
20083 (KYO); Kyoto y dare ku, Kanaido, 10 m, "Ка
zumi Tsuchiya 2121 (KYO, TD; Minami- u^ Kameoka,
Hiroe 6441 (ТМ5); Sakyo-ku, Mt. Hiei, Makino s.n.
(MAK), Okamoto s.n. (TNS); Ohhara, H. Gu et al. 412
(MO); Tanaka, Higashitakahara, Kitamura s.n. (KYO);
Shinkotari, Yamamoto 384 (KYO); Ukyu-ku, A
Kitamura s.n. (KYO); Mukou Shi, Yamamo (TI,
TNS); Otokuni Gun, Shikoutari-mura, iban 384
(TNS), 386 (TI); Tsuzuki Gun, Tanabe-tyo, Murata 8355
(KYO). Hyogo: Aioi Shi, Futatsugi to Yado through Tsu-
bakitoge, 50-140 m, Fukuoka et al. 994 (KYO); Akashi
Shi, Ohkubo-cho, 20 m, Kurosaki 13994 (KYO); Akou
Gun, Kamigoori-cho, Yaho, Kaisaka, 150-250 m, Kurosaki
^ Kato 148 (KYO); Himeji Shi, Matogatamura, Togashi
n. (A, TI; Kakogawa Shi, Ranno-cho, Saijou, Kurosaki
9716 (KYO); Hanma, Makino s.n. (MAK); Kobe Shi, Han-
ayama, 200 m, H. Gu & Noguchi 425 (MO); Hutatabidani,
Okamoto 19678 (ТІ); Fukuji, Hachiman-jinya, 180 m, Fu-
kuoka 9887 (KYO); Naka, 50 m, H. Gu & Noguchi 424
(MO); Miki Shi, Masuda, 150 m, 421 (MO); Nakasato,
m, 422 (MO); Ronhayama, 200 m, 423 (MO); Tose, 418
(MO): Sanda Shi, Aimoto, Mt. Kokuzosan, 200-400 m
Кимий 6542 (КҮО); ан: Shi, Iho-cho, 1 1585
td
utan-cho, Nojima, 20-260 m, Kurosaki 13656 (KY 0).
Arinoto s n. (GH kayama: Atetu Gun, Tetta-cho, Ka-
wanose to "Takaino, Kurse 14560 (KYO); Okayama Shi,
Nishihara s.n. (TNS); Takahashi Shi, Deguchi, Yoshino s.n.
(KYO, TD. Tottori: Tottori Shi, Tanaka 13885 (KYO).
Hiroshima: Aki Gun, Kabe, Mizuochi, Shimotomai s.n.
(MO), 385 (MO); Aburagi, 250 m, 382 (МО); Eba, Suzuki
s.n. (MAK); Saeki Gun, Yoshiwa-mura, Ishihara, 500 та
H. Gu et al. 381 (MO). Shimane: Hinokawa Gun, Kuni-
tomi-mura, Hara s.n. (TNS), Hayashigi to Gakuenji Kish-
ino s.n. (KYO); Mt. Sanbei, Kitamura s.n. (KYO); Yatsuka
Gun, Shimane-mura, Takohana, Okuyama & Utsumi
11470 (TNS). Yamaguchi: Abu Gun, ут s Ikumo,
Oka 34250 (TNS), Chomonkyo, Yunose to Akagane, Mur-
ata et al. 126 (KYO), Ikumo
377 (MO); Iwakuni Shi, Shiniwakuni, Н
(KYO), 6497 (TNS); Ошан. Tokuyama, Okamoto
s.n. (KYO, TNS); Saba Gun, Tokuji-cho, Kamimura, bond
s.n. (TI), Wado-mura, Tsuuraketoge, Migo s.n.
monoseki Shi, Ayaragi, H. Gu et al. 371 (MO), 371a c
Tokoura Gun, Houhoku-cho, Kurino-toge, 373 (MO), Na-
gatoawano, 374 (MO); Tsuno Gun, Kano-cho, Okuyama
18011 (TNS); Yoshiki Gun, Ogoori-cho, Oka 3647 (KYO,
TNS). Kagawa: Kagawa Gun, Shiyoe-cho, Kaharashimo,
H. Gu & Takahashi 386 (MO); Mitoyo Gun, Toyohama-
cho, Shirai-toge, 406 (MO); Ohkawa Gun, Hiketa-cho, 200
m, Takahashi 1702 (KYO, TI); Takamatsu Shi, Taisanji,
H. Gu & Takahashi 402 (MO); Tosannishi, 410 (MO). To-
kushima: Kaifu Gun, Shishikui-cho, Takato 311 (KYO);
Myodo Gun, Nakai 2534 (TI). Kochi: Kami Gun, Noichi-
cho, Yamanaka 54349 (KYO); Ikku, Yanagida s.n. (KYO),
Makino s.n. (MAK); Takaoka Gun, Sakawa-cho, Makino
s.n. (MAK); Tosa Gun, Tosayama-mura, Okubo s.n. (KYO);
Kuwao, Yamanaka 26157 (KYO). Ehime: Higashiuwa
Gun, Uwa-cho, Iwaki, Nomura 3 (KYO); Kamiukena Gun,
Mikawa-mura, Nanatori m, H. Gu & Takahashi 398
(MO); Matsuyama Shi, a. Furusawa s.n. (ТЇ);
Sairinji, H. Gu & Takahashi 400 (MO); 2 km N of Matsu-
yama Harbor, 403 (MO), Makino 44 (MAK); Nishiuwa
Gun, Ikata-cho, Minatoura, Nomura s.n. (KYO); Oshu Shi,
Asayama s.n. (TNS); Shuso Gun, Takasu, Yogo 3557
(TNS); Tanbara-cho, Tanogamikada, 310 m, H. Gu & Tak-
ahashi 404 (MO). Fukuoka Province: Fukuoka Shi, Mt.
Aburayama, Momiyama s.n. (ТІ); p
; Kikuhantou, Moji-ku,
Hashimoto s.n. (TI); donas Yoshioka 12 (KYO); Kyoto
Gun, Karita-cho, Oazatsutsu m, H. Gu et al. 370
(MO); Tagawa Shi, Oouchi El поља 20385 (TNS); Yu-
kuhashi Shi, Kawashima, H. Gu et al. 368 (MO). Saga:
Hikosan, Kitamura s.n. (KYO); Iki Gun, Nakamura-shi,
Sumiyoshi, Hiroe 5120 (KYO); Nagasaki Shi, Mt. Inasa-
yama, Yamasaki s.n. (TI); Mt. Tara, Kitamura s.n. (KYO).
agasaki: Fukuejuma, Minamimatuura Gun, Koumu, Fu-
kuoka 11801 (KYO); Tamauoura-cho, Arakawa, Murata &
mura, Tashiro s.n. (TNS); Maece Gun, Tara
Funazu, Murata 8415 (KYO gs Ohmura Shi, Toyama
Kurogi, Hara s.n. (TI); Yatsushiro Shi,
Mayebara 5800 (KYO, TNS). Oita: Beppu Shi, кшн
Murata 12631 (KYO); Oita Shi, Haneya, Shimada 1
(KYO). Miyazaki: Higashiusuki Gun, Katogawa-cho, 4
kasawa s.n. (TNS); Hosojima, Kitamura s.n. (KYO); Hyuga
Shi, Nagasawa s.n. (TNS). Kagoshima: Hiki Gun, Fu-
kiage-cho, Otomez uki, H. Gu et al. 349 (MO); Kagoshima
Shi, Shiroyama, Murata ee or pane Gun,
Botsu-cho, Min vam Furus ro-cho, Mt
Isomayama, 300 m
425 (TD; ا Gun, Miike, 350 m, Koyama 7541
(KYO); Kumage Gun, Kamiyaku-cho, Кизикама, Oka
37697 (TNS). е Сип, Tougou-cho, Doi 40 (KYO);
Taromizu Shi, n he 00, Kamishinmidoo to Horikiri, 50-
500 m, Миг are (TD; Yakushima, from Miyanoura
to bolo са 3486 (КҮО).
Volume 84, Number 4
1997
799
Systematics of Kalimeris
Figure 19. Geographical distribution of Kalimeris yomena subsp. angustifolia (dots) and subsp. dentata (stars).
5b. Kalimeris yomena subsp. angustifolia (Na-
kai) H. Y. Gu, stat. et comb. nov. Kalimeris
pinnatifida var. angustifolia Nakai, Bull. Natl.
Sci. Mus. Tokyo 33: 26. 1953. TYPE: Japan.
Yamaguchi: Abu-gun, Jifuku-mura, Mt. Ook-
uraga-dake, 16 Oct. 1949, T. Nakai & N. Ma-
ruyama s.n. (lectotype, here designated, TNS,
2 sheets; isolectotype, TI).
Rhizomes stoloniferous, 5-30 cm long. Stem
green, 70-180 cm tall; branches usually flexible.
Cauline leaves narrowly oblong to oblanceolate,
serrate, crenate, or deeply pinnate-lobed, 3.5-8.2
X 1.1-2.8(-3.4) cm, glabrous or rarely with scat-
tered, thick, nonglandular hairs adaxially, loosely
covered with thick nonglandular hairs and glan-
dular hairs or rarely glabrous abaxially, lobes 0.6—
15 x 0.3-0.6 cm. Capitula 40-105; peduncles
1.9-9 cm long; bracts 2-13 along the peduncle, the
lower ones 7.8-17(-21) X 1.8-2.5 mm, the upper
ones 2.5—5.5 X 0.40.9 mm; involucre 4.7-8.5 mm
broad, 4.2-7.5 mm high. Phyllaries in 3-4 whorls,
loosely or tightly imbricate; those of the outermost
whorl 2.4-3.6 х 0.8-1.7 mm, those of the middle
whorl(s) oblong or obovate, 3.4-5 X 1-2 mm, those
of the inner whorl 4-5 X 1-2.1 mm. Receptacles
1.3-2.2 mm wide, 0.6-1.3 mm high. Ray florets
14—22; corollas pale lilac or lilac, ligules 11-17.5
X 2.1-3.5 mm, the tube 1-1.6 mm long; style 2—
3.4 mm long, the branches 0.6-1.2 mm long. Disc
florets 42-82; corollas 3.2—4.5(-5.1) mm long, the
lobes subequal, the longer one 1.2-1.8(-2.1) mm
long, the shorter ones 1–1.5 mm long, both 0.4—
0.6(-0.7) mm wide, the tube 0.8-1.4(-1.7) mm
long; anther 1.1-1.5 mm long; style 2.7—4.4 mm
long, the branches 0.9-1.3 mm long. Ray and disc
pappus bristles subequal or unequal; ray bristles
(0—)0.2—0.5(-0.8) mm long; disc bristles 0.6–0.8(–
1.5) mm long. Achenes brownish or purple; margin
0.2-0.3 mm broad; ray achenes 3(—4)-ribbed, 2.2—
3 X 0.8-1.4(-1.9) mm; disc achenes 2-ribbed, 2.8—
3.5 X 1.4-2.3 mm. Chromosome number: 2n = 66,
68, 69, 70, 71, 72, 73. Figure 17D.
Phenology. Flowering period: June to late Octo-
ber.
Distribution (Fig. 19). Endemic to Japan, from
southern Honshu (from Shiga and Hyogo prefec-
tures south and west; also on Oki Island), Shikoku,
and Kyushu, to Yaku Island. It occurs predomi-
nantly at low elevations, but is also found as high
as 1200 m. Like other taxa of the genus, K. yomena
subsp. angustifolia generally occurs in sunny, rel-
atively mesic places, often around human distur-
bance. It is sympatric with K. indica subsp. indica
in southern Kyushu and with K. yomena subsp.
yomena in most parts of its distributional range.
Kalimeris yomena subsp. angustifolia has long
been called К. incisa in taxonomic treatments by
Japanese botanists (e.g., Kitamura, 1937b; Ohwi,
1965; Ito & Soejima, 1995, as Aster incisus). The
confusion has been caused mostly by its longer pap-
Annals of the
Missouri Botanical Garden
pi, which also characterize K. incisa. These two taxa
differ, however, in branching pattern and rhizome
morphology, with K. yomena subsp. angustifolia hav-
ing flexible branches up to 40 cm long and stolon-
iferous rhizomes up to 10 cm long, whereas K. incisa
has relatively stiff branches less than 25 cm long
and condensed rhizomes no longer than 1 cm.
There are two morphological forms found in this
subspecies: one with relatively narrow leaves and
phyllaries, and the other with broader leaves and
phyllaries. These two forms are not allopatric, nor
do they exhibit any habitat preferences. There are
no other character states correlated with these two
forms. Intermediates between them are often found
in the herbarium as well as in the field. They are
here treated as a single taxon.
Additional specimens examined. JAPAN. Shiga: Gam-
ou Gun, Kitahizusa-mura, Uchiike, Hashimoto 13500
(TNS); Ohtsu Shi, Hishigakaro, 4714 (TNS). Wakayama:
Nishimuro Gun, Hikigawa-cho, Fukuda s.n. (TI); Inari-
mura, Nakajima s.n. (TI); Tanabu Shi, Nakajima s.n. (ТІ).
Osaka Fu: Kongo-san, Chihaya to Fushimi-toge, 500 m,
Murata 19038 (KYO), Tada s.n. (TI). Kyoto: Makino s.n.
(MAK). Hyogo: Aku Shi, Nishiune, Kamikumi, Sakaori-
ike, 60 m, Kurosaki & Kato 22 (KYO); Ibo Gun, Mitsu-
cha, Murotsu, 20-100 A Fukuoka зү (KYO); Kasai
Shi, Aonogahara-cho, ujo, Kurosaki 9537
(KYO); Kobe Shi, неч ы boron 19677 (TI); Tar-
umi-ku, Kitamura s.n. SEED Makino s.n. (MAK); Mikata
Gun, Hanasaka-cho, Kannon-yama, 50-150 m, Murata
46619 (KYO); Miki Shi, ме 0 Murata & Nishimura
270 rete Minou Gun, Y awa-cho, Hase, 85 m, H.
uchi 420 (MO); Sanda Shi. Mt. Kokuzosan, Aim-
oto, Piso 6542 (TI); Shisou Gun, Chikusa-cho, Utsumi
(
E KYO); to Shi, in Awaji-shima, Yura-cho, Na-
rugashima, Kurosaki et al. 2705 (KYO); Takenokuchi-
hachimanjinjya m, Fukuoka et al. О);
akasago Shi, Ooishi, Ishinohoden, 20-60 m, Kurosaki
12928 (KYO); Umei, at sea novels 140001 (KYO); Taki
Gun, Sasayama-cho, Araki s.n. ( desd Saimami s.n. in
9/1968 (KYO). Oka йай, "Aida , Awakura-mura,
Shiotani, Nanba 8160 (KYO, TNS); el Gun, Uno
94 (NY): Atetsu Gun, along Nisikawa, Hubara, Kamiiti-
kam
Fukuoka 11250 (KYO); Nohara, Nishi ara s.n. (TNS); N
imi Shi, Ikura, Hara & Kurosa n. qe Ашы s.n.
(KYO). Hiroshima: Asa Gun, gone -mura, Fukuugi
Terawake III, Shimotomai s.n. (KYO); Jinseki Cun, Yuki.
cho, Senyoo, Tashiro s.n. (TNS); Hiroshima Shi, Ushita-
mura, Suzuki s.n. (KYO); Mt. Micoto, Tashiro s.n. (KYO);
Takada Gun, Mukaihara, nz = suchi,
(KYO), Kuriyama 2009 (MA i
MAK); Nima Gun, Yunotsu-cho, Matsuyama,
Takaki 327 (KYO), Yunotsu, sea coast, Miyamoto 105
(MAK); Mt. Sanbusan, Maruyama 512 ); Shimazen-
shima, Koniga, 924 (KYO), 925 (KYO); Mt. Tainosu, Ki-
tamura s.n. (KYO). Yamaguchi: Abu Gun, Ato-cho, Cho-
monkyo, between Yunose and Akagane, Murata et al. 128
(KYO); Jifuku-mura, Mt. le Oka 3671 e
Hagi Shi, Kasayama, 22925 (KYO), Kiatsu s.n. (KY
Ohtsu Gun, Mt. Ichiigadake, Hiroe 5071 (KYO); tm
S:
Gun, Hiki-cho, Ohuchiyama, H. Gu et al. 375 (MO); Shi-
monoseki Shi, Umegatou, 372 (MO); Tokuyama Shi, Kyoto
nen Okamoto s.n. (KYO, TNS); Tsuno Gun, Kano-
o, Okeyama, 500 m, H. Gu et al. 379 OR Yamaguchi
Shi. Oka 3671 (ТІ). Kagawa: Mitoyo , Yamamoto-
EUM Nitta, H. Gu & Takahashi 407 (МО); er roto Gun,
YO). Kochi: Iwata cp Saga-cho, Iyoki,
Murata A (TD, 6692 (TD; Kami Gun, Noichi-cho, from
Shinmiya to Nishino, H. Gu & Takahashi 395 (MO); To-
sayamada-cho, Kageyama, 240 m, 393 (MO); Koch Shi,
Godaiyama, 100 m, 397 (MO); Ushioe-yama, Makino s.n.
(MAK); Mima Gun, Waki-cho, Agra sia 5 Си « Так-
ahashi 389 (МО); Nangoku js и у 2 (MO); Shu-
kumo Shi, Kozukushi, Murata 6864 (TI); Sucio Shi, To-
shima, Makino s.n. te Matsuyama Shi,
Sairinji, H. Gu & Takahashi 401 (M ya Okuhira s.n.
(MAK), Uma Gun, Doi-cho, Nakamura, H. Gu & Taka-
hashi 405 (MO). Fukuoka: Fukuoka Shi, Tashima, Ма-
kajima 15499 (TNS); Itoshima Gun, Shima-cho, Keya,
m, H. Gu et al. 370a (M cui Yid
Shi, Kad 367 ppd Eo 360 МО : Hi-
gashima tsuura Gun, o, Mt. Tensan, e a
(KYO, TD; Saga Gun, cuti Sugiyama, 7 (KYO). Na-
gasaki: Kitamatsuura Gun, ra-cho, Tashiro s.n.
(KYO); Ohmura Shi, Toyama & ош 18770a (TNS);
Shimoagata Gun, Spooner Sumo, а an
(KYO); Tsushima, Shimoa
2979 (TNS), Hara s.n. (KYO), Yabe s.n. Im. Oldham 399
(K umamoto: Amakusa Gun, А usakamijima,
Ohtsu, Yamasaki s.n. y нај bh d Ushizu
Shi, Makino s.n. (MAK); ae А о, Muta, Kita-
mura s.n. (KYO); Narukawa, Hara yj; Ishinomiya-
7 to Mt. Aso, 450 m u et al. 356 (MO);
Yatsushiro Gun, Ryuhoku-cho, y и од 355 (МО); Yat-
sushiro Shi, Shimada 12993 (RTO): Oita: Mt. Aso, Shen-
; Be eppu Shi, Inonoseto, Ko-
О), 1 9 (КУО); Њени mi Gun, Nakayamaga,
Tashiro s.n. (KYO); Mt. Kujyu, Tashiro s.n. (KYO); Jijo-
hara, Tashiro s.n. (KYO); Kijimabaru, Tashiro s.n. (KYO);
Mt. Kujyu, Makino s.n. (MAK); Kusu
amiamabu Gun, Tsurumi-cho, Тазћ-
iro s.n. (KYO); Naoili Gun, Kujiu-cho, Kuninomura, Ma-
kino s.n. (MAK); Oita Gun, Yuhuin-cho, Yuhuin, Yuhuin
hot spring, 700 m, H. Gu et al. 365 (MO); Oita Shi, Mt.
Takasaki-yama, ashes s.n. (KYO); Mt. Sobosan, Tashiro
s.n. (TNS). Miyasaki: Nishiusuki Gun, Takachiha-cho,
Tashiro s.n. (KYO, TNS); Kushima Shi, Makaya, 110 m,
Koyama 7550 (KYO); Mt. Sobosan, Gokase plain, Naga-
sawa s.n. (TNS). Kagoshima: Hiki Gun, Fukiage-cho, Is-
aku-toge, 300 m, H. Gu et al. 348 (MO); Ibushuki Gun,
Kiire-cho, Makino s.n. (MAK); Ibusuki Shi, Hatusima &
Sako 27950 (MAK); Kaseda Shi, Ayukawa, 70 m, H. Gu
et al. 353 (MO); Kimotsuki бип, ће Odomari to
Sotonoura, 110 m, Yamazaki e 425 (TD; Kushikino
hi, H. Gu et al. 354b (MO), 354 | "o, 354d е Naze
Shi, Ohshima, Kimura & Furusa n. (TI); Yakushima,
Kumaba Gun, Kamiyaku-cho, Oka 37697 то» кашыр
to Sugi, along Kusukawa, 200-600 m, Murata et al. 15419
(TI).
Volume 84, Number 4
1997
Gu & Hoch
Systematics of Kalimeris
5c. Kalimeris yomena subsp. dentata (Kitam.)
H. Y. Gu, stat. et comb. nov. Kalimeris pin-
natifida var. dentata Kitam., Mem. Coll. Sci.
Kyoto Imp. Univ., Ser. B, Biol. 13: 309. 1937.
Kalimeris pseudo-yomena Kitam., Mem. Coll.
Sci. Kyoto Univ., Ser. B. 24: 53. 1957, nom.
illeg. TYPE: Japan. Kanagawa: Jinmuji, 3 Nov.
1935, 5. Kitamura s.n. (holotype, КУО; iso-
type, KYO).
Rhizomes stoloniferous, 5-14 cm long. Stem 50—
80 cm tall; branches usually stiff. Cauline leaves
linearly oblong, oblong, or oblanceolate, serrate,
4.8—7(—8.6) X 1.1–1.8(–2) cm, covered with thick
nonglandular hairs on both sides. Capitula 10—64;
peduncles 1.8-7.5 cm long; bracts 2-8 along the
peduncle, the lower ones 8.6-19 X 1.5-2.8 mm,
the upper ones (1.3-)2.1-7 X 0.4-1.5 mm; invo-
lucre hemispherical, 5.5—8 mm broad, 3.5—5.5 mm
high. Phyllaries in 3 whorls, those of the outermost
whorl 2.2-3.5 X 0.7-1 mm, those of the middle
whorl 3.5-5 X 1.1-1.5(-2) mm, those of the inner
whorl 3.5-5 X 1-1.7 mm. Receptacles 1.1-1.9 mm
wide, 0.5—1.3 mm high. Ray florets 14-22; corollas
pale lilac or whitish, ligules 6-14.5 X (1.3-)1.5—
2.8 mm, the tube 1—1.7 mm long; style 2-2.6 mm
long, the branches 0.6-1.2 mm long. Disc florets
42—70; corollas 3—4.7 mm long, the lobes subequal,
the longer one 1-1.5 mm long, the shorter ones
0.8-1.3 mm long, both 0.4—0.6 mm wide, the tube
1-1.4(-1.7) mm long; anther 0.8-1.5 mm long;
style 2.4—3.2 mm long, the branches 0.8-1.2 mm
long. Ray and disc pappus bristles subequal, 0.2—
0.3 mm long. Achenes straw-colored or brownish,
the margins 0.1—0.2 mm broad; ray achenes 3(—4)-
ribbed, 2.1-2.5 x 0.8-1.1 mm; disc achenes 2(-
3)-ribbed, 2.2-3 x 0.8-1.3 mm. Chromosome num-
ber: 2n — 40, 41, 43, 45. Figure 17E.
Phenology. Flowering period: June to late Octo-
er.
Distribution (Fig. 18). Endemic to Japan, where
it occurs in two disjunct areas: Chiba, Tokyo, and
Kanagawa prefectures of southeastern Kanto Dis-
trict, and Yamanashi Prefecture of northeastern
Chubu District; and to the southwest in Shiga,
Nara, Osaka, and Wakayama prefectures of Kinki
District. It occurs from near sea level to 850 m
elevation and, like the other two subspecies, occurs
in similarly disturbed, open habitats. It overlaps in
range with both other subspecies, but is less com-
mon than either.
Kalimeris yomena subsp. dentata is variable both
morphologically and cytologically. Judging from the
morphology and chromosome numbers, this sub-
species might have originated as a hybrid between
K. pinnatifida (n — 9) and an octoploid K. yomena
subsp. angustifolia or subsp. yomena (n — 36). The
three subspecies differ mainly in branching pattern
and the morphology of achenes and pappi (Fig. 17).
Intermediate individuals are found in areas of sym-
patry.
Additional specimens examined. JAPAN. Chiba: Hi-
чир impen, Gun, Abiko-cho, Murata & Togashi 25
(MAK, TI), 36 (MAK, TI), 38 (KYO, MAK). Saitama:
Ohmiya Shi, Okuyama s.n. (TNS); Urawa Shi, Tajimaga-
hara, Murata & Togashi 11 (MAK, TI). Tokyo: Chuo-ku,
Hamarikyu, Okuyama 23799 (TNS); Hachiouji Shi, Mt.
Takaosan, 17580 (TNS); Hatogaya Shi, Shirai s.n. (TNS);
Kamiitaba Shi, Itabashi-ku, Makino s.n. (KYO); Мба
ku. Makino s.n. (MAK); Shibuya-ku, Makino ѕ.п. (MAK);
Shiwasuda-machi (in Musashi), Okamoto 690 (A, BM, K,
KYO, TI, TNS). M To: Kyoto Shi, Sakyo-ku, Tanaka,
Kitamura s.n. (КУО
т. = Murata & Nishimuar 293 (KYO, TNS). Yaman-
33871 (KYO); Minamitsuru Gun, Misaka Mts., Oishitoge,
m s.n. (ТІ). Shiga:
54 (BM, MAK, MO, TNS). N
En A Kuragaritoge, 350 m, Mal 19662 (TNS). Wa-
kayama: Nishimuro Gun, Shirahama-cho, Shirahamagu-
chi Station, Takahashi 215 (TNS). Osaka: Mt. Kongo-san,
Chihaya to Fushimi-toge, 500 m, icons 19036 (KYO).
6. Kalimeris shimadai (Kitam.) Kitam., Acta
Phytotax. Geobot. 6: 50. 1937. Asteromoea shi-
madai Kitam., Acta Phytotax. Geobot. 2: 37.
1933. Aster shimadai (Kitam.) Nemoto, Fl. Ja-
pan Suppl. 740. 1936. TYPE: China. Taiwan:
Shinchi-ku, Chikunan, Zokyo, 50 m, Dec.
1925, ¥. Shimada 2664 c (holotype, KYO).
Aster yangtziensis Migo, Bot. Mag. (Tokyo) 56: 300. 1942.
TYPE: China. Jiangsu: аза Zi-jin-shan, 23 July
1940, H. Migo 60 (holotype, KYO).
Mtem shimadai f. pinnatifida Kitam., J. Jap. Bot. 19:
. 1943. TYPE: China. Zhejiang: Putuo, 13 Oct.
em H. Migo 49 (lectotype, here designated, KYO).
Root fibrous; rhizomes condensed, 0.1-1 cm
long, usually forming rootstocks. Stem (31—)60—200
cm tall. Rosette and lower cauline leaves covered
with long and thin nonglandular hairs. Cauline
leaves oblong or oblanceolate, crenate, serrate, or
deeply pinnate-lobed, 3.5-7.5 X (0.6–)1–2.7 cm,
densely or evenly covered with thin nonglandular
hairs on both sides and often interspersed with
glandular hairs abaxially, lobes 0.4-1 X 0.2-0.4
cm. Capitula 20-85; peduncle 1.6–9.3 cm long;
bracts 1-7 along the peduncle, the lower ones 7—
18 X 1.3-3.3 mm, the upper ones 3.1-6.1 X 0.8–
1.3 mm; involucre campanulate or rarely hemi-
spherical, 6.5-10 mm broad, 4.6—7.5(-9) mm high.
Phyllaries in 3—4 whorls, imbricate, covered with
thin nonglandular hairs and sometimes interspersed
with glandular hairs; those of the outermost whorl
802
Annals of the
Missouri Botanical Garden
lanceolate or obovate, (2.2-)3—4.7 X 0.8-1.4(-1.8)
mm, those of the middle whorl(s) obovate or spa-
thulate, 3.5—6.2 X 1.2-2(-2.5) mm, those of the
inner whorl € or spathulate, (3.3—)4—6.2(-
7.2) X 1-2(-2.3) m eceptacles convex, 2-28
mm wide, 0.7-1.5 mm high. Ray florets (11-)14—
29; supe pee lilac or light purple, ligules 13.5—
21 X —4) mm, the tube 1.2-2.2 mm; style
2-3.3 mm sins the branches 0.8-1.4 mm long.
Disc florets 48-82; corollas 3.5-5.6 mm long, the
lobes unequal, the longer one 1.2-2.2 mm long, the
shorter ones 0.4-1.4 mm long, both 0.5-0.7(0.8)
mm wide, the tube 1-2 mm long; anther 1.2-2.2
mm long; style 2.8—4.8 mm long, the branches 1—
1.4 mm long, each with a triangular or broadly lan-
ceolate appendage 0.4—0.5 mm long. Ray and disc
pappus bristles equal or unequal; the ray bristles
0.3—0.6(-1) mm long; the disc bristles 0.4—0.6(-1)
mm long. Achenes light brown or purple, broadly
obovate, covered with 4-celled biseriate nonglan-
dular hairs and rarely with glandular hairs at the
top, the margins 0.2-0.4 mm broad; ray achenes
3(4)-ribbed, 2.5-3 X 1.2-1.5 mm; disc achenes
2-ribbed, 3-3.2 X 1.7-2 mm. Chromosome num-
ber: n = 54. Figure 12G-J.
Phenology. Flowering time: June to late Novem-
er.
Distribution (Fig. 20). This species is endemic to
eastern and central China. It occurs from Shanxi,
Shandong, Henan, Shaanxi, and Gansu provinces
in the north to Anhui, Jiangsu, Jiangxi, Fujian, and
Taiwan in the southeast, and to northwestern Hun-
an, Hubei, and Sichuan in central and southwestern
China. It is distributed from low areas at sea level
to elevations up to 2800 m. It is usually found on
open slopes, dry, stony areas, or sometimes in rel-
atively mesic habitats.
This taxon is often sympatric with Kalimeris in-
dica subspp. indica and stenolepis, and with K. in-
tegrifolia in Jiangsu and Hubei. Several popula-
tions of K. shimadai and K. indica subsp. indica
were found co-occurring in Yichang Xian, Hubei
and Nanjing Shi, Jiangsu, but no intermediates be-
tween these two taxa were detected from these ar-
eas. No intermediates between K. shimadai and K.
integrifolia were found in their areas of sympatry,
either. Kalimeris shimadai and K. indica subsp.
stenolepis are sympatric in Hubei, where they are
usually distributed at different elevations. The for-
mer occurs at low elevations, mestly below 1500 m
but rarely to 2800 m, whereas the latter is often
found at higher elevations, mainly from 500 to 3000
m. Kalimeris shimadai is also sympatric with K.
lautureana subsp. lautureana in the Loess Plateau,
"Xiaofulong-shan, 120 m
where the higher elevation phenotypes of these two
taxa can be difficult to distinguish. Both tend to
have deeply pinnate-lobed or serrated leaves at
high elevations, but differ mainly by the length and
density of branches, the indumentum, and the size
of the phyllaries. Kalimeris shimadai is character-
ized by its narrower phyllaries, and leaves densely
covered with thin nonglandular hairs.
This species is very common at the southern part
of its range. Plants from Taiwan have relatively
broad phyllaries and thin rhizomes with relatively
long internodes. Smaller individuals are more char-
acteristic of the northern part of the geographic
range. The length of the pappus bristles also varies
in this species; usually they are relatively long, with
united bases, but epappose individuals are occa-
sionally found (T. P. Wang 5062, PE, WUK; and
Tang Sui Ging 5706, BM, MO).
Additional specimens examined. CHINA. Shanxi:
Hongdong, Kanasiro 4838 (KYO); M (Chieh Hsiu),
Songlinmiao, 800 m, Smith 7677 (A, W); Xiachuan
Xian, Yaoshan, 1500 m, Licent 12506 o Xia Xian, Si-
jiao, Chepuyu, Houliang, 1 m, T. W. Liu 749 (PE),
1350 m, 751 (PE); Sijiao, Xigou-dadui, Baicaigou, 1250
m, 697 (PE); “Central Shensi,” Licent 24.
gbao
m, ү! 0. Ки 40 verge WUK); Lushi Xian, Tazi-shan,
Fuping Xian, Houzhenzi-xiang, Damonghe, 1075 m, X. M.
Zhang 332 (HIB); Hu Xian, Beimenwai, У У Pai 1282
(PE); Mian Xian, SAN К. Т. Fu 3590 (PE, WUK);
uicheng Xian , 5. Y Bao 600 (PE), 1500
(PE), 1586 (PE), 587 5 Shannan Xian, Caoying, Shu-
angjian, 1 m, B. Z. Gou 3996 (PE); Shanyang Xian,
Mantou-shan, 1500 m, T. P. So 16313 (PE, WUK); Six-
i Shangtian- oa 600 m, 3626 (PE, WUK);
modit 1200 m, 2957 (PE, WUK); Taibai- фый; Liu-
jiayan, W. У Hsia 4744 (PE), 4748 (PE); Songpingsi, 1200
m, T. N. Liu € P. C. Tsoong 75 (PE); Yingtokou, У Y. Pai
K) T. P. Wang 1436 (PE); Taibai Xian,
Guaili-xiang, 620 Team 3-357 (PE); Zhouzhi Xian, 650
m, H. У. Kung 3737 (PE). Gansu: Heshui Xian, Lianjia-
bian, У H. Sun & Z. S. Jin 252 (PE, WUK); Huicheng
Xian, Taihe-gongshe to Agr ага -zhuang, 1400 m, 2. Y
Zhang 662 (KUN, WUK); Lintan, 2800 m, 7. P Wang
5062 (PE, WUK); Min Xian, 2600 m, 5045 (PE, WUK);
Taoho Xian, Toumenkuan, "m m, 5975 (PE, WUK);
Zhang Xian, com ial Team s. . Anhui: ND Xian,
, g Xian, Nanjing Forestry College Team
7800- 746 (IBSC): Jiangpu ew Tangquan, Lao-shan,
B.. Deng et al. 3726 (KUN);
Jiangyin Xian, Fus-shan, 20 m, H. B. Zhou 3021 (IBSC);
Jingjiang Shi, Kolshaff 95 (W), Zhulinsi, Migo 42 (KYO);
Jurong Xian, Mao-shan, Jinniudong, 120 m, F. B. Deng et
al. 3519 (PE); Kunshan my Migo 45 5 (KYO); Nanjing
Shi, Zijin-shan, C. R. Hua n. (ТІ), Matsuda s.n. (TI),
С. C. j
Chang 4 (E), C. N. "Chen 8621 (W). C. Y. Chiao
Volume 84, Number 4 Gu & Hoch 803
1997 Systematics of Kalimeris
С
е М
540 kr
Figure 20. Geographical distribution of Kalimeris shimadai.
14941 (PE), K. S. Chow 80259 (BM, K, KYO, NY); Suzhou
Shi, Shangfang-shan, К. Ono s.n. (ТІ); Yixing Xian, Wufu,
K. Ling 2310 (GH); Zhengjiang Shi, Feng-shan, Huadong
N). Jiangxi: Jiujiang Shi,
Carles 438 (E), Migo 61 (KYO), Sheaver s.n. (GH, K); Lu-
(KUN), Z. X. Zhao 890 (KUN).
Tung-hsiao, Ping-ting, Tamura & Koyama 32106 (KYO, TI,
. Hunan: Xiangsi Zizhiqu, Longshan, Hongyan-cun,
600 m, L. H. Liu 9357 (PE). Hubei: Jiangling Xian, Jian-
gling, Xiuhu, 20 m, Q. H. Liu 1073 (HIB); Yichang Xian,
iaoxita, H. . Chen 183 (MO), 189 (MO), Henry
2739 (G), 2739a (K), Wilson 1727 (К, W); Zaoyang Xian,
Pinglin-gongshe, Guanji-dadui, D. Y Mu 297 (HIB). Si-
chuan: from Chongqing to Chengdu, T. N. Liu 9854
(WUK); from Hanyan to Yidong, X. J. Yang 3625 (PE);
Kangding Xian, Feiyueling, 2520 m, C. S. Liu 1584 (PE).
7. Kalimeris lautureana (Debeaux) Kitam., Acta
Phytotax. Geobot. 6: 22. 1937. Boltonia lau-
tureana Debeaux, Actes Soc. Linn. Bordeaux
31: 215. 1876. Aster lautureanus (Debeaux)
Franch., Mém. Soc. Sci. Nat. Cherbourg 24:
224. 1884. Asteromoea lautureana (Debeaux)
Hand.-Mazz., Acta Horti Gothob. 12: 224.
1938. TYPE: China. Shandong: Yantai, 4 July
1860, O. Debeaux 93 (lectotype, here desig-
nated, P).
804
Annals of the
Missouri Botanical Garden
Root fibrous; rhizomes condensed, usually form-
ing rootstocks. Stem (50—)78-140 cm tall, loosely
to densely covered with thin nonglandular hairs.
Rosette leaves covered with the same type of hairs
as on the stem. Cauline leaves oblong to broadly
oblong, entire or deeply lobed, 3.2-7.7 X 0.6-2.4
cm, glabrous or sometimes evenly covered with
nonglandular hairs or only so along the veins, lobes
0.4—0.9 X 0.1-0.3 cm. Capitula 10–90; peduncle
1.6-9(-13) mm long; bracts 2-10 along peduncle,
the lower ones 6.2-15 X 0.9—4 mm, the upper ones
2.1-6 X 0.5-1 mm; involucre campanulate, rarely
hemispherical, 6.5-10 mm broad, (4—4.8—7(-7.8)
mm high. Phyllaries in 3—4 whorls, tightly imbri-
cate, glabrous or covered with nonglandular hairs;
those of the outermost whorl lanceolate, oblong, or
obovate, (2.3—)2.6—4.5 X 0.8-1.9 mm, those of the
middle whorl(s) obovate or less often spathulate,
3.2—5.8(—6.2) х 1.3–2.3(–3) mm, those of the inner
whorl obovate to spathulate, 4.7—6.5(—7.6) X (1.4—)
1.8-2.5(-3.2) mm. Receptacle convex, 1.6–2.5 mm
wide, 0.5—1.8 mm high. Ray florets 13-22; corollas
whitish to lilac, ligules 13-28 X 2.5-4(4.3) mm,
the tube 1.4-2.6 mm long; style 2-4.5 mm long,
the branches 0.5-1.1(-1.5) mm long. Disc florets
30-95; corollas 4–5.8 mm long, the lobes unequal,
the longer one 1.5-2.2 mm long, the shorter ones
0.7-1.6(-1.8) mm long, both 0.5-0.7(-0.8) mm
wide, the tube 1-1.8 mm long; anther (1-)1.4-2.1
mm long; style 3-5 mm long, the branches 0.8-1.6
mm long, each with a triangular or broadly lanceo-
late appendage 0.8-1.6 mm long. Ray and disc
pappus bristles straw-colored or brownish, unequal;
ray bristles 0.2-0.5(0.8) mm long; disc bristles
0.4—0.6(-1) mm long. Achene greenish or light pur-
ple, obovate to broadly obovate, covered 4-celled
nonglandular hairs and rarely scattered with glan-
dular hairs at the top, the margins 0.2-0.4 mm
broad; ray achenes 3-ribbed, 2.2-3 х 1.2-2 mm;
disc achenes 2-ribbed, 2.4-3 X 1.6-2 mm. Chro-
mosome number: n = 54. Figure 21.
Kalimeris lautureana is distributed mostly in east
and central China from coastal areas to montane
regions. Two subspecies can be recognized, as fol-
lows:
KEY TO THE SUBSPECIES OF KALIMERIS LAUTUREANA
la. Leaves oblong, 0.6-2 cm wide; ray florets shorter
than 25 mm; phyllaries of the middle whorl(s) to
са. 2.5 mm wide; dise corollas 4–5.1 mm long
7a. subsp. lautureana
‚ Leaves broadly oblong, 1.5-2.4 ст wide; ray flo-
rets longer than 25 mm; phyllaries of the middle
whorls to ca. 3 mm wide; disc corollas 5-5.8 mm
on 7b. subsp. mangtaoensis
—
с"
Та. Kalimeris lautureana subsp. lautureana
Stem 60-140 cm tall. Cauline leaves oblong,
rarely oblanceolate, entire to deep-serrate or less
often lanceolate-lobed, 3.2—6(—6.9) x 0.6-2 cm,
lobes 0.4—0.9 x 0.2–0.3 cm. Capitula 20-90; pe-
duncle (1.6-)2.5-10(-13) cm long; bracts 3-10
along peduncle, the lower ones 6.2-15 X 0.9-2.8
mm wide, the upper ones 2.1—5.6 X 0.5-1 mm;
involucre campanulate or less often hemispherical,
6.5-9 mm broad, 4.6–7.5 mm high. Phyllaries in
3—4 whorls; those of the outermost whorl 2.3-4 x
0.8-1.7 mm, those of the middle whorl(s) 3.6-6 x
1.6-2.5 mm, those of the inner whorl 4.8—6.5 х
(1.4-)1.8-2.6 mm. Receptacles 1.6-2.5 mm wide,
0.5-1.2 mm high. Ray florets (13—)15—22; ligules
(13-)14-22 х 2.7-3.8 mm, the tube 1.4-2(-2.5)
mm long; style 2-3.2 mm long, the branches 0.5—
1.1 mm long. Disc florets (30-45-95; corollas 4—
5.1 mm long, the lobes unequal, the longer one
1.5-2.2 mm long, the shorter ones 0.9-1.8 mm
long, both 0.5-0.7 mm wide, the tube 1-1.7 mm
long; anther 1-1.8 mm long; the branches 1-1.5
mm long. Ray and disc pappus bristles unequal;
ray bristles 0.2-0.6(-1) mm long; disc bristles 0.4—
0.8(-1.5) mm long. Achene straw-colored or light
purple; ray achenes 2.2-3 X 1.2-2 mm; disc
achenes 2.4—3 X 1.6-2 mm. Chromosome number:
n — 54. Figure 21A-D.
Phenology. Flowering period: June to October.
Distribution (Fig. 22). From Yongling Xian,
Liaoning in northeast China, west and south
through Hebei, Beijing, Shanxi, Shandong, Henan,
Shaanxi, Ningxia, and Gansu, including the Loess
Plateau and Qinling Mountains in central China,
and to Lianyun-gang in northern Jiangsu Province.
This taxon grows from low elevations (above 100
m) in the plains regions of east China up to 2200
m at Kangle Xian, Gansu (W. Y. Hsia 8848, WUK).
At lower elevations it is often found along relatively
dry roadsides, whereas at higher elevations it is
usually found near irrigation ditches, streams, riv-
ers, or sometimes on open slopes. It was mistakenly
recorded from Korea (Nakai, 1911) due to the tax-
onomic confusion with K. mongolica.
In general, populations of K. lautureana subsp.
lautureana in the plains region near the Pacific
coast have almost glabrous and entire leaves,
whereas those at higher elevations in the Loess Pla-
teau always have leaves loosely covered with short
hairs and usually with serrate margins. But excep-
tions to this pattern are not rare. Some collections
from Qingdao and Mt. Tai-shan, Shandong (near the
Pacific coast), have crenate and loosely hairy leaves
very similar to those of plants from central China
Volume 84, Number 4 Gu & Hoch
1997
Systematics of Kalimeris
Figure 21. Kalimeris lautureana. A-D. K. lautureana subsp. lautureana. —A.
Wade. Leaf. —D. Achene. E-G. K. lautureana subsp. mangtaoensis. —E. Leaf. —F. Phyllary of middle whorl.
. Achene.
Habit. —B. Phyllary of middle
Annals of the
Missouri Botanical Garden
Figure 22. Geographical distribution of Kalimeris lau-
tureana subsp. lautureana (dots) and subsp. mangtaoensis
(circled with C).
(Y C. Wang 513 and 639, PE). There are also con-
tinuous intermediates between the two extreme
forms. No taxonomic rank is here conferred upon
these two forms.
Morphologically, Kalimeris lautureana is similar
to K. shimadai, especially to populations in central
China. The main differences are that K. lautureana
is glabrous or loosely hairy, and has shorter branch-
es and peduncles and broader phyllaries, whereas
K. shimadai is always densely covered with thin
nonglandular hairs interspersed with glandular
hairs, and has longer branches and peduncles and
narrower phyllaries. These two taxa are sympatric
in the southwestern range of K. lautureana, namely
on the Loess Plateau in central China. Although we
have not seen sympatric populations in the field,
herbarium specimens of the two taxa are often col-
lected from the same localities, e.g., Lushi Xian,
Henan, and Hu Xian, Shaanxi.
The leaves of this subspecies vary from entire
to deeply serrate. Yong Ling annotated some
specimens with serrate leaves as Kalimeris lau-
tureana f. branchyloba. Since this kind of varia-
tion can been found in almost every species in
the genus without any other correlated character
states, no taxonomic rank is assigned to these
individuals, either.
Additional specimens examined. CHINA. Liaoning:
Dalian Shi, Birnie s.n. (K), Kitamura s.n. (KYO), Stuart
s.n. (BM), Yabe s.n. (PE, KYO, TNS); Dandong Shi, Ji-
jiang-shan, Houshan, W. Wang et al. 1001 (ТЕР,
kino s.n. (MAK); D
(IFP); Fu Xian, 3 km SE of
otie-s
X. Li et al. 689. P), C. Wang & Y. X.
), Sato 24 (TT), 5839 (IFP, PE); Shahekou
Station, Kobayashi 7 (TI); Tieling, Yamatsuta 111 (TNS);
Xifeng Xian, Diaoyutai, Tiehe, X. C. Jiang et al. 228
(IFP); Longjiren, 2. Е Fang 109 (IFP), 249 (IFP); Xinbing
Xian, Yongling, Xiaochegou, X. Е Tian 59 (IFP); Xinji
TNS , 200 m, S.
Liu 982 (IFP, PE
a
WUK), Fuqingsi, 570 m, K. M. Liu 3839 (PE, WUK); from
1000 m, K. S. Hao 44 n.
eastern Shansi," Licent 11379 (W); “Shensi,” Hugh s.n.
(BM). Shandong: Kunyu-shan, forestry station, Shandong
University Team 3103 (PE); Laoshan Xian, Li-cun, 500 m,
Y. C. Wang 639 (PE); Qingdao Shi, Qingdao Bot. Garden,
H. Gu & Y. Gao 289 (MO), Limmercuamv 241 (G, GH),
277 (G, GH), 283 (LE), C. Y. Chiao 2978 (GH, IBSC, K,
NY, PE); Qixia Xian, Ya-shan, 400 m, Z. X. Wang 168
(PE); Tai-shan, Clemens 1512 (E), Shinagawa 46 (TNS);
Wuchimiao, Y. C. Wang 513 (PE); Yantai Shi, Nanyue, К.
M. Liu 1728 (PE), 100 m, 1828 (PE); Taipingyan, 250 m,
1841 (PE); Qufu-shan, Faber 163 (GH), Cowdry 912 (K),
Forbes 1872 (K), Wawra 1214 (W); Yantai Shi, Debeaux
80 (P); Zhifu-dao, Ito s.n. (TNS), Williamson s.n. (E). Hen-
an: Lushi Xian, Caomu-chang, 1060 m, J. Fu 2275
(HIB, KUN, PE, WUK); Shuanghuaishu, 540 m, 919
(HIB, WUK); Mianchi Xian, Shaoshang Forestry Station,
m, gaoyuan Team 1376 (WUK); Mianshan,
Qihe River, 1040, J. Q. Fu 1201 (WUK), 1206 (WUK);
Song-shan, 570 m, K. M. Liu 5532 (PE), 5575 (PE); Xixia
Xian, Huilongsi, 1000 m, Henan Team 1563 (PE)
Shaanxi: Hu Xian, Laogu, Yinjiapo, 1700 m, K. T. Fu
14841 (WUK); Huanglong-shan, Shibao, 3131 (PE,
WUK); Lantian Xian, Qinggang-xiang, Xiaogou, 1270 m,
K); Long Xian, P. Ке 234 (WUK); Luey-
ang Xian, Baishuijiang, Maliutang, 1300 m, T. P. Wang
18561 (KUN, WUK); Qinling Mts., Baoji, Fenzel 136 (PE,
WUK); Pingli Xian, Shiyan, Nuping, Qinlizi, 1000 m,
Third Yiejin Team 384 (KUN); Taibai Xian, Guaili-xiang,
620 Team 2-357 (PE); Taochuan, Lingdanmiao, 1250 m,
K. T. Fu 9742 (WUK); Taibai-shan, Maomenshang, S. H.
Luo 140 (KUN); Yichuan Xian, Tielumiao, 1050 m
K). Ni
800 m, Y. Q. He 686 (PE, WUK);
Ziwuling, Lianjiabian, 2. W. Zhang 2855 (WUK). Jiang-
su: Lianyungang Shi, Yuntai-shan, F. X. Liu 10890
(IBSC), Kanashiro 5615 (KYO).
Volume 84, Number 4
1997
Gu & Hoch
Systematics of Kalimeris
7b. Kalimeris lautureana subsp. mangtaoensis
(Kitag.) H. Y. Gu, stat. et comb. nov. Aster
mangtaoensis Kitag., J. Jap. Bot. 9: 109. 1933.
Kalimeris mangtaoensis (Kitag.) Kitam., Acta
Phytotax. Geobot. 6: 21. 1937. Aster lautu-
reanus var. mangtaoensis (Kitag.) Kitag., J.
Jap. Bot. 13: 554. 1937. Kalimeris lautureana
var. mangtaoensis (Kitag.) Kitag., Lin. fl. man-
shur. 653. 1939. TYPE: China. Liaoning: Xia-
olongshan-dao Island, 10 Sep. 1931, M. Ki-
tagawa s.n. (holotype, TI; isotypes, IFP, PE,
TI).
Stem about 110 ст tall. Cauline leaves broadly
oblong, entire to slightly serrate, 5—7.7 X 1.5-2.4
ст, glabrous. Capitula about 20; peduncles 3.8-6
cm long; bracts 2—4 along the peduncle, the lower
ones 7.3-10 X 3—4 mm, the upper ones 4.7–6 X
0.5—1 mm; involucre campanulate, 7-10 mm broad,
6.3-7.8 mm high. Phyllaries in 3—4 whorls, imbri-
cate; those of the outermost whorl 34.3 X 1.4-1.9
mm, those of the middle whorl(s) 4.5-6.2 X 2-3
mm, those of the inner whorl 5-7.6 X 2-2.7 mm.
Receptacles 2.2-2.5 mm wide, 1.5-1.8 mm high.
Ray florets 14—19; ligules 25-28 X 3.5-4.3 mm,
the tube 2-2.5 mm long; style 3-4.5 mm long, the
branches 1.1-1.5 mm long. Disc florets about 56;
corollas 5-5.8 mm long, the lobes unequal, the lon-
ger one 1.5-2 mm long, the shorter ones 0.7-1.3
mm long, both 0.5-0.8 mm wide, the tube 1.5-1.8
mm long; anther 1.5-2.1 mm long; style 4-5 mm
long, the branches 1.3-1.6 mm long. Ray and disc
pappus bristles brownish, unequal; ray bristles 0.3—
0.5 mm long; the disc bristles 0.5-0.6(-1) mm long.
Achenes were not available from the herbarium
specimens. Chromosome number unknown. Figure
Phenology. Flowering period: July to October.
Distribution (Fig. 22). Endemic to Xiaolongshan-
dao (Snake Island) in Bohai, Liaoning Province,
China.
This subspecies occurs only on Xiaolongshan-
dao, an island east of the Liaodong Peninsula with
an area of about one km?. The island was part of
the continent and separated from the latter as a
result of the break of Liaohe and the deepening of
the Bohai Sea under the pressure of the upthrust
movement of the Himalaya Mountains (Ji, 1982).
The island is a natural conservation site now be-
cause of its endemic Pallas pit vipers, which are
reputed to have high medicinal value. The number
of vipers on the island is estimated as high as
50,000 (Ji, 1982), which makes the island unin-
habitable and traveling very dangerous. The island
has been visited only by well-equipped viper catch-
ers and researchers. The available specimens of
subspecies mangtaoensis were collected by J. Sato
in 1933 and 1937.
The main differences between the two subspe-
cies of Kalimeris lautureana are that subspecies
mangtaoensis is less branched, with more and big-
ger leaves, bigger involucres and phyllaries, and
longer pappus bristles than subspecies lautureana
(Fig. 22). The differentiation between these two
taxa may be the result of genetic isolation between
them. Although there are only 11 km separating the
small island from the closest part of the mainland,
ene flow between the two taxa is probably very
limited, since Kalimeris is dispersed mostly through
human activities. The gigantism of this taxon might
be caused by high polyploidy.
Additional specimens examined. INA. Liaoning:
Dalian Shi, Xiaolongshan- о Sato sin (IFP, PE).
. Kalimeris integrifolia Turcz. ex DC., Prodr.
5: 259. 1836 [“Calimeris”]. Aster integrifolius
(Turez. ex DC.) Franch., Mém. Soc. Sci. Nat.
Cherbourg 24: 224. 1884. Asteromoea integri-
folia (Turez. ex DC.) Loes., Beih. Bot. Cen-
tralbl. 37: 189. 1919. Boltonia integrifolia
(Turcz.) Lauener, Notes Roy. Bot. Gard. Ed-
inburgh 34: 334. 1976. TYPE: Russia. Amur:
1833, Turczaninow s.n. (lectotype, here desig-
nated, LE; isolectotypes, K, LE).
Asteromoea pekinensis Hance, Ann. Sci. Nat., Bot. sér. 4,
15: 225. 1861. Boltonia pekinensis (Hance) Hance,
J. Bot. 370. 1867. Aster pekinensis (Hance) F. H
Chen, Bull. Fan Mem. Inst. Biol. 5: 41. 1934. TYPE:
China. Near Beijing, Oct. 1860, R. Swinhoe s.n. (lec-
totype, here designated, BM; isolectotype, K).
Tap root; perennating by adventitious buds on
the upper part of main roots or on the lateral roots.
Stem (10-)40-140 cm tall. Cauline leaves narrowly
oblong to oblanceolate, entire, (1-)2—5(-6.5) X
(0.2-)0.3-0.9 cm, densely covered with appressed
uniseriate nonglandular hairs and biseriate glan-
dular hairs on both sides. Capitula usually 25-200
or as few as 5 when growing in harsh habitats; pe-
duncles 2.5-9 cm long; bracts 2-9 along the pe-
duncle, the lower ones 6.1-16 X x 1-2. 2 mm, »
upper ones 2.5—5.2 X 0.4-0.8 mm; involucre he
spherical or rarely A dbi eR (3. 84. 4-89)
mm broad, 34.6 mm high. Phyllaries in 3-4
whorls, imbricate, densely covered with nonglan-
dular and glandular hairs, sometimes purplish;
those of the outermost whorl lanceolate or oblong,
1.8-3.6 X 0.5-1 mm, those of the middle whorl(s)
lanceolate to oblanceolate, (2.5—)3—4.2(—5) х 0.8-
1.1 mm, those of the inner whorl (3-)3.5—4.5(—5.1)
808
Annals of the
Missouri Botanical Garden
X (0.6-)0.8-1.1 mm. Receptacles convex to sub-
conical, 1.2-2.3 mm wide, 0.5-1.8 mm high. Ray
florets 19-27(-29); corollas pale lilac to light pur-
ple, ligules narrowly oblanceolate or oblong to el-
liptical, (5.2-)6.2-13.5(-15) X (1.1-)1.5-2.5(-2.7)
mm, the tube 0.7-1.4 mm long; style 1.5-2.5 mm
long, the branches 0.5-1 mm long. Disc florets 42—
100; corollas 2.1-3.1(-3.4) mm long, the lobes un-
equal, the longer one (0.7-)1-1.3(-1.5) mm long,
the shorter ones (0.5—)0.7—1 mm long, both 0.4—0.6
mm wide, the tube 0.6-1(-1.2) mm long; anther
0.7-1.2 mm long; style 2-3.3 mm long, the branch-
es 0.6-0.9 mm long, each with a triangular ap-
pendage 0.2—0.4 mm long. Ray and disc yer
bristles equal or subequal; ray voe (0—)0.1—0
mm long; disc bristles 0.2—0.4(-0.6) mm a
Achenes greenish or brown, heslo obovate, cov-
ered with biseriate glandular hairs at the upper
half; margin glabrous or ciliate, 0.1-0.3 mm broad;
ray achenes 3(-4)-ribbed, 1.5-2 х 0.9-1.1 mm;
disc achenes 2(-3)-ribbed, 1.5-2.2 x 1.1-1.6 mm.
Chromosome number: n — 9, 18. Figure 23.
Phenology. Flowering period: May to October.
Distribution (Fig. 24). Kalimeris integrifolia has
a very wide geographical range. It grows in the
north along the Amur River, Amur Oblast of Far
East region, Russia, extending through eastern and
central China to Suzhou, Jiangsu Province, in the
southeast and to near Ninglang Yizu Zizhi Xian,
Yunnan, in the southwest. It occurs in the following
provinces of China: Heilongjiang, Jilin, Liaoning,
northern Nei Mongol, Hebei, Beijing, Shandong,
Henan, Shaanxi, Jiangsu, Hubei, Sichuan, and
Yunnan. It also occurs in Korea from the border
with China, through Hambook, Hamnam, Pyong-
book, and Pyongnam provinces to Mt. Jangsoo,
Неа. It grows from near sea level to 1600 т
elevation. This species commonly grows in sunny
and dry roadside and river banks, and is most often
found at margins of thickets or forests and less of-
ten along trails inside forests. Its populations are
usually large due to vigorous formation of adven-
titious buds from the main or lateral roots during
the flowering season, except for plants growing on
poor soil along dry, open roadsides or river banks.
This species is easily distinguished from the oth-
ers in the genus by its tap-root system and ap-
pressed, nonglandular hairs densely covering both
sides of the leaves (Fig. 23). Although it grows sym-
patrically with K. incisa subsp. incisa, K. mongo-
lica, K. indica subspp. indica and stenolepis, and
K. shimadai throughout much of its range, no in-
termediates have been detected so far.
Additional specimens examined. RUSSIA. Amur: Karo
1506 (P), 2767 би, МАБ). KOREA. Hambook: Hoeryong,
(KY
mi s. O). H am: Hamhung, Okuyama s.n.
(TNS). Pyongbook: Ui quot s.n. (K ongnam:
Namp'o (Chinnampo), Faurie 1073 (BM, E, KYO, W);
"ases City, H. Chung s.n. (TNS), Faurie 392 (KYO, P).
: t. Jan gsoo, Koidzumi s.n. (KYO). CHINA. Hei-
Menon Baoqing Xian, Jiefangrun, Y. L. Zhang et al. 1812
(IFP); Daxing’ ME Mts., Skvortzov s.n. (GH); Harbin Shi,
rg ар, Laoshantou, G. Z. Wang & W Wang 725 (IFP),
G. Z. Wang 834 (IFP); Hosbin я дави College, 6 (IFP),
C. S. Wang et i 33 (IFP); Songhuajiang = Litvinov 31
(NY); Taiyangdao to Miaozitaihe, G. Z. et al. 1015
(IFP), ent s Main 3698 (NY), Kitagawa а 394 ПЕР), Jett-
mar s.n. an Xian, Fuli-tun, YL Zhang et al. 1833
(IFP); iani Shi. Р Y. Fu et al. 56 (IFP); Кезһап Xian,
heh Y L. Zhang & FP, KUN); Yichin
Xian, Shuangzihe, Xingan Forestry E N. Liuet a
7802 us Yuanhiu Xian, Xigangzi, G. Z. Wang 177 (ЕР).
Jilin: Changchun Shi, Jinyuetan, W. Wang et al. 2647 (IFP);
Синана Suzuki 13 (KYO), Nakao s.n. (KYO), Кота:
s.n. (KYO); Chunhua Xian, Hadamen, C. S. Wang et al. 2424
(8 KUN), 3634 (IFP); Jiutai Xian, Tumenling, Shichang-
an, 300 FP); Tao’an Xian, Ya-
akahashi s.n. (IN . Wang 1441
(IFP); Zhenglai Xian, —— Takano 47 (TNS), Е H. Chen
579 (GH), James s.n. (К). Liaoning: Anshan Shi, Anshan
, Н. H. Zhang 272 (MO), 273 (MO), 274 me
баайан, 278 (MO); Benxi Xian, Pingding-shan, С.
Wang et al. 300 (IFP); Dalian Shi, Laohutan, Kobayashi »
KYO). Changhai, Dachangdao, G. L. Wang 630160 (IFP);
Fushun Shi, Dong-gongyuan, W. Wang et al. 548 (IFP); Fu
Xian, Luda, Delisi, C. Q. pr 693 (IFP); Fuxin Shi, Shao-
prot ia M. VE Huenhe Xian, Huenhe pec
Station, Sato 953
83 (IFP); Balicun 4150 (IFP);
Linyuan Xian, ba: Карс pesa Ser l
shun Shi, Guantou, Kitamur
12068
oyu uri Station, S. X. Li & J. L i 5471 pia
Xinshi Xian, Xinshi Railway Station, Z. E Fang
Wang 2843 (IFP); NE Xian, Shuiyuan, Yingkou E.
294 (ТЕР); Zhangwu X ggu
n. (E), Clemens 2002 (E), 4152
1 Шест 1685 (ВМ), 7761 (ВМ). Shandong: Fei Xian, 300
Y Cheo & 1. Yen 230 (BM, E, GH, P, W); Laoshan
Gu 4 Hoch 809
Volume 84, Number 4
1997 Systematics of Kalimeris
WZ
CA SAR,
CUS
Ман a
RRA
igure 23. Kalimeris integrifolia. —A. Upper branch. —B. Tap root. — С. Front view of achene. —D. Side view
r ion of achene. —F. Portion of stem and leaf. —G. Portion of stem showing indumentum.
of achene. —E. Uppe
—H. Portion of leaf showing indumentum.
810
Annals of the
Missouri Botanical Garden
Figure 24. Geographical distribution of Kalimeris in-
tegrifolia.
Xian, Licun, H. Gu 301 (MO); Mongying Xian, Hujiagou
Yamoto s.n. (BM); Tai-shan, Chaoyangdong, ni-
versity Team 19 (KUN, PE); Weihai Shi, Elipse 78 (K); Yantai
Shi, Debeaux 63 (P), 79 (P), 92 (P), Debeaux s.n. (E), Forbes
s.n. (E, BM), Wawra 1269 . Henan: Lushi Xian, Wuli-
chuan, 480—490 m, J. Q. Fu 492 (HIB, KUN, PE, WUK).
Shaanxi: Fuping Xian, Houzhizi-xian
Merrill 1507 (MO). Jiangsu: Jinjiang Xian, Carles 485 (K)
ji C. N. Chen 2058 (W); Xiaolinwei
n. (ТЇ); Zijin-shan, С. R. Huang s.n. (ТІ), T. N. Liu
493 (NY), Т: K. Nyi 3 SBI), C. N. Chen 8620 (NY, W), W.
C. Cheng 181 (P), Niederlein 6 (A); Suzhou Shi, Oka 452
(TD; Xuzhou Shi, Hori 16 (KYO); Yizheng Xian, Nong-ganx-
iao, 5. Jiang 8973 (KUN); Zhengjiang Shi, Migo 34 (KYO),
Oka 22 (TI). Hubei: Guanghua Xian, Yuhuashan Forestry
Station, G. H. Chen 193 (HIB); Wuc Shi, Mo-shan, H.
Gu & D. Chen 132 (MO), Q. H. Liu 1027 (HIB); Yichang
Xian, Henry 564 (K)
G
.
>
,
Literature Cited
Anderson, L. C. 1987. Boltonia apalachi (
a new species from Florida. Syst. Bot. 12: 133—138.
1. “ЛА
Т . Compositae. /п С. Bentham & J. D. Hook-
er, Genera Plantarum 2: 163-533. L. Reeve, Williams
kerij, Batavia.
Candolle, A. P. de. 1836. Compositae: Astereae. Prodro-
mus 5: 211-497. Treuttel € Wiirtz, Paris.
Chen, Y. L. 1986. Systematic notes on the genus Miya-
mayomena Kitam. (Compositae). Bull. Bot. Res., Harbin
6: 3746. [In Chinese.]
Cronquist, A. 1955. Phylogeny and taxonomy of the Com-
positae. Amer. Midl. Naturalist 53: 478-511.
. 1980. Vascular Flora of the Southeastern US. 1.
Asteraceae. Univ. North Carolina Press, Chapel Hill.
Davis, P. H. € V. H. Heywood. 1963. Principles of An-
giosperm Taxonomy. Van Nostrand, Princeton.
Debeaux, M. O. 1877. Florule du Tche-fou. Act. Linn.
Soc. Bord. 31: 205-239,
Gagnepain, F. 1924. Composées. /n H. Lecomte, H. Hum-
bert & F. Gagnepain, Flore générale de l'Indo-Chine 3,
5: 4 Masson, Paris.
Grau, J. 1977. Astereae—Systematic review. In V. H.
Heywood, J. B. Harborne & B. L. Turner (editors), The
Biology and Chemistry of the Compositae. I: 537-566.
Academic Press, New York.
Greuter, W., F. R. Barrie, H. M. Burdet, W. С. Chaloner,
V. Demoulin, D. L. Hawksworth, P. M. Jorgensen, D. H.
Nicolson, P. C. Silva, P. Trehane & J. McNeill. У
International Code of Botanical Nomenclature (Tokyo
Code). Regnum Veg. 131: 1-389.
Grierson, J. J. C. 1964. A revision of the asters of the
Himalayan area. Notes Roy. Bot. Gard. Edinburgh 26:
7-163
Gu, H. 1987. A biosystematic study of the genus Kali-
meris. Unpublished Ph.D. Dissertation, Washington
University, St. Louis, Missouri.
- 1989. On chromosome numbers of Kalimeris (As-
tereae, Asteraceae) and some related taxa. Cathaya 1: 1-
16.
$ M. Tara. 1990. On meiotic chromosomal dis-
junction pattern in some taxa of Kalimeris (Astereae,
Asteraceae). Cathaya 2: 11-20.
; Le , L. J. Qu, L. X. Wen & Z. L. Chen.
1994. Preliminary studies on the phylogeny of Kalimeris
mena and two other taxa using RFLP
analysis. Cathaya 6: 27-34.
Hance, H. F. 1873. Florae Hongkongensis supplementum.
A compendious supplement to Mr. Bentham's descrip-
tion of the plants of the Island of Hongkong. J. Linn.
Soc., Bot. 13: 95-144.
Handel-Mazzetti, H. 1937-1938. Plantae Sinenses a Dre.
H. Smith annis 1921-22 et 1924 lectae. 32: Compositae
2. Acta Horti Gothob. 12: 203-360.
Hatusima, S. 1975. Flora of the Ryukyus. Biological Ed-
ucation Society of Ryukyu, Ryukyu. [Second edition; in
se.]
apanese.
Hoffmann, O. 1890. Compositae. In A. Engler & К.
Prantl, Die natiirlichen Pflanzenfamilien 4(5): 161-163.
Wilhelm Engelmann, Leipzig.
Hu, S. Y. 1967. The Compositae of China.V. Quart. J.
Taiwan Mus. 20: 1-77.
Ito, M. & A. Soejima. 1995. Aster. In K. Iwatsuki, T. Ya-
masaki, D. E. Boufford & H. Ohba (editors), Flora of
Japan IIIb: 59-66. Kodansha, Tokyo.
3 & T. Nishino. 1994. Phylogeny and spe-
ciation of Asian Aster. Korean J. Pl. Taxon. 24: 133-143.
: , M. Hasebe & K. Watanabe. 1995. A
chloroplast rDNA phylogeny of Kalimeris and Aster, with
reference to the generic circumscription. J. Pl. Res.
3-96.
Ji, D. M. 1982. She-dao (Snake Island, in Chinese). Liaon-
ing Science and Technology Press, Liaoning.
Jones, A. G. 1980. A classification of the New World spe-
cies of Aster. Brittonia 32: 230-261.
Volume 84, Number 4
1997
Gu & Hoch
Systematics of Kalimeris
R. A. Smogor. 1983. Chromosome counts of and
notes on some Old World asters. Phytologia 53: 429-431.
D. A. Young. 1983. Generic concepts of Aster
(Asteraceae): A comparison of cladistic, phenetic, and
cytological approaches. Syst. Bot
R. M. & H. W. Dawson (обоо: 1975. Cassini on
ences Naturelles. 1816-1830. Vols. 1–60.] Oriole Edi-
tions, New York.
Kitagawa, M. 1936. Vegetation around Wui-hu Station.
Rep. Inst. Sci. Res. Manchoukou 1: 299
Kitamura, 5. 1937a. On yomena. Acta Phytotax. Geobot.
53. [In Japanese.]
Coll. Sci. Kyoto Imp. Univ., Ser. B, Biol. 8: 299-399.
940. Kiku. Kobundo-shobo, Tokyo. [In Japanese.]
—. 1957. Compositae Japonicae. Pars Sexta. Mem.
Coll. Sci. Kyoto Imp. Univ. „ Ser. P Biol. 24: 1-79.
. Taxonomic studies in the Composite
of Thailand 2. Acta Phytotax. Geobot. 34: 1-9.
Lauener, L. A. 1976. Catalogue of the names published
by Hector Léveillé: IX ee Notes Roy. Bot.
Gard. Edinburgh 34: 327—402.
Lawrence, G. H. 1951. Taxonomy of Vascular Plants. Mac-
millan, New York.
LHeéritier, C. L. & D. de Brutelle. 1788. Р. 27 in Sertum
Anglicum. Rud. Graeffer, London.
Linne, C. von, filius. 1782. P. 373 in Supplementum Plan-
tarum. Impensis Orphanotrophei, Braunschweig.
Ling, Y. Y. L. Chen & Z. Shi. 1985. Astereae. In Fl.
Reipubl. desir Sin. 74: 73-353. Science Press,
Beijing. [In Chin
Wid 856. Flora van Nederlansch Indie. 2:
Nakai, T. 1911. Flora Koreana. Pars Secunda. Mem. Coll.
Sci. Kyoto Imp. Univ. 31: 1-573.
. 1919. Notulae ad plantas 2 аро еі Когеае
ХХІ. Bot. Mag. (Tokyo) 33: 193-216
Genera et Species Азий: Leonardi
ürnberg.
Nishino, T. & T. Morita. 1994. 6-Phosphogluconate de-
hydrogenase (6PGD) gene кенв" in Kalimeris (As-
teraceae). Pl. Sp. Biol. 9: 91-9
Ohwi, J. 1965. ie J. G. Licet E. H. Walker (editors),
Flora of Japan. English edition. Smithsonian ж
.& T E. Raven. 1976. The genus Epilobium
in Aussi New Zealand Dep. Sci. Industr. Res.
Bull. 216: 1–321.
Semple, J. C. & L. Brouillet. 1980. A synopsis of North
American asters: The subgenera, sections, and subsections
of Aster and Lasallea. Amer. J. Bot. 67: 1010-1026.
Shinners, L. H. 1946. The genus Dichaetophora A. Gray
and its — Wrightia 1: 90—94.
Stebbins, . 1977. Developmental and "—
e
roma of the Compositae. 1: 91-110. Academic
w York.
Shiney, T. E 1990. Plant Taxonomy: The Systematic Eval-
uation of Comparative Data. Columbia Univ. Press, New
ork.
Tamamshjan, 5. С. 1959. Astereae. In Flora URSS 25:
. 1937b. Compositae Japonicae. Pars Prima. Mem...
24—290. Acad. Scientiorum USSR, Moscow and Len-
ingrad. [In Russian.]
Turner, B. L., W. L. Ellison & R. M. King. 1961. Chro-
mosome numbers in the Compositae IV: North Ameri-
can species, with phyletic interpretations. Amer. J. Bot.
48: 216-223
NUMERICAL LIST OF SPECIES
la. K. incisa subsp. incisa
lb. K. incisa subsp. macrodon
2. K. pinnatifida
3. K. mongolica
Да. K. indica subsp. indica
4b. K. indica subsp. collina
4c. K. indica subsp. stenolepis
Та. K lautureana subsp. lautureana
7b. K. lautureana subsp. mangtaoensis
8. K. integrifolia
INDEX TO NUMBERED COLLECTIONS CITED
The numbers in parentheses refer to the corresponding
species in the text and in the Numerical List of Species
presented above. — — with asterisk (*) are
discussed in more detail in
ees: 4 (4c); Amano 6806 (4a); Ando 27 (4a); Anhui
dition Team 3300 (4a); Anshun Team 665 (4a), 1376
a 1759 (4a), 1903 (4a), 3221 (4b
6 (5b), 7 (5b), 11 (5a); Balansa 840 (4a), 4498
LLA 1500 (6), 1586 (6), 1587 (6); Bao,
Yan, S. J. 2209 (4c); Beattie & Kurihara 11119
2.1 11205 (Ба); EN pud 495 (4a), 983 (4a), 1190 (4a),
564 (4a); Bodinier 1913 (4a), 2474 (4a); Bohnhof 41 (3);
а 82 (4b); Bois-Reymond 805a (4a); Boufford & Wood
19
Сао, Z. Y. 394 (4a), 855 (4a); Carles 438 (6), 485 (8),
799 ae ‘Chane, C. C. 4 (6), 5577 (4a), 6510 be 10678
(4a); Chang, H. T. 4690 (4b); ae T. C. 2906 (6); Char-
ette 1454 (5a); Chavet 804 (6); Chen, B. Y. et al. 221 (4b);
Chen, C. N. 2058 (8), 8603 (4a), a e. 8621 (6); ne
F. H. 328 (3); 579 (8); Chen, G. H. 193 (8); Chen, Q. L
65 (4b); Chen, S. 1829 (4a); Chen, S 70. 1472 (4Ь), 11800
4007 , 16312 (4b), 16760 (4b);
. C. 65 (4a);
Y. 2978 (7a), 14941 (6); Chiao. C. Y. & Fan,
(4a); Chien, S. S. 5320 (4a), 5948 (4a); Ching, R. C. 3629
Ар ge су: 10015 (4a), 10260 (4a), 10462 (4a); Chou,
593 (4c), 26886 (4c), age К 40593 (3);
59 (la); Chow 33
4c)
18412 (4a); Chow, K. 5. 802
(4a), 1823 (4a), 2076 (4a); Chung 473 (Ла), 3101 (3), 3107
(la), 9288 (Та); Chung, H. H. 1702 (4c), 2326 (4b), 2402
(4b), 3815 (4b), 7596 (4c), 9129 (4c); Clemens 1512 (7a),
2002 (8), 4152 (8), 4153 (3); Cowdry 106 (8), 119 (3),
120 (8), 912 а Cui, С. & Lin, С. 1483 (Ла), 1499 (1a),
1528 (3), 1
= = x TE 7. H. 425 (4c), 922 (4c); Dai, L. Y.
. 2366 (4c); Dai, T. L. 102239 (4a), 102260 (4a),
102532 (4a), 103190 (4a), 106939 (4a), 107099 (4a);
812
Annals of the
Missouri Botanical Garden
Dann 54 (1a); David 2000 (8), 2029 (3); ra 63 (8),
79 e 80 (7a), 92 (8), 120 (4a); Deng, F. B. e
(6), 3726 (6); Deng, L. 2062 (4b), 5081 (4b), 7027 (4b),
7108 (4b), 7972 (4b), 8419 (4b), 8732 (4a), 8736 (4a);
ee Team 507 (4a), 772 (4a), m м 892 (4а),
999 (4а); Doi, У. 40 (Ба); Dong, N. С. 1472 (4b); Dorsett
& Dorsett 3698 (8); Ducloux 289 (4a), 1585 (4a
Eberhardt 3284 (4b); Ehara 48 (5a); Elipse 78 (8); El-
liott 144 (2)
pa 9 (Та); Fan, С. 5. & Li, У. У. 133 (4a); Fan,
H. N. 9557 (4b), 9588 (4b), 9610 (4b), 9623 (4b), 9654
(4b), Ser (4b), 9728 (4b), 9784 (4b); Fang, M. Y. 24783
(4c), 24785 (4c), 24915 (4c); Fang, W. P. 1442 (4a), 12360
(4a); Миа W. Z. T bs Le. (4a); Mou Z. F. 109 (7a), 249
(7a); Fan g, Q. S. 2843 (8); Faurie 226 (4a),
392 (8), 303 ke Mes (la), 404 (la), 1014 (2) 1073 P
1141 (4a), 5158 (2), 5159 (2), 6743 (2); Feng, К
0290
А 5 (4a); Forrest
3063 (4a), 4005 (4a), 8312 (4a), 11539 (4a), 13025 (4a),
22064 (8) 22424 (4a), 28457 (4a), 30584 (4a); Fortune
124 (4a); Fu, G. X. & Zhang, Z. G. 259 (4c), 260 (4c),
464 (3), 1145 (4a), 1584 (4c); Fu, J. Q. 40 (6), 365 (4a),
A (8), 565 (4c), 919 (7a), 1201 (7a), 1206 (7a), 2275
Im Kk T
4c
(4a), 8758 (4c), 9742 (7a),
(Та), 1 сукњу Fu, P. У. 1754
(8), 2649 (8); Fu, Р. У. & Zhang, Z. Н. 451 (3); Fu, P. У.
et al. 56 (8), 959 (3), 2039 (8), 2180 (1a), 4568 (3); Fu,
S. X. 110814 (4c); Fudan University Team 50299 (4a),
), 2664 (5b), 7518 (2), 8265
(2). 9887 (5a), 11250 (Sb), 11801 (5a), 12298 (5b); Fu-
kuoka et al. 994 (5a); Fung, H. 21109 (4b); Furuse 27
(2), 13944
Gadlceau 3944 (1a); Gao, У. 2. et al. 10 (4b); Ging, T.
S. 5233 (4b), 5399 (4b), er (4a), 6657 (4a), 7143 (4a);
Gu, H. 301 (8); Gu, H. and Chen, D. 121 (4c), 122 (4a),
hs (4a), дн > 134 а 183 ©. 189 (6), 195 (4a),
00 (4a); Gu, H. & Gao, Y. 289 (7a), 292 (4a), 293 (4a),
Eh (4a), 295 (4a); Gu, de & Han, 0. 205 (4c), 206 (4a)
208 (4a), 219 (4c); Gu, H. € "А
(4a); Gu, Н. & Li, 0. 305 a Gu, H. & Li, Z. 306 А
307 (4Ь), 308 (4Ь), 309 (4Ь), 310 чь) 314 (4b); Gu, Н
& Jiang, М. 304 (4a); Си, Н. & Noguchi 418 (5a), 420
(5b), 421 (Ба), a (5a), 423 ph 424 А (Sa), 425 (5a); Gu,
. & Takahashi 386 (5a), 389 (5b), 392 (5b), 393 (5b),
pes (5b), 397 ch 398 (5a), 400 (5a), 401 (5b), 402 (5a),
403 (5a), 404 (5a), 09 x 406 ( has үү (5b), 408 (5b),
410 (5a), 411 (5b); G 15 (2), 322 (2), 323
(2), 325 (2), 326 (2), goes (2), 327 а, 328 e» hig e»
426 (2), 427 (2), 428 (2), 429 (2), 430 (2); С
Zhang, H. 272 (8), 273 ыз. 274 (8), 278 (8), 283 | s E.
(3), 286 (3), 286 (3); С ang, W. 276 (la), 279
(la), 280 (la); Gu, H. et ч 249 (8), 251 (8), 257 (3), 263
(3), 264 (3), 265 (8), 266 (8), 266a (la), 267 (3), 268 (8),
269 (3), 270 (1a), 348 (Sb), 349 (5a), 350 (5a), 351 (5a),
352 (5a), 353 (5b), 354b (5b), 354c (5b), 354d (5b), 355
415 (2); Guam, K.
J. 1963 (4a), 74457 (4a); Guan, К. J. & Wang, W. T. 1
(4a), 2264 (4a), 2629 (4a); Guan, K. J. et al. 725 robe
854 (4a); Guangxi Team 3760 (4a); Guangzhou Qisi Team
4956 (4b); Guizhou Team 3581 (4b), 3947 (4b), 4107 (4a),
7063 (4a), 7412 (4b), 8053 ed ا . 2. 48 (Ас), 1839
У 3996 (6); Guo, Q. F. 1400
andel-Mazzetti 2694. un. de Коб а),
ex (4a); Hashimoto 754 (Ба, 5c)*, 4670 (2), 4714 (5b),
1 (5 Пи 17309 (4a); Hatusima & Sako
Eoi с 3088 7 (4a); Hautefenille 161 (4a); Не, 5. B.
7 (4c), 735 (4c); He, X. Y. 24938 (4a), 25882 (4a),
Res (4a), tenet (4a), 29842 (4a); He, Y. Q. 686 (7a),
1102 (4a), 14897 (4a); Heihuangdi Team 15 (8); Heilong-
jiang "acts Team 142 (1a), 151 veel 205 (1a); Hem-
mi 355 (2); Henan Team 1563 (7a), 2080 (4a); Henry 70
(4c), 89 (4a), 216 (4a), 384 (4a), 564 (8), 835 (4c), 891
(8), 1051 (4c), 1343 (4c),
48 (4c); Hori 16 (8); Hotta 10181
(2), Howell 95 (4a); а W. Y. 1945 (3), 4744 (6), 4748
(6), 6338 (4a), 9949 (7a); нан, Т. М. 539 (4а), 559
(4a), 616 (4a); Ни, О. М. 2966 (4a), 3375 (4b), 8 (4a),
5297 ry 5522 9 осы (4); Ни, 0. 7. 3 (4а); Ни,
ie & He, Z. 12026 (4a); Hu, Y. Y. & Wen, S. K.
0599 (4a), 58071 T p a); Huadong Field Station Team
А (6), 3170 (4а), 3773 (4с), 3776 (6), 4274 (4с), 4488
©), 4795 (4a), uo ); Huang, C. 165222 (4a); Huang,
R. H. 56 0 (4c); Huang, Z. 37257 (Ab), 42103
вы, 42350 чы, pum (4b); Huanghe and Ganyi Teams
835 (4c), 1939 (4c); Huangtu-gaoyuan Team 368 (7a),
x (4c), 4 (4с), 1376 PU. a (6); Hubei Expedi-
tion Team 24197 (4a); Hugh 44
Inoue & Im 2252 (5a); Ito, H. 2); Ito, Т. 510 ene
A og 2 (4a); Iwatsuki 3486 oe atsuki & Коу
37 (2); Iwatsuki et al. Т 9428 (4
оба 5. 8973 (8); Jiang, Х. C. et al. 228 (7a); Jiang,
M. 35676 (4a); Jiangxi Team 835 (4a), 2419 (4a); Ji-
gxi Teachers’ College Team 12037 (4b); Jilin Teachers’
Done т 788 (3)
Капа! 419 (2), 6481 (2), 731271 (2); pus 18 “x =
3354 (8), yA» (3), 4757 (4a), 4838 (6), 5615 (Та); K
black 273 (4a); Karo 182 (la), 1506 vs тэт по, 9767
а Каю 5 5184 d Ke, P. 234 (7a
Кабан, К.
(7a hk ain S. 3485 it 4204 0),
13126 (2); viter H. 27236 (Ба), 65094. (Ба); Koid-
zumi, С. 94955 (2); Кокапо 8 (5b), 19 (5b); Kolshaff 95
(6); Komarov 1507 (1a), 1507b (3), 1510 (3); Кота 11564
(2); Koyama 220 (2), 407 (2), 775 (2), 1492 (2), 2979 (5b),
7311 (2), 7541 (5a), 7550 (5b); eec 14382 (3), 14389
(3; Kuan, K. C. & Chen, 4 (Та); Kung, Н. W.
2845 (4c), 3737 (6); Kurachi: ass pé Кипћага 99 (5a);
Kurihara et al. 160 (2); Kuriyama 2009 (5b); Kurosaki
2116 (2), 2298 (2), 2607 (5a), 6542 (5b), 8019 (2), 9537
(5b), 9716 (Ба), 11017 (Ба), 11104 (2), 11365 (2), 11432
(2), 11585 (5a), 11893 (2), 12069 (2), 12928 (5b), 13656
(5a), 13994. (5a), 140001 (5b), 14560 (5a); Kurosaki &
Kato * — 148 (5a); cutem et al. 2705 (5b
5841 (4a); Lai, S. S. 1651 (4a), 1797 (4b), 3134
(4c); Lai, S: S-S "e M. = 3809 (4a); Lai, 5. 5. et al
; . 664 (4b), 2152 (4b); Lau, S. К.
4097 (4b), 4157 an. 28542 (4b); Lau, S. Y. 20551 (4a);
Law, Y. W. 1
272 (4a), 1081 (4a), 3446 (4b); Li, C. L. 1938 (4a); Li,
G. L. 63217 (4a), 63279 (4a); Li, H. J. 4767 (4c), 4791
Volume 84, Number 4 Gu & Hoch 813
1997 Systematics of Kalimeris
Ас), sar (4c), 5486 (4c); 5757 (4c), 6246 (4c), 6493 asawa 66 (4a), 523 (4a); Nagata 2457 (4a); Nakai
Ас), 6623 (4c), 7229 (4с), 8739 (4c), 8900
3308 (4a), 4223 44 , 4843 (4c), 5501
4a), 5560 (4a), 5620 (4a), 5963 (4a), 6799 (4a), 6813
4a), 6909 (4a), 7004 (4a), 7056 (4a), 7211 (4a), 7330
a), a),
4a), 7454 (8), 7574 (4a), с (4а), 8153 (4а), 8439 (
8512 (4а), 8631 (4а), 8723 (4а), 8797 (4а), 9101 (4a),
9178 (4a), 9427 (4a), d E aer (4a), 9976 (4a),
11118 ru 109010 (4a); Li, R 159 (8); Li, S. 760
(la); Li, S. et al. 4655 (1a); Li, S. HE 167 (8; Li, S. X. &
Jak. hane 5471 (8); Li, S. X. et al. 2913 (8), 6893 (7a);
Li, S. Y. 428 (7a); Li, X. С. 202385 (4b), 202806 (4b);
Li, Y. et al. 821 (1a); Li, Y. K. 401235 (4b); Liang, B. H.
12882 (4a); Liang, C. F. 30624 Ја
(3), 1685 (8), 2422 (6), 7739 (8), 7761 (8), 11379 (7a),
12506 (6), 12526 (4c); Limmercuamv 241 (7a), 277 (7a),
283 (7a); Lin, C. Q. 693 (8), 783 (7a); Ling, K. 2310 (6);
Ling, Y. 2174 (4b), 2965 (4b), 3248 (4b), 3281 (4b); Lipuo
hen 2249 c о а (1a), 3115 (8); Liu, С.
584 (6); Liu, F. Та); Liu, J. M. 10477 (4a);
үм К. М. 1728 (Та), по (Та), 1841 (Та), 3821 (Та),
3839 (Та), 4842 (Аа), 5532 (Та), 5575 (Та), 8574 (Ас),
8692 (4c), 8995 (4c), 9039 (4c), 9048 (4c), 10342 (Та),
10457 ae у. (7a); Liu, К. R. 393 (4с), 774 (4с);
an ч "4 Os MN L. H. & He, G. Z. 16483 (4a);
; Liu, Q. F. 30164 (4b); Liu, Q. H.
T Pw © (^ gi S. 3685 (1a); Liu, S. et al. 1282
hee dy shee 1802 "à 8298 уз Liu, E: L. et al. 18
(4
(3); TA T. W. 697 (6), 749 (6). 751 (6); Liu, X. Q. 25137
(4b), 28542 (4b); Liu, Y. С. 2538 (4b); Li
T. P. 1439 (4a), 1781 (0), 1983 (4a), 2697 (4a); Long, Х.
4c); Luo, S. ке 140 (Та)
8 B22 (4a),
peed 50 (4a), 351 (4a), 454 (4a), 525
68 (4a); Makino 2 (2), 28 (2), 44 (5a), 149 (2), 169 (4a);
a 4176 (2); Денеа 512 (5b), 924 (5b), 925
(5b); Mayebara 5800 (5a); Mengning Team 1150 (3); Mer-
rill 1507 (8), 1509 (4a), 1526 Де 10021 (4a); Michaelis
78 (4c); Midorikawa 508 540a (2), 540b (2), 661 (2),
1028 (2), 1368 (2), 1729 (2), 2002 (2); Migo 34 (8), 35
da) 36 ба), 37 ба) 38 (а), 41 (a), 42 (6), 43 (4a), 44
4a), 66 (4a); Mills Це
4008 (Ба); уай 7277 (4a); орно 80 (5b), 105 An
2 (2); Momiyama 18 (2);
Mori 53 0), 195 (3), 282 (3), 335 (4а); Ми,
297 (6); Murai 33 (5b); Murata, С. 6716 (2), 8355 qn
11818 (2), 12631 (5a), 12681 (5a), 13430 (2), 13232 (2),
eid (5a), 14351 (2), 17491 (2), 18437 (5a), 19036 (5c).
38 (Sb), 1966 gis (Ба), 20083 (Ба), 27445 (2), 27540 (2),
ca (2), 466 sar Murata, G. & Koyama 41306 (2),
. & Nishimura 270 (sb), 293 (5c);
; Миг
5c), 36 (5c), 38 (5c), 39 (2); M
128 (5b), 33895 (2); Murata, J. 6075 (2),
(5b), 6864 (5b), 8415 (5a), 9908 (2), 10069 (5a), 11496
(4a), 114890 (5a); Murata, J. & Ohba 5382 (2); Murata,
J. et al. 4496 (5a), 6257 (2), 15419 m. 33871 (5c);
њама К. 1610 (2)
= Sa), 2773 (la), 2785 (la), 3592 (la), 6519 (1b),
6520 (4a), 6522 (1b), 13218 (la); Nakajima 86 (5b),
10998 (5b), 15499 (5b), Nakashima 16552 (5a); Nanba
se b Nanjing Forestry College Team 7800-447 (4a),
6); Naruhashi 1856 (2); Nie, M. X. 4681 ~
eg m Nie, M. X. & Chen, S. L. 7418 (4a); Nie, M
X & Lal, 5.5. 3508 (4a), 4152 (4a), 4681 (4a), 5172 (Aa);
Nie, M. X et al. 3031 (4a), 9650 (4b), 9757 (4b); Nied-
erlein 6 (8); Ningxia Team 1-0228 (4a); Nitta & Mabuchi
12012 (2); Nomura, N. 53 (1a); Nomura, Y. 3 (5a); Nyi,
rk
8 (2); Ohashi, H. et al. 1546a (2); Ohba 48
as n jS 652000 (5a), 718022 (2); Ohba & Aki-
yama 1987 (2), 2559 (5a); Ohwi 1002 (5a); Oka, K. 3647
(5) 3671 (5b), 6496 (5a), 6497 (5a), 22925 (5b), 34250
Ce 37697 Ge); Ока, 5. 22 (8), 452 (8); Okamoto, K. 690
*; moto, 5. 16067 (la), 16068 (la), 19677
(5b). 19678 Ga): A 17476 (5a), 17580 (5c), 18011
(5a), 23799 (5c); Okuyama & Utsumi 11470 (5a); Oldham
244 (4a), 399 (5b); Ou, L. S. 150 (4b); Oouchi & Okuyama
20385 (5a)
Pai, Y. Y. 1168 в) 1282 (6), 1340 (4a), 1399 (6); PE
Team 1585 (3); Р 8083 (4a); Peng 46699 (4a); Peng,
D. Y. 45389 ен gom = p Ping, T. K. 1884 (4b);
Бру; e M E
Gian 0 shat Team 1781 (4a),
2329 ab ag do, p (4a), 3065 (4a), 3723 (4a);
Qiannan Team 2329 (4a); Qiao, Y. L. 108 (4a), 353 (4a);
Qiu, B. Y. 50787 T 60231 (4a), 77126 (4a), 77566 (4a),
86 (la); Rechinger 2260 (4b); Rock 41 (3)
— 613 (1a), 770 (1a); Saimi 5857 (5a); Saito 270
4150 n^ 5836 (1b), 5839 (7a), 5842 (3),
(3), 9537 (8), 10219 (3), 10240 (3), 10283 (8), 10287 (3);
Schindler 308a (4а); Schnack 135d (3); Second Humeng
Team 336 (8); бари Unversity Team 19 (8), 3103
(Та); ee 284-B (8); gnongjia Team 21777 (4c);
Shi, G. L. 12093 (4b), 13241 db) Shigeru 200 (5a); Shi-
Team 172 - t
(4c); 620 Team 2- dee iim. 3-357 es чнч, Н. UN «a
6429 (4c), 7677 (6); Steward & Cheo, E: C. 1077 (4b);
Steward et al. 51 (4a), 450 (4a); Su, G. D. 60 (7a); Sun,
2 (4a), 830 (4c), 849 (4a), 1239 (4a), 1259 (4a);
ч un, S. С. & Chang, К. 667 (4a), 1170 (4a), 1206 Saa
1417 (4a), 1772 (4a); Sun, S. L. 982 (4a); Sun, Y
Jin, Z. S. 252 (6); Suzuki, M.13 (8); Suzuki, T. 12068 »
Tagawa 106 (2), 2989 (2), 6748 (5a); Tagawa & Iwatsuki
1991 (2), 2442 (4a); rra H. 2838 (2); Takahashi,
K. 215 (5c); Takahashi, M. 547 (3), 860 (3), 1702 (5a);
Takahashi, N 102 (2); Takaki 327 pp Takano 47 (8);
Takato 311 (5a); Takenaka 198 (2), 2
Koyama 32106 (6); Tan, P. X.
59944 (4b), 60363 (4b), 60368 (4b), 60369 (4b); Tanaka
93 (4a), 13885 (5a); Tang, С. С. & Ма, S. E. 2906 (4b);
€ a L. 399 (4c), 561 6 976 (4а); Tang, 5. С. 5706
(6); T. sia, 96 (4a); Taquet 226 (4a), 950
(1b), 951 (1b), 952 (1b), ob. 99] (1b), 992 (1b) 6236
(1b), 9679 (4a); Tateishi et al. 4363 (2), 4496 (5a), 4563
(2); Tatewaki 31540 (3); Tawada 3 (4a); Teng, 5. W. 233
(4c), 90735 (4b); еен 728 (5b); Third Yiejin Team
384 (7a); Tian, X. F. 59 (7a); Togashi 2641 (8); Tomioku
5972 (2); Torii 2388 (2), 2392 (5a); Toyama & Okuyama
un
814
Annals of the
Missouri Botanical Garden
18770a (5b); Tsai, H. T. 53509 (4a), 53573 (4a), 60729
(4a), 61066 1066 (4a), 62482 3 (4а) 62827 (4a); Tsang, W. T. 7
(4b), 20844 (4a), 21400 (4b), 21594 (4b), 22759 (4b),
22994 (4b), 23247 yen 28036 (4b); Togashi 2641 (8);
Tsiang, Y. 1189 (4b), d eA 10223 (4a), 10770 (4a);
siang, Y. ang, sor Tsoong, P. C. 26 (4b),
39422 (4a); ея : 1 (Ба); Tsui, T. M. 696 (4b);
beers S. 25505 (5a), #7560 (5a), 27532 (5a); Tung, У.
aed mura 6 (5b); Uno 94 (5b), 22976a (la)
Wang, C. 40255 (4b); Wang, C. & Liu, Y. X. 982 (7a);
(3); Wang, C. гу А (4а); у; g“ =з 45 (4a), 2247
a
(3); Wang, н. 41433 (Да); Wang,
(4a); Wang, J. W. & Wu, F. T. 120 (4c); 7
(4a), 302 (4c), 306 (4c), 370 Чо Wang, Т. 1671 (1а),
2465 (la), 2572a (la); Wang, Т. et al. 2182 (Ла), 2
а
(8), 548 (8), 1001 (Та), 2647 (8), 2656 (3); Wang, X. W.
et al. 192 (4b); Wang, Y. C. 513 (7a), 639 (7a); Wang, Z.
Х. 168 А Wawra 1214 Ber 1269 bred 196 (8);
Wei, Z. 047 (4c), 1966 (4c), 2 4a),
12199 (4 e e on 1887 (4a); tan d (4c), 1703
(4a), 1727 (6); Weight 142 (4a); Wu, K. M. 60165 (4b),
а = ђ)
n University Team 333 (4b); Xing, J. 0. 1075
(Ac), 7992 (Aa), 9342 (Да), 9660 (4c), 10063 (4a), 10308
(4a), 10555 (4c), 10652 (4c), 10901 (4c), 10975 (4c),
11073 (4c), 11273 (4c), 11679 (4c), 12096 (4c), 12619
(4c), 12738 (4c), 12820 (4c), 13056 (4c), 13089 (4c),
17789 (4c), 17990 (4c); Xinjin Bot. Exp. Team 9 (7a), 56
(7a); Xiong, J. H. & Chou, Z. Y. 91840 (4a), 92444 (4a),
009 (4a); Xiong, S. 7109 (4a); Xiong, Y. G. 10070 (6);
Xu, X. H. & Lin, H. F. 153866 (4b); Xu, Y. B. 10051
4b)
Yamamoto 384 (5a), 386 (5a), 3446 (5a); Yamanaka
26157 (5a), 54349 (5a); Yamasaki 8986 (2); Yamatsuta 13
(2), 80 (3), 84 (3), 85 (3), 87 (1a), 93 (1a), 94 (3), 98 (8),
99 (1a), 104 (3), 105 (3), 111 (7a), 114 (3), 115 (3), 277
(3), 290 (3), ete => 4299 А Yamazaki et al. 425
Yang, G. H. 56939 (4a), 65084 (4a); Yang, J. X. 144
oa keen (4c), 598 (Та); Yang, X. : 3625 (6); Yang, +
6 (4a); Yang, 2. В. & Yao, J. 1164 (4c); Yao, K.
ane Yos Yao, Z. W. 4042 (4a); us H. H. 462 (Ab); Yi,
W. Q. 60-1238 (4a); Yingkou Team 294 9 dr 3557
(5a); Yoshioka 12 (5a); Vues 523 (4a); Y
4a), 992 (4a); Yue, J. S. et al. 1706 (4b), 2181 (Aa), 2826
A 3865 (4a), 3865 2 4894 (4 a)
han, X. X. et al. 1067 (4a); Zhang et al. 1864 (3);
ow б. Ge et ak а (4Ь); Хћапг, Н. Per 1929 (4b);
Zhang, R. M. 679 (4c); Zhang, R. Z. 25652 (4c), Zhang,
S. Y. 1052 (4a), 4194 (4a), 6828 (4a), 3107 (4a); Zhang,
X. M. 162 (8), 332 (6), 332 (8); Zhang, X. Y. 1131 (4a);
Zhang, Y. & Wang, S. 657 (1 He ути У. et al. 1226 (1a),
2117 (la); Zhang, Y. L. & Liu, T. S. 8 (8); Zhang, Y. L.
et al. 945 (3), 1812 (8), 1833 (8), 2291 (8), 2292 (3);
Zhang, Z. R. 25884 (4c); Zhang, Z. W. 2855 (7a); Zhang,
Z. Y. 189 (4a), 662 (6), 1237 (4c), 3541 (4c), Е (4c),
3789 (4c), 6366 (4c), 6573 (4c), 6839 (4c), 7065 (4c),
7094 (4c), 7884 (4c), 8827 (4c), 9240 (4c), бен (4с),
11802 (4с), 13029 (4с), 13732 (4с), 14791 (4а), 16908
а);
9121 (4с), з
4c); Zheng, 7. e Ê ); Zho:
60092 (4b), A60821 (4b), A60881 (4b), A62622 ae
u, H. B. 3021 (6); Zhou, T. P. & Liu, Z. F. 2421 (4b);
Zollinger 895 (4a), 899 (4a)
Mesum damnet len eri. m mm |
____ - JN
A REVIEW OF THE GENUS
CYDISTA (BIGNONIACEAE)!2
Warren D. Hauk?
ABSTRACT
vmm (Bignoniaceae) is a genus of six primarily lowland Ang ranging from central ¿and southern wg nde to
lin
guay and eastern Brazil. All species are lianas with showy
Par
to Maur abinde fruit, and winged seeds. This paper баа > оп eu work of the ‘wig 7 ең m Gentry and ин a ап
extensive database
ced from his investigations. Descriptions, separate keys to flowering and fruiting material,
pr
species ин graphs of flowering and fruiting phenology, and an illustration are presented.
The genus Cydista (Bignoniaceae) belongs to
tribe Bignonieae, which is composed almost exclu-
sively of lianas. All six Cydista species are lianas
that produce showy white to magenta, campanulate-
funnelform flowers, frequently with conspicuous
nectar guides (Fig. 1). As one of several closely
related genera, Cydista differs from its allies in its
cymose inflorescences, 2-foliolate leaves, simple
tendrils, linear-lepidote ovary, and medium reticu-
late pollen (Gentry, 1978; Gentry, Flora de Colom-
bia ms., unpublished; Gentry & Tomb, 1979). Other
characters that are useful to distinguish Cydista are
stems with eight phloem arms in cross section and
the absence of both a nectariferous disk and inter-
petiolar glandular fields. Cydista has no established
infrageneric classification. One species, C. aequin-
octialis (L.) Miers, has two recognized varieties.
The late Alwyn H. Gentry treated Cydista in sev-
eral regional floras, but his monographic work did
not encompass Cydista or any other genera of the
liana tribe Bignonieae. This paper utilizes the vast
amount of information chronicled by Gentry in his
studies of Bignoniaceae, compiles these sources of
information on Cydista into a single treatment, and
maintains the fundamental species concepts em-
ployed by Gentry, because there is ample evidence
to support his taxonomic hypotheses. The database
established by Gentry’s investigations has allowed
production of detailed maps of geographic distri-
bution (Figs. 2-6) and graphs of flowering and fruit-
ing phenology (Figs. 7-14). The new distribution
and phenology data presented here provide an im-
portant foundation for future investigations of tax-
onomy and speciation in Cydista.
HISTORY
John Miers first described Cydista in 1863,
based on Bignonia aequinoctialis L., and trans-
ferred Bignonia diversifolia Kunth to Cydista. At
least five other species names were published that
are now regarded as synonymous with C. aequin-
octialis [C. amoena Miers, С. incarnata Miers, С.
seemannii Miers, C. spectabilis (Vahl) Miers, and C.
pubescens S. F. Blake], and one that is synonymous
with C. diversifolia (Kunth) Miers [C. vargasiana
(DC.) Miers]. In 1919, Loesener transferred Arra-
bidaea potosina into Cydista, and in 1940 Seibert
described C. heterophylla from material collected
in the Yucatán Peninsula of Mexico. Until the work
of A. H. Gentry, no other lasting additions to Cy-
dista were made. Gentry (1977b) transferred Ane-
mopaegma decorum S. Moore into Cydista, and the
following year described material from Venezuela
as a new species, C. lilacina A. H. Gentry. Histor-
ically, species of Cydista have been placed in nu-
merous other genera (i.e., Bignonia, Arrabidaea,
Temnocydia, Levya, Апето , Clytostoman-
thus, and Pleonotoma), a fact that illustrates the
difficulty surrounding delimitation of generic
boundaries in Bignoniaceae. At least two other cur-
rently recognized species of Bignoniaceae, Mansoa
difficilis (Cham.) Bureau € K. Schum. and Roent-
genia bracteomana (K. Schum. ex Sprague) Urb.,
were originally described as species of Cydista.
SYSTEMATICS
As traditionally circumscribed, Cydista is one of
a group of closely related genera that possess an
s paper is number 4 of the GENTRY INVITATION SERIES, in acknowledgment of the contributions to the
коду а of the Bignoniaceae made by Alwyn H
ntry.
21 thank Peter Raven and the Missouri Botanical Garden
contribution to this work. 1 thank Scott Mori for helpful comments on the manuscript.
guidance was an invaluable
. 1 am especially grateful to William С. D'Arcy, whose
Susan A. Moore provided ds illustration. Financial support was provided by the National Science F oundation (grant
DEB-9509270
).
3 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S
ANN. MISSOURI Bor. git 84: 815-840. 1997.
816
Annals of the
Missouri Botanical Garden
Figure 1. Cydista potosina. ;
Alcorn 2172. —С. Enlargement showing lateral branching and quadrangular stem. —D. Seed
array of somewhat overlapping characters (Table 1;
Gentry 1977a, 1997; Gentry & Tomb, 1979; Tomb
& Gentry, unpublished). The genus most similar to
Cydista is Roentgenia, which Gentry (1978) consid-
ered “barely separable from Cydista.” Roentgenia
differs from Cydista in its trifid tendrils, racemose
inflorescence, and colpate pollen according to the
generic concepts established by Miers (1863) and
—A. Inflorescence and leaves after C. Cowan 3022. —B. Fruit with seed after Lh
.
Urban (1916), and later maintained by Macbride
(1961) and Gentry (1977a, 1978, 1982). Gentry
(1978) suggested that Cydista lilacina “links” the
genera Cydista, Roentgenia, and Clytostoma by its
elongate racemose-paniculate inflorescence and
overall leaf appearance. Clytostoma differs from
Cydista in its distinctive echinate fruit and glan-
dular-warty ovary, although Clytostoma has a “Cy-
a
һм мы. ҺА A | A #& YY A A AC A ОУУ _—
W _ A ~ AA "ÁO n
» d
Volume 84, Number 4
Hauk 817
1997 Review of Cydista
30
У a N
у in
20N У y n NN ~,
У
10N У
У Кам
у vL 3.
у т 7
v, ^. y М
0 > v У ý
wy" y "Ue wow
vY, ы T Y У Y, ~
y, > v of x
105 ¥ „т ы y
“~
wv" v
У
y v
20S
110W 100W 90W 80W ТӨМ 6 50W 40W
Figure 2. Distribution of Cydista aequinoctialis var. aequinoctialis.
dista-type" flower morphology (Gentry, 1978) and
lacks a nectariferous disk. Gentry and Tomb (1979)
suggested that the genera which lack a nectarifer-
ous disk and exhibit a tendency toward “multiple-
bang" flowering phenologies (Cydista, Roentgenia,
Clytostoma, and Phryganocydia) compose a natural
group. All four genera possess glandular-lepidote
corollas (Seibert, 1948; Gentry, 1977a), and Cydista
and Roentgenia are characterized by conspicuous
calyx glands (Seibert, 1948). Another closely allied
genus is the monotypic Potamoganos, which shares
the *Cydista-type" flower and cupular calyx, but
lacks glandular fields and possesses a well-devel-
oped nectariferous disk (Gentry & Tomb, 1979).
Palynologically, Cydista species have medium-
reticulate pollen (Gentry & Tomb, 1979), with pol-
len of C. potosina (K. Schum. & Loes.) Loes. and
C. heterophylla inaperturate, that of C. aequinoc-
tialis inaperturate or pericolpate, and that of C. de-
cora (S. Moore) A. H. Gentry, C. lilacina, and C.
diversifolia pericolpate. The latter three species
may exhibit a pollen type intermediate to the col-
pate pollen of closely related genera (e.g., Roent-
genia). Clytostoma shares with Cydista inapertura-
te, reticulate pollen but is distinguished by a
medium to coarse-reticulate exine (Gentry & Tomb,
9).
Goldblatt and Gentry (1979) reported chromo-
some counts of 2n = 40 and 2n = 40 + 1-2B for
Cydista aequinoctialis. The only other chromosome
count reported for Cydista is 2n — 40 for C. div-
ersifolia (Venkatasubban, 1944). Based on a survey
of 23 genera from tribe Bignonieae, these counts
are consistent with a base number for the tribe of
x — 20 (Goldblatt & Gentry, 1979).
DISTRIBUTION
Cydista species are found primarily in lowland
neotropical regions (Figs. 2-6) below 500 m in al-
titude, with collections reported from up to 2000 m
for C. aequinoctialis and C. diversifolia, to 1200 m
for C. heterophylla, to 950 m for C. potosina, to 900
m for C. lilacina, and to 3000 m for C. decora.
Among Cydista species, C. aequinoctialis (var. ae-
quinoctialis) is by far the most widely ranging (Fig.
2), extending from the western coast of Mexico
through the southern West Indies to the eastern
edge of Brazil and northern Paraguay. All other
species are restricted to portions of this area. Cy-
818 Annals of the
Missouri Botanical Garden
N
к
à
\
AS Ж
ә 2 Um
: j
| v
ram
А
МЕ A v
10N MS RS .
|
0 >
110W 100W 90W 80 70W
Figure 3. Distribution of Cydista aequinoctialis var. hirtella.
dista potosina (Fig. 6) inhabits the southeastern half
of Mexico and reaches central Nicaragua with a
single collection known from southern Costa Rica.
Cydista heterophylla (Fig. 5) has a range similar to
that of C. potosina, except that it reaches the west-
ern coast ‘of Mexico and extends through Panama
into the northern coastal regions of Colombia. Cy-
dista diversifolia (Fig. 4) has restricted represen-
tation along the coasts of central Mexico, is fre-
quent in the Yucatén Peninsula and Cuba, and
extends along the Pacific side of Honduras, Nica-
ragua, and Costa Rica, through Panama to central
Colombia and northwestern and north-central Ven-
ezuela. The remaining two species, С. decora (Fig.
4) and С. Шаста (Fig. 6), are restricted almost
exclusively to South America. Cydista lilacina is
distributed throughout South America north of Par-
aguay. Cydista decora has a more restricted distri-
bution than С. Шаста and inhabits two disjunct
regions, one in western Ecuador and a second, larg-
er area encompassing portions of Bolivia, Paraguay,
and southwestern Brazil.
PHENOLOGY
Cydista aequinoctialis and C. diversifolia possess
what Gentry (1974) referred to as “multiple-bang”
flowering phenology. Multiple-bang species have
numerous, synchronized, short flowering periods
(ca. 3 days) that may occur at any time of the year.
The absence of a nectariferous disk (and presum-
ably nectar), coupled with conspicuous visual and
olfactory attractants, indicates that pollinator deceit
may be the ultimate pollination strategy. The short,
repeated floral bursts may serve to lure novice pol-
linators that effect pollination through visits to only
a few flowers, after which they seek a more ample
nectar source. Visits by potential pollinators are in-
frequent, presumably because pollinators learn
quickly = the flowers offer no nectar reward
(Gentry, 19
Cydista s eiue) has a reduced multiple-
bang flowering strategy with only a very few (1—
2) bursts of flower production per year (Gentry,
1974). Detailed observations of flowering pat-
terns in C. lilacina, C. decora, and C. potosina
are not available.
ETHNOBOTANICAL AND ECONOMIC USES
Ethnobotanical reports include the use of Cy-
dista branches for basket weaving in Mexico's
Yucatán Peninsula, the incorporation of C. ae-
y Y
^
AA A AA €
а = ње es лави 77
"тн mmn m РЕР е р ДЬ ч тенч ОВР
Моште 84, Митбег 4
1997
Hauk 819
Review of Cydista
20N
10N
10S
20S
110W 100W 90W
Figure 4. Distribution of Cydista decora (cross-circle) and Cydista diversifolia (triangle).
quinoctialis into an infusion used as eye drops,
and the use of Cydista extracts to treat skin ail-
ments and ringworm (Gentry, 1992). Bignonia-
ae are known to produce naptho[1,2-b]furan
ie naptho[2,3-b]furan quinones, which have cy-
totoxic effects on neoplastic cells and exhibit po-
tential utility as trypanosomicidal and virucidal
agents (Ferreira et al., 1990).
MATERIALS AND METHODS
Gentry established a private database that com-
piled label information from herbarium specimens
he collected and from specimens that he personally
examined at other herbaria. Gentry's private data-
base has been incorporated into the Missouri Bo-
tanical Garden database management system TRO-
PICOS, which now contains label information for
all Cydista specimens housed at MO. For this treat-
ment, all specimens entered into TROPICOS were
assumed to have been examined by him. Specimens
examined by the author are indicated by a “!” in
superscript in the lists of representative specimens
and the Index to Exsiccatae. All types were as-
sumed to have been seen by Gentry unless other-
wise noted. Only a single type, that of C. lilacina,
was examined for this treatment. Gentry did not
always designate types as “holotype,” “isotype,” or
“syntype,” and the designations presented here are
based upon inferences drawn from Gentry’s work
and the original literature; these type designations
were not based on personal verification of speci-
mens at the various herbaria.
Data used for mapping and phenology were
downloaded from TROPICOS. For records with no
latitude/longitude coordinates in TROPICOS, co-
ordinates were obtained from gazetteers produced
by the U.S. Board on Geographic Names, Office of
Geography, Dept. of the Interior. Distribution maps
were produced using the computer program VER-
SAMAP 1.51 (C.H. Culberson, Newark, DE, 1991—
1995). Graphs of flowering and fruiting phenology
were generated using the computer program Quat-
tro Pro 7.00 Corel Inc. (1996). Phenology is re-
ported as the number of flowering specimens col-
lected during each month of the year and does not
imply that detailed studies of flower production (per
plant, per population, per species, or per time pe-
riod) have been conducted. Amounts of precipita-
tion used in the graphs of phenology were obtained
from Agroclimatological Data for Latin America
and the Caribbean (FAO, 1985).
820 Annals of the
Missouri Botanical Garden
| |
=з
y
b,
\
ui ~
7 ۲ dou "
v v¥
нн 8 eS s
¥ vA
110W 100W 90W 80 70W
Figure 5. Distribution of Cydista heterophylla.
TAXONOMIC TREATMENT
Cydista Miers, Proc. Roy. Hort. Soc. London 3:
191. 1863. TYPE: Cydista aequinoctialis (L.)
Miers.
Levya Buresi ex Baill., Hist. Pl. 10: 28. 1888. TYPE:
nicaraguensis Bureau ex me = Cydista ae-
qu E var. hirtella (Benth.) A. H. Gentry
Clytostomanthus Pichon, Bull. Soc. Bot. | DAR 92: 224.
.TYPE: into decorus (S. Moore) Pi-
chon = Cydista decora (S. Moore) A. H. Gen ntry.
Lianas, stems woody with 8 phloem arms in cross
section, branches terete to tetragonal, hollow or sol-
id in cross section, interpetiolar glandular fields
lacking, vegetative portions lepidote to puberulent;
pseudostipules inconspicuous to foliaceous, gener-
ally eglandular. Leaves opposite, petiolate, estipu-
late, simple or 2(4)-foliolate with leaflets oppositely
arranged, often with a simple, terminal tendril (or
tendril scar); petioles and petiolules glabrate to
conspicuously pilose or lepidote; leaflets entire, ve-
nation brochidodromous to actinodromous, second-
ary veins pinnate, midrib and secondary veins
prominent, occasionally with glandular fields in ax-
ils, margins plane to slightly undulate. Inflores-
cences cymose, terminal or axillary, several- to
many-flowered; rachis and peduncles minutely or
conspicuously bracteolate. Flowers ovoid in bud,
apices straight or curved; calyx campanulate, api-
cally truncate, mucronate teeth present or absent,
margin split or intact, eglandular, glabrate to dense-
ly lepidote; corolla conspicuously zygomorphic,
campanulate-funnelform, white to pink or purple,
glabrous to lepidote; corolla lobes 5 (2 upper and
3 lower), short-orbicular; fertile stamens didyna-
mous, a single staminode present, stamen and
staminode adnate to corolla; fertile anthers with two
spreading thecae, included, glabrous; disk wanting,
ovary cylindrical, usually densely lepidote, stigma
bipartite, the divisions laterally flattened, included.
Fruit a compressed, woody, elongate to elongate-
linear septicidal capsule, valves parallel to the sep-
tum, midline inconspicuous, surface smooth to stri-
ate or wrinkled, glabrate to lepidote or puberulous,
many-seeded; seeds oblong, flattened, bialate, oc-
casionally subhyaline toward margin, body ovoid,
frequently bipartite.
According to Gentry (1977a, 1982) there are six
species and two varieties ranging from Mexico and
бонна
Volume 84, Number 4 Hauk 821
Review of Cydista
30
+y
ays "
M \
XT ucc
20N
Y vg Y boss YS pus
уг? Ч
M
10N +
eo
" e
e ®
0 >
Ф
E Es
e
ev 9, 7
105 . суч x
8 e o
205 e
| 110W 100W 90W 80W ТОМУ 6 SOW 40W
Figure 6. Distribution of Cydista lilacina (cross-circle) and Cydista potosina (triangle).
the West Indies through Central America to Brazil 5'. Older and younger stems drying
and Paraguay еды ovules 4-seriate; bud apices
ay 5. C. lilac
KEY TO FLOWERING SPECIMENS
KEY TO FRUITING SPECIMENS
L pre drying bicolorous apically, pu t
m long; pseudostipules linear-triangular, n E — generally 3 cm wide or great
жш and overlapping ioe- snas wiles se osina nchlets terete; capsule surface "minkled:
1’. Calyces dui Bir iim dark apically, generally sane mostly 5-14 ст wide ....... 5. C. lilacina
7 mm long; pseudostipules ovate-foliaceous or 2’. Branchlets tetragonal; vw surface
inconspicuous, paired, and never overlapping smooth; leaflets mostly 2-5 cm wide ..........
2. Branchlets tetragonal; pseudostipules con- о 6. C. potosina
cipum po E LI . C. decora
3'. Inflorescences minutely bracteate; bran-
chlets hollow in cross section ---------------
ы ME 3. 6 e
2'. Branchlets terete to ruinas pseud
stipules inconspicuo
4. Plants ee while essentially leaf-
less; glandular fields in axils of seco ndary
veins infrequent and inconspicuous .....
4. С. ee
. Plants flowering with S glandular
fields in axils of secondary veins frequent
and (relatively) conspicuous.
ounger
stems drying dark; ovules 2-seriate;
bud apices straight 1. C. aequinoctialis
1'. Capsules generally 2 ст wide or less
Branchlets hollow in cross section; capsule
valves 1.0-1.5 ст wide; seed body darker
i 3. C. diversifolia
3'. Branchlets solid in cross section; capsule
valves pm 1.7-2.0 cm wide; seed body
same color as
4. ls conspicuous and folia-
ceous; олн tetragon o DEM Mp C. decora
> oe are poorly
eloped; buone | terete to qu иза
,
~
ins
Leaflets usually pinnately veined at
petiole apex; fruit lacking submargin-
al ridges or margins inconspicuously
uM SL su . aequinoctialis
. Leaflets with 3 principal veins ae
at petiole apex; fruit with 2 para
submarginal ridges ... 4. C. не
3
Annals of the
Missouri Botanical Garden
No. of Specimens or Precipitation
6 T
Month
ES Flower — —e- Fruit >< Managua - + - Caracas |
A
ai N
о о
/
E
UM ~
x
~
~
%
N
Ф,
M.
4
ГА
/
ў
ГА
x
1
~
EA 4
4
1
1
Мо. of Specimens ог Precipitation (ст)
0 Mea H і і | ; H ct
P 2 T $ 6 г 8 8. "m 1 x
Month
M Flower — -e- Fruit -x- Managua |
30 x
25 l- ms i
No. of Specimens or Precipitation (cm)
1772 5 4 5 6 PO
Month
ES Flower -œ Fruit NM Managua |
YE
M
te P oO e
Volume 84, Number 4
1997
Hauk 823
Review of Cydista
1. Cydista aequinoctialis (L.) Miers, Proc. Roy.
Hort. Soc. London 3: 191. 1863. Bignonia ae-
quinoctialis L., Sp. Pl. 623. 1753. Temnocydia
aequinoctialis (L.) Mart. ex DC., in A. DC.,
Prodr. 9: 155. 1845. TYPE: Plumier, Pl. Am.,
tab. 58 (holotype, P).
Lianas, older stems terete and drying gray,
younger stems subtetragonal and drying dark often
with four distinct light-colored ridges, solid in
cross section, occasionally sparsely to densely len-
ticellate, glabrate to pilose; pseudostipules incon-
spicuous, lepidote. Leaves 11-21 cm long, once-
pinnate with two primary leaflets and often a sim-
ple, terminal tendril; petioles 2-4 cm, lepidote or
pilose; petiolules 1—4 cm, lepidote or pilose; leaf-
lets entire, 7-14 X 3-7 cm, mostly narrowly to
broadly ovate or ovate-elliptical, sometimes orbic-
ular, apices acuminate to acute-acuminate, bases
obtuse or rounded, occasionally oblique or ine-
quilateral; venation actinodromous or brochidod-
romous basally, brochidodromous apically, 4—5
vein pairs, often with dark, abaxial confluent glan-
dular fields in axils of secondary veins, veins gla-
brous or conspicuously pilose. Inflorescences to 14
cm long, several-flowered, peduncles 1.5—6.0 ст,
the rachis and peduncles minutely bracteate, ped-
icels 5-18 mm long, sulcate, lepidote. Flowers
ovoid in bud, apices straight; calyx 4—5 X
mm, drying uniformly dark, margin intact to shal-
lowly split, teeth present and minute, lepidote with
glandular fields apically or irregularly overall; co-
rolla funnelform-campanulate, exserted 35 mm
above level of calyx lip, 2-3 mm wide at calyx
mouth, 15 mm wide at mouth, densely lepidote;
corolla lobes 20-22 X 15 mm, irregularly elliptic-
orbicular; stamens unequal, 9 or 15 mm long,
shorter filaments inserted 3 mm and longer ones
8 mm above level of calyx lip, the staminode 3
mm long, inserted at level of calyx lip; ovary 3
mm long, lepidote, ovules 2-seriate, style 25 mm
long. Capsule elongate-linear, 40-45 X 1.7-2.0
cm, submarginal ridges lacking or inconspicuous,
the central ridge more prominent; seeds flattened,
1-2 х 3-4 ст, oblong with lateral membranous
wings, each 2-2.5 cm long, hyaline margin 2 mm
wide, body ovoid, 1 X 1.8 cm, bipartite, not well
differentiated from wings. Figures: Schultz (1792:
t. 81), Descourtilz (1822: t. 100), Rohrhofer (1931:
t.3), and Gentry (Flora de Colombia ms., unpub-
lished).
Cydista aequinoctialis is one of the most common
and widely distributed species of the genus (Figs.
2, 3), extending from Mexico to Paraguay and east-
ern Brazil. It grows in tropical dry forests and in
edaphically drier regions of tropical moist forests
(Gentry, 1973b).
Morphologically C. aequinoctialis is the most
variable of Cydista species. The leaflets range
from narrowly to broadly elliptic, and venation
varies from basally actinodromous to brochidod-
romous. Pollen in C. aequinoctialis varies from
inaperturate to pericolpate and encompasses the
range of pollen types in Cydista. Thus, C. ae-
quinoctialis appears to be a cache for collections
that do not fit clearly into the more narrowly de-
fined limits of the other five species. А prominent
field character is the presence of glandular fields
in axils of main lateral veins on the undersurface
of leaves (Gentry, 1973b), although these are not
present on all collections and somewhat similar
glandular fields can be observed on leaves of C.
lilacina.
Two varieties of C. aequinoctialis, one with gla-
brate foliage and one with pubescent foliage,
were recognized by Gentry (1973a), with *some
hesitation as to the extent of genetic difference"
between them. Gentry (1973a) reported that the
glabrous and pubescent varieties grow in close
proximity with otherwise identical features. A
few individuals with intermediate pubescence
are found. However, nearly all herbarium collec-
tions can be easily attributed to either the gla-
brous or pubescent variety. These two taxa ap-
pear to correlate with ecological parameters; the
pubescent variety is found only in tropical dry
forest, whereas the glabrous variety occurs in
both tropical dry and moist forests, but more
commonly in the latter (Gentry, 1973a). The two
varieties have been recognized as distinct spe-
cies under a number of different names.
Gentry (1977a) noted that Plumier’s illustrations
were seen by. Linnaeus in 1738.
E
Figures 7-9. Flowering and fruiting phenology. —Figure 7 (top). Cydista aequinoctialis var. aequinoctialis, for the
entire range of the variety (see Fig. 2). Based on 827 flowering and 260 fruiting specimens. Precipitation in mm is
plotted 0.5X for Managua, Nicaragua, and 1X for Caracas, Venezuela. —Figure 8 (middle). Cydista aequinoctialis var.
aequinoctialis, from Central America. Based on 214 flowering and 82 fruiting specimens. Precipitation in cm is plotted
for Managua, Nicaragua. —Figure 9 (bottom). Cydista aequinoctialis var. hirtella. Based on 41 flowering and 20 fruiting
specimens. Precipitation in cm is plotted for Managua, Nicaragua.
Annals of the
Missouri Botanical Garden
N
о
=>
сл
Мо. of Specimens ог Precipitation (cm)
on о
о
meu
N
о
~
a
со
о
E
о
AEN
=>
=>
N
| => Flower -e- Fruit
No. of Specimens of Precipitation (cm)
>
о
6 T
Month
EE Flower -€e- Fruit
>< Managua -+- Maracaibo |
60
~
о
N
о
pee
ЕМ
Xe.
=
о
No. of Specimens or Precipitation
о
о
|
Џ
R
o
—— w
—
2 3 4 7
Month
10
To
+ Flower -@ Fruit
-x- Managua —— Tocumen |
us о“ —" — "QM — а" —Q' —-O ~ "о —
—— 2 Р ч y c
|
=
Volume 84, Number 4
1997
Hauk 825
Review of Cydista
KEY TO VARIETIES OF CYDISTA AEQUINOCTIALIS
1. Leaflets glabrous or nearly so _______________
1 en. Mgr cmn var. aequinoctialis
. Leaflets conspicuously pubescent, especiall
along main veins ben
LC aequinoctialis var. hirtella
lA. Cydista aequinoctialis (L.) Miers var. ae-
quinoctialis
Bignonia spectabilis Vahl, Symb. Bot. 3: 80. 1794. Cydista
spectabilis (Vahl) Miers, Proc. Roy. Hort. Soc. Lon-
: . 1863. [ones apenaii (Vahl)
Mart. ex DC., in A. DC., Prodr. 9
ae oH би ићи “ex India” (H. West?) (holotype?,
ype?, P-JU).
Binion villosa Vahl, Eclog. Amer. 2: 44. 1798. TYPE:
Colombia. Santa Marta: von in 21 (holotype. C).
Bignonia pica. Kunth, in Humb., Bonpl. & Kunth, Gen.
Sp. Quarto ed. 3: 136, Folio Р 3: "106. 1819. TYPE:
Mentel Orinoco: Humboldt & Bonpland 1078
(holotype
Bignonia hostmannii E. Mey., Nova Acta Phys.-Med.
s. Leop.-Carol. Nat. Cur. 12: 719. 1825.
E: тен Sieber s.n. (isotypes?, М, W).
Bignonia nitidissima DC., in rodr. 9: 160. 1845.
TYPE: Venezuela. ee Vargas 244 (holotype, G-
DC).
Arrabidaea isthmica Standl., J. Wash. Acad. Sci. 15: 46
E: Panama. Canal Zone: Pittier 2576 d
юйре, US; isotypes, GH, MO, NY).
Description as for Cydista aequinoctialis except
younger stems, petioles, petiolules, veins of leaflets,
and calyces glabrous or nearly so.
Although Cydista aequinoctialis var. aequinoc-
tialis flowers throughout the year, the peak for all
collections is July-August (Fig. 7). However, when
the Central and South American collections are
separated, two distinct peaks are observed: (1) Cen-
tral American collections peak in April (Fig. 8), (2)
South American collections climax during July—Au-
gust. The differences in flowering collections in
Central and South America may indicate that local
populations have adapted to regional climatic pat-
terns. Fruiting collections did not exhibit a marked
peak, although Central American fruiting collec-
tions were most numerous from December to Jan-
uary.
Representative specimens. MEXICO. Colima: W of
Manzanillo Bay, i. W of Santiago, 19*00'N, 104^00' W,
90-150 m бека 15708 (MICH). Guerrero: Baqueta,
Langlassé 506 (F). Jalisco: Mun. La Huerta, Estación de
Biología Chamela (UNAM), 19°30'N, 105°03’W, Lott 1873
(МО). Nayarit 15 km SE of San Blas, near Miramar,
22°52'N, 105°06' W, 0 m, Feddema 955 (MICH). Oaxaca:
12 km N de carretera Pinotepa—Pto. Escondido, Dtto. Jam-
poy 16?17'N, 97°49'W, 0-3 m, Torres & Cedillo 629
МО). Sinaloa: Villa Unión, Villa Unión, Mazatlán,
25°00'N, 107?30'W, 10 m, Gonzdlez-Ortega 5448
(MEXU). Tabasco: 10-40 km W of Huimanguillo, La-
guna Ocuapan, 17°51'N, 93°23'W, Barlow 30/179 (MICH,
WIS). Veracruz: Laguna de Sontecomapan—Río Coscom-
pan, 1971, Calzada 461 (CHAPA, F, MEXU, MO'). GUA-
TEMALA. Izabal: Río Oscuro, 0-8 km SW of Lake Iza-
bal, 14%05'N, 90?36' W, 0-600 m, Jones & Facey 3491 (F,
TEX-LL, MICH). Petén: Río Santa Isabel between El
Porvenir € mouth of Río Sebol, 100 m, Steyermark 45875
. San Marcos: Ocos, 14°31'N, 92711", 1-2 m, Stey-
ема 37855 (F). HONDURAS. Colón: road to Castillo,
0.2 mi. E Trujillo, 15%55'N, 86°00'W, Saunders 170 (MO').
mayagua: Las Limas, 14%08'N, 87748", 1000 m, Ed-
wards P-128 (F). Copán: La Florida, Pittier 8488 (US).
Cortés: Entre Cofradia у Cusuco, 15%30'N, 88°00'W,
200-1300 m, Molina 7299 (F, TEX-LL). Gracias a Dios:
La Mosquitia, 15%00'N, 84720", 1973, Clewell & Cruz
4002 (MO). Morazán: Cerca de Guaimaca, 14^30'N,
—_
14°45’ N, 86?00'W, 400
bara: Ulua River between Пата and Gualala, 15?10'N,
88?20'W, 500 m, Molina 22042 (F). BELIZE. Belize: 6—
7 km NW of Belize, Northern Hwy., 17°30’N, 8812'W,
1973, Dwyer 10710 (MO). Stann Creek: 16%58'N,
88713", Schipp 771 (F, MICH, МО). EL SALVADOR.
Ahuachapan: El Imposible, San Benito al S del Río Ar-
enal, 13?49'N, 8956", Sandoval & Chinchilla 332
MO’). Chalatenango: Carretera a la Palma, 14^19'N,
89^11'W, Montalvo & Vargas 3168 (MO). Santa Ana: Var-
gas & Montalvo 3216 (MO'). NICARAGUA. Chinandega:
Entre Corinto y Paso Caballos, 12°29'№, 87°10'W, San-
dino 2591B (MO). Chontales: Río Mayales valley, near
Сиара, Llano Grande, 12°13’N, 85%25'W, 200 m, Neill
7431 (MO). Estelí: Salto la Estanzuela, 13%01'N,
86°21'W, 1000 m, Moreno 21153 (MO'). León: Km 55
carretera Nueva a León, 12?18'N, 86°42’ W, 40-60 m, Mo-
reno 9870 (MO). Managua: road between El Crucero and
house of aep "me Julia, 11°59’N, 86°19’W, 600-900
m, Standley 1 (F). Matagalpa: Las Palomas, 11 km
W de Río atem Matiguas, 12°52’N, 85°15! W, Mo-
reno xc i
NE del Jícaro, 1 13°45 N,
Río San Juan: San Miguelito, 11°23'N, 84°54’ W, 50-80
m, Sandino & Martinez 3868 (MO). Rivas: Isla Ometepe,
Volcán Concepción, 11°32'N, 85°34'W, 50 m, Robleto
1594 (MO). Zelaya: Harbor of Bluefields, 12%00'N,
8410'W, Marshall & Neill 6522 (МО). COSTA RICA.
Alajuela: San Mateo, 09*56'N, 84*31'W, Brenes 3647
(CR, F, NY). Guanacaste: Finca la Pacifica, 2-3 mi. N
he bears 10°26'N, 85°06'W, Gentry 1113 (F, MO). Li-
n: Cerro Coronel, along Río Colorado at Laguna Danto,
10742! N, 83°39'W, 7 m, Stevens 24012 (MO). Puntar-
„~
be
Figures 10-12. Flowering and fruiting phenology. —Figure 10 (top). Cydista decora. Based on 46 flowering and 5
iting specimens. Precipitation in cm is plo tte d for Pedro Juan Caballero, Paraguay. —Figure 11 (middle). Cydista
aracaibo, Venezue
Precipitation in cm is plotted for Managua, Nicaragua,
120 flowering and "i games speci-
mens. етар in mm is plotted 0.2X for Managua, Nicaragua, and 0. x for Tocumen, Panam
826 Annals of the
Missouri Botanical Garden
n
=
Ф
Е
о
Ф
а.
[77]
5
о
z
0 T T MO e WO T T T
1 3 3 4 5 6 7 8 B. 410. 14 + d
Month
M Flower -&- Fruit |
50 ^ >
S 40 X 4 а
9 i “J
— / Ж
30 4
о 1
Ф ,
E 20 X. ў
; 2 / | ІА SS
0 \ 4
O Y
510 ا
> ~“ bcne
0 و o o o o Ls و
1 2 3 4 6 7 8 # wW IT 12
Month
+ Flower -@ Fruit — Veracruz -x- Belize |
Figures 13, 14. Flowering and fruiting phenology. —Figure 13 (top). Cydista lilacina. Based on 25 flowering and
12 fruiting specimens. —Figure 14 (bottom). Cydista potosina. Based on 95 flowering and 11 fruiting specimens.
Precipitation in cm is plotted 1.25X for Veracruz, Mexico, and 0.2X for Belize, Belize.
Morphological/anatomical characters and character states for Cydista and four genera that are presumably
)
able 1.
closely related (after Gentry, 1977a, 1997; Gentry & Tomb, 1979; Tomb & Gentry, unpublished).
Phlo-
em Nectar Corolla Ovule Fruit Pollen Pollen
Tendrils arms disk pubescence organization surface Seed aperture exine
Cydista simple 8 absent lepidote 2(-A)-seriate smooth bialate inaperturate/ reticulate
icolpate
Roentgenia rifid 8 sent lepidote 2-seriate smooth Ыајаіе 3(4)-colpate reticulate
Phryganocydia simple 8 absent lepidote 2-seriate moo оку inaperturate reticulate
Clytostoma simple 8 absent lepidote 2(4)-seriate echinate corky ^ inaperturate reticulate
Potamoganos _ trifid 4 present glabrous/ 4-seriate ? 3-colpate reticulate
lepidote
Volume 84, Number 4
1997
Hauk 827
Review of Cydista
enas: Osa Peninsula near Rincón, 09%55'N, 84^13'W,
Gentry 1263 (MO). San José: Tabarcia, Bajo de los Bus-
tamante, 09°51'N, 8414", 840 m, Solís 543 (CR-
24222). PANAMA. Bocas del Toro: Río San Pedro,
08°49'N, 81733", Gordon 80C-a (MO). Canal Zone:
Barro poen Island, 09?11'N, 79°57'W, Croat 11097
(SCZ). Chiriquí: Puerto Armuelles, 08°17'N, 82%52'W,
Croat Vilis (МО). Coclé: vicinity of El Valle, 08°37'N,
"W, 600-1000 m, Allen 1780 (F, GH, MO', NY, US).
Colón: along roadside ca. 2 mi. E of Fort Sherman,
09*22'N, 79°57'W, Gentry 733 (МО). Darién: El Real,
08°08'N, 77743", Gentry 4564 (МО). Herrera: 0.5 mi.
Е of Las Minas, 07748'N, 80745", Gentry 3138 (МО).
Los Santos: Punta Mala, 07728'N, 80700", D'Arcy &
Croat 4215 (МО). Panamá: Tocumen, 09%01'N, 79?23' W,
Dwyer 5130 (MO). San Blas: mountains above Puerto Ob-
aldía, 0840'N, 77?25'W, Gentry 1477 (MO). Veraguas:
Islas Contreras, va ial 07°51'N, 81°47'W, 0 m,
Churchill 5711 (MO
АТП. Bayeux, Bord-de-la-mer, 19°49’N, 72°26’W,
Ekman H2669 (MO). DOMINICAN REPUBLIC. Distrito
Nacional: Arroyo Tosa, 1.5 km from La Victoria on road
to Mata Mamon, 18?36'N, 69*50'W, 10-20 m, Mejía «€
Zanoni 9017 (MO?. El Seibo: Laguna El Limón, 38 km
of Miches, 18°58’N, 68°51'W, 0-5 m, Zanoni et al.
15883 (BSD, MO). Espaillat: 8 km W de Gaspar Her-
nández, 1938'N, 70°10' W, 10 m, Gentry & Zanoni 50609
ez: Llanura de Nagua, 29.4
Thomas: Mag Bay.
99'W. Кё per (EL Ly GUADELOUPE. 16*01'N,
тея 1395 (МО). GRENADA. W. Ind. Woburn
61^44'W, Broadway 1004 (1). MARTINIQUE. Environs
e la case Pilote Mars, 14?35'N, 61%00'W, Hahn 1407
(MICH). TRINIDAD AND ТОВАСО. Trinidad: entrance
to Caroni Swamp National Park, pen Port of Spain,
10°32’N, 61730", Harriman 17563 (MO
COLOMBI BIA. Amazonas: Leticia, 04°09'S, 69°57'W,
1974, Gentry 12735 (COL, MO). Antioquia: Mun. de
Sabanalarga, З km de Sabanalarga, 06^51'N, 75°49’ W,
1100 m, Callejas et al. 2238 (MO'). Atlántico: Barran-
quilla Las орем, 10%58'N, 74%54'W, 1937, Dugand
1135 (Е, МО). Bolivar: near Cartagena, 10°28'
75°32'W, 10 m, Gentry & Cuadros 47618 (MO). Boyacá:
El Yopal, Llanos Orientales, 04°44'№, 72°15'%, Blyden-
stein & Saravia 1251 (COL). Caldas: 14-21 km N of La
Они а on road to San Miguel, 05°27'N, 74°40’W, 330
m, Gentry et al. 18150 (COL, MO!). Caqueta: Cuernan,
Rio Caquetá, Pabon 533 (ARAR). Cesar: 5 km W о
Manaure, 10°22’N, 73?08'W, m, Gentry et al.
60737 (JBGP, MO). Chocó: Ríosucio, Parque Natural
Nacional Los Kaytos Cacaricas, 07°25'№, 7710" W, 120
m, León 338 (COL, МО). Córdoba: road from Fresquillo
to Tierralta, 08°05’N, 76°10’W, 100 m, Gentry & Cuad-
wa
15149 (COL, MO). Стани
67%06'W, 120 m, Gentry & Stein 46475 (MO). Outta
re: San José del Guaviare, 02?35'N, 72?38'W, 240 m,
Cuatrecasas 7473 (COL). Magdalena: Parq. Nac. Tai-
rona, 50 m, Gentry & Cuadros 47502 (JBGP, MO). Meta:
Mun. La Macarena, Río Guayabero 02*10'N, 74?06' W,
310 m, Callejas & Marulanda 7118 (МО). Nariño: Tu-
maco, Mun. Espriella, 01°49’N, 78?46' W, Romero-Cas-
tafieda 2793 (COL). Norte de Santander: Río del Oro,
18271 (COL, MO). Santa Cruz: 5 km W
Vista, 17°27'5, 63°42'W, 350 m, Nee et al. 36128 (MO).
Santander: 51 km E of Barranca Bermeja toward Bu-
caramanga, 07%03'N, 73%52'W, 200 m, Gentry & Forero
15351 (МО). Suere: Coloso to Finca Sirena, along Que-
brada El Salto, 09?30'N, 75?21'W, 300 m, Gentry &
Cuadros 68250 (МО). Tolima: Mariquita, 05^12'N,
°54'W, 600—650 m, Uribe-Uribe 3003 (COL). Valle:
Armenia. turnoff from Cali-Medellín hwy., S of Zarzal,
04?25'N, 76°05'W, 1000 m, Gentry & Juncosa 40922
(MO). Vaupés: Río iniri, vus in Palito, Schultes
& Cabrera 13126 (COL). Vichada: near Humaypia, Río
Vichada, 04%55'N, 67°50’W, Giovanni s.n. (COL). EC-
UADOR. Napo: 2-6 km above Puerto Bolívar, Río Cuy-
abeno, 00?06'S, 76?10'W, 300 m, Brandbyge et al.
33731 (AAU, МО). Pastaza: Via Auca, 115 km al S de
Coca, cerca del Río Tiguino, 01715'5, 76%55'W, 320 m,
Rubio 121 (МО). Sucumbíos: Cuyabeno, 00^16'S,
75°53’ W, 265 m, Paz & Mino 81001 (МО). PERU. Ju-
nín: Chanchamayo Province, Hacienda La Genoa, Finca
Italia, 11?05'S, 75?25'W, 1150 m, Gentry et al. 73439
MO). Loreto: Requena, Sapuena, Jenaro Herrera,
04*50'S, 73745", 170 m, Vásquez et al. 10040 (МО).
Madre de Dios: Tambopata, ca. 5 km from Puerto Mal-
donado, 12735'5, 69?09'W, 200 m, Gentry & Revilla
16281 (MO). Paseo: Cabeza de Mono, Río Iscozacin,
10°20’S, 75718", 320 m, Gentry et al 41685 (MO).
Puno: Río Tavara, ridge top across from mouth of first
major tributary, 13°21'S, 6940", 500 m, Gentry et al.
76897 (МО). San Martín: Tocache Nuevo, 08^10'S,
76°32'W, 450 m, Gentry et al. 25480A (МО). Ucayali:
Coronel Portillo, Bosque Nacional de von Humboldt,
08°40'S 75?00' W, 270 m, Gentry & Horna 29508 (МО).
BOLIVIA. Beni: Yacuma, San Borja, 50 km hacia San
Ignacio де Mojos, 14?49'S, 66*51'W, 250 m, Beck 13215
a Paz: Alto Beni, 14%00'S, 65°30’ W, Seidel &
Mundo, Campamento 18, 18 km
66°46'W, 160 m, Gentry et al. 77759 (MO
Perseverancia, NW of Santa Cruz on Río Меро, 14?38'S,
62°37'W, 100 m, Gentry & Mostacedo 73715 (МО).
VENEZUELA. Amazonas: banks of the Río Manapiare,
close to San Juan de Manapiare, 05%04'N, 66703", Ber-
ry 1593 (MO). Anzoátegui: km 227 on Caracas—Bar-
celona Hwy., 16 km Е of Boca de Uchire, 10708'N,
65*26' W, Gentry & Berry 14833 (МО). Apure: Ditto. San
Fernando, 06°16'N, 67°31'W, 55 m, Davidse & González
1603 (МО). Aragua: 8—9 km from the гедота at Сага
ach, SW towards Cuyaga, 1029'N, 68°42’W, 400 m,
pus et al. 2289 (МО). Barinas: 10–15 km W of Ваг-
inas, 08°38'N
07°35'N, 62°58’W, 270 m
Carabobo: Bahia de ном 10°26'N, 67°55’W, De-
lascio Chitty 2417 (MO). Delta Amacuro: Dept. Ped-
"№, 62703", 50 m,
Guaira, Est. Teleferico del Avila, 1
Plowman 7664 (MO). Falcón: Dtto. Zamora, Cerro
828
Annals of the
Missouri Botanical Garden
pee qp 11?27'N, 69?17'W, 400 m, González 1051
0. Guárico: banks of Río Orituco, S of Calabozo,
ies N, 67°27'W, Gentry 10263 (МО). Lara: Jiménez
Parq. Nac. Yacambu Qda. Honda, 09'41^N, 69*30'W,
700 m, Davidse & González 21329 (MO'). Mérida: Santa
María de Caparo, Dtto. Arzobispo Chacón, 08°30'N,
71°10'W, López-Palacios & Bautista 3299 (МО). Mir-
anda: Cerros del Bachiller, 10 km of Cupira, 10?09'N,
65°48'W, Steyermark & Davidse 116368 (MO'). Mona-
gas: Río San Juan, Dtto. Benítez, Estado Sucre у Dtto.
Maturin, Monagas, 09%24'N, 63%02'W, Marcano- Berti
Táchira: Ойно. Capacho, 07%52'N, 72719", 1250 m,
Bono 4950 (МО). Trujillo: Sabana Libre, 09°21'N,
70?39' W, Christ 75 (VEN). Zulia: Опо. Perijá, Carretera
San Ignacio-Barranquitas, 10%00'N, 72?30'W, 125-150
m, Bunting 5460 (MO'). GUYANA. Demerara: Demer-
ara-Mahaica Region, along road from Cane Grove to La-
may Meri 06°35'№, 57750", 1-10 m, Hahn et
al. 3817 (МО). ero ier W Demerera Region, W of
0'N, 58
merara River, 06? ;
Visserijzwamp, 05*?01'N,
5°42' W, Reijenga 57 (0. “FRENCH GUIANA. Cayenne:
Riviere de Kaw, 09'W, Granville 6848 (MO',
U). Inini тачан, Ed ifie en amont de Saut Bad-
jere, 04?39'N, 52?20'W, Granville B-3728 (CAY, MO).
Saül: 0338'N, 53712", 220 m, Gentry et al. 62974
(MO)
1B. Noct aequinoctialis var. hirtella (Benth.)
. H. Gentry, Ann. Missouri P Gard. 60:
s 1973. Dino nia sarmentosa Bertol. v.
hirtella Benth., Bot. Voy. Sulphur 128. 1845.
TYPE: Nicaragua. Realejo: Hinds s.n. (iso-
type?,
Bignonia sarmentosa Bertol., Fl. Guatimal. 25. 1840. Cy-
quintla: Velásquez s.n. (holotype?, BOLO).
Levya nicaraguensis Bureau ex Baill., Hist. Pl. 10: 28-29,
MSS. IX Nicaragua. Levy 38 (holotype?, P; is-
огуре КА.
Abe RU K. Schum. & Loes.,
Bot. Jahrb. Syst. 23: 129. 1896. TYPE: ich
Bernoulli & Cario 2056 (holotype, K).
Arrabidaea pseudochica Kraenzl., Repert. Spec. Nov. Reg-
ni Veg. 17: 19. 1921. (fide Sandw., Kew Bulletin 22:
403—420. 1960.) TYPE: Mexico. Michoacán: Lang-
lassé 137, 506 (syntypes, K).
Anemopaegma tonduzianum Kraenzl., Repert. Spec. Nov.
Regni Veg. 17: 116. 1921. TYPE: Costa Rica. Guan-
acaste: Tónduz 13912 (isotypes?, K, P).
Cydista pubescens S. F. Blake, Contr. U. S. Natl. Herb. 24:
2. TYPE: Honduras. Copán: Pittier 8488 (ho-
lotype?, US).
Description as in C. aequinoctialis except that
the younger stems, petioles, petiolules, and veins
of leaflets are conspicuously pilose.
Variety hirtella is restricted primarily to Mexico
and Central America (Fig. 3); only three collections
e known from South America. The number of
flowering specimens of C. aequinoctialis var. hirtel-
la peaks in May (Fig. 9), whereas in variety ae-
quinoctialis the peak is April (Fig. 8). Both varieties
initiate flowering before the onset of the wet season
in May, but flowering collections of variety hirtella
drop sharply after May, and flowering collections of
variety aequinoctialis remain common through Au-
e These apparent differences in phenology m
consequence of ecological divitias
SEX 1973a). However, few stable characters are
available to distinguish the two varieties, and rec-
ognition at a higher taxonomic rank is not warrant-
ed at this time.
Representative specimens. MEXICO. Chiapas: Acala,
Laughlin 850 (DS, F, MEXU). Guerrero: Montes de Oca,
Vallecitos, Hinton 10219 (MO'). Oaxaca: 5 km N of Ma-
tias Romero, King 808 (MICH). башк Pinotepa,
16719"М, 98701", Galeotti 7060E (MO). Veracruz: Es-
tación de Biología Tropical Los Tuxtlas, 18°34'N,
9509", 450 m, Colín 144 (MO). GUATEMALA. Jutia-
pa: Río Paz, Heyde 6363pp (MO). HONDURAS. Comay-
agua: Humuya e Hazlett 1099 (MO). EL SALVA-
DOR. Аћџасћарап: 0-2 mi. МЕ of San Francisco
Menéndez, 200-450 m, Croat 42068 (MO'). NICARA-
GUA. Carazo: between Amayito and Barranco, 11?40'N,
86°18'W, 30-300 m, Stevens 22734 (MO'). Chontales: 0.6
km NE of Hwy. 7 on road to Comalpa, 12?10'N, 85*33'"W,
160 m, Stevens et al. 17177 (MO'). Estelí: Paso León, 3.9
km NNE of Hwy. 1 at Estelí, 13*08'N, 86°20’W, 815 m,
Stevens et al. 15487 (МО). León: 4.0 SW of La Paz Cen-
tro, 12°18’N, 86°42’W, 35 m, Stevens et al. 17243 (MO).
atagalpa: carretera vieja a Jinotega, 800 m, Moreno
22917 (MO. Nueva Segovia: El Jícaro, 13?45'N,
86°06'W, 650—700 m, Moreno 1693 (MO?. Rivas: Potrero
Largo, 11°05'N, 85°42'W, 10-20 m, Morales 2935 (MO).
io San Juan: San Miguelito, Sandino & Martínez 3868
(MO). COSTA RICA. Guanacaste: Isla San José,
10°53’N, 85755", 50 m, Janzen 12474 (MO). PANAMA.
D леу Golfo de San Miguel, Gentry 3957 (MO). Vera-
ca. 11 mi. W of Зога, D'Arcy 5279 (MO).
У COLOMBIA. Сћосб: Bahia Solano, Gentry & Fallen
17173 (МО). Santander: Los Santos, Sabana de Torres,
328 m, Renteria 202 (MEDEL). VENEZUELA. Zulia: 13
de Embalse, 10%25'N, 70°49’W, 550—600 m, Bun-
ting et al. 11268 (MO!
2. Cydista decora (S. Moore) A. H. Gentry, Sel-
byana 2: 42. 1977. Anemopaegma decorum S.
Moore, Trans. Linn. Soc. London, Ser. 2. 4:
421. 1895. Clytostoma decorum (S. Moore) Bu-
reau & K. Schum., in Mart., Fl. Bras. 8c: 151.
1896. Arrabidaea decora (S. Moore) Hassl.,
Spec. Nov. Regni Veg. Repert. 9: 49. 1910
Clytostomanthus decorus (S. Moore) Pos
Bull. Soc. Bot. France 92: 224. 1945. TYPE:
Brazil. Mato Grosso: S. Moore 980 (holotype,
BM; isotype, NY).
Lianas, stems tetragonal with four conspicuous
p ч
y nina Wadi
Volume 84, Number 4
1997
Hauk 829
Review of Cydista
ridges, solid in cross section, drying gray to brown,
glabrous; pseudostipules foliaceous, 2.5 X 2.0 cm,
elliptic-orbicular, entire, glabrous. Leaves 8-13 cm
long, bifoliolate with a simple, terminal tendril (or
tendril scar); petioles 2-4 cm, sulcate, glabrate to
puberulent, petiolules 1—4 cm, sulcate, glabrate to
puberulent; leaflets 6-9 X 3—7 cm, mostly narrowly
to broadly ovate or ovate-elliptical, apices acute to
acute-obtuse, bases obtuse or rounded, occasionally
oblique, venation actinodromous basally and bro-
chidodromous apically, 4-5 vein pairs, midrib and
secondaries prominent, major veins puberulous be-
neath, isolated and scattered glands often in axils
of secondary veins, margins slightly undulate, char-
taceous. Inflorescences to 20 cm long, several-flow-
ered, peduncles 3.0—7.5 cm, the rachis and pedun-
cles conspicuously bracteate, the bracts 2-10 mm
long, pedicels 5 mm long, sparsely lepidote, pu-
berulent. Flowers ovoid in bud, apices straight; ca-
lyx 6-7 X 5 mm, margins intact, apically truncate
with five prominent mucronate teeth, 0.5-1.0 mm
long, the whole with minute raised glands; corolla
funnelform-campanulate, exserted 45 mm above
level of calyx lip, 3—4 mm wide at calyx mouth,
15-20 mm wide at mouth, glabrate with short hairs
in region of ovary; corolla lobes 20-22 Х 15-20
mm; stamens 15 or 22 mm long, shorter filaments
inserted 1 mm and longer filaments inserted 5 mm
above the level of the calyx lip, the staminode 5—
6 mm long, inserted at the level of the calyx lip,
anthers included; ovary 3 mm long, sparsely to
densely lepidote, style 32 mm long, included. Cap-
sule elongate-linear, 19-36 X 1.2-2.0 cm, many-
seeded; seeds flattened, 1-2 X 3—4 cm, oblong with
lateral membranous wings, each wing 1-1.5 cm
long, margin subhyaline, 5-8 mm long, body ovoid,
X 1.2 ст, not conspicuously bipartite, not well
differentiated from wings. Figure: Gentry (1977a:
fig. 10)
Cydista decora is one of only two Cydista species
restricted primarily or exclusively to South Ameri-
ca, where it inhabits two disjunct regions: (1) west-
ern Ecuador, and (2) portions of Bolivia, Paraguay,
and southwestern Brazil (Fig. 4). The two C. decora
populations differ in that the Ecuadorian collec-
tions possess conspicuously though somewhat in-
consistently bracteate inflorescences (Gentry,
1977b). The peak in flowering collections occurs in
March and appears to be correlated positively with
rainfall (Fig. 10), but collections of fruiting speci-
mens are too few to determine fruiting phenology.
The generic affinity of Cydista decora was un-
certain before fruiting material was known. Once
fruiting material was available, Gentry (1977b)
transferred it from the monotypic genus Clytosto-
manthus to Cydista and suggested that it may be
closely related to Cydista diversifolia, although it
differs, in part, by: (1) having larger flowers with a
narrower calyx, (2) producing larger inflorescences,
and (3) lacking glandular fields in the axils of the
secondary veins beneath (although scattered, iso-
lated glands are not uncommon). Cydista decora
has more prominent calyx teeth than all other spe-
cies in the genus.
Representative specimens. ECUADOR. Chimborazo:
Rfo Chanchan, from Naranjapata to below Huigra,
u
14468 (МО). Los Ríos: рате, 01?30'S, 79°25’ W, ~
m, Dodson et al. sane? е оси of A
Blanca, ca. 15 of Puerto Lépez, , 80°4 AT W,
90 m, Hekker & Hekking 10259 (MO, “OCA, U). BOLIV-
IA. La Paz: Alto Madidi, ide ca 7 km NE of camp,
13°35’S, 68?46'W, 300 m, Gentry
(MO). Santa Cruz: San Juancito, 30 kms №
nacio, 1548, $, 67°16'Ұ, 400 т, Вес
umba, Hoehne
22%43'S, 53710", 246 m, Ee а 5642 (MBM).
3. Cydista diversifolia (Kunth) Miers, Proc. Roy.
Hort. Soc. London 3: 192. 1863. Bignonia div-
ersifolia Kunth, in Humb., Bonpl. & Kunth,
Nov. Gen. Sp. Quarto ed. 3: 133; Folio ed. 3:
104. 1819. Pleonotoma diversifolium (Kunth)
Bureau & K. Schum., in Mart., Fl. Bras. 8:
274. 1897. TYPE: Mexico. Campeche: Hum-
boldt & Bonpland s.n. (holotype, P).
Anemopaegma vargasianum DC., in A. DC., Prodr. 9: 190.
1845. Cydista vargasiana (DC.) Miers, Proc. Roy.
Hort. Soc. London 3: 192. 1863. TYPE: Venezuela.
Caracas: Vargas s.n. (holotype?, G-DC).
Bignonia sagreana DC., in A. DC., Prodr. 9: 148. 1845.
TYPE: Cuba. Sagra s.n. (holotype?, G-DC; isotype?,
K).
Lianas, stems tetragonal, hollow in cross section,
drying light to dark brown with four distinct and
light-colored ridges, puberulent; —0 „а
liaceous, to 1.0 X 1.5 cm, elliptic-orbicular, р
entire. Leaves 8-12 cm long, once-pinnate Le two
rimary, opposite leaflets and often a simple, ter-
minal tendril; petioles 2-4 cm, sulcate, sparsely
Annals of the
Missouri Botanical Garden
lepidote, petiolules 1-3 cm, sulcate, sparsely lep-
idote; leaflets 5-8 X 4—7 cm, mostly ovate or ovate-
elliptical, sometimes orbicular, apices acuminate to
mucronate, bases truncate-rounded to slightly cor-
date, venation actinodromous (perfect, reticulate,
basal) basally and brochidodromous apically, with
3-5 major veins, brown glandular fields often in
axils of secondary veins, margins slightly undulate,
chartaceous. Inflorescences to 20 cm long, several-
flowered, peduncles 3.0—9.0 cm, the rachis and pe-
duncles minutely bracteate, the bracts 1 mm long,
pedicels 5 mm, sparsely lepidote-puberulent. Flow-
ers ovoid in bud, apices straight; calyx 4-5 X 5-6
mm, generally drying uniformly dark, margin intact
or shallowly split, apically truncate with five minute
teeth; corolla funnelform-campanulate, exserted 35
mm past calyx lip, 4-5 mm wide at calyx mouth,
15 mm wide at mouth, glabrous except at fusion of
filament and corolla tube where short, glandular
hairs reside; corolla lobes 20-22 X 15 mm; sta-
mens unequal, 13 or 16 mm long, inserted 1 mm
below level of calyx lip, the staminode 6 mm long,
inserted 1 mm below level of calyx lip, anthers in-
cluded; ovary 3 mm long, lepidote, style 18 mm
long. Capsule elongate-linear, 24-35 X 1.0-1.5 cm,
reddish brown with _ inconspicuous longitudinal
seeded; seeds flattened, 1.0 X
with lateral membranous wings, each wing 1.2-1.8
cm long, body ovoid, 1.0 X 0.6 cm, clearly darker
than wings, hyaline margin lacking.
Cydista diversifolia is a common liana, extending
from the coastal regions of central Mexico to the
Yucatán Peninsula and Cuba, and south through
the Pacific side of Central America (Fig. 4). South
American collections are restricted to central Co-
lombia and northwestern and north-central Vene-
zuela. Although it favors tropical dry forest, it may
also occur in premontane moist and more infre-
quently tropical moist forests (Gentry, 1973b).
Gentry (1974) reported that C. diversifolia pos-
sesses a multiple-bang pollination syndrome and,
like other species of Cydista, C. diversifolia appar-
ently does not produce nectar and probably relies
on deception of pollinators to effect pollen transfer.
en r of flowering collections peaks in July
or August (Fig. 11), and there are no significant
phenological differences between Central and
uth American collections. Unlike other Meso-
american Cydista species (C. aequinoctialis, C. het-
erophylla, and C. potosina), C. diversifolia initiates
its peak flowering period after the onset of the wet
season; flowering collections peak during the “ver-
nillo” or lull of the wet season. Fruiting collections
peak between November and January (Fig. 11)
Cydista diversifolia differs from other species of
Cydista in its hollow branchlets, a character not
previously reported for this species (perhaps be-
cause this character is readily detectable only in
cleanly cut branchlets). Venation in C. diversifolia
is nearly palmate, with the five principal veins join-
ing at the base of the blade.
Representative specimens. MEXICO. Campeche: 28
km e Escarcega, rumbo a Champoton, 18%37'N,
90°43'W, Cabrera et al. 2045 (MO). Chi
Angostura, 45 km from Tuxtla, 16°34’N, 92°48’W, 700 m,
Breedlove 37440 (DS, MO'). Guerrero: El Pusulmiche, 3
km del Entronque de la Brecha a Corinto, Mun. Tecpan
de Galeana, 17%15'N, 100%41'W, 60 m, Tenorio et al. 1348
(МО). Jalisco: La Huerta Est. de Invest., Exp. y Dif. Cha-
mela, 19%32'N, 105°05'W, Magallanes 3150 (MO). Mi-
choacán: 29 km al NE de Spe me 18?31'N,
0221", Soto Núñez 3644 (МО). Oaxaca: Cerro Ar-
enal, Tehuantepec, 16°20" N, 95°14’ W 1000 m, Mac-
Dougall s.n. (1971) (F, MO'). Quintana Roo: 6 km N де
Xel-Ha, 20°12'N, 87?20'W, Téllez & Cabrera 3229
(MEXU, МО). Yucatán: Chichén Itzá, 20°40'N, 88°34’ W,
Gentry 537 (MO). GUATEMALA. Chiquimula: between
Chiquimula and La Laguna, 14°48'N, 89°33’ W, 500-1000
m, Steyermark 30680 (F). Izabal: 25 km W of El Estor,
15°32'N, 89°21'W, Harmon & Dwyer 4319 (МО). Petén:
ong road оп N shore of Lake Petén, 16°59'N, 89°50'W,
Croat 24725 (MO'). BELIZE. Belize: W of Benque Viejo
near border with Guatemala, 17?06'N, 89%08'W, Gentry
8264 (F, MO'). Cayo: Xuanantunich, Maya ruins just NW
of Benque Viejo, 17%05'N, 89°08'W, Ugent 13 (MO, WIS).
Toledo: 2—4 ті. W of San José Rd., 16°26’N, 89°02’W,
00-366 m, Gentry 8188 (МО). HONDURAS. Cholute-
“ vic. of седење 13°18'N, 87?12'W, 20 m, Standley
4406 (Е). Comayagua: Los Manjas Station, 7 km N of
Сауан 14223'N, 87°37'W, 600 m, D'Arcy 18146
(MO). Copán: 4 mi. E of Copán, 14%50'N, 8909", 800
m, Croat 42501 (MO). Cortés: Quebrada El Encanto,
Lee ana la Cumbre, 15°30’N, 88°00’W, 200 m, Molina
94 (F). Morazán: Río de la Orillo, 14*30'N, 87%00'W,
: S of El Paraíso,
00 m, Molina 18422 (F). Valle:
veo de San ено; 13°30'N, 87°35'Ұ, 0 m, Molina
96 (F). Yoro: vic. of Coyoles, near Medina, 15°15'N,
ems) W, 200 m, Yuncker et al. ‚ги (Е). EL ЗАТУАРОВ.
Libertad: 6 mi. NW of La Libertad, 13°29’N,
89°19’ W, Wunderlin et al. 743 (MO). NICARAGUA. Boa-
co: 4 km al S de Boaquito, Fr E oes 12?26'N,
85°44’ W, 200 m, Moreno 18011 (МО). Carazo: between
Amayito and Barranco Bayo, 11°40'N, 86°18’ W, 30-100
B
—
1215 (МО). A Laguna
8
00 atagalpa: Las
Playitas de Moyua NE de Lago Moyua, 12°37'N, 86°04’ W,
А Rag ary d
а.
$" cil” до
мо NR
Volume 84, Number 4
1997
Hauk
Review of Cydista
475 m, Sandino € Guzmán 844 (MO'). Rivas: El Coyol
Tola—Las Salinas, 11?23'N, 85?58'W, 35 m, Stevens 9745
(МО). COSTA RICA. Alajuela: Llanos de Turrúcares,
09754" М, 83°33'W, Pittier 16436 (CR, US). Guanacaste:
Paloverde, OTS Field Station, 8 km W of Bagaces,
10°32'N, 85°18'Ұ, 100 m, Gentry et al. 71450 (MO). Li-
món: La Bomba—Cahuita, 09%44'N, 82?50'W, 20 m, Gó-
mez & Hampshire 20130 (MO). Puntarenas: Isla del
Caño, 17 km NW of Pta. Llorona, 08°37'N, 83^44' W, Jan-
zen 11641 (MO). San José: La Garita-Alajuela en pro-
piedad del ICE, 09%59'N, 84?20' W, Pontes & FcoCiccia
4172 (WIS). PANAMA. Canal Zone: Boy Scout Camp
road near Ma dden Lake, 09%15'N, 79°35) W, Gentry 2480
CUBA. Cienfuegos Central Soledad, 22°09’N, 80°27'W,
Howard 4785 (B, C, CLEMS, L, TEX-LL, MEXU, MI). La
abana: Sierra de Anafe, 22°55'N, 82°40'W, 200 =
Gentry & Hammel 71228 (MO). Las Vi ma
Bauao, 20?28'N, 76°22'W, León & Luna 21956 Tr
Matanzas: Playa on road from Buryvaea, 23%02'N
81734", Britton et al. 556 (CM). Oriente: Sabanaso,
20°48'N, 76^42'W, Ekman 7433 (AAU, B). Pinar del
Rio: Sierra de Anafe, Ekman 13029 (B, MO)
COLOMBIA. Antioquia: Currulao Nueva Antioquia,
11 km E-NE de Turbo, 08?06'N, 76°43’W, 45 m, Callejas
et al. 5011 (COL, MO). Atlántico: Usiacuri, camino de
Isabel López, 1045'N, 74?59'W, 100 m, Dugand & Gar-
cía-Barriga 2323 (COL). Bolivar: Mun. Cartagena, Isla
de Tierrabomba, EAR ur d W, 5-30 m, Cuadros
4338 (MO). Cesar Pto. Nacional, 10°22’N,
73708", 70 m, Uribe- Uribe 2409 (COL). Cundinamar-
са: 14 km S of Melgar, 04°12'N, 74°39’ W, 400 m, Gentry
et al. 8972 (COL, MO). Guajira: Maicao, near El Ocho,
11%23'N, 72713", Bunch 212 (MO). Huila: 17.5 km N
of Neiva, 03*02'N, 75°19’ W, 640 m, Croat 55260 (MO).
Magdalena: Fundación, Santa Rosa, 09*25'N, 74°05'W,
200 m, Romero- Castañeda 11055 (MO'). Norte de San-
tander: Río Peralonso, alrededores de Santiago, 07°52'N,
72°43'W, 120 m, Araque & Barkley 18NS104 (COL). San-
tander: 17 km NE of Socorro toward San Gil, 06°29'N,
73°16'W, 1250 m, Gen
Corozal, Palmitos, 09?19'N, 75°
9356 (COL). Tolima: Flandes, Finca 5
04'18'N, 74?49' W, 350 m, {жилы et al. 158 (МЕР). Val.
le: Cali, 03°27'N, 76°31'W, 900 m, Forero & Нета em
1555 (MO). VENEZUELA. Ar ejeria-La Encru
јада, 10%15'N, 67°10'Ұ, preci 4311 (МО, VEN).
Carabobo: Carretera de Maracay a Valencia, 10%05'N
68*05'W, 400 m, Williams 12193 (VEN). Cojedes: Via
hacia Tinaco, cerca puente los Monos, 09%42'N, N, 68°26’ W,
retera Carenero-Chirimena, 2 km NW
10732'N, 66%07'W, 0-5 m, Steyermark € Bunting 102300
(NY, VEN). Portuguesa: 20 km al SO de Guanare,
09%03'N, 69*45' W, 200 m, Aymard & Ortega 2844 (МО).
Táchira: Matorrales, 07°52'N, 72°19’W, 1000 m, Bono
5044 (MO). Trujillo: El Cenizo, 09°25'N, 70°30'W, 30
m, Lasser 2832 (VEN). Yaracuy: Hacienda Гоа, near
Guama, 10°20'N, 68745", Pittier 11158 (VEN US). Zu-
ia: Оно. Mara, Guasare-Socuy, 10%52'N, 72°29’W, 100
m, ‘Bunting 10319 (МО).
4. Cydista heterophylla Seibert, Publ. Carnegie
Inst. Wash. 522: 417. 1940. TYPE: Mexico.
Yucatán: Lundell & Lundell 7350 (holotype,
MICH; isotypes, A, F, US).
Bignonia lepidota Seem., Bot. Voy. Herald 179. 1854, non
Kunth. TYPE: Panama. Isla de Iguana: Cuming
1262 (isotype?, K).
Lianas, stems terete, solid in cross section, dry-
ing gray to brownish, inconspicuously lenticellate,
glabrous; pseudostipules inconspicuous. Leaves 11—
17 cm long, once-pinnate with two primary, oppo-
site leaflets and often a simple, terminal tendril;
petioles 2–3 cm, sulcate, sparsely puberulent, pet-
iolules 2-3 cm, sulcate, sparsely puberulent; leaf-
lets 7-12 X 4–7.5 cm, ovate to ovate-elliptic, api-
ces acuminate to acute, bases obtuse or rounded,
venation actinodromous basally (three principal
veins sil اھ and brochidodromous apically, 3—
4 vein pairs, glandular fields in axils of secondary
veins és сном and infrequent, margins slight-
ly undulate, chartaceous. Inflorescences to 14 cm
long, several- to many-flowered, peduncles 0.3-1.0
cm, the rachis and peduncles minutely bracteate,
bracts 1 mm or less, pedicels 11 mm, lepidote.
Flowers ovoid in bud, apices straight; calyx 4-5 X
4–5 mm, generally drying uniformly dark, margin
shallowly split, sometimes appearing bilabiate,
teeth lacking, densely lepidote; corolla funnelform-
campanulate, exserted 40 mm above level of calyx
lip, 2-3 mm wide at calyx mouth, 15 mm wide at
mouth, glabrous; corolla lobes 15 X 15 mm; sta-
mens unequal, 11 or 16 mm long, inserted 5 mm
above level of calyx lip, the staminode 3 mm long,
inserted 4 mm above level of calyx lip, anthers in-
cluded; ovary 3 mm long, lepidote, style 28 mm
a included. Capsule elongate-linear, 20-35 X
2.0 cm, brown with minute dark glands and
two conspicuous darkened submarginal ridges,
many seeded; seeds 1.5 X 7 cm, oblong with lateral
кран wings, 3.0-3.5 cm long, becoming ћу-
e 5-8 mm from margin, body ovoid, not well
poa from wings, not obviously bipartite
but the dark midline extending % length of body.
Figures: Gentry (1973b: fig. 10, p. 841), Gentry
(1982: fig. 10, p. 101).
Cydista heterophylla is a common liana of trop-
pe dry forests and drier areas of moist forests, oc-
urring less commonly in moist regions (Gentry,
19735). Its range extends from central Mexico to
Annals of th
Missouri E si Garden
the Yucatán Peninsula, south through Central
America, and to the extreme northwestern portions
of Colombia (Fig. 5).
Flowering is initiated prior to the onset of the
wet season, and flowering collections peak in May,
after which numbers of flowering collections de-
crease markedly (Fig. a Fruiting collections peak
slightly from November to February (Fig. 12).
Seibert (1940) distinguished Cydista heterophyl-
la by its conspicuously bilabiate calyces and pro-
duction of flowers during the dry season while es-
sentially leafless. Gentry (1973b) reported that C.
heterophylla has “a propensity to simple leaves,”
and that the leaflets possess “а basal pair of arcuate
veins with glandular fields in their axils.” Many
plants, especially juveniles, often exhibit two ter-
minal pairs of simple leaves that mimic a whorl of
four (Gentry, 1973b)
Representative specimens. MEXICO. Campeche: El
Tormento, 5 km W of Escarcega, 18°37'N, 90°43’ W, Held
et al. FC86 (U). Chiapas: 5.6 mi. E of Chiapa de Corzo,
16°42'N, 93°00'W, 833 m, Breedlove 9559 (DS, Е
. Guerrero: Zihuatenejo, rumbo Isetla,
17?38'N, 101%33'W, Ge
Puerto Vallarta, 20°37'N, 105?15'W, 1
(МО). Nayarit: 6 mi. E of San Blas, 22^52'N, peo W,
dp 242- 73 (MO). Oaxaca: Mun. Matías Romero, 8
de Esmeralda, 17%06'N, 94°48'W, 150 m, Wendt et
at 4809 : tana Roo: En los alrederores de el
cruz: Ban
IE езт", 2 а
Petén: Santa Ele ni;
Y
Id
P|
8907", љета P al. 3152 (МО). Pre" e Walk: Indian
Church, 17* ^40'W, Arnason & "E Linker 17341
(МО). Жеш Creek: Stann ба Railway, 16°58'N,
88°13'W, 33 m, Schipp 224 (F, MICH, МО). Toledo: 2—
4 mi. W of San José road, 16^15'N, 89°02'%, ac
m, Gentry 8192 (MO). HONDURAS. Choluteca: 5
NW sh San riis 13*33'N, 87^16' W, Lent 598 (MO)
Comayagua: a, San Louis, 14°25’N, 87°37’ W, 833
m, Edwards P-597 E MICH). Copan: Entre San Nicolas
y Trinidad, carretera Santa Rosa de Copán, 14%57'N,
88745", 100 m, Molina 11725 (F). Cem e rod
radia y Montana Cusuco, 15°30’N, 88°00'W,
lina 7302, (F). Morazan: Zamorano, PAM ВТО" у
m, Molina 115 (F). Yoro: Entre Yor
15°15'N, 87°15'W, 700 m, MNT 6925 (F). ÉL: SALVA:
DOR. La Unión: Carretera guna de Olomega,
13°19'N, 88°04'W, Lagos 625 MO NICARAGUA.
5
22435 (МО). Granada: Casa Tejas, 11^46'N, 85°54’ W,
40 m, Moreno 17090 (МО). León: Quebrada of Las Rue-
das, NW of El Tránsito, 12%05'N, 86°43’W, 15-30 m, Ste-
vens et al. 20138 (MO). Managua: Carretera a Monteli-
11°49'N, 86°31'W, 10 m, Guzmán et al. 438 (MO).
asaya: P. N. Volcán Masaya, 11%59'N, 86710", 300 m,
Neill 3135D (MO). Matagalpa. Puertas Viejas, 1-2 km
camino a San José de los Remates, 12%35'N, 86"01'W,
430—470 m, Moreno 16263 (MO). Nueva Segovia: El
Jícaro, Casa Viejas, 13°44’N, 86%05'W, 600 m, Moreno
13505 (MO). Rivas: Isla Ometepe, Volcán Maderas, Mé-
rida, 11°27'N, 85°33'W, 240 m, Robleto 3978 (MO?. Ze-
aya: Kurinwacito, 13%08'N, 84^55'W, Moreno 2397.
(MO). COSTA RICA. Alajuela: between Caldera and Or-
otina near Sitio Huacas, 09°54'N, 83°33’ W, 150 m, Gentry
et al. 71439 (MO). Guanacaste: Parq. Nac. Santa Rosa
road to Estero Real, 10%50'N, 85%35'W, 150 m, Huft et
al. 2103 (MO). Puntarenas: Barranca от Му of
Miramar turnoff, 10%06'N, 84°44’W, Gentry 2 (MO).
PANAMA. Canal Zone: i Colorado oar 09.11N,
79.57W, Croat 8206 (МО, SCZ). Darién: El Real,
08°08'N, 77°43'W, Gentry 4575 (МО). Herrera: 12.5 mi.
E of Las Minas, 07°48'№, 80°44’ W, Gentry 3143 (MO).
Panamá: Río Espavé, 09°14’N, 78°46'W, Gentry 3700
~
(МО).
CUBA. La Habana: 23*08'N, 8222", Sagra 89 (MO,
COLOMBIA. Bolivar: Mun. Turbaco, La Cantera de
Cimaco, near ее 10*20'N, 75225" М, 100 m, Gen-
try et al. MO). Sucre: Estación de Primatos,
09930"М, 75°30’ W, 300 m, Gentry & Cuadros 68146 (MO).
5. Cydista lilacina A. H. Gentry, Mem. New York
Bot. Gard. 29: 277. 1978. TYPE: Venezuela.
Bolívar: Gentry, Morillo & de Morillo 10673
(holotype, MO'; isotypes, МО’, ۴
Lianas, stems terete, solid in cross section, gen-
erally drying dark brownish green to dark gray, gla-
brous; pseudostipules inconspicuous. Leaves 20-25
cm long, once-pinnate with two primary, opposite
leaflets and frequently a simple, terminal tendril;
petioles 2.56 cm, inconspicuously sulcate, lepi-
dote, petiolules 1.54 cm, conspicuously sulcate,
lepidote; leaflets 11-19 X 5-14 cm, mostly broadly
ovate-elliptical, occasionally ovate, apices obtuse-
mucronate to acuminate, bases obtuse or rounded,
occasionally oblique, venation actinodromous ba-
sally and brochidodromous apically, 4—5 vein pairs,
midrib and secondaries prominent abaxially with
dark confluent or isolated glands in axils of sec-
florescences to 19 cm long, several-flowered, pedun-
6.0-11.5 cm, the rachis and peduncles mi-
nutely bracteate, pedicels 5 mm, lepidote. Flowers
ovoid in bud, apices curved; calyx 6-7 X 4.5 mm,
drying uniformly dark, margin shallowly split, teeth
five and minute, Reyes ciliate with uniseriate
hairs; corolla exse mm above level of calyx
lip, 2-3 mm wide at јеж tip, 15 mm wide at
mouth, stalked glandular hairs overall; corolla lobes
Se, A — — — — — — —
Volume 84, Number 4
1997
Hauk
Review of Cydista
833
20-22 X 15 mm, irregularly elliptic; stamens un-
equal, 13 or 20 mm long, inserted 1 mm below
level of calyx lip, the staminode 5 mm long, in-
serted 1 mm below level of calyx lip, anthers in-
cluded; style 25 mm long, included; ovary 3 mm
long, lepidote, ovules 4-seriate. Capsule elongate,
16-32 X 3-4 cm, surface wrinkled, many-seeded;
seeds flattened, 1-2 X 4—5 cm, elliptic-oblong with
lateral membranous wings, each 1.5-2.0 cm long,
hyaline margin 3-5 mm, body ovoid, not clearly
distinct from wings, bipartite with distinct longitu-
dinal ridge. Figure: Gentry (1978: fig. 125).
Cydista lilacina is one of two almost exclusively
South American Cydista species; a singl i
from Costa Rica is known (Fig. 6). Collections are
infrequent and distributed throughout South Amer-
ica north of Paraguay. Cydista lilacina is relatively
common in only two regions: (1) northern Bolivia,
eastern Peru, and western Brazil, and (2) eastern
Venezuela. Flowering collections are few and ap-
pear to peak in October (Fig. 13). If this peak rep-
resents the true peak in flowering for the species,
С. lilacina would be the latest-flowering species of
the genus (precipitation was not plotted with phe-
nology data because the paucity of and wide dis-
tribution of fertile collections precludes compari-
sons to any localized region).
Cydista lilacina differs from all other Cydista
species in its 4-seriate ovules and curved bud api-
ces. Capsules of С. lilacina and С. potosina are
generally wider than those of all other species.
However, the wrinkled surface of Cydista lilacina
capsules differs from the smooth surface typical of
capsules of C. potosina (Gentry, 1978). Vegetatively,
C. lilacina resembles wide-leaved specimens of C.
aequinoctialis but possesses round rather than sub-
tetragonal branchlets.
Representative specimens. COSTA RICA. Puntaren-
as: Reserva Biologica Carara, 09*46'N, 84°29'W, 20 т,
Zúñiga 90 (CR, MO).
COLOMBIA. Bolívar: Santuario Nacional de Los Co-
lorados, Mun. San Juan Nepomuceno, 09°58'N, 75710",
230-250 m, Gentry et al. 60654 (JBGP, MO). ECUADOR.
Pastaza: 2 km del pueblo de Villano, 01*25'5, 77°20'W,
400 m, Траг et al. 485 (МО). PERU. Huánuco: carretera
nández et al. 151 (МО, USM). Loreto: Ditto. Calleria
Bosque Nac. Alex. Humboldt, 250-300
(AAU, MO). San Martín: 07°11'S, 76°33’ W, 350 m, Gen-
44983 (МО). Ucayali: Bosque von Hum-
^00 50 m, Gentry et al. 29527
of Río Enlatagua, 13235'5, 68°46’W, 280 m, Gentry &
19589 (MO). SURINAME. Nickerie: Kabalebo Dam
area, 03?34'N, 55°59’W, 30-130 m, Lindeman & Roon
741 (МО). VENEZUELA. Amazonas: Dept. Atures, San
Juan de Manapiare, 05?18'N, 66°03’W, 150 m, Huber
1056 (MO'). Barinas: Carretera La Yuca—Barinitas,
08°45'N, 70?25'W, Marcano-Berti et al. 112-980 (IPA,
MBM, МО, U). Bolívar: 2-15 km E of La Paragua-Ciu-
dad Piar hwy., on road to San Pedro, 06°50'N, 63°20'W,
300 m, Gentry & Berry 15063 (MO. Delta Amacuro: E
of El Palmar, 08?20'N, 61?40'W, Gentry & Berry 14987a
taro, 10%43'N, 62
121011 (MO). BRAZIL. Acre: on Río Branco Santa Rosa,
0814'S, 73°13'W, Lowrie et al. 440 (МО). Amazonas:
basin of Río Jurua, near mouth of Río Embira, 07°30'S,
70°15'W, Krukoff 5046 (MICH, MO', US). Maranhao: St.
Luzia, Reserva Florestal de Buriticupu, 03°08'5, 42°54’ W,
dos Santos et al. 70 (MO). Mato Grosso: Barra до Bug-
res, entre Denise e Arenapolis, 15%05'S, 57°11'W, Saddi
4323 (CH). Minas Gerais: W side of Río Aripuana, belo
Salto dos Dardanelos, 20°41'S, 46715", Berg et al.
P18531 (CH, MO). Pará: Parauapebas, Reserva Biologica
da Serra dos Carajas, 07°45’S, 51°30'W, 600 m, dos San-
tos et al. 218 (MO). Rondénia: Río Jaru, estrada Porto
Velho-Cuiaba, 10°05’S, 61759", Duarte & Appa 7038
(MO', RB, INPA). Roraima: SEMA Ecological Station,
Ilha de Maraca, 03?22'N, 61°25'W, Ratter et al. R5373
MO).
~
6. Cydista potosina (K. Schum. & Loes.) Loes.,
Herb. Boissier 3: 618. 1895. TYPE: Mexico.
San Luis Potosf: Seler 616 (holotype, B not
seen by Gentry).
Clytostoma mayanum Standl., Carnegie Inst. Wash. Publ.
461: 86-87. 1935. TYPE: Guatemala. Petén: Yaxha—
Remate Road, Lundell 4008 (holotype, F; isotype, 5).
Lianas, stems tetragonal, solid in cross section,
drying gray with four distinct and often light-col-
ored ridges, glabrous; pseudostipules linear-trian-
gular, numerous, overlapping, usually less than 8
mm long. Leaves 6-14 cm long, once-pinnate with
two primary, opposite leaflets and often a simple,
terminal tendril; petioles 14 cm, inconspicuously
sulcate, puberulent, petiolules 14 cm, sulcate, pu-
berulent; leaflets 4-10 X 2-5 cm, mostly ovate to
ovate-elliptical, apices acuminate, bases rounded to
obtuse, venation brochidodromous, 4-5 vein pairs,
veins puberulent-pilose beneath, solitary glands in
axils of secondary veins occasionally present, mar-
gins slightly undulate, chartaceous. Inflorescences to
10 ст long, ca. 4—8-flowered, peduncles 0.5—8.5
cm, the rachis and peduncles minutely bracteate,
pedicels ca. 5 mm, lepidote. Flowers ovoid in bud,
apices straight; calyx 2-7 X 5-6 mm, apically trun-
cate with five minute teeth, drying dark basally and
lighter apically, lepidote with dense aggregations of
hair; corolla funnelform-campanulate, exserted ca.
Annals of the
Missouri Botanical Garden
35 mm above level of calyx lip, 2-3 mm wide at
calyx mouth, ca. 15 mm wide at mouth, lepidote;
corolla lobes 16 X 20 mm, irregularly elliptical;
stamens 5, unequal, 10 or 17 mm long, filaments
paired, distinct, adnate to corolla tube, shorter fil-
aments inserted 2 mm and longer ones 5 mm above
level of calyx lip, the staminode 3 mm long, in-
serted at level of calyx lip, anthers with two spread-
ing thecae, subterminal on the filament; disk want-
ing, ovary 3 mm long, cylindrical, lepidote, style
ca. 30 mm long, included, stigma bipartite, divi-
sions laterally flattened. Capsule elongate, 20-24 X
2.8—3.2 cm, drying dark, surface smooth, many-
seeded; seeds flattened, 1.5 X 6 cm, oblong with
lateral membranous wings, each 1–2 cm wide, fre-
quently unequal, margin hyaline, 1 mm or less,
body ovoid, not obviously bipartite but with dark
medial ridge extending % length of seed
Cydista potosina (Fig. 1) has the most restricted
distribution of all Cydista species (Fig. 6); it ranges
from the southeastern half of central Mexico to the
Yucatán Peninsula and Belize, with sporadic col-
lections from Guatemala, Honduras, El Salvador,
and Nicaragua. A single collection is known from
Costa Rica. Gentry (1982) reported a “multiple-
bang” flowering phenology in C. potosina. Flower-
ing collections peak in June and decline sharply
by August (Fig. 14). Like C. aequinoctialis and C.
heterophylla, C. potosina appears to initiate flow-
ering before the onset of the wet season. Fruiting
collections are evenly distributed from September
to March (Fig i
Cydista potosina differs from all other Cydista
species in its distinctive, bicolorous (when dried),
ciliate-margined calyx. Cydista potosina and C. lil-
acina have capsules that are wider than those of
other Cydista species. The branchlets of C. potosina
and C. decora are tetragonal and frequently dark
with four light-colored, longitudinal ridges. The leaf-
let venation of C. potosina more closely resembles
the uniformly brochidodromous type of Clytostoma,
rather than the basally actinodromous type of many
Cydista species. The pseudostipules of C. potosina
are reminiscent of the bromeliad-type clusters typ-
ical of Clytostoma, which further suggests a rela-
tionship between the two genera. However, the
smooth, lepidote ovary and smooth fruit of C. po-
tosina indicate a closer affinity to Cydista than to
Clytostoma.
агонии specimens 20. Campeche: Но-
Wendt et а 3618 (МО). Puebla: М. Sn. Diego,
18%50'N, 98200", Ramírez 258 (MEXU). Quintana
Roo: 1 km NW of Puerto Morelos, 20%50'N, 86%52'W, 5
m, Davidse et al. 20022 (MO'). San Luis Potosi: San
Antonio, La Ladrillera, 22°08'N, 100°59’W, 300 m, Alcorn
3199 (МО). Tabasco: Cerro las Campanas, З km E of
Teapa, 50 km S of Villahermosa, 17°33’N, 92°57’ W, 50-
100 m, Conrad & Conrad 2813 (MO', MEXU). Veracruz:
Cerro El Vigía Estación de Biología Tropical Los Tuxtlas,
450 m, Sinaca-Colin 144 (MO). Yucatán: Miei village
8 km from Xuilub, 25m, Mogensen 1038 (MO
MALA. Escuintla: slopes of the Río Cóyolite; 13?57'N,
91*19'W, Harmon 2379 (MO). Izabal: across bay from
Puerto Barrios, 15?43'N, 88°36’W, 20-50 m, Steyermark
39853 (F) Petén: Remate, 13 km NE of Village,
17%00'N, 89%42'W, Contreras 927 (MO', TEX-LL). BE-
LIZE. Belize: W of Spanish Lookout, road to ferry,
17°13'N, 88?59'W, Dwyer et al. 461 (MO). Cayo: near
Teakettle, 17°14’N, 88°51'W, Whitefoord 3317 (МО). Co-
rozal: 18°24'N, 88:24", Gentle 181 (MICH). Stann
Creek: Cockscomb Mts, 16?48'N, 88?37' W, 100-166 m,
Gentry 8001 (MO. Toledo: Southern Hwy., 14 mi. N of
Punta Gorda-San Antonio, 16?15'N, 89°02’W, 100-166
m, Gentry 8228 (F, MO). HONDURAS. Atlántida: E of
Tela near Yoro Trail, 15?44'N, 87°27'W, Yuncker 4973 (F,
MICH, MO). Yoro: near the village of Los Flores,
15?15'N, 87°15'W, 950 ft, Yuncker et al. 8163 (МО).
NICARAGUA. hostales: 2.6 km NW of Cuapa,
12°16’N, 85?23' W, 500 m, Stevens & Montiel 17441 (MO).
Jinotega: 21 km del Valle del Cua, NW de El Cedro,
13730'N, 85°38’ W, 700 m, Moreno 944 (МО). Nueva Se-
govia: Río Diplito, E de Ciudad Ocotal, 13°37'N,
86°27'W, Sandino 2995 (МО). COSTA RICA. Puntar-
enas: Parque Nacional Corcovado, Pavo Forest, 08°27'N,
8333", 0-150 m, Kernan 573 (CR). EL SALVADOR.
uachapán: El Imposible, cerca de los Enganches,
13*52'N, 89759" W, Sermeno 175 (МО). La Libertad: Ha-
cienda La Argentina, 13%40'N, 89?20'W, 500 m, Montalvo
4267 (ITIC).
DuBious OR REJECTED NAMES
Bignonia variabilis Seibert ex E. Mey., Nova Acta
Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur.
12: 779. 1825. Pro syn., non Jacq. (1797).
Bignonia incarnata Aubl. sensu Splitg., Tijdschr.
Natuurl. Gesch. Physiol. 9: 7. 1842, non Aubl.
(1775).
Cydista amoena Miers, Proc. Roy. Hort. Soc. Lon-
don 3: 191. 1863. Nomen nudum.
Cydista analoga Miers, Proc. Roy. Hort. Soc. Lon-
on 3: . 1863. Nomen nudum.
Cystida blanda Miers, Proc. Roy. Hort. Soc. London
3: 192. 1863. Nomen nudum.
OPES eximia Miers, Proc. Roy. Hort. Soc. London
: 192. +863. Nomen nudum.
: Linus incarnata sensu Miers, Ptoc. Roy. Hort.
. London 3: 192. 1863. Nomen nudum.
Cydista priscis Miers, Proc. Roy. Hort. London
Soc. 3: 192. 1863. Nomen nudum.
Cydista орно Miers, Proc. Roy. Hort. London
Soc. 3: 192. 1863. Nomen nudum.
mM" A MN аи
pm АНИ
Volume 84, Number 4
1997
Hauk 835
Review of Cydista
Cydista seemannii Miers, Proc. Roy. Hort Soc. Lon-
don 3: 192. 1863. Nomen nudum.
Literature Cited
Descourtilz, M. E. 1822. Fl. Méd. Antilles, i 00.
O (Food and Agriculture Organization of dá United
Nations). 1985. Ааваас Data for Latin
America and the Caribbean, Rome.
түз pá T AN. ге & М. до С. Pinto. 1990. NBS
чү new пучини type reaction in the chemistry of
quinones. Anais Acad. Brasil Ci. 62: 329-333.
Gentry, A. H. 1973a. Generic delimitations of Central
6-242.
; oodson, Jr. &
R. W. Soay: угине Flom of Paddle Ann. Missouri
Bot. eris 60: 781–97
1974. Flowering phenology and diversity in trop-
ical Bighoiüiscese: Biotropica
. 19 Fam. 178. руа In: С. Harling
& B. е Flora of Ecuador. Opera Bot. 7: 1–173.
977b. New species of Leguminosae, Lauraceae,
and Monimiatedé: and new combinations in Bignonia-
ceae ve western Ecuador. Selbyana 2: 39-45.
8. The botany of the Fecha pce Big-
noniaceae. Mud New York Bot. Gard. 29: 245-283.
982. Bignoniaceae. /n: Flora ^ denar In-
Айшә Nacional de Investigaciones sobre Recursos Bió-
ticos. Xalapa, Veracruz, Mexico
— ———. 1992. А synopsis of Bignoniaceae ethnobotany
and economic botany. Ann. Missouri Bot. Gard. 79: 53—
. Bignoniaceae. Pp. 403491 in J. A. Stey-
örin; E E: e B olst, Flora of the Vene-
zuelan Guayana, Vol. 3. Missouri Botanical Garden, St.
Louis
& А. 5. Tomb. 1979 [1980]. Taxonomic implica-
tions of Bignoniaceae palynology. Ann. Missouri Bot.
rd. 66: 756-777.
Goldblatt, P. & A. H. Gentry. 1979. Cytology of Bignoni-
aceae. Bot. Not. 132: 47 2.
Loesener, Th. 1919. Mexikanische und zentralamerikan-
Bignoniaceae. Repert. Spec.
Macbride, J. F. 1961. Bignoniaceae. Pp. 3-103 in: Flora
of Peru. Field Mus. Nat. Hist., Bot. Ser. t XIII, Part
E
o.
Miers, Ј. 1863. Report on the plants collected by Mr. bs
especially the Bignoniaceae. Proc. Roy. Hort Soc
202.
don 3: 179—
Rohrhofer, J. 1931. Morphologische Studien an den Stam-
inodien der Bignoniaceae. Oesterr. Bot. Z. 80, t. 3, figs.
Већина, F. J. 1792. Abb. Baume, Staud. Strauch. Oesterr.
E Qk
Seibert; R. J. 1940. Botany of the Maya area: Miscella-
neous papers XXI, The Bignoniaceae of the Maya area.
Carnegie Institution of Washington, Publication No.
522: 375-434.
948. The use of glands in a taxonomic consid-
eration fd the family Bignoniaceae. Ann. Mi ssouri Bot.
Gard. 35: 123-136.
Urban, I. 1916. Uber Ranken und poe der Bignoni-
aceen. Ber. ge Bot. Ges. 34:
Venkatasubban, K. R. 1944. Clg чы sts in Big-
noniaceae. нане Univ., Annam
INDEX TO EXSICCATAE
Collections are listed alphabetically by the principal
ed by collection number, and a boldface
number (1-6) designating the species collected. All spec-
imens entered into TROPICOS мег h
been examined by A. H. Gentry. Spec
the author were limited to duplicates housed a , and
are indicated by a “!” i
nd
Index to Exsiccatae
is as follows: la = Cydis
tialis, 1b = Cydista aequinoctialis var. hirtella, 2 = Cy-
dista decora, З = Cydista diversifolia, 4 = Cydi
erophylla, 5 = Cydista lilacina, and 6 = Cydis
Abbott 1446 la; Acevedo 2810' la, sni la, 3443 la;
Acuna et al. 24151 3; Agostini & {Аме 1052 1а ; Agui-
lar 259 4, 40 6, s.n. З; Alain 1417 3; p жыз 2172: 6;
Allen 127 la, 173 Ла, 3250 Та, 1756'4, 1964 Та, 2458
IM 2503 Та, 2641 Та, 3646 Та, 6520 la, 6589 3, 858'
Та; Almeda et al. 3115 3; Alston 5609 Та, 8815 1, 8865
4, 88654 Та; Alvarez et al. 812' Та; Amaral et al. ЗАГ
Та; Americo 12671' Та; Ansouin 621 3: Antonio 4241 Та;
Apolinar 383 З; Araquistain 110' 3, 261' 3, 2973 3, 306
3; Araquistain & Moreno 530' 3; Archer 8333 la; Ariste-
guieta 2259 Та, 4312 3, 4313' 3, Mod la, 6438' 3; Ar-
nason 1773% 3; Arnason & Lam 17710' 3; Arturo &
Magallanes 3150 3, 3563 Та, 828 T Arvigo 133' 3; Ar-
. 557 4; Atwood & Neill AN207 3; Aubréville
7326' Та; Aviles X-1' Та,
Та; Ayala 2376' Та, 2504 la, 3717 la; Aymard 3395 1а;
Aymard & Delgado 8454' Та; Aymard & Опера 2844' 3;
унше et al. 10212 5, 370' 1a, 3942' la.
Bach s.n. MG4157 Та; Bachbett s.n. 6; Badillo 2024
MV la; Bailey 227 3; Bailey & Bailey 132 la, 610 Та;
Balee 1981 5, 2596 5; Balick et al 3153 6, 1488' la,
3152: 4; Balsev 97298 Та; Balsev et al. 97298 Та; Ban-
gham 422 Та, 605 Та; Barbier s.n.' Та; Barclay s.n. la;
Barkley et al. 17M755 3: Barlow 32718 6, 32118A 6; Barr
& Mason 62-327 Та; Barrier 4027 Та; Bartlett 12439pp
6; Stergios & Delgado 12893 Та; von Bayern 332 M 1a;
Beck 3798: Та; Beck et al. 20381' 5; Benitez de Rojas 475
nardi 849 la, 1500 Та, 19222' 2, 19506 2, 19582 2,
2553: 3, Bernoulli & Cario 2056 la; Berry 1036' Та,
1080 Та, 1588: 5, 922' Та, 1185 la, 1205' Та; Beuther
102:3; Billiet & Jadin 1060 la, 1791 la, 4814 Та; Black
48-2937 la, 48-3435 Та, 48-8432 la, 49-8432 Та;
Black & Roi 48-3382 la, 48-3405 la; Black & Ма-
galhaes 51-12886 la; Black et al. 54-16530 Та, 54-
17579 la, 54-17623 la; Blackmore 3722 4, 4020 la;
Blanco 1285 1a; Вит & Dwyer 2130' 1a; Boege 3241 3;
Bond et al. 205 1a; Bonet 29 1a, 31 3; Bonpland 39416'
la; Boom 4758 la, 7156' Та, 8520' Та; Boom & Grillo
6148 Та, 6545' Та; Boom & Marín 10365 Та; Bourgeau
14071 la; ee & Asanza 31 405. la; Bravo 460 6;
Bredemeyer 180 W ; n. reedlove 24607
lb, 24694 1b, 26015: 3, 27312 la, 36550 3, 37262 3,
: 46 1b; Brenes 15567 4,
15598F 4, 15607 4, 22461 du Садр 4, 22464 ја,
Та, 51 55 Та; fer 58 la; edo
676 la, 7621' la, 7825 Та, s.n. la; Brunt 2244 6; Buch-
tien 1322 5; Bunting 11621 Та, 4297 Та, 5319' Та, 5333'
3, 5514A! la, 5514F la, 5620 3, 7866 3, 8817 la,
9019: 3, 9152' 1a, 9428' Та; Bunting & Alfonzo 8018' ће.
Annals of the
Missouri Botanical Garden
9783' 3, 9787 3; Bunting & Fucci 7991' 3, 7996' 3; Вип-
ting & León 12573 Та; Bunting & Licht 1286 Та; Bunting
& Licht 825 la; Bunting & Stoddart 9191 3; Burger &
Burger 7862 lb, 8461 la; Burger & Gentry 9167 la;
Burger & Stolze 4861 4, 4.
Cabrera 11453' 6, 4121'6, 4262! 4, Cabrera & a
15221 3; Cabrera et al. 2575' 6; Callejas et al. 4444
5036' Та, 5125' Та; Calzada 640 4, 788' Та, 978: 4; Cal.
zada et al. 6693 З, 6769 З, 6792 3; Camby s.n. Та; Camp
E-3812'2, E-3892 2; Canela s.n. Та; Cantin s.n. Та; Cár-
denas de Guevara 1532 3, 2156 la; Cardona 1134 la,
150 la, 164 la, 1669 Та, 2376 la, 4054' 3, 428 la,
Та, 919 Та, 920 la; Carlson 1508 1a; Carnevali & Lobo
' la; Carrasquilla & Taymes 140' la; Castellanos
26930 2; Castillo & Tapia 658' 4; Castillo 1484 3; Cav-
alcante 1901 Та, 444 la, 869 la, 901 la; Cedillo &
Torres 1089 Та; Chacón & Chacón 2031 15; Chaffanjon
153 la; Chaing 234 6; Chan 1145 3, 1393 3, 1823 3,
191 6; Chan & Burgos 1417 6, 1452 6, 336 4; Chan &
Ucan 839 6, 942 3; Chanek 201 6; Chavarria 1001 3,
189'3, 261' 3, 781'4, 825'3; Chiang 340'4, 726' Та; Cid
& Nelson 2936' 5; Cid et al. 110' la, 197 Та, 2025 Та,
507 Та, 5139' Та, 8467 la; Cisneros 81'3; Clewell & Cruz
4028' Ya; Colella et al. 1046 Та, 1048' Та; Combs 332' 3;
Сопвтур 120(40) Та; Сопігегаѕ 4364 4, 5859 3, se
8583 4, 900 4, 933'4, 965' 6, 9743'3, s.n. 4; Conz
et "al 3134 Та; Coradin & Cordeiro 651' Та; Córdoba 390
la; Cornelio 95 3; Correa & Escobar 1788' la; Cowan
3022' 6; Cowell 189 Та, 78 la; Cremers 5025 la, 5227
la, 6678 la, 6916 Та, 7193' la, 7194! la, 7270! la,
7276' Та, 7361' Та, 7533! Та, 7537 la, 7811 la, 8092!
‚ 8410 Та, 8445' la; Croat 10324' Та, 10798: Та,
spe la, 12268 Та, 12645' Та, 12796' Та, 12957 Та,
13487 Та, 14482' 4, 14499' 4, 15356' la, 25188: la,
42276' 3, 42506 З, 43672' 3, 43749 3, 4764 Та, 5119
Та, 5162' Та, 5208! Та, 5426' 4, 5439' Та, 55333! 3, 5589!
la, 5611' Та, 5700 Та, 6143' Та, 6155 la, 6169 la,
7176 Та, 8310' Та, 8552' 1а, 8755' la, 9514 la, 9566'
la, 9776' 4, 9846' Та; Стога! 300' Та, 856C' Та; de la
Cruz 1455' la, 1866' la, 1992! la, 3210' la, 3798' la,
4242' Та; Cuadros 1109'3, 1384: 3, 1683 la, ЗООбА: la
3597 Та, 4492' la; Cuadros & Gentry 30064 la, 3050
Та, 3083' la; Cuatrecasas 7491 la; Cuevas & Guz
©
2933 Та; Ситапа 397 Та; Cuming 1262рр 4; pret &
Haman 13 a.
Daly & Cardoso 3918' Та; Daly et al. 1406' Та, 4332!
Та, 7156' Та; Danin 76-14-4' 3; Darwin 2369' 3; Dauben-
mire 20) 3, 701] 4, 711] 4, 770] 4; Davidse 15318 la
- taz & Jaramillo 1515
Та, 1517 Ја, 1518 la; Diaz & rei 1526: la; Diaz
et al. 1037 la; Dodge 10715' la, 16570! 4, 3507 la;
Dodson & Gentry 9934' 2; Dodson 11520' 2; Dodson et al.
$E
N
Qo
2
"a
N
E
8
B
a
сл
~
en
>
; де Duchassaing 1851 Та; Ducke s.n
la; Dugand 1107 3. 434 la, 4553 3, 5034 3, 6052 Та,
6338 Та, 6412 Та; Dugand & Jaramillo 3278 Та, 3433
Та, 3434 la, 3435 Та, 3500 Та; Du gand & Seibert 1111!
3; Duke 10993 Та, 12268' Та, 14256 Та, 4696' la, 4922'
la, 4978' 1b, vine ies 553I' la, 9777 Та, 9700: Та;
Dwyer & bagi wt 14' 6, 10144App 6, 10214' 6; Dwyer
et al. 146 6
Ebinger 288 за, а Р-148 Та; Edward Р-359 3,
Р-597 4, P-630 3; Eggers 13121 3, 13128 3, 13546 3,
15070 2, 2631 la, 310B la; Egler & Irwin 46031 la;
Egler & Pires 47765 Та; Ek 648: Та; Ekman H11175 la;
Elias 1484 3, 540 3; Enriquez 558 4, 598 6, 739 3, 775
4; Erlanson 38 4, 485 1a; Ernst 1656 la, 1839 1a; Es-
pina 1016 3, 535 Та, 535 по Езрта & Mosquera 2115
Та, 2118 la; Eyerdam 886
Fager & Ribos 639 2; sts 1934 Та, 2028 Та;
Fernández 293: 3; Kotütndn & Susanna 8437 la; Fer-
nández Casas & Molero 6147 2; Fernández 307: Та, 359
3, 4127 Та, 6944' Та, 906' 5; Fernández-Alonso 8134 3;
Fernández-Pérez 20127 la; Feuillet 1583' Та; Fiebrig
4814 2; Fleury 631' Та; Flora Falcón 709 Та, 733! la;
Flores & Puch 9052 6; Flores & Yam 9142 3; Flores s.n.
6; Florschutz & Maas 2618 C Та; Foldats 9220! Та; Fol-
som 1653' Та; Fonnegra & Torrez 758 la; Fonseca 351
la; Fonseca & Onishi 1279' Та; Forero 1762 Та; Forero
933 la; Fournet 246 la; Frankie 408 lb, 441 3; von
Friedrichsthal 546 Та; Froes 1839 Та, 28509 Та, 30991
а, 3099la la, 31914 Та, 32347 la, 32537 la, 32729
la, 34466 Та, 34607 la, 34610' Та, 35607 Та; Froes &
Black 24517 Та, 24633 Та, 27486' Та, 27635 Та; Fryxell
T ii ode 3444' 6; Funck 294 la; Funck & Schlim 711
Qo s.n. la; Gandara & Dorantes 16' 4; García 51'
3; García-Barriga 10761 Та, 12166 Та, 13410 3, 13462
3, 6466 Та; García-Barriga €: Jaramillo 20561 Та; Gar-
cía-Barriga & Lozano а la; Garibaldi 235 Та; Gar-
| rnie A, 3; Сайы 1966: 3, 23818 3,
к=н 6. 24024 6, pee 3, 24298 3, 24345 6, 725pp
6, 751' 4; Gaumer et al. 725' 3, 795: 3; Gentle 1080 6,
3
SF
©
ce
а, 1604' Та, 16119' Та, 1619 4, 16281
Та, 1779 4, 1817 la, 19592 la, 2055 la, 2198 la,
2204 4, 2219 4, 2229 la, 2256 4, 2267 la, 2274 4,
2359 4, 24801' Та, 2562: Та, 2564 4, 2615 4, ae la,
28186' Та, 29508 la, 301 4, 306 3, 306A
40922 la; 40959 la, 4116 la, 41235 a 41326 5,
41685 Та, 4206 4, 431' Та, 4410! Та, 4420 4, 44983 5,
45101 5, 4539' 4, 4564 Та, 4580 Та, 46257 5, 46475
Та, 47302 Та, 47478' Та, 47492' 3, 47502 Та, 47618'
la, 47656 3, 482 4, 48611 Та, 4967 4, 4970! La, 50384'
la, end la, 51222' Та, 5126' 4, 51421 la, 537 3,
, 54102 la, 56373' Та, 5726 Та, 5758! la,
59285 2, pe 2, 60581 З, 60654' 5, 60737 Та, 61652
5, 62974' Та, 63360' Та, 63956' Та, 6494! Та, 6609 Та,
66161' 2, 68146 4, 68250 Та, 68253: 3, 68623 5, 68924
5, 69106 Та, 69145 Та, 69814 Та, 70274: э, 70503 5,
70713: 2, 70828: Та, 70878: 5, 710 la, 71485A la,
71952 3, 72062 3, 72132: 5, 72368: 2, 72633 2, 73008:
2, 73009 2, 73439' la, 7454: 4, 75599 5, 7585' la, 7597
la, 7620' 4, 766 4, 77391' Та, 7815 6, 7851' 4, 79290
Volume 84, Number 4
1997
Hauk
Review of Cydista
4, 79383: 6, 794 3, 79610 2 Та, 798 4, 812 Та, 815 la,
8150' 4, 8177 6, 826 la, 8262: 6, 8270 3, 829 3, 8311'
6, 8318 З, 8339 6, 836 la, 8360: 6, 841 la, 844 3,
847 3, 860 4, 863 3, 880 3, 881 3, 884 Та, 887 4, 888
3, 892 1а, 9404! Та, 9423' Та, 9429 Та, 9447 Та, 9744'
la; Gentry & Aronson 25049 Та; Gentry & Berry 14516'
Та, 14527 Та, 14698' Та, 14771'3, 14917 5, 14987 1а;
Gentry & Cuadros 47478 Та, 47492 3, 47656: 3, 63946
Та, 68253 3; Gentry & Curso de Розртадо 68623' 5,
68924' 5; Gentry & Daly 18914 Та; Gentry & Dwyer 3390
la, 4827' la; Gentry & Estensoro 70503 5; Gentry & For-
его 15359 Та, 15396' Та, 15460! 3, 15462' 3; Gentry &
Cuadros 47502' Та; Gentry & Gilly 10724 4; Gentry &
Jaramillo 57887 5; Gentry & Josse 72368 2; Gentry &
Josse 72633 2; Gentry & Juncosa 40959 la; Gentry &
Núñez 66031'5, 69610' Та; Gentry & Pinheiro 13096' Та;
Gentry & Puig-Ross 14247 Та, 14267 1а; Gentry & Puig-
Ross 14364: Та; Gentry & Ramos 12925' Та, 13339 Та;
Gentry & Revilla 16667 Та, 61119 Та; Gentry & Smith
45101' 5; Ge
Zardini 50384 la; Gentry et al. 10427 la, 10452' 5,
5, 10644 la, 10673' 5, 10737 5, 10922' la,
10978' 3, 11042: la, 11116! la; 11160' Та, 18077 Та,
18095: la, 18097: 3, 18155' Та, 18662' Та, 25213' Та,
25960! la, 27217 5, 32311' 6, 34799A' 4, 37593 la,
37648' 5, 38145 la, 51222 la, 51421' Та, 54049 Та,
5410' la, 56373 la, 60581' 3, 61652' 5, 63360 la,
69106' Та, 71485-а la, 73884' 2, 7526' Та, 76931' Та,
77759 Та, 8972: 3, 8973: 3; Gillespie 1198' la, 2125' Та,
2188' la, 2348' la; Gilmartin 663 2, 665 2; Ginés 4518
la, 5100 la, 5150 la; Glaziou 9951 la; Goeldi s.n.
MG398 Та; Goldman 1861 Та; Gómez-Pompa & Riba 196
6; González & Ortega 1358 Та; González 74 4; González-
Ortega 1176 Та; Gorts - van Куп et al. 478' Та; Grández
860!
et al. 10010 la, 9590 Та, 9650' Та, 9877 la, 9952' 1;
Grayum 4251 3, 4636' 4; Gregg 1033' Та; Grenand 2074
la, 2088: Та, 220 Та; Grewal & Persaud 128 1a; Grewal
& Persaud 179 1a; Grijalva 1024' 3, 1220' 3, 2151 3,
857 3; Guagliumi 13 3; Guánchez et al. 4216' la; Gudino
111% Та, 1351 Та; Guedes MG2293 la; Guedes 7 la;
Guillen 2404Z la; Guillet 711 la, 712 la; Guppy & Lin-
man 80 la; Gustafson 1450 3; Gutiérrez 25 3; Guzman
& Montiel 337 1b; Guzmán et al. 1079 3, 1104 3, 343:
93, 370.3, 382 3, 5758.
Haenke 1407 Та; Hahn 13292, 666 4, 3524 la; Hahn
et al. 1329 2, 1333: 2; Hammel 4579 Та; Hansen et al.
1549'6; Harley et al. 10760' Та; Harmon 2524 6; Harmon
& Dwyer 3826 З; Harris 1135' la, 1171' la; Harvard
Course in Tropical Botany 67 3; Hassler 10022 2, 10022a
2, 10432: 2, 5409 2; Hatschbach 46153' 2; Hatschbach &
Zelma 49084' 2; Haught 2249 la, 2295: 3, 3497 2, 3603'
la, 6577 3; Hayden 20' Та; Hayes 126'4, 412pp Та, 587
la, 590 Та; Hazlett 742 3; Heithaus 1 73 4, 300' 3; Heller
4447 la; Henderson 612: Та; Henderson € Ribamar 578'
la, 612 la; Henrich 108' 4, 181' 4; Henrich & Stevens
181 4; Hensen 150! 2; Herald & Clark 390 la; Heriberto
40 Та; Hernández 1308' 6, 310! 3, 358 1а; Hernández
Dorantes 1812' 4; Hernández 122 1а; Heyde 395 Та, 580'
; & Lindeman 107 5, 258' 5, 299 12; Higgins
203' Та, 207 la; Hinds 1841 Та; s.n. la; Hinton et al.
11732 Та, 16213 3; Hitchcock & Stanford 6901' 6; Hoff
5711' La; Hoff et al. 6614 Та; Hoffman 1226 la; Hoffman
& Benjamin 575' la; Hoffman et al. 739' Та; Hoffman-
nsegg s.n. la; Holm & Iltis 281 3; Holm & Пиз 669 Та;
Holm-Nielsen 19964 Та, 19967 la, 19989 Та; Holm-
Nielsen et al. 19441 Та, 19964 Та, 19967' Та, 19989' Та,
21304 1a, 19941' Та; Holst 2289 Та, 2305' Та; Holst et
al. 2305 la; Holton 604 3; Hopkins et al. 675' Та, 749'
la; Horner 115 la, 148 la, 189' Та; Horner et al. 115'
Та, 148' Та, 189 la, 244' Та, 363' Та; Hostmann 701 Та,
s.n. Та; Hostmann & Kappler 1051' Та; Huber s.n. Та, s.n.
МС1160 Ла, s.n. MG267 la, s.n. MG3826 la, s.n.
MG3836 Та, s.n. MG782 Та; Huertas & Hernández 4082
Та; Huft et al. 2113 1b; Humboldt & Bonpland 1078 Та,
л
Ibarra-Manríquez & Sinaca 1752' 6; Idrobo 1516 3,
1508 З; Idrobo et al. 1319 Та; Ijjasz-Madriz 478 Та; Iltis
27284a 3; Пиз & Nee 1469 Та; Irwin et al. 48056 la,
47425 Та, 47575 Та, 48055 Та, 48056 Та, 48789 Та,
48790 la, 53349 Та, 55529 Та, 55544 Та; Isert s.n. Та.
Jacquemin 2098: Та, 2344 la, 2543 la; Јаћп 1231
4, 12307 Та, 12318 4, 12321 la, 12329 4, 12335 1b,
12348 lb, 12352 1b, 12362 4, T-31 3, T-89 3, s.n. 4;
Jaramillo 219 3, 6889 1a, 6892 Та, 9070 Та; Jaramillo
$: Coello 2941' la, 4149pp la, 4227 Та, 4419pp Та,
4442 la, 4469 la; Jaramillo et al. 1214 la, 417 la;
Jaramillo-Mejía & Palacios 7870 1a; Jaramillo-Mejía et
al. 8061 la; Jardim 19257. 2, 24947. Та; Jiménez 1093
3; 655-34298 la, 3355 la, 508 la, 545 la, 587 la,
972 3, s.n. 4; Johnson 242-73 4; Johnston 812' 4, 831
4; Judziewicz 4565 Та.
Kant 1' 3; Keeler 171 3; Kelly 133 6; Kenoyer 533 Та;
Khan et al. 1226 Та; Killeen 54327, Та; Killip 33955 1а;
Killip & Dugand s.n. 3, 2465EPA 3, 37532 1a; Killip &
Smith 14079 3, 27481 la, 30469 la, 30569 Та; Killip
& Tamayo 37057 3; Killip et al. 38200: 3; Kimber 1576
la; King 635 1a; Kinloch 249 1a; Klug 4268' Та, 4310
la, 594 Та; Knight 69-87 Та; Koch 4908 3; Kral 25028
6, 69355: 3; Krebbs s.n. la; Krukoff 1784 la, 4942' la,
6482! Та, 6945' Та, 7068' Та, 8837 1а; Kuhlmann 1383
2: Kukle & Boom 60! Та; Kvist et al. 384 Та.
Langenheim 3242 3; Langlassé 137 Та, 506' Та, 739
P la; Larpin 958' Та; Lasseigne 4382' la; Lasser 4330
la; Lasser & Foldats 3008 Та, 3042 3, 3050 3, Laughlin
1631 3,871 1b, 872 4; Lawesson et al. 43463 Та, 44265
la, 44346 Та; Lazor 5460 Та; Leavenworth 205' 6; Lel-
linger & de la Sota 605 la; Lent 103 la, 1202 3; León
368: la, 18078 3, 3156 3, 6861 3; Levy 38 la; Lewis
1865! Та; Lewis et al. 1865 Та, 6533 la; Licata et al. 47
3: Liebmann 11 4, 58 6, 7986pp 4, 7992' 6, 8019 la,
8020' 6, 8026 4; Liesner 1369 la, 1 63654 5, 2243 3,
2342 la, 2458 3, 2800 3, 3386 3, 4427 lb, 4596 3,
4801 la, 4907 4, 5749 la; Liesner & Funk 16365 5;
Liesner & González 10695' Та, 11068' Та, 5816' la, 749
la; Liesner et al. 8201' 3, 8314: 3, 8321' 1а, 8442' 1а;
Lima 388-68 la; Lima 38868 Та; Linares 1316 3; Lin-
deman 6358 la; Lindeman et al. 96 5; Linden 219 4,
220 6; Liogier 19353 la, 9043-1 Та; Liogier et al. 33809
la; Lizot 75 1a; Long 115 1a; López 548' 4; z et al.
1006 6; López-Palacios 1784 la; Lott 1025'4, 372'6; Lott
& Barth 1166 3; Lott et al. 456' Та; Lugo 2588' Та; Lun-
dell 1489 З, 15949 4, 16087 6, 3447 4, 4007 6, 4008
$6, 6141 6, 6971 3, 839 3, 841' 3; Lundell « Lundell
7347 6, 7600 6, 7893 ' 3, 8000 3, 80771 3.
M 7781 la; Maas & Maas 2479 Та; Maas et al. 5457
la, 6805 Ла, 7728 la; MacDougall 1146 Та, H436 3,
s.n.(1950) 3, Magallanes 828' 3; Maguire 23797 Та,
24700 Та; Maguire & Wurdack 35763 Та; Mamani 5717.
2; Manara s.n.' 3; Manriquez 1752 1b; Marcano-Berti 553
Annals of the
Missouri Botanical Garden
la; Marcano-Berti et al. U 5; Marcks & Marcks 980 6;
Marín 101' Та; Martinelli 7078: Та; Martinez 89 3; Mar-
tínez-Calderón 1405: 6, 1425 4, 1934 6; Martino 467 la;
Martius 2608 la, 2679 Та, s.n. la; Marulanda 767 la;
Mathias & Taylor 5978' Та, 6054' Та; Matuda 15631 lb,
16589 6, 16956 3, 16959 3, 17571 lb, 17776 6, 17823
1b, 17857 la, 3278'6, 3528'6, 5631 la, 5652 6; Maury
6199 6; Maxon & Harvey 4; McDaniel 11860' Та;
McDowell 2131' Та, 2197 la, 3270' Та, 3334 Та, 3339
Та, 3712' la; McKearin 65' 3; McPherson 10913 la;
19014 Та; Miller 6480' 1a; Milliken et al. 95' 1a; Miranda
5229 4, 5484 3, 7214 6, 7748 lb, 7749 lb, 7983 4,
З, 6657 lb, 6935 6, 7469 3, 7927 lb, s.n. 3; Molina €
Molina 24691 3, 30745 3, 30872 1b, 30881' 3; Molina
& Barkley s.n. 3; Montalvo 3290! 3, 3506 3, 3716 3:
Moore 980 2; Moraes 990' Та; Morales 1659' 3, 2560' Та,
2576' Та, 3278 3, 3351 6, 3353 4, 4029 4, 4126 1a,
4127 4, 4131 4, 4153 Та, 4710 4; Moreno 10017 3,
10054 3, 10098' 3, 10405' 4, 10679' 3, 10798' 3, 11253:
4, 1205' 3, 13167 Та, 13295' Та, 131714' 1а, 16643: 3
16950' 3, 1762' 3, 1788: 3, 17923' 3, 17961' 4, 18388: 3,
18546 3, 18673' 3, 18722' 3, 18733' 4, 18741 3, 18755:
4, 1935' 3, 21153 la, 21449' 4, 21492' 4, 22515 3,
22534 3, 22613' 3, 22788' 3, 22887 3, 23533: 4, 23675:
la, 2385' 3, 23946' Та, 24243' Та, 24428' 3, 24479 1b,
24480' 4, 3038' 3, 3464' 3, 3533' 3, 3626 3, 3641 3,
3789' 4, 3816' 3, 4210' 3, 4599' 3, 4625'4, 4670 4, 4772:
‚ 5084 3, 5375 4, 5623' 3, 6603' Та, 6797 3, 9310' 3,
9400' 3, 9911' 3; Moreno & Henrich 8458' 4; Moreno &
Robleto 22887 3; Moreno & Sandino 13295 Та, 6190! ме
Mori & Kallunki 1785: Та; Mori & Gracie 22099! 1a;
$
r
3; Mrs. J.J. Ricksecker 426! =
Murillo & Jaramillo 21 9 3: Mutis 4180 Та, 5638 Та, 929
la.
Narváez 33 3, 83 3; Narváez & Puch 448 6, 462 6;
Nassar 57 3; Nee 10289 Та, 11404 Та, 11774 la, 27551
4, 28266' 3, 33480! 2, 33835' 2, 37959' 5; Nee & Miller
27551 4; Neill 2411' 3, 2471' 3, 2645' 3, 3184 4, 443!
Та, 4635'4, 58 ` pes 7304' Та, 814 3; Neill et al. 8264:
la, 8315' la; n 634: la; Nicolson о la; Núñez
10167 Та, 12796 5. 1280» 5; OTS s.n. 3.
Oersted 7957 4; 7960 3, 7965 3, s.n. 3, s n. la,
la, s.n. 4; Oldeman 1375 Та, 2220 Та, 2368 1 la, 3028
la, B-1107' Та, B-1305' Та, B-2515' Та, В-3406: la, B-
524 Та, В-780 Та, B524 Та, T-42 Та; Oliveira 1209 da.
1669 Та, 3424 Та, 3772 Та, 37772 Та, 4192 Та, 4337
la, 4926 la, 503 la, 5243 Та, 5755 la, 998 la; Oll-
gaard et al. 57183 Та; Opler 156 4, 1671 3, 1738 lb,
1745 4, 1764 4, 634 3, 642 1b, 832 1b, 969 3; Ortiz
1175 6; Otero M-19 Та; der Outer 958 L la; Ozment et
al. 453 6.
Palacio & Bautista 3299 Та; Palmer 48pp Ла, 587 la;
Pannier 835 3; Para 80' 1a; Paul 236 1a; Paul et al. s. n.
(779) Та; Pavez s.n. Та; Paz y Mino 81002! Та; Pedersen
12281' 2; Pena 130 la, 156 la; Pérez 1678: 3; Pérez-
Arbaláez & Cuatrecasas 6357 Та, 6400 la a; Pére Duss
3065' la; Philcox et al. 3327 la; Philipson et al. 2020
la; Pinto et al. 328 Та; ]
3, 12454 Та, 15117 la, 15539 Та, 15577 la, 15585 Та,
16105 Та, 2576pp Та, 2629 3, 2773 Та, 4766 4, 6056
7832 la, 7900 3, 7901' Та, 7946' 3, 7959' 3, 8095 Та,
733 Р 78' Та; Porter 1144
3; Prance 22452. DR 29875 hid s.n. la; Prance & Pen-
nington 1335' la; Prance et al. 11893 Та, 13980' la,
atia la, 18216 5, 25895' Та, 7513'5, 7947 5, P25895
s.n. la; Prevost 1080' Та, 1395' Та, 628' Та, 795' la;
Pringle 3692' 6; Procter 4785' Та; Proctor 30216 6; Pruski
et al. 2781 la; Рита 78 3; Puch & Narváez 304 3;
us 8791 4
de Queiroz 803: Та; Quezada 76' 3; Quintero 333 Та;
Quiros 733 3; Quistgaard 58 1a.
abelo & Penafort 1453 Та; Ramirez 13' Та; Ratter
& Milliken R6116' Та; Ratter et al. R5703' Та; Rentera
et al. 1701' Та, 3581' la, 4282' Та; Revilla 1207 la,
1208' Та, 703' Та, 778' Та; Reyes 126' Та; Ribeiro 148
la; Ribeiro & Pinheiro 1185: la; Ricksecker s.n. 1896:
la; Rivet 5446 3; Robertson & Austin 198: 1а; Robles
138 4; Robleto 1089' Та, 1220 lb, 1594 Та, 1716: la,
189' 4, 1967 Та, 2099 1b, 3974: 1а, 400 1b, 827! la;
Roe et al. 1332 3; de Rojas 1127' Та, 311' 1a, 641 la,
705 3, 843 la; Roldán et al. 1462: la; Rombouts 781'
a; Romero-Castafieda 10093 3, 11140 1a, 11142 1a,
11229 1a, 11236 3, 113 1a, 1161 1a, 1742 1a, 240
3. 360 Та, 6363 la, 9028 Та, 93 9669' 3,
Та, 9769 3, 9796 3, 9810 3, 9940 Та; Rosa 2487 la.
099 Та; Rosa & Cardoso 2540' Та; Rothschild 370: 3
Rovirosa 149 la; Squires 317 1а, 317a Та; Rutkis 176
la; Rubio 1831' 2, 1832 2; de la Rue s.n. Та; Rue
2547 Та, 2635 Та, 2856 3; Rugel 349 3, 86 3; Кизђу
& Pennell 128 3, 379 3; Rusby et al. 647 Та; Ryan
s.n. (455b) Та; Rzedowski 21146 3, 7459 6; Њеним
& McVaugh 1422 3.
Sabatier 720 Та; Saddi & Mattos Filho 3432 2, 3579
5, 3667 5; Saer 251 Та, 522 3, 651 la, 658 3; Sagot
402 la; Sagra s.n. 3; de la Sat 33892 3; Saldias
3607 Та, 3817 5; ow 68 la; Sanabria et al. 178
a; man a; о 9838: 6; Sandino
P366 La, 1396 3, 1872 3, -— 3, 2591B' la, 2995 6,
E
1 la; Sastre 1431' 1a, 8114' la;
Sastre et al. 4143! la; Sastre & Sastre 4158' Та; Sauer
& Gade 3185 ~ clio v 53 la;
~
ES.
y
e
- 4; Seibert 1664' Та; Seidel 117: 2; Seidel
& Beck 213! 2; Seler 5551 6; Serv. For. Cayenne 3803
la, 7781 Та; Sessé & Mocifio 2395 Та, 2406 3; Shan-
non 5048 la, 5049 la; Shattuck 1040! Та, 1147 la,
784pp' 4; Silva 159 Та, 713 la, 134773 le; 1619 la,
2326' Та, 2359 Та, 2586' Та, 2908 Та, 933 Та; Silva
et al. ASI02' Та, 7109: Та; нисте 2572 Та, 2658:
Та, 3675 Та; ems -Sopkin 2658 la, 5556' la,
6138' Та; Sinclair s.n. Та; Sintenis 1817 la, 5616 1a,
945 Та; Skinner 51 3, 53 3, 60' Та, 66' 3; Skog et al.
7439' Та; Smith 2537 Та, 2710' 1a, 3382' Та, 1386 4,
1143' la, 2515' la, 351! 3, 352! La, 355 8, 738 la,
s.n. 3, 1721 la; Smith & Hodgdon 3194! 3; Solomon
2446 3, 610 3; Solomon et al. 7016 2; Solymosy 7111
3; oe 17 la; Soria 2613: 2; Soto Núñez & Sites
510' 3; Sousa 1759 6, 293pp Ла, 4490 4, 7285'
ehe 1089 Та, 130 la, 309 la; 358 Та; Stahl
0 la; Standley 22061 4, 2211 9 1b. 27944 la,
е
Volume 84, Number 4
1997
Hauk 839
Review of Cydista
28628 la, 28642 la, 31183' Та, 40027 Та, 64106 6,
73829 3, 9208 3, 9361 Та, 9362 4, 97 3; Standley &
Chacén 5239 3, 5945 3; Standley & Padilla 3047 3,
3206A 3; Standley & Valerio 46564 1a; Steere 1061 3,
1366' 3, 1606 6, 2055 3, 2273 3, 2338 3, 2340 3,
3007 3, 7704 3; Steggerda 18 3; Stergios 10051 1a,
10776 la, 10780 la, 11069 la, 6952' 3, 7029' Та;
Stergios & Aymard 4182' Та; Stergios & Delgado 12926
la; Stergios & Stergios 11333 Та, 8440' Та; Stergios et
al. 2687 Та, 3341' la, 3914 16, 3921' Та, 3940 Та,
5006А' Та, 5285' la, 6034 Та, 6041' Та, 6109' la,
6115' Та, 6144 la, 6151' 1a, 6160 la, 6166 Та, 6184'
Та, 6256' Та, 8169B' Та, 8702' 3, 9378 la, 9392 la,
9627: la; Stern 96' la; Stern et al. 755 la, 882 la;
Stevens 11274' 3, 13597 3, 17418' la, 21785' 3,
22764B 3, 23050 3, 24012 la, 24013 la, 25020 la,
2702: 3, 306 3, 307 3, 309 3, 317 3, 3872' 3, 4431
3, 51 3, Ћи: 53 3, 5352' 3, 54 4, 5431 3, 9537 3,
! 8; Stevens & Montiel 17142! 4; Stevens et al.
16887 4, 17229 Та, 17286' 4, 20164' 4, 25020 Та,
Steyermark 108400! Та, 39099 la, 39686 1а, 45805
6, 87224 Та, 87729 Та, 88143 Та, 93143 Та, 98879
ч A la; Steyermark & Aristeguieta 113 3; Stey-
ark & Berry 111806' 3; Steyermark & Brewer
05275 la; Steyermark & Delascio 129176' Та; Stey-
mark & Liesner 127419% 1a; Steyermark & Manara
110761: 3, 110966' Та; Steyermark et al. 101516 Та,
102105 Та, 114627 la, 114668 Та, 114721' la,
114743' la, 114807 la, 120157' la, 123146' la,
123161' 3, 123406' Та; Stork 2795 Та, 4009 3; Strud-
wick et al. 3932' 1a; Suringar s.n. L 1a; Svenson 417
4; Swallen 2546 6, 2547 6.
Tamayo 2803 Та, 2853 Та, 3640 Та, 4426 Та; Taylor
11156' Та; Taylor 4421 3; Téllez 10365' Та, 10369 4;
Téllez 2055 6, 2223' 6, 2389 6, 2418' 6, 2470' 6, 2478'
6, 2518! 4, 2691' 6, 2863! 3, 3289 6; Téllez & Cabrera
e Ti
Timaná & E Јали 2386 5; Tonduz 13826 1b, 13912,
883 2779 Та; Torres & Torres 210' 6; Torres
540 la; von Tuerckheim 2527 la;
1175:6, 12116, 1764 4, 18436, 1907 6, 1910 4, 1986
3, 2616 6, 292 3; Tyson 1089 Та, 2683! 1а, 6638 Та,
а.
Ucan 108 3, 2315 6, 307 6, ps 3; Ucan & Burgos
6 3; Underwood &
2
11564 4, 12675 4, 12757 4, 13976 4, 20799' 3, 3728
4, 7750 4, 8077 4, 9423 4; Ventura & López 3690' la:
Villiers & Feuillet 2022 Ya; Vogel 1269 Та, 1 389 3, 391
la, 790 3, 973' 3, 982 Та, s.n. la, s.n. 389 3; Vreden
i la.
Warscewicz 51 Та; Webster & Kennedy 16879' Та; Web-
ster et al. 12722: 3; von Wedel 2388' Та, la, 2449'
Та, 2451' Та, 2827 la, 554 Та; van der We ;
van der We: nzález 5345 la; West 3561 3; West 15
la, 25/2 Та; White 146 4; Whitmore 29 3; Wilb
3; Wilbur & Wilbur 1595 Та; Williams 10308' 3, 10643
a, 10845 Та, 10847 Та, 11446 Та, 12192 Та, 12531,
la, 12648 la, 13599 la, 15346 la, 15506 la, 16097
la, 9267 la, 9833 la, 9875 1b, s.n. 4; Williams et al.
26473 la; Williams 14119 4; Williams & Molina 12577
lb, 14681 3; Williams & Williams 18306 3; Williams 228
la, 624 5; Woodbury 3403 1a; Woodson 1196' 1b; Wood-
son & Schery 1309' Та, 1651' Та, 1679 la, 811A’ Та;
Woodson et al. 119 1b; Woytkowski 35018' Та, 6264' Та;
Wright 3049: 3; Wullschlaegel 2005 la, 2005 Та, 2505
photo la; Wurdack & Adderley 43075 Та; Wurdack &
Monachino 41276 Та.
Yero 1046 3, 238 3; Ујазг 478 Та; Yuncker 8262 6.
Zambrano 1389 1a; Zamora 2118 3; Zanoni & Mejía
9017 Та; Zanoni et al. 21237 Та, 34268-A Та; Zardini
4125: 2: Zarucchi 3859 Ла, 4064 Та, 4126 1а, 5500 Та,
5 Та: Zarucchi & Betancur 6465' Та; Zarucchi &
Cuadros 3859: Та, 4064! Та, 4126' Та; pr et al.
a; Zaruma 599! Та; Zul:
w/o pe de 123 la; w/o collector 55 la: wh collector
48: Та.
сл
INDEX TO SCIENTIFIC NAMES
Anemopaegma A in 815, 829
Bun р uL, oi 815, 828
RUNE A o DEN 828
Vargasianum он
——— засе ир ou oa 815, 833
cci: A САО м2 ai HUMUM UE 828
md oo o o nue cip аа 825
potosina e 815, 833
pseudochica = 828
ee 815, 823, 825, 828, 829, Е 834
dO Ural ipae LD E 23
diosa оа ce in 829
mau 251.5 24 aruis desiit
incarnata aaa
lepidola n = — — — ———————— 831
мањине au ene ———— 825
pia Eu и 825
s B t 2555 QU LU i esu pe 829
sarmentosa 2 A a
A a es ON
иара О аар иат 834
hu. (d а E
Clytostoma —....-.------ 816, 817, 826, 828, 833, 834
сү н
E a И
Clytostomanthus -.....--------------- 815, 820, 828, 829
eons. A ی ا denne 820,
Cydista aequinoctialis -..
aequinoctialis var. aequinoctialis ....... 817, 823, 825
aequinoctialis var. hirtella ........... 818, 820, 823, 825
amoena A 815, 834
MAN — —— TTL
Hd з LL emere
834
déco. eo 7-821, 825, 828, 829, 834, 835
dcm с 815, 817-819, 821, 825, 830, д
incarnata нс
lilacma == 815-819, 821, 823, 826, 82085
magnifica A LL
potosina ...... 816, 817, 818, 821, 826, 830, 834, ins
pubescens — ————— 815, 828
840 Annals of the
Missouri Botanical Garden
rubicunda 834 Phryganocydia 817, 826
sarmentosa 828 leonotoma 815, 829
seemannii 815, 835 diversifolium 829
spectabilis 815, 825 Potamoganos 817, 826
vargasiana 815, 829 Roentgenia 815-817, 826
Lev туз --- 815, 820, 828 bracteomana 2815
nicaraguensis 820, 828 Temnocydia 815, 823, 825
Mansoa .. 815 iui rn 823
d са 5ш cuu Мы E 815 spectabili 825
SYSTEMATICS OF ELEUSINE
GAERTN. (POACEAE:
CHLORIDOIDEAE):
CHLOROPLAST DNA AND
TOTAL EVIDENCE!
Khidir W. Hilu? and John L. Johnson*
ABSTRACT
Indian cereal, E. coracana subsp. corac
4 Ed oracana, a taxon with a
site variation is used here to elucidate the phylogenetic relationships among species of El
s
genus. The present investigation provides additional support for the origin of the crop from the tetraploid E. coracana
subsp. africana and substantiates the monophyly of the two subspecies of E. coracana and E. indica. Evidence presented
points to the African origin of the vicariant Е. tristachya and an earlier introduction to a d further differentiation in
South America.
Eleusine Gaertn., Poaceae subfamily Chloridoi-
deae, is comprised of four annual and five perennial
species (Phillips, 1972). Polyploidy (both euploidy
and aneuploidy) has played a role in the evolution
of the genus with the evident presence of diploids
and polyploids based on basic chromosome num-
bers of x — 10, 9, and 8 (Hiremath & Chenna-
veeraiah, 1982; Hiremath & Salimath, 1991). Clay-
ton and Renvoize (1986) indicated that active
speciation has made the genus taxonomically dif-
cult. Eleusine has a center of diversity in East
Africa where eight of its nine species occur. The
exception is E. tristachya (Lam.) Lam., which oc-
curs from South America north to the southwestern
United States, and as a rare adventive in East Af-
originally on th
(Phillips, 1972; Hilu & deWet, 1976a). However,
1 We thank D. M. Porter for comments on à draft of the manu
va) for providing the chloroplast DNA clones,
of the seed collections used in the study, and two anonymous
d under grant No. DHR-5 |
he Science Advisor, U.S. Agency for International Development, and
(University of Gene
Regional Plant Introduction Station for providing some
reviewers for their comments. This research was supporte
Science and Technology Cooperation, Office of t
isozyme information (Werth et al., 1994) points to
close affinity between E. multiflora and other spe-
cies of Eleusine. The remaining annual taxa include
E. coracana (L.) Gaertn. subsp. coracana, an im-
portant East African and Indian crop known as fin-
ger millet. The crop is believed to have been do-
mesticated from E. coracana subsp. africana
(Kennedy-O’Byrne) Hilu & deWet (Chennaveeraiah
& Hiremath, 1974; Hilu, 1988). Isozyme informa-
tion, on the other hand, raised the point that sub-
species africana may not be the direct ancestor of
the crop (Werth et al., 1994). This hypothesis and
the unknown identity of one of the diploid parents
leave unresolved questions about the origin of tet-
raploid E. coracana.
Among the perennials, E. semisterilis S. M. Phil-
lips has been described from a single specimen col-
lected from the southeastern part of Kenya, near
Mombasa (Phillips, 1972), and possibly has be-
come extinct. The remaining perennial species
have wider geographic distributions in East Africa.
seript, T. H. N. Ellis (John Innes Institute) and A. Day
s. the U.S. Department of Agriculture Southeastern
U-VU-1Vi .
The International Board of Plant Genetic Resources grant 85/4
2
Department of Biology, Virginia Polytechnic Institute
3 Department of Biochemistry, Virginia Po
and State University, Blacksburg, Virginia 24061, U.S.A.
lytechnic Institute and State University, Blacksburg, Virginia 24061, U.S.A.
ANN. MISSOURI BOT. GARD. 84: 841-847. 1997.
842
Annals of the
Missouri Botanical Garden
Table 1. Species, chromosome numbers, plant collections used in the study, and sources of the material. The PI
numbers refer to the U.S. Department of Agriculture collections (USDA), and the KH numbers designate K. Hilu
collections. Voucher specimens are located at VPI.
Species Collection number
mosome
Chro
Geographic origin number (2n)
Eleusine coracana
subsp. coracana
E. coracana
subsp. africana
E. indica
E. tristachya
E. multiflora KH258
E. jaegeri KH221
E. floccifolia USDA, PI196853
Dactyloctenium aegypticum Hilu, KH263
USDA, PI231130
USDA, PI315700
USDA, PI231130
Fort Portal, Uganda 36
Pretoria, South Africa 36
Nilgiri Hills, India 18
ruguay 18
Rift Valley, Kenya 16
Narok, Kenya 20
Addis Ababa, Ethiopia 18
Eldorat, Kenya —
Phylogenetic relationships among the perennial
species have not been examined.
The objectives of this chloroplast DNA study are
to elucidate the phylogenetic relationships among
species of Eleusine, evaluate the proposed infra-
generic grouping of the species, provide informa-
tion on the origin of the New World species E. tris-
tachya, assess the affinity of E. multiflora to
Eleusine, and provide additional insight into the or-
igin of polyploid E. coracana.
MATERIALS AND METHODS
Chloroplast DNA (cpDNA) was isolated from
plants grown in the greenhouse. The species, seed
collections, and sources of material for Eleusine and
the outgroup Dactyloctenium aegyptium (L.) P.
Beauv. are listed in Table 1. Dactyloctenium Willd.
and Eleusine are members of the subtribe Eleusi-
ninae and are considered to be taxonomically
linked (Clayton & Renvoize, 1986). Plant material
for E. intermedia S. M. Phillips and E. kigeziensis
S. M. Phillips was not available. A previous study
on the cpDNA variation in Eleusine (Hilu, 1988)
demonstrated the lack of infraspecific variability;
thus one collection per taxon was used here.
Seeds were grown in flats and leaves were har-
vested from 5—7-inch seedlings, frozen in liquid ni-
trogen, and stored at — 70°С. DNA was isolated fol-
lowing the procedure of Saltz and Beckman (1981)
as modified in Hilu (1988). The DNA was digested
with the restriction. endonucleases Aval, Avall,
BamHI, Већ, Ва, Dral, EcoRI, Sspl, PstI. The
DNA fragments were resolved electrophoretically
on 0.8% agarose gels, stained in ethidium bromide,
and photographed in UV light. For the Southern
hybridization, DNA was transferred to Zetaprobe
nylon membranes (BioRad Inc.) using the alkaline
procedure (Reed & Mann, 1985). The membranes
were baked in an oven at 65°C for 2 hours and
stored at 4°C.
To examine the restriction sites in the cpDNA,
ten Pstl cloned cpDNA fragments of barley (pro-
vided by T. H. N. Ellis and A. Day) covering 98%
of the genome (Day & Ellis, 1985) were used se-
quentially as hybridization probes. The probes were
labeled with *P using the nick translation kit of
Bethesda Research Laboratories Inc. (BRL). The
membranes were prehybridized overnight at 65°C
in 3X SSC, 20 mM phosphate buffer pH 7.0, 796
SDS, 10х Denhardts solution, and 100 mg/ml
salmon sperm DNA. Identical conditions were used
for probe hybridization. Membranes were exposed
to Kodak XAR-5 film to visualize homologous
bands. Probe stripping was carried on after each
hybridization by washing the membranes three
times, 20 min. each, in 0.1X SSC and 0.5% SDS
at 95°C. DNA fragment sizes were calculated by
comparison to Lambda HindIII and a 1-kilobase
fragment ladder marker (BRL).
Phylogenetically informative sites (ie., those
found in two or more but not all species) were
scored as present-absent. Small deletions and ad-
ditions unique to particular taxa were excluded
from the analysis. The data were polarized in re-
lation to the outgroup species Dactyloctenium ae-
gyptium, transformed into NEXUS format using
MacClade 3.0 (Maddison & Maddison, 1992), and
analyzed by the Wagner arsimony method in
PAUP version 3.0 (Swofford, 1990). The parsimony
analyses were conducted using the exhaustive
search method with MULPARS, TBR branch swap-
ping, and CLOSEST addition to estimate relation-
ships and tree topology. The bootstrap method with
100 replications and the branch-and-bound search,
Volume 84, Number 4
1997
Hilu & Johnson 843
Systematics of Eleusine Gaertn.
Table 2.
Restriction site mutations detected in the cpDNA of Eleusine species. The position of the restriction site
on the chloroplast genome is identified by the probe used (P1-P7) and the restriction enzyme (see Day & Ellis, 1985,
for probes map). When more than one restriction site is revealed by a probe, the sites are designated by an alphabetical
etter.
EcoRI Sspl BamHI Aval Avall Dral
PI. P2 P2 РЭБ: РА. Pl Ра РЬ РЬ Pl. P2 ОР РЊ PS .Pl 22
coracana 1 0 0 0 0 1 0 0 0 0 0 0 1 1 1 0
їпйїса 1 0 0:-:07. 0 1 ЈЕ 40 0 о: 0 Oroa 1 1 0
tristachya 1 0 0.5. Eo. 0 1 E O0 E :U 1 1 1 1 m
multiflora 0 1 1 0 1 0 1 1 0 1 1 0 1 1 1 1
jaegeri 0 1 1 15-0 1 E 0 o 1 1 I ~ У 1 1
floccifolia 0 1 0" 01 1 | "о | 1 1 1 1 1 050
Dactyloctenium 0 1 0 1 1 0 1 1 1 1 1 O 509 0 1
and the decay analysis were performed in PAUP to
determine relative support for the clades (Felsen-
stein, 1985; Bremer, 1988).
RESULTS
Hybridization of the cpDNA clones to the restric-
tion digests of the Eleusine species revealed 28 re-
striction sites. Sixteen sites were phylogenetically
informative, while the remaining 12 were present
in only one species (Table 2). Ten of the unique
sites were characteristic of the E. multiflora ge-
nome, one was found in E. floccifolia (Forssk.)
Spreng., and the other occurred in Dactyloctenium
aegyptium. Among the cpDNA clones used, P8 did
not resolve informative or unique sites. This clone
covers the inverted repeat region of the chloroplast
genome, a region less likely to produce informative
sites because of its highly conserved nature. Small
addition-deletions were also observed. The exhaus-
tive search evaluated 945 trees of 23 to 39 steps
in length and retained a single, most parsimonious
tree of 23 steps. The consistency index (CI) and
retention index (RI) for the most parsimonious tree
were 0.70, reflecting the relatively low homoplasy
on the tree. The bootstrap and the decay index val-
ues for the different clades are given in Figure 1.
DISCUSSION
EVOLUTION OF THE Е. CORACANA-E. INDICA-E.
TRISTACHYA COMPLEX
1. Evolution of Tetraploid Species
The evolution of the tetraploid E. coracana
subsp. coracana and subsp. africana and its ge-
nomic relationship to the diploid E. indica has been
the focus of various studies (see introduction). This
cpDNA study did not resolve restriction site differ-
ences between the two subspecies of Е. coracana.
A similar finding was also obtained in a previous
cpDNA study (Hilu, 1988) that focused on the two
subspecies of E. coracana, E. indica, and E. tris-
tachya. These studies thus provide evidence in
support of the direct origin of finger millet (E. cor-
acana subsp. coracana) from E. coracana subsp.
africana. Additional evidence in support of this
theory comes from restriction fragment variation in
the intergenic spacer region (IGS) between the 17S
and 255 ribosomal genes (rDNA). Hilu and Johnson
(1992) showed that the domesticated subspecies is
quite homogeneous in IGS pattern and that its
rDNA phenotype is identical to one of the IGS phe-
notypes detected in subspecies africana. In con-
trast, recent isozyme data (Werth et al., 1994) dem-
onstrated the presence of alleles in domesticated
finger millet, subspecies coracana, which were not
shared with the proposed wild ancestor subspecies
africana. Consequently, the study questioned the
possibility of a direct origin of subspecies coracana
from subspecies africana. This disagreement has
two possible explanations. One, the two tetraploid
taxa had different origins, sharing only one common
diploid genome; the donor of the second genome
might have contributed the unique alleles reported
in subspecies coracana. This hypothesis is incon-
sistent with cytogenetical information (Chennaveer-
aiah & Hiremath, 1974; Hiremath & Salimath,
1992) that demonstrated complete genome homol-
ogy between the two taxa. Two, the tetraploid sub-
species africana is genetically quite variable due
to high diversity incurred by polyploidization and
possible multiple origin, and subspecies coracana
was derived from a limited number of populations
of subspecies africana, a situation typical of crops.
(RAPD) markers (Hilu, 1995). The rDNA and
RAPD studies demonstrated that the DNA patterns
Annals of the
Missouri Botanical Garden
E
a
=
Q
5 S
© ©
5 Е =
= »
Q > x
3 Е
~
ГЕ GNC == GNS
а pw PU
ie SN = >
LA LC == LA
ID IR 77 IS
s+
E
34% (1)
racemose.
(phenotypes) of subspecies coracana are present in
and can be derived from those of subspecies afri-
cana, and that these phenotypes represent only a
subset of the genetic variation in that tetraploid
wild taxon. Due to the demonstrated genetic vari-
ability in subspecies africana, the isozyme data
might represent only part of that variation. There-
fore, the origin of the domesticated taxon from one
or a few genotypes of the wild tetraploid is a more
likely explanation for the incomplete concordance
between the isozyme data and the other molecular
; LA, ; PW, palea winged; PU
by pericarp; SN, seed nacked (i.e., seed free of pericarp)
з
З = =
$ 5
E 5 3 S
E 3 5
А
GNC
LD A
PW 7 GNC
SP LD
LA 69%(1) PW
9 А =
нў 100%(6) 3
40% (1) T '5
4
10
14
, palea unwinged; SP, seed surrounded
; ID, inflorescence digitate; IS, inflorescence subdigitate-
information. This explanation is strongly supported
by the genomic homology and interfertility between
the two taxa (Hiremath & Salimath, 1992). A more
extensive isozyme study that includes a large sam-
ple of subspecies africana from across its range of
distribution might shed some light on this disagree-
ment
Eleusine indica shares the same restriction sites
with the two tetraploid subspecies of E. coracana,
indicating the presence of a common chloroplast
genome among the three. This study thus further
Моште 84, Митбег 4
1997
Hilu & Johnson 845
Systematics of Eleusine Gaertn.
supports a previous cpDNA investigation (Hilu,
1988) that pointed to E. indica as the “A” genome
donor of the tetraploid E. coracana. Originally, the
diploid species E. indica was considered as the ge-
nomic donor of finger millet (Greenway, 1945; Ken-
nedy-O’Byrne, 1957; Jameson, 1970). Based on
lack of chromosome pairing in a synthetic hybrid
between E. coracana subsp. coracana and E. in-
dica, Chennaveeraiah and Hiremath (1974) con-
cluded that the latter species did not contribute any
of the genomes of finger millet. That study, however,
was based on a single interspecific hybrid. In a
more recent cytogenetic study based on more than
one hybrid, Hiremath and Salimath (1992) found
an appreciable amount of chromosome pairing be-
tween the E. coracana subsp. coracana and E. in-
dica genomes, confirming the genomic contribution
of E. indica to the tetraploid E. coracana as pro-
posed by the cpDNA study (Hilu, 1988).
2. Origin of E. tristachya
To address the question of the origin and dis-
persal of the vicariant Eleusine tristachya, three
points have to be considered. First, the monophyly
of E. tristachya and E. coracana and E. indica is
substantiated by information from this study (100%
bootstrap, decay index of 6, and five unambiguous
mutations) as well as from previous molecular, bio-
chemical, and cytogenetic work (Hilu et al., 1978;
Hilu & Johnson, 1992; Hiremath & Salimath,
1992: Werth et al., 1994). Second, all the other
species of Eleusine are native to East Africa and
are widely distributed in that region, even when
they are found on other continents. Third, E. tris-
tachya has relict populations in the Sudan area of
Africa. Considering these points, a South American
origin of E. tristachya is not likely. The species, at
its incipient stages of differentiation from the com-
mon ancestor of the annuals (species in the termi-
nal clade, Fig. 1), must have moved to South Amer-
ica during the early stages of continental drift.
Clayton’s (1981) study of the geographic distribu-
tions of grass genera promoted the possibility of a
Tertiary spread of grass genera across the Atlantic
when the latter was a relatively narrow water pas-
sage. Dispersal during the post-Columbus trading
times is less likely because of the very limited dis-
tribution of E. tristachya in northeastern Africa.
INTRAGENERIC SYSTEMATIC RELATIONSHIPS
In her revision of the African species of Eleusine,
Phillips (1972) asserted that the genus can be di-
vided into two groups of species on the basis of the
annual and perennial habit. She indicated that
within each group, the differences between species
are often small, and that among the annuals in par-
ticular, introgression is frequent. In addition to the
annual and perennial habit, Phillips cited differ-
ences among the two groups of species in spikelet
morphology, such as the number of nerves and the
presence of a keel in the glumes and lemma, and
in the presence of a keel in the palea. These mor-
phological characters are mapped on the cpDNA
cladogram (Fig. 1). On the basis of inflorescence
and spikelet characters, the annual E. multiflora
occupies an isolated position in relation to the an-
nual species and the genus as a whole. Its raceme-
like inflorescence is atypical of the digitate-spike
arrangement in Eleusine. Phillips (1972) also noted
that E. multiflora can be distinguished from the
other African species of Eleusine by the short,
broad spikes. The lemma keel of E. multiflora ex-
tends into a cusp or a mucro, unlike other species
of Eleusine where the keel does not extend at the
lemma tip. The seed of E. multiflora ruptures from
the membranous pericarp before it is dispersed
from the spikelet, whereas in the other species the
seed remains enclosed in the pericarp after dis-
persal.
The most parsimonious tree based on the cpDNA
data showed the three annual species Eleusine cor-
acana, E. indica, and E. tristachya as a terminal
lineage strongly supported by six unambiguous re-
striction sites, a decay index of 6, and 100% boot-
strap value (Fig. 1). The fourth annual species, E.
multiflora, appeared in an individual clade situated
between the two perennial taxa. Among the peren-
nials, E. floccifolia emerged as a sister species to
the annual species assemblage of E. coracana, E.
indica, and E. tristachya. The E. floccifolia clade
was supported by 40% bootstrap and one unambig-
uous restriction site mutation. The other perennial,
E. jaegeri, formed a basal clade in the genus, di-
verging after the outgroup Dactyloctenium aegyp-
tium. The position of the E. floccifolia clade as a
sister taxon to the three annual species receives
support from chromosome number and meiotic
chromosome behavior (Chennaveeraiah & Hire-
math, 1973; Hiremath & Salimath, 1992). These
three annual species and E. floccifolia are diploids
or polyploids based on x — 9, in contrast with the
basic number of x — 10 for E. jaegeri and x — 8
for E. multiflora. Crosses between E. floccifolia and
the annuals E. tristachya and E. coracana subsp.
coracana revealed a good amount of genome ho-
mology, with a mean of 7.6 to 8.6 bivalents (Chen-
naveeraiah & Hiremath, 1973; Hiremath & Sali-
math, 1992). The high affinity between E. floccifolia
and the three annual species was also demonstrated
846
Annals of the
Missouri Botanical Garden
in the isozyme study of Werth et al. (1994). The
basal position of E. jaegeri in Eleusine was previ-
ously demonstrated on the basis of isozyme infor-
mation (Werth et al., 1994). The chromosome num-
bers for the two other perennials E. intermedia and
E. kigeziensis were reported to be 2n = 18 and 38,
respectively (Hiremath & Salimath, 1991). The 2n
= 18 is indicative of the diploid nature and a basic
number of x = 9 for E. intermedia, a species that
appears to be morphologically intermediate be-
tween the perennial Е. jaegeri and the annual com-
plex of E. coracana-E. indica (Phillips, 1972). The
phylogenetic arrangement of the Eleusine species
could point to a descending order of aneuploid
chromosomal evolution from х = 10 to both х = 9
in the annual species E. coracana, E. indica, and
E. tristachya and the perennial E. intermedia, and
x = 8 in E. multiflora. The chromosome count of
2n — 38 for E. kigeziensis needs to be verified since
it appears as an aneutetraploid when compared
with the basic numbers x — 8, 9, and 10 found in
Eleusine. Hiremath and Salimath (1982) proposed
that x — 9 not x — 10 as the primitive number in
Eleusine, from which other basic numbers were de-
rived.
This cpDNA study unequivocally supports the
monophyly of the three annual species Eleusine cor-
acana, E. indica, and E. tristachya (Fig. 1). It also
substantiates the placement of the annual E. mul-
tiflora within the genus with three unambiguous
mutations, but in a lineage distinct from the clade
of the other annuals. Consequently, the results in-
dicate that the annual condition appears to have
arisen twice in Eleusine. The perennial species did
not emerge as a monophyletic group. The internal
placement of E. multiflora among the perennials is
probably due to the exclusion from the analysis of
the 10 mutations that are unique to this species.
When the decay analysis was performed, the three
basal clades representing the two perennials and E.
multiflora in the most parsimonious tree collapsed
into a polytomy with one additional step. Forcing
E. jaegeri and E. floccifolia into a monophyletic
clade with the Constraint option, the new tree was
only two steps longer and the CI index slightly low-
er (0.64 vs. 0.70). It is to be noted that the basal
branch that represents the perennial E. jaegeri is
supported by only one, homoplasic apomorphy (Fig.
1). These analyses show low support for the basal
nodes and imply a weak resolution at the base of
the tree.
Further information on the systematics of Eleu-
sine comes from previous biochemical, molecular,
morphological, and cytogenetical studies. Hilu et
al. (1978) surveyed flavonoid variation in the four
annual species of Eleusine, the perennial E. floc-
cifolia, and Ochthochloa compressa (Forssk.) Hilu
(a taxon closely allied to Eleusine). Although the
study could not be used to draw a conclusion con-
cerning the perennials since only one species was
represented, it highlighted the similarities among
the annuals and underscored a closer affinity of E.
multiflora to Eleusine than to Ochthochloa. Eleusine
multiflora shared three flavonoids common to all
other Eleusine species but lacking in O. compressa.
The isozyme study of Werth et al. (1994) substan-
tiated the genetic similarities among the annual
species as a group and confirmed the taxonomic
affinities of E. multiflora to Eleusine. Information
from restriction site variation in the ribosomal in-
tergenic spacer region (IGS) of six species of Eleu-
sine revealed a similar pattern of affinities (Hilu &
Johnson, 1992). The study showed the annuals, ex-
cept for E. multiflora, to share similar IGS restric-
tion sites. Eleusine multiflora had distinct IGS re-
striction sites, but displayed phenotypes that are
found in Eleusine. The perennials E. jaegeri Pilger
and E. floccifolia differed in restriction sites but
had comparable IGS variants.
In a phenetic study based on 37 vegetative and
reproductive morphological characters of Eleusine
species and one species each of related Dactyloc-
tenium and Ochthochloa, Hilu and deWet (1976a)
showed the segregation of the annuals (except for
E. tristachya and E. multiflora) in a distinct cluster
linked at a correlation coefficient value of about
0.54. The two subspecies of E. coracana and E.
indica formed a tight cluster. Eleusine tristachya
formed a cluster with Ochthochloa compressa. All
five perennial species formed one well-defined
group with two subgroups; one included E. inter-
media and E. semisterilis, whereas the other con-
tained E. floccifolia, E. kigeziensis, and E. jaegeri.
Eleusine multiflora formed a group with D. aegyp-
tium that was last to cluster with the Eleusine spe-
cies. Therefore, the morphological study confirms
the taxonomic affinities among the perennial spe-
cies and underscores the distinct position of E.
multiflora. The spikelet morphology of the annual
E. tristachya has possibly led to the separation of
this species from the remaining annuals.
It is evident from the cpDNA and the above stud-
ies that the annual species Eleusine coracana, E.
indica, and E. tristachya represent a monophyletic
group of closely related species and that the re-
maining annual E. multiflora is a morphologically
and genetically distinct taxonomic entity in the ge-
nus. The raceme-type inflorescence of this species
in a predominantly digitate-type genus could raise
the question of whether E. multiflora is a member
As
Volume 84, Number 4
1997
Hilu & Johnson 847
Systematics of Eleusine Gaertn.
of Eleusine. The inflorescence of E. tristachya and
some collections of E. coracana subsp. africana
show а tendency toward the raceme-type (Hilu 4
deWet, 1976b). Removing E. multiflora from Eleu-
sine will only create another monotypic genus, as
the species would not fit well in other related gen-
era. The taxonomic treatment of the perennial spe-
cies remains problematic. None of the morpholog-
ical treatments is phylogenetically based and, thus,
it is difficult to compare them with the cpDNA re-
sults. Available evidence, therefore, points to a pos-
sible paraphyletic origin of these taxa from an an-
cestral stock of Eleusine, as the monophyly of the
genus is well demonstrated in this investigation.
Literature Cited
Bremer, K. 1988. The limits of amino acid sequence data
in angiosperm phylogenetic reconstruction. Evolution
95-803.
етан. M. S. & S. C. Hiremath. 1973. Genome
relationships d Eleusine tristachya and E. floccifolia. J.
Cytol. Genet. 7
&
1974. Genome analysis of Eleusine
coracana (L.) Gaertn. Euphytica 23: 489-495.
Clayton, W. D. 1981. Evolution and distribution of grass-
es. Ann. Missouri Bot. Gard. 68: 5-1
A. Renvoize. 1986. Canon Graminium.
HMSO, ове
Day, А. & T. H. N. Ellis. 1985. Deleted forms of plastid
DNA in albino plants from cereal anther culture. Curr.
Genet. 9: 671-67.
ерене», J. 1985. Confidence limits on phylogenies:
An approach using the bootstrap. Evolution 39: 783-
Greenway, P. J. 1945. Origin of some East African food
plants. E. Afric. Agric. J. Kenya 10: 177-1
Hilu, K E 1980. Eleusine tristachya (Lam.) Tan (Po-
aceae). Мадгоћо 27: 177-1
988. Identification of the “A” genome of finger
millet chloroplast DNA. Genetics 118: 163-167.
. 1995. Evolution of finger millet: Evidence from
random amplified polymorphic DNA. Genome 38: 232—
-&). M. J. deWet. 1976a. Domestication of Eleu-
sine coracana. Econ. Bot. 30: 199—2
1976b. Racial evolution of finger mil-
let Eleusine coracana. Amer. J. Bot. 63: 1311-1318.
& J. L. Johnson. 1992. Ribosomal DNA variation
in finger millet and wild species of Eleusine (Poaceae).
Theor. Appl. Genet. 83: 89
. M. J. deWet € D. Seigler, 1978. Flavonoids
and the systematics of Eleusine. Biochem. Syst. & Ecol.
6: 247-249.
Hiremath, S. C. & M.
netical studies in wild and с
sine —— Caryologia 35:5
ath. 1991. ce nuclear
DNA فاا in Eleusine (Gramineae). Pl. Syst. Evol.
178: 225-233
S. Chennaveeraiah. 1982. Cytoge-
ultiv. m "nme of Eleu-
2. The 'А' genome donor of Eleu-
sine coracana v Gaertn. (Gramineae). Theor. Appl.
Genet. 84: 747-7
Jameson, J. D. 1970. Agriculture i in Uganda. Oxford Univ.
Press, Oxford.
Kennedy-O'B уте, J. 1957. Notes on African grasses.
XXIX. A new species of Eleusine from tropical and
South Africa. Kew Bull. 11: 2.
D. R. Maddison. 1992. MacClade:
of DNA from miagatone са to d membranes. Nuc
Acids m 13: 6207-7221.
Saltz, Y. & J. Beckman. 1981. Chloroplast DNA prepa-
ration DIA Petunia and Nicotiana. Pl. Molec. Biol. News-
lett. 2: 73—74.
Swofford, D. L. 1990. PAUP: Phylogenetic Analysis Using
Parsimony. Version 3.0, Illinois Natural History Survey,
Champaign, Illinois.
Werth, C. R., K. W. Hilu & C. A. Langner. 1994. Isozymes
of Eleusine (Gramineae) and the origin of finger millet.
Amer. J. Bot. 81: 1186-1197
THE FRUITS OF JASMINUM
MESNYI (OLEACEAE), AND
THE DISTINCTION BETWEEN
JASMINUM AND MENODORA!
Jens G. Rohwer?
ABSTRACT
The fruit and seed morphology and anatomy of Jasminum mesnyi are described here for the first time, as part of a
larger comparative study on the fruit and seed structures in the Oleaceae. The evidence presented here weakens the
a ee ee
Jasminum is by far the largest genus within the
Oleaceae, although recent estimates as to its num-
er of species differ widely, from ca. 200 (Green,
1994) to ca. 450 (Mabberley, 1987). The vast ma-
jority of the species have white flowers and opposite
leaves, whereas the eight species of section Alter-
nifolia DC. (Green, 1961) are characterized by yel-
low flowers and alternate leaves. Only two species,
Jasminum nudiflorum Lindl. and J. mesnyi Hance,
combine yellow flowers with opposite leaves. The
fruit of Jasminum is usually described as a double
berry, which arises through independent growth of
the two carpels of the ovary, while the septum grows
much in thickness but very little in extent. In ear-
lier papers (Rohwer, 1993, 1994, 1995b) it was
shown that the fleshy part of the fruit originates
from the seed coat rather than the pericarp, and
that the seed structure allows one to distinguish the
sections of the genus. Jasminum nudiflorum turned
out to be unusual in that its fruit does not become
black at maturity but rather dries out and fragments
irregularly, releasing the seeds. The fruit of the
closely related Jasminum mesnyi, however, ге-
mained unknown, even though the species is fre-
quently cultivated as an ornamental. Fletcher
(1916) reported that “J. primulinum Hemsl.,” as
the species is usually called in the horticultural
trade, had to be introduced to England as whole
' I thank Р. S. Green (Kew) for providing literature unavailable in German
and U. Wagenfeld processed the photos
plants, because the plant collector E. H. Wilson,
working for Veitch’s nurseries, had searched in vain
for seeds. Fletcher claimed that he had obtained a
few seeds of J. mesnyi from pollination with both
its own pollen and that of J. nudiflorum, but gave
no details about the fruit. Otherwise, the species
has obviously been propagated only vegetatively so
far. Green (1965) commented on the apparent ste-
rility of the cultivated plants, citing different chro-
mosome counts (2n = 24, 26, 39) and suggesting
that at least some clones were triploid.
MATERIALS AND METHODS
In the botanic garden of Heidelberg, attempts to
achieve fruit-set in Jasminum mesnyi by artificial
pollination (either geitonogamously, since all plants
within reach were from the same clone, or with pol-
len from Ј. nudiflorum) failed for several years.
Only after these attempts had been abandoned were
four young fruits found in April 1996. One of them
was fixed in FAA (5 ml formalin, 5 ml glacial acetic
acid, 56 ml ethanol 96%, aqua dest. ad 100 ml) on
May 24, at an obviously immature stage. The other
three were bagged in nylon nets, as a precaution
against loss due to premature abortion, with the
hope that at least one of them would develop until
maturity. The bags were inspected at intervals of
у. S. Ball stained and mounted the sections,
* Institut fiir Spezielle Botanik, Universitit Mainz, 55099 Mainz, Germany.
ANN. MISSOURI Bor. Garp. 84: 848-856. 1997.
=
Volume 84, Number 4
1997
ohwer
Jasminum mesnyi (Oleaceae)
two to three days. The fruits slowly became larger
and paler, but did not look quite mature even on
June 17. On June 19 they were found open, with
the seeds still attached in one of them but shed
into the bag in the others. All available material
was fixed in FAA. A flowering voucher specimen
(Rohwer 135) of the investigated plant had been
deposited in HEID earlier.
For sectioning, the material was transferred to
70% ethanol for at least a day, then embedded in
a 2-hydroxy-ethyl methacrylate resin (Kulzer’s
Technovit 7100; for details, see Igersheim, 1993),
sectioned with a rotary microtome (Leica RM 2145)
at a thickness of 3—6 jum, stained with Giemsa so-
lution for 2 hr. (2.5 ml Gurr’s improved R66 in 100
ml aqua dest.), differentiated in aqua dest. with two
or three drops of acetic acid and subsequently in
96% ethanol, each for about 5-20 sec. according
to the intensity of the stain, transferred to xylene
via isopropanol, and enclosed in Vitro-Clud em-
bedding medium. The microphotos were taken with
a Zeiss Axioskop on Agfapan APX 25 film.
RESULTS
In the young fruit of Jasminum mesnyi (Fig. 1),
it is obvious that both carpels of the ovary had start-
ed to grow up to form separate mericarps. In all
four fruits, however, one of the two mericarps ended
its development early, while the other continued to
grow. In this case the developing mericarp shifts to
a position almost in continuation of the pedicel,
displacing the aborted mericarp and the remnant
of the style sideways. Early on, a longitudinal ridge
with a shallow furrow in its middle becomes visible
close to the style (Fig. 1), becoming more and more
distinct and stretching more toward the distal end
of the developing mericarp with maturity. At ma-
turity, not only the fertile mericarp but the whole
fruit opens along this suture, splitting the style in
half (Fig. 2). In one of the three mature fruits the
dehiscence continued in a straight line along the
carpel median, over the tip down to the base of the
fertile mericarp. In the other two the splitting was
straight only slightly beyond the point where the
suture becomes indistinct, followed by an irregular,
branched crack above.
In the immature fruit (Figs. 3, 4), the pericarp
consists of an outer epidermis of small, isodia-
metric to slightly flattened cells with a rather
thick cuticle, several at least initially parenchy-
matous layers, and an inner epidermis of small,
thin-walled, slightly flattened cells. Numerous
small, weakly developed vascular bundles are
found embedded among the inner parenchyma-
tous layers. These layers, about four to six in the
distal part of the mericarp, cannot keep up with
the growth of the fruit in the areas between the
vascular bundles, so that they are destroyed early
in fruit development. The outer epidermis, some
outer parenchymatous layers (one or two distally,
more toward the base), and the inner epidermis
remain distinct for most of the fruit development.
At maturity, only the outer epidermis and its hy-
podermal layer are still recognizable in the distal
part of the mericarp (Figs. 5, 6). Slightly below
the tip, the fruit wall becomes thicker on both
sides of the dehiscence line, and polygonal cells
with slightly thickened, pitted, lignified walls ap-
pear (Fig. 7). The most distal ones of them are
more or less isolated, but soon they aggregate to
form woody strands along the suture. Further to-
ward the base, these strands not only become
thicker but also extend further around the meri-
carp (Fig. 8). Even at the base, however, there
are about as many parenchymatous as sclerified
cells in the abaxial wall of the mericarp (Figs. 9,
10). The outer and the inner epidermis, the (usu-
ally no more recognizable) inner parenchymatous
layers, and several layers forming the weakness
zone within the suture are never lignified. Oth-
erwise, the thickness of the cell walls and the
degree of lignification increase from the abaxial
side of the mericarp toward the suture, and from
the outside inward.
The seeds are elongate, about 8-10 mm long
and 3.5-4.5 mm wide. Their outline in cross sec-
tion (Fig. 11) is thickly plano-convex when both
ovules of a locule develop; otherwise it is elliptic.
The two seeds face each other with their raphal
sides. The seed coat consists of numerous cell
layers, of which the inner ones are gradually de-
stroyed by the growing endosperm. As in other
species of Jasminum, but in contrast to most oth-
er Oleaceae, there is no trace of an endothelium
(jacket layer) at any time during development.
The two outermost layers are very different from
the rest of the seed coat, and unequally devel-
oped on the two sides of the seed (Figs. 11, 12).
On the antiraphal side (Fig. 13), the outer epi-
dermis (exotesta) consists of small, transversely
elongate cells with thickened, lignified walls,
more massive on the outside than on the inside.
The cells of the hypodermal layer are much larg-
er, with massive, lignified thickenings occupying
almost their entire anticlinal walls, and much
smaller thickenings on the periclinal walls. All
further layers are parenchymatous and entirely
collapsed at maturity. On the raphal side (Fig.
14), the cells of the exotesta are much larger,
850
Annals of the
Missouri Botanical Garden
4 77
Sm) 00
qt. А $ a
Figures 1—4. Fruits of Jasminum
with the seeds (S) still attached. —3. Cr
young pericarp and outer layers of young testa.
pericarp, R = raphe bundle, 5 = seed, SE =
3 = 1 mm, in 4 = 200 pm.
forming a palisade of hexagonal cells. The lig-
nified wall thickenings are much weaker and al-
most entirely restricted to the anticlinal walls,
usually to the corner where three cells meet. The
cells of the hypadermal layer are longitudinally
elongate and smaller than those of the exotesta,
though still larger than on the antiraphal side.
They have (mostly two) band-shaped wall-thick-
enings in their anticlinal walls, occasionally also
in their outer periclinal wall. When the seed
dries out, the two outer layers separate from the
rest of the seed coat, tearing apart the thin inner
end of the anticlinal cell walls in the hypodermal
layer. In the area where this separation occurs,
several of the following cell layers have band-
shaped wall thickenings as well, much smaller
than in the hypodermal layer and becoming still
smaller and more irregularly distributed toward
the inside (Fig. 14). Below them follows the ra-
phal bundle, which is the only vascular bundle
of the seed, and several parenchymatous layers
that are increasingly collapsed toward the inside.
About three to five (only apically and basally
more) layers of endosperm surround the embryo,
which has very large, flat cotyledons with well-
developed, usually two-layered palisade paren-
Volume 84, Number 4 Rohwer
; 851
Jasminum mesnyi (Oleaceae)
Tes
ADD Ly yy yy
de
pu Ур YD ^^
M M SPT
РАИ
NNI >
4
+ 2 imb
9 we".
-
of Jasminum mesnyi. —5. Membraneous, distal part. —6-8.
> below the apex of the mericarp (Fig. 6) to slightly below the
middle (Fig. 8). —9. Basal part of the fertile mericarp, slightly above the level where the two mericarps become
separate. —10. Pericarp on abaxial side, same section as in Figure 9. A = remains of aborted carpel, D = dehiscence
line, F = remains of funicular tissue. Scale bars in 5, 6, 7, 8, 10 = 200 jum, in 9 = ] mm.
pericarp and seeds with a fleshy testa. Dry but ir-
regularly fragmenting fruits, and seeds with an al-
most identical structure as in J. mesnyi, have re-
cently been described in another well-known
ornamental species, J. nudiflorum (Rohwer, 1993,
1994, 1996). The development of a median suture
can even be observed in young fruits of this species
(Fig. 15), but none of the open fruits observed so
far had actually dehisced along this line. In addi-
tion, the pericarp of J. nudiflorum is almost ho-
chyma. Starch was not found in either the en-
dosperm or the embryo.
DISCUSSION
The fruit of Jasminum mesnyi agrees with that of
the other species of Jasminum in the development
of two separate mericarps. With its loculicidal de-
hiscence, however, it goes beyond the current cir-
cumscription of the genus, which has normally in-
dehiscent fruits with a membranous to coriaceous
852 Annals of the
Missouri Botanical Garden
Figures 11-14. Seeds of Jasminum mesnyi. —11.
Testa and endosperm on antiraphal side, longitudinal section. —14. Edge of ra
cotyledon, E = endosperm, R = raphe bundle. Scale bars in 11, 12 = 1 mm, i
Cross section. —12. Longitudinal section, chalazal end. —13.
phal side (above), cross section. C =
n 13 = 100 jum, in 14 = 200
Volume 84, Number 4
1997
Rohwer 853
Jasminum mesnyi (Oleaceae)
Figures 15, 16. Comparable fruit structures in other species. —15. Jasminun nudiflorum, very young stage of fruit
development. —16. Meno
dora africana, mature fruit, the mericarp on the left open and empty, the one on the right
still closed, showing the difference between the thicker proximal part and the thinner distal part (hexagonal wall
thickenings of the seed coat shining through). A = Aborted mericarp, D? = ridge with longitudinal furrow, similar to
5
the dehiscence line in J. mesnyi. Scale bars in 15 = 1 mm, in 16 =
mogeneously membranous throughout, with only a
few stone cells near the base, so that it lacks the
mechanism for an orderly dehiscence. All other
characters, however, whether vegetative, floral, or
seed characters, show beyond doubt that the two
species are very closely related. Still the differ-
ences appear important enough to reject the view
expressed by Henry (1904), who thought that “J.
primulinum” was just a variety of J. nudiflorum.
Fruits with dry, dehiscent mericarps are, within
the Oleaceae, the most important diagnostic char-
acter for Menodora, a small genus (22 spp.; Stey-
ermark, 1932; Turner, 1991, 1995) of low shrubs
or, more frequently, suffruticose to almost entirely
herbaceous plants. In this genus, however, the de-
hiscence of the fruits is not longitudinally loculi-
cidal but normally circumscissile (Fig. 16). In M.
africana Hook. at least, where the fruits and seeds
have been described in detail recently (Rohwer,
1995a), the pericarp is thicker below the equatorial
suture line, but does not contain stone cells, or any
other lignified elements, except in the very base of
the fruit. No trace of a median suture line was
found in this species, at any stage of development.
n exception within the genus, Steyermark
(1932) described the presence of a median suture
in Menodora spinescens A. Gray, but he never found
fruits that had opened along this line. The shape
of the fruit is likewise unusual. The mericarps are
elongate and spreading in M. spinescens, whereas
they are subglobose and more or less appressed
mm.
against each other in the other species. Jasminum
mesnyi has similarly elongate mericarps, and this
suggests that they would most likely spread if both
were developed. In J. nudiflorum, in contrast, they
are roundish and adjoining. This character seems
to be of minor significance, since there are many
parallel cases and intermediate forms known from
Jasminum, often in closely related species (e.g.,
roundish/adjoining in J. bignoniaceum Wall. ex G.
Don, intermediate in J. humile L., elongate/spread-
ing in J. odoratissimum L.). More important is the
aberrant number of ovules in M. spinescens. While
the other species of Menodora have four ovules per
locule, M. spinescens has only two, like most other
Oleaceae, including the yellow-flowering species of
Jasminum, whereas the majority of the white-flow-
ering species of Jasminum have only one. This
again bridges the discontinuity between the two
genera. In Menodora, the information given by
Steyermark (1932) and Turner (1991) suggests that
M. spinescens is the only species with basically
white flowers (often tinged with brownish purple),
whereas the basic flower color of the other species
is yellow (sometimes tinged with red to purple).
Inasmuch as the fruits of Jasminum mesnyi and
Menodora spinescens suffice to undermine the tra-
ditional distinction between Jasminum and Meno-
dora, it may be worthwile to examine other char-
acters that could possibly separate the two genera
(summarized in Table 1). Habit (shrubs and climb-
ers in Jasminum versus at most subshrubs in Men-
Annals of the
Missouri Botanical Garden
*peururexo sarads ивоџошу YHON Ајио 18} ос,
‘apis үецавл по [влолов ‘apts [ецделцив uo oup) ə
‘apis [ецавл uo eysajoores ‘apis [ецделцив uo вәә рәрем-ҹәтці "реше p
"влофив popn[our цим әст вјјолоо Buoy e sey *pz0pouapy [eoIdA) e злодовлецо 19470 [је ur мој иој Diopouay Ing »
‘qnays зепдол в st 275пд01 D40pOU9]y q
"иоове ут пв ио SI asuaurmis шпитшую[ y
Zz umouyun uMouxun 6e '92 "vc Zs 9% 9 9g ис зәдшпи ошовошолу“)
зер A1qeqoad rey ey Tey ey ey о8влој5 aBBIOIS 10 WH suopo[4107)
umouyun e e ,[enboun ,[enboun [919498 [81998 I sguruaxorq [Ем
чим sia dR] [е1вә1овәр
umouyun + de + + — — — 8159} шуим dex)
UMOUNUN pəuəyəry} “еше ралловола jou ppenboun ppenboun рә[[әә-[үешв 2189100108 8189102188 в]вәохә Jo o1njonujs
umouyun I 1 I I I ќивш I peas/saypung .re[noseA
—— — ]euonounj-uou јеиоцопиј Алејиоштрпл -— — == ainyns UBIPIJA
ојгввтовштоло 9[rssrosumouto Jenou [purpniruo[ 1e [naut — — — sduvouour Jo aousostyaq
qsıpunoı qsıpunoı ayesuoja oje3uo[o цѕгрипол 9[qeueA I[GBLIBA 9[qeueA зфавоџош jo adeys
Kip *oped Kip ‘ayed Kip *oped Kip 'ојеа Aap ‘aped 39e1q Lysay 19814 ‘Aysoy xoerq ‘Aysay Ayunjyeu 18 pinay
v v ^A A 7, e т Ауепеп z A[[ensn ојпоој sad so[na()
peuesxo/noqs pauesxo/uous рәмәѕхәдлоцѕ pepn[ourZuo[ рорпјош/виој рорпјошудиој popnjour/2uo[ рерпјошувиој siou]ue/aqn] e[[0107)
MO[[9Á мо[әќ usum мо[әќ мо[әќ MO[[9Á HYM зицм лојоо ломој4
o[duirs pajoassip ој шта HPOJ ejerojpuj "иша 10 "оуд əjdwıs/ тој ayeuurd валво/]
aytsoddo "ye 10 ‘ddo отеилоује aytsoddo ansoddo 9jeuoj[e ayisoddo aysoddo JUQUISFURLUE je]
48q19u y squay sqniys sqniys sqniys sqnuus «SI9QUII/SQNAYS злодшо иден
‘dds олороигуј pupoufo suagsauids usa wnaojfipnu отојћилоту опуђођоћиј y -14], ununusD f
SOW D40pouapy DIOPOUI jy штитшрј штитиој 29s штитшврој 51098 umunuspf ‘pes штитшврј
"мороигуј pue wnunusof jo exe) ројоојов Jo волтвој Витуетаице(ј "ү AQEL
Volume 84, Number 4
1997
Rohwer 855
Jasminum mesnyi (Oleaceae)
odora) is rarely used as a diagnostic character (or
only as a supplementary character, as by Verdoorn,
1956, 1963), but is frequently mentioned in de-
scriptions. However, M. robusta (Benth.) A. Gray
and M. spinescens are quite robust, erect shrubs,
whereas J. parkeri Dunn is a prostrate to decum-
bent shrublet and J. siamense Craib is (always?) a
weak subshrub, both species not more than 30 cm
tall. More frequently used as a diagnostic character
is the relative length of the corolla tube and the
position of the anthers (e.g., Kupicha, 1983). Co-
rolla tubes that are about as long as the corolla
lobes or longer, with included anthers, are char-
acteristic of Jasminum, whereas exserted anthers
and corolla tubes much shorter than the lobes are
supposed to characterize Menodora. This character
fails, however, in M. longiflora (Engelm.) A. Gray,
a North American species, which has long corolla
tubes and included anthers, but otherwise is typical
of Menodora. Taylor (1945) and Johnson (1957) use
chromosome number (n = 11 in Menodora vs. n =
13 in Jasminum) as a supplementary character in
the diagnosis of the genera. However, it cannot yet
be accepted as proven that this is a real disconti-
nuity. While Jasminum appears to have been ade-
quately sampled by now, with counts of n = 13
from numerous species belonging to different sec-
tions of the genus and coming from different regions
of the world, the same cannot be said about Men-
odora. To my knowledge, neither the aberrant M.
spinescens nor any South American or South African
species has been examined so far.
The seed structure of Jasminum mesnyi and J.
nudiflorum differs from that of all other species of
both Jasminum and Menodora so far investigated.
In the other species of Jasminum, the seeds may
be plano-convex in cross section when there are
two per locule, but they are never even nearly as
unequal-sided as in J. mesnyi and J. nudiflorum.
Our knowledge about seed structure in Menodora
is still inadequate. Only two species have been ex-
amined so far (Rohwer, 1995a), M. spinescens and
M. africana, only the latter in all stages of devel-
opment. This species has an exotesta of small cells,
in which the outer wall becomes so much thickened
as to occlude the lumen (almost) completely, even
more than on the antiraphal side in J. mesnyi.
a consequence of this thickening, which occurs
long before maturity in M. africana, the exotestal
cells can neither enlarge nor divide any further as
the seed grows, so that the exotesta becomes dis-
continuous at maturity. The following layers are
similar to those near the edges of the seed in J
mesnyi. The hypodermal layer consists of very large
cells with conspicuous thickenings in the anticlinal
walls, followed by a few layers of much smaller
cells with much smaller, more irregularly distrib-
uted thickenings. A gap between the outer two lay-
ers and the rest of the seed coat, as on the raphal
side in J. mesnyi, can even be observed in M. af-
ricana, all around the seed but more distinct on the
antiraphal side. In M. spinescens, however, the sep-
aration occurs deeper in the seed coat, between the
outer layers with thickenings and the inner paren-
chymatous part. The fate of the exotesta, unfortu-
nately, could not be followed in the relatively poor
material available of this species. In any case, the
seed of M. spinescens appears more similar to that
of M. africana than to those of any species of Jas-
minum.
In summary, the distinction between Jasminum
and Menodora is not as straightforward as most
treatments of the family imply, and the gap is fur-
ther closed by the data presented here for J. mes-
nyi. Kim and Jansen (1993) found that Jasminum
and Menodora share a 21 kb inversion in the chlo-
roplast genome, and that Menodora and some spe-
cies of Jasminum agree in the loss of clpA gene
introns. This would suggest that Jasminum may be
paraphyletic with respect to Menodora. Unfortu-
nately, their results were published as an abstract
only, so that a full comparison with their data is
still impossible. A similar conclusion was reached,
however, through a detailed study of the fruit and
seed structures of the Oleaceae (Rohwer, 1996).
With the addition of J. mesnyi, it becomes clear
that the discontinuity between Jasminum and Men-
odora is not greater than the gaps encountered
within Jasminum, namely among the species with
(a) alternate leaves and yellow flowers (sect. Alter-
nifolia), (b) opposite leaves and yellow flowers (J.
mesnyi and J. nudiflorum), and (c) opposite leaves
and white flowers (sects. Jasminum, Trifoliolata,
and Unifoliolata). Demoting Menodora to the rank
of a section under Jasminum would perhaps make
the system of the Oleaceae slightly more natural,
but this small gain is overcompensated by the dan-
ger of losing information on the few species of Men-
odora among the hundreds of Jasminum. The most
logical alternative solution, i.e., giving generic rank
to all four groups, should wait until the details of
the molecular analysis become available; it would
also require new generic names for groups a and b.
For the morphological interpretation of the rela-
tionships within the Jasminum-Menodora complex,
it would be most important to know whether the
dehiscent fruits of J. mesnyi were a primitive or an
advanced feature. Loculicidal dehiscence as such
is certainly plesiomorphic within the Oleaceae
(Rohwer, 1996), but it is questionable whether this
Annals of the
Missouri Botanical Garden
is true also for J. mesnyi. On the one hand it ap-
pears unlikely that such a functional character
should be at first lost and then secondarily regained
within a family, but on the other hand the construc-
tion of the mechanical tissue in J. mesnyi is entirely
different from that in the other Oleaceae. In these
we find a sharply delimited endocarp, consisting of
fibers that are stretched transversally to diagonally
on the flanks of the fruit, but longitudinally along
the suture. This pleads clearly in favor of an in-
dependent origin. It is tempting to add that it would
be difficult to understand how and why three dif-
ferent modes of dehiscence, loculicidal, irregular,
and circumscissile, would have evolved within in
the Jasminum—Menodora complex when one of
them had already been established. This, however,
is a weak argument, because the presence of a non-
functional suture in the species with irregular de-
hiscence (J. nudiflorum, M. spinescens) shows that
such a development must have occurred anyway.
Considering the morphological differences within
Menodora and the widely disjunct distribution of
the genus (southwestern North America, southern
South America, and South Africa), it may be asked
whether Menodora is even monophyletic. For the
bulk of the genus, there is little doubt. The com-
bination of herbaceous habit, four ovules per loc-
ule, and separate, circumscissile mericarps is too
unusual to have arisen more than once. Menodora
spinescens, however, shows none of these character
states, so that its placement within this genus (e.g.,
as opposed to Jasminum) may appear questionable.
However, its minute leaves, short corolla tube, ex-
serted anthers, and separate, dry mericarps sharply
distinguish it from Jasminum, and make it far more
similar to the typical Menodora species than to any
other taxon. In its habit it resembles the more ro-
bust species of this genus, especially M. robusta.
Therefore, it makes little sense to remove M. spi-
nescens from the genus, unless it can be shown that
it is not the closest sister group to the remaining
species. Further studies are needed to elucidate its
relationships within this complex.
e unexpected morphological and anatomical
data presented here, in a supposedly well-known
species, thus raise a number of questions that pres-
ently cannot be answered with confidence. The pur-
pose of the present paper is to bring them to notice,
for consideration in future studies.
Literature Cited
ah pad’ = B. 1916. Jasminum primulinum. In: Scientific
C e Report, Feb. 22, 1916. J. Roy. Hort
: xliv.
Green, P. 5. 1961. Studies in the genus Jasminum 1. Sec-
tion Alternifolia. Notes Roy. Bot. Gard. Edinburgh 23:
55—384.
5. Studies in the genus Jasminum III. The
species in cultivation in North America. Baileya 13:
137-172.
———— 1994. 80. Oleaceae. Pp. 327-334 in A. E. Or-
chard & A. J. С. Wilson (editors), Flora of Australia,
vol. 49, Ocio Islands 1. Australian Government Pub-
lishing ue Canberra.
904. Jasminum primulinum (Hemsley). Fl. €
9.
Igersheim, A. 1993. The character states of the Caribbean
monotypic endenie Strumpfia (Rubiaceae). Nordic J.
Bot. 13: 545-
Johnson, L. A. 4 ғ А review of the seed Oleaceae.
Contr. New سو Wales Natl. Herb. 2: 3 8.
R. K. Jansen. 1993. Phylogeny of Oleaceae
based on ndhF sequence i and chloroplast ge-
cts, XV International Bo-
5)
1983. Oleaceae. Pp. 300-327 in E. Lau-
nert (editor), Flora Zambesiaca, и 7, part 1. Flora
Zambesiaca Managing Committee, London
Mabberley, D. J. 1987. The Plant-book. Cambridge Univ.
Press, Cambridge.
Rohwer, J. G. 1993. A preliminary survey of the fruits and
seeds of the Oleaceae. Bot. Jahrb. аам ма 271-291.
994. Seed characters in Jasminum (Oleaceae):
Un звања support for De Can dolle’s | sections. Bot.
Jahrb. Syst. AE 299-319.
. Fruit and seed structures in Menodora
(Oleaceae): A comparison with Jasminum. Bot. Acta
108: 163-168.
. 1995b. Seed characters in Jasminum (Oleaceae).
IL rae from additional species. Bot. Jahrb. Syst.
117: 299-315.
— ———. 1996. Die Frucht- und Samenstrukturen der Ole-
aceae—eine vergleichend-anatomische Untersuchung.
Biblioth. Bot. 148: 1-177.
mares J. A. 1932. A gh owe of the genus Menodora.
n. Missouri Bot. Gard. 1 —176.
пала Н. 1945. mo x phylogeny of the Ole-
aceae. Brittonia 5: 337-367.
Turner, B. L. 1991. An overview of the North American
=> of Menodora (Oleaceae). Phytologia 71: 340—
56
995. Menodora gypsophila (Oleaceae), a new
species from near Galeana, Nuevo León, Mexico. Phy-
tologi 8-9.
Vélos I. C. 1956. The Oleaceae of Southern Africa.
Bothalia 6: 549-640.
963. Oleaceae. Pp. 100—128 in R. A. Dyer, L.
E Codd & H. B. Rycroft (editors), Flora of sae het
Africa, Vol. 26. The Government Printer, Pretoria.
REVISION DEL GENERO
GALIANTHE SUBG. EBELIA
STAT. NOV. (RUBIACEAE:
SPERMACOCEAE)!
Elsa L. Cabral? y Nélida M. Bacigalupo?
RESUMEN
Se subordina al género Galianthe (Rubiaceae: Spermacoceae) el subgénero Ebelia (Rchb.) E. L. Cabral & Bacigalupo,
a las especies que presentan frutos de mericarpos
indehiscentes. Se reconocen nueve especies: Galianthe
ehis
bogotensis (Kunth) E. L. Cabral & Bacigalupo, Galianthe brasiliensis (Spreng.) E. L. Cabral € Bacigalupo, Galianthe
э» dh J
1:
Sucre & С. С. Costa) Е. L. Cabral, Galianthe dichotoma
cymosa (Cham.) E. L. Cabral & Bacigalupo, G
(Willd. ex Roem. & Schult.)
cr
E. L. Cabral & Bacigalupo, Galianthe hispidula (A. Rich. ex DC.) E. L. Cabral &
Bacigalupo, Galianthe polygonoides sp. nov., G. humilis sp. nov. y G. vaginata sp. nov.; las tres tiltimas viven en Brasil.
Todas se describen e ilustran.
ABSTRACT
A new subgenus, Ebelia (Rchb.) E. L. Cabral & Bacigalupo, is proposed within the genus Galianthe (Rubiaceae:
Spermacoceae). The species included therein are characterized by their fruits with indehiscent mericarps. Nine species
are recognized: Galianthe bogotensis (Kunth) E. L. Cabral & Bacigalupo,
Cham.) E. L. Cabral & Bacigalupo,
& Bacigalupo, Galianthe cymosa (
Galianthe brasiliensis (Spreng.) E. L. Cabral
Galianthe dichasia (Sucre & C. G. Costa) E. L.
Cabral, Galianthe dichotoma (Willd. ex Roem. & Schult.) E. L. Cabral & Bacigalupo, Galianthe hispidula (A. Rich. ex
iy
DC.) E. L. Cabral & Bacigalupo, Galianthe polygonoides sp. nov., G. humilis
. nov., and G. vaginata sp. nov.; the
last three are found in Brazil. All the species are described and illustrated.
La delimitacién de algunos géneros de la tribu
Spermacoceae (Rubiaceae) ha planteado dificulta-
des que se manifiestan en el distinto criterio de los
autores sobre este tema.
Schumann (1891) reconoció en esta tribu 18 gé-
neros, 13 de los cuales estaban representados en
América. Posteriormente a esta lista se sumaron
nuevos géneros de 1 6 2 especies: Diacrodon
Sprague, Diodella (Torr. & A. Gray) Small, Micro-
sepalum Urb., Tortuella Urb., Spermacoceodes Kun-
tze, Tobagoa Urb., que no han sido totalmente
aceptados. Sin embargo la mayoría de los autores
en estudios florísticos regionales han seguido en
término generales el criterio de Schumann, introdu-
ciendo sólo cambios menores. Verdcourt (1975:
301) señaló la dificultad en reconocer ciertos gé-
neros casi tinicamente por la dehiscencia de los
frutos, sobre todo ante material florífero y además
por el escaso valor que él le asigna a este carácter
unitario. Este autor amplía los límites de Sperma-
coce L. y subordina a éste las especies de Borreria
G. Mey. y Diodia L., si bien resalta que Diodia
virginiana L., especie americana y tipo del género,
tiene frutos de estructura diferente a las especies
de Diodia del área africana. Verdcourt (1975) sefi-
aló que Steyermark (en litt.) concordó en gran parte
con su opinión, pero a pesar de ello reconoció a
estos tres géneros separadamente, ya que admitió
que se podían diferenciar bien, evitando de este
modo numerosos cambios nomenclaturales. En el
tratamiento de las Rubiáceas de Venezuela, Stey-
ermark (1974) mantuvo este criterio. Actualmente
algunos autores se inclinan por apoyar con varian-
tes el criterio de Verdcourt, así Chaw y Sivarajan
(1989), Fosberg et al. (1981), Howard (1989) y Bur-
he American Genera of the Tribe Spermacoceae
| Est iet i rte del trabajo Delimitation of t а
sta comunicación constituye una pa J Systematics of the Rubiaceae (1993), St. Louis,
(Rubiaceae) presentado durante la International Conference |
i rera que realizó las diagnosis latinas
(MBM, PACA, R, RB, SP, SPF), Estados Unidos (F, MO, NY, US) y A
facilitaron sus colecciones.
2 Facultad de Ciencias Exactas,
Argentina.
3 Instituto de Botánica Darwinion—C.C. 22, 1642 San I
Físicas y Naturales y Agrimensura (UNNE+IBONE, С.С.
on the
y a los curadores de los herbarios de Brasil
rgentina (BA, BAB, CTES, LP, SI) que nos
209, 3400 Corrientes,
sidro, Buenos Aires, Argentina.
ANN. MISSOURI BOT. GARD. 84: 857-877. 1997.
858 Annals of the
Missouri Botanical Garden
Tabla 1. Caracteres del Ebelia subg. Ebelia y del subg. Galianthe.
Subg. Galianthe
Subg. Ebelia
Fruto de mericarpos dehiscentes
Flores siempre heterostilas
Semillas a veces complanadas, con bordes aliformes
Tallo nunca alado
Cromosomas x = 8
Cerca de 40 sp., América del Sur
10-358
Fruto de mericarpos indehiscentes
Flores casi siempre heterostilas, excepto en G. dichotoma
adas
Semillas nunca complan
Hábito erecto, con frecuencia xilopodio muy desarrollado Hábito variado, erecto, postrado, trepador
alado
Tallo casi siempre
Cromosomas x = 14, 15
9 sp., Centro y Sudamérica
20°N-35°S
ger y Taylor (1993) reconocieron Diodia y Sper-
macoce y subordinaron en este género las especies
de Borreria.
De ahí que un estudio global de los géneros de
Spermacoceae analizando el mayor número de car-
acteres de las especies que los representan, es ne-
cesario para fundamentar los límites y las rela-
ciones de los mismos. Con este fin se ha continuado
con el estudio de los géneros Diodia y Borreria. De
estos se han separado algunas pocas especies que
por sus caracteres se han asimilado al género Gal-
ianthe (Cabral, 1991; Pire € Cabral, 1992), pero
del que se diferencian por los frutos de mericarpos
indehiscentes. Por esto se propone ampliar los lím-
ites del mismo e incluir las especies estudiadas en
un nuevo subgénero: Galianthe subg. nov. Ebelia
(Rchb.) E. L. Cabral & Bacigalupo.
MATERIALES Y MÉTODOS
Este estudio se ha realizado con material de los
herbarios nacionales y extranjeros cuyas siglas se
registran de acuerdo con Holmgren et al. (1990)
(BA, BAB, BR, CTES, F, HAS, ICN, G, K, LIL, LP,
MBM, MNES, MO, NY, P, PACA, R, RB, SI, SP,
SPF, US).
TRATAMIENTO TAXONÓMICO
En el estudio emprendido de las especies amer-
icanas de los géneros Spermacoce, Borreria y Diod-
га se han separado unas especies que no reunen
los caracteres de estos géneros: Borreria dichasia
Sucre & С. С. Costa; Diodia brasiliensis Spreng.,
D. brasiliensis var. angulata (Benth.) K. Schum., D.
cymosa Cham., D. dichotoma (Willd. ex Roem. &
Schult.) K. Schum. y D. hispidula A. Rich. ex DC.;
y Spermacoce bogotensis Kunth.
De esta lista, Diodia dichotoma fue descrita er-
róneamente en el género Knoxia L., ya que este
género es considerado actualmente en una tribu in-
dependiente, Knoxieae, que se caracteriza por pre-
sentar flores con óvulos de placentación apical,
péndulos, de micrópila sápera. Este error fue en-
mendado y la especie fue reubicada sucesivamente
en los géneros Spermacoce, Borreria, y Trio
por distintos autores y finalmente como Diodia di-
chotoma (Schumann, 1889).
Borreria dichasia fue descripta como Borreria a
pesar de no haber contado con material con frutos.
Diodia cymosa y D. hispidula fueron descritas den-
tro del género Diodia por sus frutos de mericarpos
indehiscentes. Spermacoce bogotensis fue reubicada
en el género Diodia o Borreria por distintos autores.
Diodia brasiliensis Spreng. fue separada por De
Candolle (1830) junto con D. anthospermoides Cham.
& Schltdl. y D. polymorpha Cham. & Schltdl., en un
nuevo género, Triodon. Este autor caracterizó a dicho
género por el hábito sufruticoso, las inflorescencias
en fascículos espigados y los frutos de mericarpos in-
dehiscentes y lo nominó haciendo alusión a los tres
diminutos dientecitos, correspondientes a restos de
hacecillos vasculares, persistentes en el ápice de los
pedicelos al caer los frutos. Posteriormente iro
agregó dos nuevas especies, THodon a.
México y T. laxum de ки. Este género по fue
aceptado por algunos autores (Schumann, 1888; Stan-
Феу, 1930; Steyermark, 1974) у posteriormente sus
especies se reconocieron dentro del género Diodia,
como uno, dos 6 tres taxones distintos, por tener frutos
de mericarpos indehiscentes. De Candolle al describir
Triodon по registró el dimorfismo floral, a pesar de
que incluyó como basónimo де 7 polymorphus a
polymorpha Cham. & Schltdl. En esta especie
sus autores diferencian variedades y dan una des-
cripción detallada de las flores brevistilas у longistilas
que tienen, si bien no usan estos términos. Schumann
(1889) consideró a las tres especies citadas por De
Candolle, sinónimos de Diodia polymorpha, a pesar
de que registró a una especie anterior D. brasiliensis
Spreng. (1825) entre sus sinónimos.
grupo de especies aquí estudiado se correspon-
de con los caracteres del género Triodon, pero este
nombre genérico está invalidado por un homónimo
anterior (Tabla 1) y en su lugar debe usarse Ebelia
Volume 84, Number 4
1997
Cabral y Bacigalupo 859
Galianthe Subg. Ebelia
Rchb. Pero Ebelia es muy similar a Galianthe Griseb.,
género rehabilitado recientemente (Cabral, 1991; Pire
& Cabral, 1992). Ambos presentan inflorescencias
amplias de ramificación cimosa, flores dimorfas y di-
fieren por los frutos, de mericarpos dehiscentes en
Galianthe e indehiscentes en Ebelia. Por lo tanto se
propone ampliar los términos de la definición de Gal-
ianthe y subordinar Ebelia como: Galianthe Griseb.
E Ebelia (Rchb.) E. L. Cabral & Bacigalupo.
En este nuevo subgénero se incluyen los siete tax-
ones mencionados y tres especies nuevas de la flora
ism Galianthe humilis E. L. Cabral & Bacigal-
po, Galianthe polygonoides E. L. Cabral & Bacigal-
oa y Galianthe vaginata E. L. Cabral & Bacigalupo.
Galianthe Griseb., Abh. Kónigl. Ges. Wissensch.
Góttingen 24: 157. 1879. Borreria G. Mey.
sección Galianthe (Griseb.) K. Schum., en
Martius, Fl. Brasil. 6(6): 40-42. 1888. Borrer-
ia G. Mey. subg. Galianthe Sopa Standl.,
Field Mus. Hist. Bot. Ser. 8 (5): 392. 1981.
TIPO: Galianthe fastigiata Griseb.
Se amplian los límites de Galianthe y se recon-
ocen dos subgéneros sobre la base de la diferencia
de dehiscencia de los frutos:
subg. Galianthe
la. Frutos de mericarpos dehiscentes ..
subg. Ebelia
lb. Frutos de mericarpos indehiscentes -...
Galianthe Griseb. subg. Ebelia (Rchb.) E. L. Ca-
bral & Bacigalupo, comb. et stat. nov. Ebelia
pa in Bot. Herb. Buch. 74. 1841. Trio-
Prodr. 4: 566. 1830, non Baumg.
Tb. TIPO: Triodon polymorphus DC. [=
Galianthe brasiliensis (Spreng.) E. L. Cabral &
Bacigalupo] (lectótipo, aquí designado).
Hierbas perennes, sufrátices o pequefios arbus-
tos, erectos, trepadores o postrados. Tallos tetrágo-
nos, en general alados. Hojas con la nervadura
marcada o apenas visible. Vaina estipular fimbria-
da, breve, rara vez tubulosa, prolongada por encima
de la inserción del par de hojas correspondiente.
Inflorescencias complejas, tirsoides o cimoidales,
inflorescencias parciales en dicasios más o menos
congestos, en fascículos o cabezuelas subglomeri-
formes, rara vez inflorescencias simples, en cimo-
ides laxos. Flores tetrámeras, heterostilas, pilosas
en el interior del tubo corolino. Frutos de dos mer-
icarpos indehiscentes, con línea de dehiscencia
preformada sobre la línea media de la cara adaxial.
Semillas reticuladas o foveoladas con estrofíolo en
la cara placentar. Granos de polen isopolares, ra-
diosimétricos, entre 25—50 jum, oblato-esferoidales
o prolato-esferoidales, estéfano-colporados, de exi-
na semitectada-reticulada, con retículo complejo o
simple (Galianthe bogotensis y G. dichotoma) (Pire,
1996). 2n = 28 (С. bogotensis); n = 15 (G. bras-
iliensis subsp. angulata). [Los otros detalles bajo G.
brasiliensis subsp. angulata.]
CLAVE PARA DIFERENCIAR LAS ESPECIES DE GALIANTHE
SUBG. EBELIA
la. Vaina estipular Марини prolongada рог ка
de la inserción del par de hojas correspondien
2a. Vaina pilosa; tallos di ángulos че» Brasil
9. G. vaginata
2b. Vaina glabra; tallos sin alas; Brasil ...........
. polygonoides
lb. Vaina estipular breve, no sobrepasando la inser-
de hojas correspondiente
4a. mee simple
axo, nudos floríferos con Кабык
ias; Ecuador, Perú ............ 5. G. dichotoma
4b. Uie rr compuestas, nudos flor-
íferos con flores en fascículos o glomé-
rulos
5a. Cáliz de 4 sépalos; PAET i
compuestas сито ida inflores-
cencias parciales тег.
mes; Colombia ........... gotensis
; Cáliz de 2 ó 4 sépalos; inflorescen-
id
сл
=
formes о cim
cias parciales en fascículos; México
entina p
a
3b. Hojas con nervios зое зигсадов е
п inflorescen-
arciales congestas, неча"
Ta. ` Tallos de a potione ala-
ie hojas m lat.; Brasil,
raguay, UNE > Argentina c
. dichasia
ludere нета e Be cymosa
6b. Inflorescencia cimoidal de ramas cin-
cinoides, con inflorescencias parciales
paucifloras, fasci culadas.
8a. Tallos simples; ето sub-hemisfér-
tx de — ong.; Brasil,
raguay, Argentina .. 6. G. hi.
8b. ie деседе frutos turbina-
os de 5 mm long.; Brasi
ad hos 4 7. G. humilis
1. Galianthe bogotensis (Kunth) E. L. Cabral &
Bacigalupo, comb. nov. Basónimo: Spermacoce
bogotensis Kunth, in Humb., Bonpl. & Kunth,
Nov. Gen. Sp. Quarto ed. 3: 347, Folio ed. 3:
271. 1819. Diodia bogotensis (Kunth) Cham.
& Schltdl., Linnaea 3: 350. 1828. Borreria bo-
gotensis (Kunth) Standl., Publ. Field Colum-
bian Mus., Bot. Ser. 7: 160. 1930. TIPO: Co-
lombia. Crescit juxta urbem Santa Fe de
у" alt. 1370 hex, Humboldt & Bonpland
n. (holótipo, B destruido; foto F860, CTES,
SE lectótipo, aquí designado, P). Figura 1.
Annals of the
860
Missouri Botanical Garden
Volume 84, Number 4
1997
Cabral y Bacigalupo
Galianthe Subg. Ebelia
Borreria anthospermoides DC., Prodr. 4: 550. 1830. TIPO:
Colombia. In Amer. austr. prope Sta.-Fe de Bogotá,
colector desconocido, ex herb. Delessert, 1816 (hol-
биро, С; foto #6689, CTES, SI).
Sufrútice muy ramificado, semipostrado a postra-
do, formando una extensa mata, de tallos radican-
tes, estrechamente alados, glabros, escabriúsculos
o pilósulos hacia el ápice de los entrenudos, con
brotes axilares que dan apariencia verticilada a los
nudos foliares. Hojas de 6-18 X 1. mm, an-
gostamente elípticas u ovales, de base y ápice agu-
dos y margen revoluto, sólo con la vena central
manifiesta, subglabras, escabriúsculas sobre el
margen, a veces algo sobre el haz y la vena media
en el envés. Vaina estipular de 1-2.5 mm long.,
pubescente, con 5-7 lacinias de 2.54 mm long.
Inflorescencia terminal cimoidal, con inflorescen-
cias parciales, subglomeriformes, a veces en ramas
laterales reducidas a 1 sola cabezuela. Cáliz con 4
segmentos triangulares y algún dientecito interca-
lar, sólo en los bordes escabriúsculos. Hipanto tur-
binado de 1.5-2.5 mm long., glabro a subglabro.
Corola blanca, micropapilada en la superficie ex-
terna; disco entero. Flor brevistila: corola de 3.5-
5 mm long., con lóbulos iguales o más cortos que
el tubo corolino en su interior con pelos monilifor-
mes cortos y delgados sobre el tubo y pelos gruesos
sobre el tercio inferior de los lóbulos; anteras de
1-1.5 mm long. y filamentos estaminales de 0.6–
1.2 mm long.; estilo de 1.5-3 mm long., de ramas
estilares filiformes de 0.5-1.8 mm long. Flor lon-
gistila: corola de 4-5.5 mm long., de lóbulos más
cortos que el tubo, de superficie interna con pelos
dispersos sobre el tubo y pelos más densos, gruesos
y largos en la base de los lóbulos; anteras de 1–
1.3 mm long. y filamentos de 0.5-1 mm; estilo de
4-5 mm y ramas estilares de 2.5-0.9 mm long.
Fruto de 2-3.2 mm long., glabro, de mericarpos
indehiscentes. Semilla de 2-2.3 mm long., super-
ficie foveolada, con ancho surco longitudinal en la
línea media ventral, cubierto parcialmente por el
estrofíolo.
Distribución (Fig. 11). Habita en el páramo o en
área vecina en Colombia, entre 2200-3575 m s.m.,
en borde de ciénagas, matorral subserial, pastizal
о rastrojos.
Material examinado. COLOMBIA. Nueva Granada,
1857, Triana 75 (NY, P). Boyacá: Cordillera Oriental,
>
Páramo de Belén, 6 mayo 1959, Barclay & P. Juajibioy
7573 y 7661 (MO, NY); Socha, 8 nov. 1944, Fosberg
22225 (NY); Santuario de Yguaque, 26 mar. 1981, Me-
lampy 1313 (MO). Cundinamarca: Boquerón de Chi-
paque, 16 mar. 1939, Killip 34201 (F, NY); Páramo de
Guasca, 15 dic. 1938, Balls 5696 (K); Páramo de Siberia,
19 Feb. 1944, Fosberg 214
Bogotá, 13 set. 1917, Pennell 1934 (NY); Sabana
of Bogotá, supra hill, 4 Feb. 1945, Schiefer 402 (F); Bo-
gotá, Nov. 1852, Holton 417 (K), 29 Oct. 1975, André 728
(K); above La Cita, 10 mayo 1946, Schultes 7122 (F); près
de Bogotá, Triana 3123 (P); Boquerón de Bogotá, 21 nov.
1975, André 72 (K); Bogotá, 20 set. 1913, Apolinar 74 (F);
Montecillo E of Guatavita, 29 mayo 1947, Haught 5774
(F); Paipa, ene. 1938, Jimenez 68 (F).
Esta especie se parece por sus inflorescencias a
G. cymosa y G. dichasia, pero se diferencia de am-
bas por sus hojas más pequeñas y subenervias. Su
área está restringida a los páramos de Bogotá y
zonas circunvecinas, mientras que las otras dos es-
pecies son del SE del Brasil, la primera citada sólo
para el estado de Paraná y la segunda desde Paraná
llega hasta el Paraguay, Argentina y Uruguay; am-
bas de terrenos bajos y húmedos.
2. Galianthe brasiliensis (Spreng.) E. L. Cabral
& Bacigalupo, comb. nov. Basónimo: Diodia
brasiliensis Spreng., Syst. Veg. 1: 406. 1824.
: Brasil, Sello (holótipo, B destruido);
Brasil. Sin loc., afio 1828, Pohl s.n. (neótipo,
aquí designado, С; foto F6728, SI).
Subarbusto de 0.30-0.70(-1) m de alto. Tallos te-
trágonos, de ángulos marginados a estrechamente ala-
dos, glabros a hírtulos. Hojas opuestas, o en general
seudoverticiladas, 3-35 X 0.7-11 mm, ovadas o elfp-
ticas, agudas, atenuadas en seudopecíolo, escabriús-
culas en el margen y los nervios en el envés a glabras,
nervadura poco visible, con excepción del nervio me-
dio; vaina estipular 3-7 laciniada, glabra o hispídula.
Inflorescencia frondosa a bracteada, tirsoide, espici-
forme a pleiotirsoide con inflorescencias parciales
también espiciformes о cimoidal, de ramificación di-
casial a monocasial, siempre con flores congestas, fas-
ciculadas en los nudos floríferos. Flores brevemente
pediceladas. Cáliz 2 6 4 partido, con dientecitos in-
tercalares. Hipanto turbinado, glabro o hírtulo. Corola
A A A RS
x
Figura 4. Galianthe a. —A. Planta. —
gu cymos - Vaina estipular. —C-E. Flor brevistila. F-H. Flor longistila. —I.
Fruto. A-E, Tesmann et al. 749; F-I, Dusén 2695.
|
|
a ————M
Volume 84, Number 4
Cabral y Bacigalupo 867
Galianthe Subg. Ebelia
SUE
` Figura 5 ianthe dichasia. —A.
Gal. Planta
EL "Flora longistila. —J. Fruto. —K. Mericarpo. A-F, J,
en resalto en el envés. Vaina estipular pilosa, 3-5
mm long., 5—7 lacinias glabras, de 2-10 mm long.
Inflorescencia cimoidal, dicasial a monocasial, con
inflorescencias parciales subglomeriformes. Cáliz
de 4 segmentos aleznados, 1.6-2.8 mm, a veces
__В. Vaina estipular. —C. Corte del tallo. —D-F. die залай
К, Krapovickas et al. 44888; G-L Imaguire
ligeramente menores, opuestos, a menudo con dim-
ab
glabro. Corola blanca, externamente micropapilada,
con papilas notables en el dorso apical de los lób-
868
Annals of the
Missouri Botanical Garden
Ws a
"4
00 У ۵
PITA A
А
Volume 84, Number 4
1997
Cabral y Bacigalupo 869
Galianthe Subg. Ebelia
ulos, estos más cortos que el tubo, superficie inter-
na con dos anillos de pelos moniliformes, uno sobre
el tubo y otro de pelos algo más gruesos sobre la
base de los lóbulos; disco entero. Flor brevistila:
corola 4—5.2 mm, anteras 1-1.2 mm, filamentos es-
taminales 0.5-1.5 mm, estilo 2.5—4.7 mm, ramas
estigmáticas de 2-2.2 mm de largo, con papilas
ensas. Flor longistila: corola 3.7-5 mm, anteras
0.7-1 mm, filamentos estaminales 0.2 mm y estilo
de 4–5.8 mm long. Fruto 2.5-3.5 mm long. Semilla
2 mm de largo, plano-convexa con estrofíolo cub-
riendo parcialmente la cara ventral.
Distribución (Fig. 11). Esta especie vive en ter-
renos bajos, pantanosos, orilla de vertientes. Su
área abarca el SE del Brasil (Paraná, Sta. Catarina
y Rio Grande do Sul), Paraguay (San Pedro,
Guairá), Uruguay (Rivera) y Argentina (Misiones).
Material examinado. BRASIL. Paraná: Rincao, 9
feb. 1947, Hatschbach 624 (CTES, SP); Curitiba, 28 mar.
1950, Tessmann 67 (MBM); 5 km E de Curitiba, 2 feb.
1973, Krapovickas et al. 23093 (CTES, LP), 14 feb. 1978,
Krapovickas et al. 33657 (CTES); Piraquara, 3 mar. 1971,
Imaguire 2781 (ICN), 31 ene. 1967, Dombrowski 2391
TES). Santa Catarina: Сасадог, 22 dic. 1956, Smith
et al. 9073 (R). Rio Grande do Sul: Torres, Lagoa dos
Quadros, 21 feb. 1950, Rambo 45992 (CTES, LIL); Far-
roupilha, 7 feb. 1950, Rambo 45708 (CTES); Caxias do
Sul, 8 feb. 1955, Rambo 56689 (PACA), 1 ene. 1949, B.
Rambo 30882 (PACA), 1932, Augusto s.n. (PACA); S. Leo-
1 ; Bom Jesús, 13 ene.
1941, B. Rambo 8521 (PACA), 15 ene. 1942, Rambo
4.
(РАСА); Cambará, feb. 1948, Rambo 36712 (PACA).
PARAGUAY. San Pedro: 10 km W de San Estanislao,
13 ene. 1941, Rambo 3973 (PACA). ARGENTINA. Mi-
siones: Gral. Belgra
G
tín, Ao. Garuhapé, Mroginski et al. 378 (C ;
Galianthe dichasia guarda estrecha relación con
G. cymosa de la que se diferencia por sus tallos
más robustos, de ángulos bien marcados a mani-
fiestamente alados y sus hojas anchas.
5. Galianthe dichotoma (Willd. ex Roem. 4
Schult.) E. L. Cabral & Bacigalupo, comb. nov.
Basónimo: Knoxia dichotoma Willd. ex Roem.
& Schult., Syst. Veg. 3: 532. 1818. TIPO:
América meridionalis. Humboldt (holótipo, B
no visto, foto B-W 2676, F 862, CTES, SI).
Figura 6.
Diodia glabra Willd. ex Roem. & Schult., Syst. Veg. 3:
532. 1818, non Pers. (1805).
Spermacoce dichotoma Willd. ex Kunth, in Humb., Bonpl.
& Kunth, Nov. Gen. Sp. Quarto ed. 3: 348; Folio ed.
3: 272. 1819. Borreria dichotoma (Willd. ex Kunth)
Cham. & Schltdl., Linnaea 3: 340. 1828. Diodia di-
chotoma (Willd. ex Kunth) K. Schum., in Mart., Fl.
brasil. 6 (6): 11. 1888. TIPO: Perú. Prope Peruvi-
anorum et in declivitate montis ignivomi Tungura-
uae, alt. 1400-1600 hex, Bonpland (holótipo, P,
foto F37152, CTES, SI).
Triodon laxum [Triodia laxa] Bentham, Pl. Hartw.: 194.
1840. TIPO: Juxta ponte Guapulo, prope Quito, Har-
tweg 1068 (holótipo, K no visto).
Sufrátice con ramas postradas de 10-30 cm
long., tallos tetrágonos con alas estrechas, de mar-
gen escabriúsculo. Hojas ovales, de 7-20 X 2-7
mm, ápice acuminado y base atenuada en corto
seudopecíolo, margen revoluto, escabriúsculo, dis-
coloras, haz escabriúscula a glabra, con vena media
surcada, poco conspicua, envés escabriúsculo sobre
la vena media sobresaliente, y a veces sobre los 2—
3 pares de nervios poco conspicuos. Vaina estipular
1.7-3.5 mm long., pilosa, con 3-7 lacinias glabras,
de 1.54 mm long. Inflorescencia cimoide, dicasial,
con brácteas foliáceas en los nudos basales, dis-
minuyendo a linear-subuladas, diminutas en nudos
apicales. Cáliz bipartido, de 2 segmentos triangu-
lares, carnosos, de bordes escabriásculos, casi
siempre alternando con algunos dientecitos inter-
calares; hipanto obcónico, 1–1.2 mm long., esca-
briúsculo en la mitad superior. Corola 4(—5)-lobada,
2538 mm long. de lóbulos más cortos que el
tubo, micropapilada por fuera, con papilas notables
en el dorso apical de los lóbulos y con pelos mon-
iliformes en su interior. Anteras 0.7–0.8 mm long.
y filamentos breves. Estilo 0.5-1.2 mm long., con
ramas de 0.5 mm long. Fruto turbinado 3-3.5 mm
long., escabriúsculo a subglabro. Semilla 2.5-3 mm
long., subobovoide, de superficie foveolada, con es-
trecho surco en la cara ventral cubierto parcial-
mente por el estrofíolo.
Distribución (Fig. 11). Habita en zonas altas, en-
tre los 2300-3340 m s.m. en Ecuador hasta el norte
de Perú.
Material examinado. ECUADOR. El Oro: Hacienda
Chepel, headwaters of Río San Luis, 22 km E of Zaruma,
13 Feb. 1945, Fosberg & Giler 23007 (NY). Loja: upper
Malacatos Valley, 15-20 km 5 Loja, 6 July 1944, Prieto
36 (NY); Paso de Cajanuma above Pueblo Nuevo, 9 feb.
1945, Fosberg & Giler 22889 (P); Loja, W of town, 3 Oct.
ll RE ОБИ dut кї Dedi mE у a
€—
Figura 6. Galianthe dichotoma. —A. Planta.
—H. Fruto. —1. Mericarpo. —J.
Barclay et al. 8454.
—B. Vaina estipul
Base del fruto. —K. Semilla no madura, cara
ar. —C, D. Flor longistila. —E-G. Flor brevistila.
ventral. A-D, H-K, Cerón 2253; E-G,
870
Annals of the
Missouri Botanical Garden
1955, Asplund 17894 (G, P); San Pedro Loja, 26 abr.
1946, Espinosa 246 (NY). Pichincha: Cantén Quito, Par-
roquia Calacali, Reserva Geobotánica Pululahua, alrede-
dores de Pailon, 25 sep. 1987, Cerón 2253 (CTES, MO);
ad margen viae ad flumen Machángara pr. Quito, 16 abr.
1920, Holmgren 522 (G); cóté de Guápulo, 20 mar. 1930,
Benoist 2195 (P); in Andibus Ecuadorensibus, 1861, R.
(G, P); Quito, sin fecha, Jameson 792 (G).
PERU. Sin localidad, sin fecha, Mattheros 1502 (BR).
nas: Leimabamba, 13 dic. 1962, Woytkow.
az
(MO). Cajamarca: camino a la
timbamba, Marcabal, 26 feb. 1949, Infantes Vega 1902
(LIL), 18 ago. 1952, Infantes Vega 3738 (LIL).
En esta especie no se manifiesta un claro di-
morfismo floral, pero se incluye en el subg. Ebelia
por sus inflorescencias de ramificación cimosa y
sus frutos de mericarpos indehiscentes.
6. Galianthe hispidula (A. Rich. ex DC.) E. L.
Cabral € Bacigalupo, comb. nov. Basónimo:
Diodia hispidula A. Rich. ex DC., Prodr. 4:
565. 1830. TIPO: Brasil. Brésil Leandro do
Sacramento 103, año 1819 (holótipo, P no vis-
to, foto F37153 CTES, SI). Figura 7.
Diodia paradoxa Cham., Linnaea 9: 216. 1834. TIPO:
Brasil. Brasilia, Sellow (holótipo, B destruido, foto
F869, CTES, SI); Brasil. Minas Gerais: 3-1-1846,
Widgren 1025 (neótipo, aquí designado, BR).
Sufrútice rizomatoso, de tallos simples, de 30—
60 cm de alt., tetrágonos, alados, glabros o rala-
mente pilosos sobre los ángulos o con escasas
emergencias o lacinias con pelo apical. Hojas 35—
100 X 15-35 mm, discoloras, vinosas al envejecer,
elípticas u ovales, de ápice agudo o acuminado y
base obtusa, atenuada en seudopecíolo, con 5-8
pares de nervios secundarios, ligeramente curvos y
paralelos, haz glabra o escabriúscula sobre toda la
lamina o sólo sobre los nervios y bordes; vaina es-
tipular escabriúscula o pilosa, con 5-9 lacinias
hasta 10-13 mm long., glabras o algo escabriús-
culas en la base. Inflorescencia cimoidal, mono-
casial a pleiocasial, con paracladios en gran parte
cincinoides, con inflorescencias parciales contrai-
das, fasciculadas. Flores hipanto turbinado, esca-
briúsculo. Cáliz 4-partido, de 2 sépalos de 2.5-3
mm long, triangular-subulados, alternando con
otros 2 sépalos menores y algún dientecito inter-
calar, glabro, de borde escabriúsculo, o rara vez
también con alguna papila en el dorso de los sé-
palos. Corola blanca, 4-lobada, 3.8-4.6 mm long.,
de lóbulos más cortos que el tubo. Flor brevistila:
superficie interna de la corola con pelos dispersos
sobre el tubo y base de los lóbulos, anteras 0.8—
1.2 mm long., estilo 0.8—1.6 mm long. y ramas es-
tilares de 1 mm. Flor longistila: superficie interna
de la corola, con pelos moniliformes gruesos sobre ,
los lóbulos y delgados sobre el tubo; anteras 0.8
mm long., estilo 3.5—4.5 mm long., ramas estilares
de 1-1.2 mm. Fruto subhemiesférico, 1.6-2 mm
long., de superficie escabriúscula a glabra. Semilla
de color castaño oscuro, 1.4—1.6 mm long., con es-
trofíolo rodeado por surco profundo.
Distribución (Fig. 11). Hierba umbrófila, habita
en bosques y selvas del sur del Brasil, Paraguay y
NE de la Argentina. Florece y fructifica desde la
primavera hasta entrado el огоћо.
Material seleccionado. BRASIL. Minas Gerais: Cal-
das, Regnell I 178 (P), 1845, Widgren 202 (K); Bandeira
do Sul, 20 ene. 1980, Krapovickas et al. 35405 (CTES).
Paraná: Porto Helena, 1 feb. 1949, Schwarz 7425
(CTES); Campo Nov. Laranjeiras do Sul, 8 dic. 1968,
Hatschbach 20563 (NY). Rio de Janeiro: Nova Friburgo,
morro da Caledonia, 8 jun. 1977, Martinelli 2538 (RB).
Rio Grande do Sul: Reserva Nonoai, 7 dic. 1974, Porto
1951, Rambo 49879 (US). PARAGUAY. Alto Paraná:
die. 1950, Montes 11101 (CTES, LP). ARGEN-
A. Misiones: in distr. urb. Posadas, praecipue in vi-
cin. coloniae Bonpland, sin fecha, W. Lillieskold s.n. (G);
L. N. Alem, Alem, Camping Municipal, 22 nov. 1986,
Cabral et al. 496 (CTES); Gral. Belgrano, San Antonio,
Soberbio, 6 dic. 1983, Hunziker et al. 10801
SI); Iguazú, Reserva Apepú, 4 mar. 1982, Ferraro 2463
(CTES); Ayo. Uruguaí, nov. 1949, M. Crovetto et al. 5777
(BAB, SI); Parque Nacional Iguazú, 11 ene. 1972, Mro-
ginski et al. 300 (CTES), 1 dic. 1993, Vanni et al. 3082
(CTES), 15 nov. 1976, Guaglianone 174 (51); 8 km del
cruce de Libertad, Salto Uruguaí, 9 dic. 1983, Hunzi
=,
54548), 28 nov. 1884, Nied-
erlein 366 (BA); Ayo. Uruguaí, km 10, 14 ene. 1963, Par-
tridge s.n. (ВА 61693); Ођега, 24 dic. 1970, Mroginski
80 (СТЕ i
(LP); Salto Tabay, 14 oct. 1977, Cabrera et al. 28771 (SD;
San Javier, S. Javier, nov. 1962, Mi
(SD, 8 di
993, Rodríguez et al. 557 (CTES). Corrientes:
Santo Tomé, Ea. Timbó, 8 dic. 1981, Tressens et al. 1598
(CTES).
А Galianthe hispidula se la reconoce fácilmente
por ser esta una hierba sufruticosa, estolonífera, de
tallos simples. Comin en sotobosque.
Youn 84, Number 4 Cabral y Bacigalupo 871
Galianthe Subg. Ebelia
ум“
(Ег; 7
— ==
=
M
N WIDE,
Y A
Ca
ATT
RN m
" d
Figura 7. Galianthe hispidula. —A. Planta. — B. Corte transversal del tallo. —C. Vaina estipular. —D, E. Flor
brevistila. —F, G. Flor longistila. —H. Fruto. — ]. Semilla, cara ventral. —J. Corte transversal de semilla. A-C, F-J,
Martínez Crovetto et al. 5777; D, E, Martínez Crovetto et al. -
7. Galian umilis : Suffrutex caulibus tetragonis, glabris, anguste alatis,
^ пен И кам & dein" alis glabris vel scabriusculis. Folia sessilia 25—45 m
sp. nov. TIPO: Brasil. Minas Gerais: Fouso longa, 7-9 mm lata, elliptica, apice acuto, base attenuata,
Alegre, 2 mayo 1927, F. C. Hoehne 19357 пегүіѕ secundariis ibus, conspicuis, supra gla
(holótipo, SP). Figura 8. vel scabriuscula. Vagina folii 3 mm longa aliqui pilosa,
872
Annals of the
Missouri Botanical Garden
Figura 8. Galianthe humilis. —A. Rama. —B-D. Flor longistila. —E. Fruto. A-D, Souza León 11; E, Hoehne
19357.
3-7 laciniata. Flores in cymis compositis dispositi. C ve
tetrasectus, 1.2-2 m segme
linus sequantibus, intus
dense perno Flos A ай corolla 3.6 mm lo
ва, antheris 1 mm longis, stylo 2 mm longo, ramis stig-
matibus 0.6 mm longis. Flos лй ignotus. Fructus
5 mm longus, subturbinatus, lateraliter WEAR com-
pressus, сет ашан: eolatus іп ѕіссо. Semen іп-
maturum m longum, A pa A
faciem бисте in parte tegen
Hierba sufruticosa de tallos tetrágonos, glabros,
con alas angostas, glabras o escabriúsculas. Hojas
sésiles, de 25-45 X 7-9 mm, elípticas, de ápice
agudo y base atenuada con 3-4 pares de nervios
secundarios, marcados en la haz y en resalto en el
envés, glabras o algo escabriúsculas en la haz sobre
todo hacia el margen, éste recurvado. Vaina esti-
pular 3 mm de long., algo pilosa con 3-7 lacinias
glabras, hasta de 4 mm long. Inflorescencia cimo-
idal con inflorescencias parciales fasciculadas, par-
е ee
Figura 9. Galianthe polygonoides. —A. Ram
—G.
-—
a. —B. Vaina estipular. —C, D. Flor brevistila. —E, F. Flor longistila.
Fruto. —H. Semilla no madura, cara "n A-D, G, H, Braga 1741; E, F, S 190.
ucre 4
873
Cabral у Bacigalupo
Galianthe Subg. Ebelia
RL ҺӘМ. Ц Ala
Volume 84, Number 4
1997
о
Annals of the
874
Missouri Botanical Garden
Volume 84, Number 4
1997
Cabral y Bacigalupo 875
Galianthe Subg. Ebelia
vifloras hasta flores solitarias en algunos nudos.
Cáliz 4-partido, 1.2-2 mm long., de segmentos
triangulares, 2 ligeramente mayores, con algún
dientecito en los senos intercalares, glabros o de
borde escabriúsculo. Hipanto glabro, subturbinado,
1.8 mm long. Corola blanca, 4-lobada, 3.6 mm
long. de lóbulos tan largos como el tubo corolino,
con densa pubescencia en su interior. Flor longis-
tila: corola 3.6 mm long., anteras 1 mm long., estilo
2 mm long., ramas del estigma de 0.6 mm de largo.
Flor brevistila: no vista. Fruto 5 mm long., subtur-
binado, algo comprimido lateralmente, glabro, os-
curamente reticulado-foveolado al secarse. Semilla
inmadura, 3 mm long., reticulado-foveolada con es-
trofíolo cubriendo parcialmente la cara ventral.
Distribución (Fig. 11). Se ha coleccionado hasta
el momento sólo en Brasil, en el Estado de Minas
Gerais, en campos altos, de suelos rocosos.
Material examinado. BRASIL. Minas Gerais: Car-
angola, 1400 m s.m., 20743'5, 42729", 22 jul. 1988, L.
Souza Leon 11 (SPF); sin localidad, 1858, Weddell s.n. (С).
Se asemeja a Galianthe cymosa, siendo ésta más
grácil, de tallos finos, minutamente papilosos, con
inflorescencias parciales densas y corolas de lób-
ulos más cortos que el tubo.
8. Galianthe polygonoides E. L. Cabral 4 Ba-
cigalupo, sp. nov. TIPO: Brasil. Rio de Janeiro:
Petrópolis, Vale do Bonsucesso, 750 m s.m.,
15 nov. 1969, D. L. S. Braga 1741 (holótipo,
RB; isótipo, CTES). Figura 9.
Suffrutex scandens, caulibus fistulosis, glabris, ramis
briusculo. Vagina stipularis glabra conspicuissima 7-13
mm de largo, 6—7 laciniis, inaequalibus 1-5 m go
Inflorescentia th monochasia, dichasia, vel pleo-
» .
chasia); inflorescentiis partialibus cymosis contractis, fas-
ciculatis, interse distinctis (cincinndeis) hypanthio sca-
i ] ntis
briusculo 1.5-2 mm де la
5 mm, 2 mericarpis
indehiscentibus, sepala persistente. Semina + plana-con-
vexa, strophiolo faciem ventralem in parte in linea media
tegente.
Sufrútice trepador, tallos fistulosos, subcilíndri-
cos a tetrágonos, glabros, entrenudos de 6-9 cm de
largo, brotes axilares escasos a nulos. Hojas de 40-
60 X 10–14 mm, elípticas a lanceoladas, de ápice
agudo a acuminado y base atenuada en breve pseu-
dopecíolo, glabras, con bordes revolutos, escabriús-
culos, levemente discoloras, con 3—4 pares de ven-
as secundarias subopuestas. Vaina estipular muy
conspicua, tubulosa, glabra, 7-13 mm long., con
6-7 lacinias. Inflorescencia cimoidal (monocasial,
dicasial o pleocasial), con paracladios cincinoides.
Cáliz con 4 segmentos triangular-subulados de 1.5-
2.5 mm long., escabriúsculos, a veces con dientes
intercalares; hipanto obcónico, escabriúsculo, 1.5—
2 mm long. Corola blanca, externamente micropa-
pilada, con papilas más densas y más grandes en
el dorso apical de los lóbulos; superficie interna,
con pelos moniliformes sobre el tubo hasta la base
de los lóbulos; disco bipartido papiloso. Flor brev-
istila: corola 3-3.2 mm long., lóbulos iguales o más
cortos que el tubo corolino; filamentos estaminales
1-1.5 mm long., anteras 1 mm long., estilo 2-3 mm
long., bifurcado en su extremo. Flor longistila: cor-
ola 3-3.5 mm long., lóbulos más cortos que el tubo
corolino, filamentos estaminales muy breves y an-
teras 0.7-1 mm, estilo 3.54 mm long., ramas es-
tilares filiformes 1 mm, densamente papilosas. Fru-
to subturbinado, escabriúsculo, 2-2.5 mm long.,
separándose en dos mericarpos indehiscentes, sé-
palos persistentes. Semilla más o menos plano-con-
vexa, 1.7-2 mm long., cara ventral cubierta par-
cialmente por el estrofíolo en la línea media, cara
dorsal foveolada.
Distribución (Fig. 11). Esta especie fue encontra-
da hasta el presente, sólo en rasil, en el Estado
de Rio de Janeiro, Petrópolis a 750—800 m s.m.
Material examinado. BRASIL. Rio de Janeiro: Pe-
trópolis, 24 nov. 1968, D. Sucre et al. 4190 (RB); Caetetu,
1 jul. 1943, G. Goés et al. 240 (RB); Carangola, 29 ago.
1943, G. Goés et al. 500 (RB); Grota do Jacó, 800 m s.m.
2 nov. 1968, D. Sucre et al. 4013 (US), 2 dic. 1971, J.
Barcía 349 (R); Araras, Ma. Comprida, 30 ene. 1971, Ur-
bano 9927 (K); Serra da Estrella, 1844, Weddell 44 (P).
Galianthe polygonoides se individualiza por su
hábito trepador y por la vaina estipular muy nota-
ble, tubulosa por encima de la inserción de las ho-
jas, a modo de una ocrea, similar a la que presenta
G. vaginata de la que se distingue por tener esta
vainas pubescentes y tallo marcadamente alado.
9. Galianthe vaginata E. L. Cabral & Bacigalu-
po, sp. nov. TIPO: Brasil. Minas Gerais: Sierra
do Itatiaia, ad marginem viae, ca. 1800 m s.m
25 mayo 1902, P. Dusén 109 (holótipo, R). Fi-
gura 10.
ES sudden MN POR
+=
Figura 10. Galianthe vaginata. —A. Rama. —B. Vaina estipular. —C. Bot
109.
ón. —D-F. Flor longistila. A-F, Dusén
876
Annals of the
Missouri Botanical Garden
| = = = = rj 50 40
= rs [ 110
9 мав)
30 Y w.
a A ITE О e З
ae eea ass “л RSI RRS ВН ПА БН a B e cii
кы “бш pm 2 a de
-— a о =e FEX. Bites
i: "а
AM
Ы / Ола; 1
ма" x + []
" ~ е
"4 +т =
\
P \ E 10
4 2
&
10) Md
C A A
‘So, p c
" А À b
„“ AS |
У;
; :
Ea
* с bogotensis | 2 7 E
O ©. brasiliensis 4 E ; X c
~ 5 z :
О С. brasiliensis subsp. angulata M :
10 т С. cymosa — run
E i 4 "I BEN
ж G. dichasia Yo Ea 1 |
i £T А
А G dichotoma A j
k A. u^ ТФ
* С. hispidula de A E. CM ILS : y 4
ee 4 AAA
20} Dm
зг С. polygonoides 4 E
i А 1
T © G. vaginata ~
{ ~
М lo
O 200 400 600 800 1000km j
је
0 100 200 300 400 500 600 miles if
100 90 80 "UN 60 50 40
subtus scabriuscula plerumque in nervis et mar:
u 5 mm longus, ramis 0.8-1 mm
longis. Flos brevistylus et fructus haud visus.
Sufrútice de tallos tetrágonos notoriamente ala-
dos. Hojas de 30-90 х 10-24 mm, discoloras,
elípticas, de ápice agudo o acuminado y base aten-
uada, con 4-5 pares de nervios secundarios, haz
glabra con diminutas papilas sobre los nervios, en-
vés escabriúsculo esencialmente sobre los nervios
y los bordes; vaina estipular tubulosa, prolongada
por encima del par de hojas correspondiente, 6-12
mm de largo, pubescente, de bordes laciniados. In-
florescencia cimoidal con inflorescencias parciales
fasciculadas, distanciadas entre sí. Flores heteros-
tilas, hipanto turbinado, escabriúsculo. Flor longis-
tila: cáliz de 4 segmentos triangular-subulados, 2
de 1.5-2 mm long. alternados con 2 ligeramente
menores. Corola blanca, 4.54.7 mm long., de lób-
ulos más cortos que el tubo, con pelos dispersos en
el interior del tubo corolino hasta la mitad de los
lóbulos. Anteras 1-1.2 mm long., filamentos muy
breves. Estilo 3.5—4.5 mm long., ramas estilares de
-8-1 mm long. Flor brevistila y frutos no vistos.
Distribución (Fig. 11). Esta especie es brasileña
y habita en los Estados de Minas Gerais y Sáo Pau-
lo.
aterial examinado. BRASIL. Sáo Paulo: Campos
do Jordáo, in campestris dumetosis, ene. 1944, E. Frider-
Ц
Volume 84, Number 4
1997
Cabral у Bacigalupo
Galianthe Subg. Ebelia
ich S. J. s.n. (РАСА 27764). Minas Gerais: prov. Mina-
rum Ad Caldas, 1867, А. Regnell 1: 178 (BR).
ESPECIES DUDOSAS
Triodon polymorphus var. intermedius DC., Prodr.
: 566. 1830. In Brasil meridionali, Sellow.
TIPO: B, destruido. Probablemente existan du-
plicados no localizados. Su autor la caracteriza
por sus hojas de tamaño intermedio.
Diodia polymorpha var. floribunda K. Schum., in
Mart., Fl. Bras. 6(6): 13. 1888. TIPO: no de-
signado, Brasil. Los ejemplares citados: “in
provincia Rio de Janeiro prope Canta Gallo:
Peckolt; Glaziou 3028; in silvis prope Gongo
Soco: Bunbury; loco haud accuratius indicato:
Sello 3433” B, destruidos. Duplicados no lo-
calizados. Según su autor se corresponde con
plantas de hojas pequeñas, 5-13 X 14 mm,
cáliz 2-mero.
Diodia polymorpha var. lasiodisca K. Schum., in
Mart., Fl. Bras. 6(6): 12. 1888. TIPO: Brasil.
“Habita in provincia Minas Geraes prope
Sumidorio: de Langsdorff” (holótipo, В des-
truido). No se han localizado duplicados.
Diodia polymorpha var. densa Zahlbr., Anz. Kaiserl.
Akad. Wiss. Wien, Math.-Naturwiss. K1. 12:
82. 1923. TIPO: Brasil. Sáo Paulo (no visto).
Duplicados no localizados.
Literatura Citada
Bentham, G. 1840. та С. Pamplin (editor), Plantae Наг-
twegianae. Londre
Burger, W. & C. Taylor. 1993. Rubiaceae. En: W. Burger
(editor), Flora nd Fieldiana, Bot. 33: 124.
Cabral, E. L. . Rehabilitación del género Galianthe
(Rubiaceae). y Soc. Argent. Bot. 27: 235-249.
Candolle, A. P. de. 1830. Rubiaceae, Spermacoceae. En:
Treutel € Wiirtz (editores), Prodromus systematis na-
turalis regni vegetabilis 4: 538-578. Paris.
Chaw, 5. M. & У. Sivarajan. 1989. Seed coat micromor-
phology of some Asiatic Spermacoceae (Rubiaceae).
Bot. Bull. Acad. Sin. 30: 15-24.
Fosberg, F. R., M. H. Sachet € R. L. Oliver. 1981. Ru-
biaceae. Flora of Micronesia. Smithsonian Contr. Bot.
44—
Grisebach, A. 1879. Symbolae ad Sm argentinam.
. 153-159. dressed Gótting
gren & L C. Barnett. 1990.
tation, Univ.
. M. 1997. its Galianthe subg. Ebelia (Rubi-
aceae: Spermacoceae): Estudio palinológico. Ann Mis-
souri Bot. Gard. 84: 878-887.
abral. 1992. El valor del polen en la
revalidación de — (Spermacoceae—Rubiaceae).
pe ЗНА Зрегтасосеае. Еп: С. Е.
Ру , A. W. Eichler & I. Urban (editores),
M ons, 6(6): 6-398; 401-409. Fleischer,
Leipzig.
. 1891. Rubiaceae, Spermacoceae. En: A. Engler
& K. Prantl, Die qon Pflanzenfamilien 4, 4:
146. Engelmann, Lei
Sprengel, C. 1825. Systema Vegetabilium
Standley, P. C. 1930. The Rubiace - d Colomba Publ.
Field petywi Mus., Bot. Ser
938. Rubiaceae. En: uta Ф боне Rica. Publ.
[5 же Hist., Bot. ic 18: TE E
Macr 74. Rubiac e: Spermacoceae. En:
Lasser ај Flora де марас 9(3): 1834-1978.
Inst. Botánico, Caracas
Verdcourt, B. 1975. Studies i in the Rubiaceae—Rubioideae
| the Flora of Tropical East Africa: I. Kew Bull. 30:
—326.
Nec Hire T. G. 1938. Rubiaceae. En: A Contribution of
the Flora of Honduras. Field Mus. Nat. Hist., Bot. Ser
17: 287-407.
GÉNERO GALIANTHE SUBG. Stella Maris Pire?
EBELIA (RUBIACEAE:
SPERMACOCEAE): ESTUDIO
PALINOLOGICO!
RESUMEN
Se estudia el polen de las especies de la tribu Spermacoceae conocidas actualmente como: Borreria anthospermoides
DC., Diodia brasiliensis Spreng., D. cymosa Cham., D. hispidula A. Rich. ex DC., Knoxia dichotoma Willd. ex Roem.
& Schult. у D. brasiliensis var. angulata (Benth.) Standl., y se señala su afinidad con el polen del género Galianthe
Griseb. Este estudio palinológico refuerza el criterio sustentado por Cabral y Bacigalupo de ampliar los límites del
género Galianthe e incluir estas especies en el nuevo subgénero Ebelia (Rchb.) E. L. Cabral € Bacigalupo, junto con
С. dichasia (Sucre € С. С. Costa) E. L. Cabral, G. humilis E. L. Cabral € Bacigalupo у С. polygonoides E. L. Cabral
& Bacigalupo, por sus frutos de mericarpos indehiscentes.
ABSTRACT
The pollen of the species of tribe Spermacoceae currently known as Borreria anthospermoides DC., Diodia brasiliensis
Spreng., D. cymosa Cham., D. hispidula A. Rich. ex DC., D. dichotoma (Willd. ex Roem. & Schult.) K. Schum., and
brasiliensis var. angulata (Benth.) Standl. is studied, and its affinity with the pollen of the genus Galianthe Griseb.
is indicated. This palynological study reinforces the suggestion of Cabral and Bacigalupo to extend the boundaries of
the genus Galianthe to include the species listed above, because of their fruits with indehiscent mericarps, in the new
Galianthe subg. Ebelia (Rchb.) E. L. Cabral € Bacigalupo, together with С. dichasia (Sucre & С. С. Costa) E. L.
Cabral, G. humilis E. L. Cabral € Bacigalupo, and С. polygonoides E. L. Cabral & Bacigalupo.
Este trabajo es parte de un proyecto más amplio
que comprende el estudio palinológico de toda la
tribu Spermacoceae. Esta es una de las tribus de
Rubiaceae que incluye varios géneros conflictivos
o críticos cuya delimitación es discutida, como
Spermacoce L., Diodia L., Borreria G. F. W. Meyer,
etc. (Steyermark, 1974; Verdcourt, 1976; Fosberg
et al., 1981; Howard, 1989; Burger & Taylor, 1993,
entre otros). El estudio palinológico pretende apor-
tar datos que ayuden a la redefinición de los gé-
neros, emprendimiento encarado junto con N. Ba-
cigalupo y E. Cabral.
Hasta el momento se ha estudiado el polen de
145 especies representantes de la totalidad de los
géneros (16) de las Spermacoceae citadas para
América. Al realizar este estudio se encontró que
algunas especies del género Diodia L. poseen gra-
nos de polen con retículo complejo, ésto es con un
suprarretículo y un infrarretículo a un nivel más
bajo que el anterior. En Rubiáceas, este tipo de
ornamentación, hasta el momento, sólo ha sido des-
cripto para Galianthe Griseb. (Pire & Cabral,
1992). Este género ha sido recientemente revali-
dado por Cabral (1991).
Diodia, en sentido amplio, es un género que in-
cluye taxones de caracteres muy dispares que no
se corresponden con su definición genérica. Estos
caracteres involucran al fruto, inflorescencia, mor-
fología floral y también al polen. Bacigalupo y Ca-
bral (inédito) reconocen, hasta el momento, 4 es-
pecies como válidas de Diodia (D. virginiana L.,
D. kuntzei K. Schum., D. macrophylla K. Schum. y
D. saponariifolia Cham. €: Schldl.), principalmente
por presentar frutos indehiscentes, y proponen la
reubicación de las restantes especies en otros gé-
neros. El primer paso para la reubicación de las
mi agradecimiento a las siguientes personas
arios mencionadas en este trabajo
gen
* Facultad de Ciencias Exactas y Naturales y Agrimensura (U.N.N.E.)-PRINGEPA, C.C. 128, 3400-Corrientes, Ar-
tina.
ANN. MISSOURI Bor. GARD. 84: 878-887. 1997.
Volume 84, Number 4 Pire 879
1997 Estudio Palinológico
Cuadro 1. Combinaciones nuevas propuestas por Cabral у Bacigalupo (1997) con sus nombres actuales equivalen-
tes.
Nombres actuales
Combinaciones nuevas
Borreria anthospermoides DC.
Borreria dichasia Sucre & С. С. Costa
Diodia brasiliensis Spreng.
Diodia brasiliensis var. angulata (Benth.) Standl.
Diodia cymosa Cham.
Knoxia dichotoma Willd. ex Roem. & Schult.
Diodia hispidula A. Rich. ex DC.
Galianthe bogotensis (Kunth) E. L. Cabral & Bacigalupo
Galianthe dichasia (Sucre & C. G. Costa) E. L. Cabral
Galianthe brasiliensis (Spreng.) E. L. Cabral & Bacigalupo
Galianthe brasiliensis subsp. angulata (Benth.) E. L. Ca-
bral & Bacigalupo
Galianthe cymosa (Cham.) E. L. Cabral & Bacigalupo
Galianthe dichotoma (Willd. ex Roem. & Schult.) E. L.
Cabral & Bacigalupo
Galianthe hispidula (A. Rich. ex DC.) E. L. Cabral & Ba-
cigalupo
Galianthe humilis E. L. Cabral & Bacigalupo
Galianthe polygonoides Е. L. Cabral & Bacigalupo
Galianthe vaginata Е. L. Cabral & Bacigalupo
especies lo constituye la inclusién de Diodia bras-
iliensis Spreng., D. cymosa Cham., D. hispidula A.
Rich. ex DC., Knoxia dichotoma Willd. ex Roem.
& Schult., Diodia brasiliensis var. angulata (Benth.)
Standl., en Galianthe (Cabral & Bacigalupo, 1997,
en este volumen). Al mismo tiempo se incorpora
Borreria anthospermoides DC.; todas estas especies
poseen, como Galianthe, frutos dehiscentes, flores
heterostilas о dimorfas e inflorescencias casi siem-
pre complejas, tirsoides о cimoidales, con inflores-
cencias parciales en dicasios más o menos conges-
tos. Si bien, a diferencia de Galianthe que posee
frutos con mericarpios dehiscentes, los mericarpios
de estas especies son indehiscentes. Por esta razón
se las ha agrupado en un nuevo subgénero: G. subg.
Ebelia (Rchb.) E. L. Cabral & Bacigalupo (1997).
Dentro de este subgénero estan comprendidas ade-
más, Galianthe dichasia (Sucre & С. G. Costa) E.
L. Cabral, G. humilis E. L. Cabral & Bacigalupo,
С. polygonoides E. L. Cabral & Bacigalupo y б.
vaginata E. L. Cabral & Bacigalupo.
En el Cuadro 1 se da las equivalencias entre los
nombres conocidos y las combinaciones nuevas
propuestas por Cabral y Bacigalupo, de las espe-
cies comprendidas en Galianthe subg. Ebelia.
ANTECEDENTES
De este grupo de especies son pocas las que han
sido estudiadas anteriormente desde el punto de
vista palinológico. Galati en su Tesis Doctoral (Fa-
cultad de Ciencias Exactas y Naturales, Universi-
dad de Buenos Aires, 1988, inédito), al estudiar la
embriología de la tribu Spermacoceae, analiza el
polen de 7 géneros y 19 especies, entre ellas Diod-
ia brasiliensis y ya señala la coincidencia de car-
acteres palinológicos y embriológicos entre esta ез-
pecie y otras dos, Borreria fastigiata (Griseb.) K.
Schum. y B. laxa Cham. & Schldl., que ahora se
consideran dentro de Galianthe (Cabral, 1991).
Esta coincidencia se refleja también, en la tipología
del megagametófito (Galati, 1991).
En un trabajo anterior (Pire & Cabral, 1992), al
analizar el valor del polen en la revalidación del
género Galianthe, se hace referencia a la morfología
polínica de G. dichasia y de Borreria anthosper-
moides. Esta última especie no fue incorporada
dentro de Galianthe por Cabral (1991), por presen-
tar frutos con mericarpos indehiscentes y granos de
polen con retículo simple, quedando pendiente, en
ese momento, su clasificación genérica. А Borreria
dichasia Sucre € С. G. Costa se la incorporó como
Galianthe dichasia (Sucre & С. С. Costa) E. L. Са-
bral, a pesar de sus frutos de mericarpos indehis-
centes, porque en ese momento no se disponía de
material fructificado y el resto de los caracteres
coincidían bien con Galianthe.
MATERIAL Y MÉTODOS
Para el estudio de los granos de polen se utilizó
anteras de botones florales bien desarrollados, ob-
tenidos de material de herbario. Los ejemplares tes-
tigos se encuentran depositados en los siguientes
Herbarios: CTES, MBM, MO, NY, SI, SP y RB.
El material palinológico, para su observación con
microscopio óptico, fue tratado de acuerdo a la téc-
nica de acetólisis de Erdtman (1966), utilizándose
gelatina-glicerinada como medio de montaje. Di-
chos preparados se hallan depositados en la Pali-
noteca de la Universidad Nacional del Nordeste
(PAL-CTES). Para la observación con microscopio
electrónico de barrido (MEB) se utilizó polen ace-
tolizado y en algunos casos polen natural, en ambos
880
Annals of the
Missouri Botanical Garden
casos se los lavó varias veces con agua destilada у
se los montó en un trocito de “papel España.”
De cada especie se estudió el polen de flores
ейн у brevistilas. Sobre un minimo де 30
granos, se midieron los parámetros: eje polar y diá-
metro ecuatorial; en un menor número de granos se
midieron: el espesor de la exina, largo y ancho de
los colpos, diámetros mayor y menor de las en-
doaperturas, distancia entre colpos. Además se cal-
culó la relación entre longitud del colpo o ectoap-
ertura (ECA) y longitud del grano o eje a (P) y
se clasificó los colpos en: cortos (ЕСА/Р = 0.20-
0.35), medianos (ЕСА/Р = 0.36–0.45) y paa
(ЕСА/Р = 0.46–0.65). Estas observaciones se efec-
tuaron utilizando un microscopio óptico Leitz Or-
thomat. Otras mediciones como, ancho de los muros
del retículo, diámetro de los lúmenes, largo y ancho
de las espínulas, se realizaron sobre la base de fo-
tomicrografías tomadas con el microscopio elec-
trónico de barrido JEOL-JSMT 100 del Servicio de
Microscopía Electrónica de la Facultad de Ciencias
Naturales y Museo de la Universidad Nacional de
ata.
La terminología utilizada es básicamente la de
Erdtman (1966) y se consultó también Kremp
).
A continuación se cita el material examinado
bajo el nombre de la nueva combinación propuesta
por Cabral y Bacigalupo y entre barras el basónimo
correspondiente.
MATERIAL EXAMINADO
Galianthe bogotensis d E. L. Cabral & Bacigalupo
[Borreria anthospe rmoi DC.]. Flores brevistilas: CO-
LOM oyacá: Soc ti F сву 22225 (NY) [PAL-
CTES 3792]. NUEVA GRANADA. J. J. Triana 75 (NY)
a CTES E Flores И COLOMBIA. Воу-
: Сога. ntal, Páramo de Belén, Н. Barclay & Р.
Jua 7661 (мо) [PAL-CTES 3120]. Cundinamarca:
dillera Oriental, Páramo, H. García Barriga 16136
(NY) [PAL-CTES 3790].
heres i bare (Spreng.) E. L. Cabral & Baci
galupo [Diodia brasiliensis Spreng.]. Flores мез
PARAGUAY. Encarnación, oy-caé, Bertoni 2401
(CTES) = ane 3175]. Flores longistilas: ARGENTI-
: Candelaria, Cerro Сога, Bertoni 2604
(CTES) [PAL-CTES 3174].
lianthe brasiliensis subsp. angulata arse E LE
Cabral & Bacigalupo [Triodon gm Benth.]. Flores
brevistilas: Pringle 10803
о [PAL-CTES 3404]. Flores as КАД. MEXIC
ra ле El Candeje, Gimate 852 (CTES) [PAL-CTES
OMM cymosa (Cham.) E. L. Cabral & ae ee
[Diodia cymosa Cham.]. Flores brevistilas: BRASIL. Р
rana: Curitiba, Barigui, Ferreira 186 (MO) [PAL-CTES
3794
Galianthe dichasia (Sucre & C. G. Costa) E. L. Cabral
[Borreria dichasia Sucre & C. G. Costa ]. Flores brevis-
tilas: BRASIL. Paraná: Curitiba, Tessmann 67 (MBM)
[PAL-CTES 3077]. Flores longistilas: BRASIL. Paraná:
Ringao, Hatschbach 624 (SP) [PAL-CTES 3076].
Galianthe dichotoma (Willd. ex Roem. & Schult.) E. L.
Cabral & Bacigalupo [Knoxia dichotoma Willd. ex Roem.
& Schult.]. Flores brevistilas: ECUADOR. Azuay-Loja:
Nudo de Cordillera Occidental y Cord. Oriental, entre Оћа
y Rancho Ovejero, Barclay et al. 8454 (MO) [PAL-CTES
3114]. Flores longistilas: ECUADOR. Pichincha: Cantón
Quito, C. Cerón 2253 (MO) [PAL-CTES 3953]. PERU. Ca-
јатагса: camino a laguna Chamis, Sánchez Vega 759 (SI)
[PAL-CTES 3407].
COMME hispidula (A. Rich. ex DC.) E. L. Cabral &
Bacigalupo [Diodia pucr b ме ex DC.]. Flores
rt ARGENTINA. Misiones: L. N. Alem, Cabral
et al. 496 (CTES) [PAL-CTES 3393). ^ lores longistilas:
А INA. Cor a.Timbó, Tres-
sens et al. 1 598 (СТЕ) | [PAL.CTES не Мијо Зап
Ignacio, Schwarz 1699 (CTES) [PAL-CTES 3954].
ianthe humilis Е. L. Cabral & Bacigalupo. Flores
longistilas: BRASIL. Minas Gerais: Pouso Alegre, Hoeh-
ne 19357 (SP) [PAL-CTES 3392].
Galianthe polygonoides Е. L. Cabral & Bacigalupo. Flo-
res brevistilas: BRASIL. Rio de Janeiro: Petrópolis,
Caetetú, D. C. Goés et al. 240 (RB) [PAL-CTES 3788];
Carangola, Goés et al. 500 (RB) [PAL-CTES 3955]. Flores
a BRASIL. Rio de Janeiro: Petropolis, Sucre
4190 et al. (RB) [PAL-CTES 3391].
DESCRIPCIÓN GENERAL DE LOS GRANOS DE POLEN
Las especies de Galianthe subg. Ebelia consti-
tuyen un grupo bastante homogéneo desde el punto
de vista palinológico. Los granos de polen presen-
tan las siguientes características: isopolares, ra-
diosimétricos, tamaño mediano (25-50 um), oblato-
esferoidales o prolato-esferoidales. De acuerdo a
as aperturas son zono-colporados: 6—7 colporados
en Galianthe brasiliensis y Galianthe dichasia
(flores brevistilas) y 8—9(-10) colporados en el resto
de las especies. Los colpos generalmente son largos
y las endoaperturas lalongadas. La exina, de 2.5—
3.5 jum de espesor, es semitectada-reticulada, con
retículo complejo diferenciado en un suprarretículo
(SR), psilado, y un infrarretículo (IR), espinulado,
a un nivel más bajo que el anterior. Se exceptúan
G. bogotensis y G. dichotoma que presentan retículo
simple, de muros espinulados
El SR puede presentar mallas cerradas o abiertas
(con muros interrumpidos), que, generalmente, au-
mentan de tamaño hacia los polos.
El IR se puede encontrar distintamente difer-
enciado; por lo general, se lo observa como puen-
tes, simples o ramificados, que atraviezan los lú-
menes del SR (1), o a veces, reducido a espínulas
ubicadas lateralmente en la base de los muros del
SR (G. brasiliensis y G. humilis) (2), o bien, está
totalmente ausente y en este caso las espínulas se
localizan sobre el SR (G. dichotoma y G. bogotensis)
(3). De este modo, se podrían reconocer 3 estadíos
Volume 84, Number 4 Pire 881
1997 Estudio Palinológico
1 2
Figura 1. Representación gráfica de los estadíos de modificación de la exina entre retícul plejo (1-2) y retículo
simple (3).
de modificación de la estructura de la exina, entre
retículo complejo y retículo simple (Fig. 1), cuya
secuencia evolutiva no puede ser establecida aún
con certeza.
Por otra parte, las flores longistilas y brevistilas
presentan un dimorfismo polínico poco marcado
que está dado por finas diferencias en la ornamen-
tación de la exina y por un tamaño levemente may-
or de los granos de polen en flores brevistilas.
DIFERENCIAS ENTRE LAS ESPECIES
Las principales diferencias entre las especies es-
tan dadas por:
(1) ORNAMENTACIÓN DE LA EXINA
De acuerdo a ella se pueden agrupar las especies
en: con retículo simple o con retículo complejo.
Con retículo simple hay 2 especies: G. bogotensis
y G. dichotoma, que presentan diferencias entre sí.
En la primera el retículo posee lúmenes profundos
y amplios (de hasta 4.5 jum de ancho), entre los
que se intercalan otros menores (0.2 шт) y muros
rectilíneos o suavemente curvilíneos, provistos de
1 sola hilera de espínulas (Fig. 2A, C). En la se-
gunda especie, los lúmenes son comprimidos, de
contorno muy irregular y de menor tamaño (де 2-
um, a veces 3.5 шп de largo X 0.6-1 jum de
ancho), los muros son curvilíneos con 1 6 2 hileras
de espínulas (Fig. 2B, D).
Con respecto al retículo complejo, las variantes
observadas corresponden a los 3 tipos básicos ya
descriptos para otras especies de Galianthe (Pire &
Cabral, 1992):
TIPO I: SR completo en todo el grano; IR poco
visible;
TIPO II: SR interrumpido formando mallas
abiertas en los mesocolpios; IR parcialmente visi-
ble;
TIPO III: SR reducido a fragmentos pequefios en
la zona ecuatorial; IR parcial a completamente vis-
E
e.
El TIPO I es el más frecuente, el TIPO II se
presenta en G. brasiliensis subsp. angulata y en G.
cymosa (Fig. 3A-D) y el TIPO III, sólo en flores
brevistilas de G. polygonoides (Fig. 3E-G). En el
TIPO I los muros del SR son + rectilíneos у los
lámenes poligonales, isodiamétricos o alargados.
En los TIPOS II y III, los muros son curvilíneos y
los lúmenes tienen forma irregular. Con respecto al
IR en algunas especies se observaron variaciones
en la distribución de las espínulas: en б. bras-
iliensis subsp. angulata (flor brevistila) los muros
son pobremente espinulados; en mesocolpios po-
seen 1 6 2 espínulas y hacia los polos tienden a
ser psilados (Fig. 3C); en 6. brasiliensis, y flores
longistilas de G. humilis, у G. polygonoides, las es-
pínulas se localizan, ya sea sobre los muros que
atraviezan los lúmenes de SR o ya sea lateralmente
en la base del SR (Figs. 2G, 3E-G, 4).
(2) LONGITUD DEL COLPO O ECTOAPERTURA
En todas las especies, excepto 6. bogotensis, los
granos de polen poseen colpos largos (ECA/P may-
or de 0.45). Sin embargo, algunas especies, como
G. dichotoma y las flores longistilas de G. bras-
iliensis subsp. angulata y G. dichasia, pueden te-
ner, además, granos con colpos medianos (ECA/P
= 0.40-0.51). En G. bogotensis los colpos pueden
ser cortos o medianos (ECA/P = 0.33-0.43).
Otras diferencias respecto al número de apertu-
ras, forma y tamaño de los granos, diámetro de los
lámenes del SR y ancho de los muros del SR e IR,
se exponen en el Cuadro 2.
DiscusióN Y CONCLUSIONES
En el Figura 5 se resumen, en forma esquemá-
tica, los caracteres exomorfológicos y caracteres
palinológicos de los géneros Galianthe (subg. Ebe-
Annals of the
Missouri Botanical Garden
Volume 84, Number 4 Pire 88
3
1997 Estudio Palinológico
E F,
Ed
s en vista ecuatorial y detalle d
angulata f b. gud 10803). —B, D. G. cymosa,
ocolpio (Goés et al. 240). Las escalas equivalen a 10 jum еп A,B,Ey2 pme
. Mes
£1, flor Конь)
е la боша del apocolpio. —A, С. Galianthe gurges subsp.
Ferr Apoc
a 186). E-G. G. polygonoides, f.b. — ›осо1р1о.
n C, D, F, G. (f.b., i brevistila;
— A, C. Galianthe bogotensis, f. b.
E. f.b. peg 2401). —
o A,B, E,F ya
Granos en vista ecuatorial y detalle de la escultura del apocolpi
2. 1
(Е, „р 22225). —B, D. С. dicet. Е]. (Sanchez Vega 7. G. G. emis —
ы eg 2604). —Е, x^ G. his a, f.l. (Tressens et al. 1 598). je escalas equivalen a 10 pm e
m en C, D, G, H. (f.b., flor и: f.l., flor longistila.)
884 Annals of the
Missouri Botanical Garden
Figura 4. Grano en vista ecuatorial y detalle de la escultura en mesocolpio. —A, B. Galianthe humilis, flor longistila
(Hoehne 19357). Las escalas equivalen a 10 jum en A y
Cuadro 2. Galianthe subg. Ebelia: diferencias respecto al número de aperturas, forma y tamaño de los granos,
diámetro de los lúmenes del SR y ancho de los muros del SR y IR. Referencias: obl-esf., oblato-esferoidal; prol-esf.,
prolato-esferoidal; P, eje polar; E, diámetro ecuatorial; SR, suprarretículo; IR, infrarretículo. Las medidas estan en jum.
Respecto al tamaño de los granos se indica las medidas promedio y de los lúmenes del SR, el diámetro mayor.
Especies
Aperturas
Forma Tamaño Р х E SR muros//lúmenes IR muros
G. bogotensis
fl. long. 8-9 obl-esf. 38.6 X 40.8 0.4–0.5//1(2.6)4.5 —
fl. brev. (7)8–9 obl-esf. 43.0 X 46.5 0.6—0.9//1.4(3.4)4.5 -—
G. brasiliensis
fl. long. 6(7) рг Ê 25.7 X 24 0.3-0.4//0.3(1.5)2.9 0.15-0.4
fl. brev. : (6)7 obl-esf./prol-esf. 2653x272 0.4—0.7//0.7(2.0)3.3 0.35–0.6
>. brasiliensis subsp. angulata
fl. lon 8)10 obl-esf. 263 X 315 — es
E ‘ 9 prol-esf. 36.8 x 36 0.3–0.6//0.2(1.3)2.8 0.3-0.4
G. cymosa
fl. brev. — 8(9) prol-esf. 37.6 X 36.7 0.3-0.4//1.4(2.8)5.8 0.3-0.35
G. dichasia
А. long 8 subprol. 42.7 X 36.2 0.4—0.5//0.5(1.5)2.0 0.35
fl. brev. 7(8-0) subprol. 41.5 х 35.3 0.5—0.6//1.0(1.5).18 0.3-0.5
G. dichotoma
fl. long. 9(10) obl-esf./prol-esf. 34.0 X 34.2 0.5-0.7//0.8(2.5)3.5 vus
|. brev. 9 obl-esf. 41.5 X 35.3 0.5—0.6//1.0(1.5)1.8 0.3-0.5
G. hispidula
. long. (7)8 obl-esf. 33.0 x 34 0.3–0.4//0.4(1.2)2 0.2
|. brev. (8)9 obl-esf./prol-esf. 34.7 х 34 0.4—0.5//1(1.2)3 0.3-0.4
G. humilis
fl. long. _ 8 prol-esf. 37.3 X 36.5 0.4–0.5//1(2.1)3.7 0.3-0.4
G. polygonoides
fl. long. (78 prol-esf. 28.0 x 26.3 0.3—0.5//0.4(1.4)2.2 .2-0.35
fl. brev. (7)8-9 obl-esf. 34.9 x 36.0 0.35//0.9(1.4)3.8 0.35-0.4
885
ire
Estudio Palinológico
Volume 84, Number 4
1997
“01x31 [ә ua иотовоцахо 194 ‘soorsoyouryed
валојовлво SNS А (o13 Á лор *erouaosa10pur) воотојојошохо волојовлео sns ap ase вј a1qos “Hs "s ipo] А омошод “9YTUDIJDA) so19u93 soj әциә зорвршуе А serouarop(] сс BINAL]
"] міаота
19Á9|N
мясо мнзннов
PFUs
tirn "n 1 pan wr
NS
әціиецео '1295
|qesu5 3HLNVIVD 7 ^ ^
ellaq3 1295
886
Annals of the
Missouri Botanical Garden
lia у subg. Galianthe), Borreria y Diodia 8. str.3,
que se consideran de mayor valor taxonómico:
(1) tipo de inflorescencia: tirsoide, con inflorescen-
cias parciales + congestas o empobrecidas, ci-
moides versus glomeriforme o sólo de 1-2-3
flores;
(2) tipo de flor: heterostila (longistila y brevistila)
versus homostila;
(3) tipo de fruto: parcial o totalmente dehiscente
versus indehiscente;
(4) tipo de polen: semitectado-reticulado, con col-
pos largos versus tectado-perforado con colpos
cortos.
Como se puede apreciar, en las especies del gé-
nero Galianthe, las inflorescencias son terminales
y tirsoides y las flores heterostilas. En el subgénero
Galianthe se agrupan las especies con frutos de
mericarpos dehiscentes y en el subgénero Ebelia se
reúnen las que tienen frutos de mericarpos inde-
hiscentes. En el género Borreria las especies pre-
sentan frutos como los de Galianthe subg. Galian-
the, pero se separan bien por sus flores homostilas
y por sus inflorescencias glomeriformes, terminales
y/o axilares. En el género Diodia s. str. las flores
son isomorfas, axilares, solitarias o en inflorescen-
cias paucifloras y los frutos totalmente indehiscen-
En cuanto al polen, la estructura de la exina es
el carácter más importante para distinguir Galian-
the de los otros dos géneros; en segundo lugar se
encuentra la longitud de los colpos (Fig. 6).
En Galianthe se ha estudiado la morfología del
polen de más de 30 especies del subgénero Gal-
ianthe (Pire € Cabral, 1992; Pire, inéd.) en las
culados, con retículo complejo y zonocolporados,
con colpos largos o medianos (ECA/P — 0.40—0.65)
(Fig. 6A). En G. subg. Ebelia, los granos de polen
presentan un retículo complejo o, con menor fre-
cuencia, un retículo simple, siendo los colpos pre-
dominantemente largos.
En Borreria se analizaron cerca de 50 especies
(Cabral, 1985; Pire, inéd.*) que presentan gran var-
iabilidad en sus caracteres palinológicos; sin em-
bargo la estructura de la exina, tectada-perforada,
se mantiene casi constante en todas ellas. En este
género los granos de polen pueden ser colporados
o porados; cuando son colporados, los colpos son
* Diodia s. str. comprende a las cuatro especies recon-
ocidas por Bacigalupo y Cabral: D. virginiana L., D.
kuntzei К. Schum., D. macrophylla К. Schum. y D. sapon-
ariifolia Cham. & Schldl.
‘Trabajo presentado en la II International Rubiaceae
[
ајо
Conference, Meise, Bélgica (septiembre, 1995).
о toons RAIDER
Figura 6. Granos en vista ecuatorial. —A. Galianthe
centranthoides (flor brevistila). —B. Borreria terminalis.
—C. Diodia virginiana. Las escalas equivales a 5 jum.
Volume 84, Number 4
1997
Pire
Estudio Palinológico
generalmente cortos (ECA/P = <0.35) (Fig. 6B).
Las especies con granos de polen longicolpados po-
seen exina tectada-foveolada y espínulas localiza-
das principalmente alrededor de las aberturas
En cuanto a Diodia s. str. (D. virginiana L., D.
kuntzei K. Schum., D. macrophylla K. Schum. y D.
saponariifolia Cham. & Schldl.) posee granos de
polen tectado-perforados y con colpos cortos, sim-
ilares a los ya mencionados para Borreria (Fig. 6
Con este estudio se demuestra que las especies
comprendidas en Galianthe subg. Ebelia presentan
caracteres no sólo exomorfológicos sino también
palinológicos que son típicos de Galianthe, los que,
a su vez, permiten diferenciarlas perfectamente de
las de Borreria y Diodia. Esto apoyaría el criterio
de Cabral y Bacigalupo (en este volumen) de am-
pliar los límites de Galianthe creando el nuevo sub-
género, G. subg. Ebelia (Rchb.) E. L. Cabral & Ва-
cigalupo.
Literatura Citada
Burger, W. & C. M. Taylor. 1993. Rubiaceae, Flora Cos-
taricencis. Fieldiana, Bot. 33: 124.
Cabral, E. L. 1985. Valor taxonómico del polen en las
especies argentinas del género PA LS (Rubiaceae).
Bo EU Bot. 24 (1-2): 169-178.
Rehabilitación del i Galianthe (Ru-
biacéae): ^d Soc. Argent. Bot. 27 (3—4): 235-249.
N. Bacigalupo. 1997. Revisión del género Gal-
ianthe subg. Ebelia stat. nov. (Rubiaceae: Spermaco-
ceae). Ann. Missouri Bot. Gard. 84: 857-877.
Erdtman, G. 1966. Pollen Morphology and Plant Taxono-
my. Angiosperms. Haffner, New York.
Fosberg, F. R., M. H. Sachet & R. L. Oliver. 1981. Ru-
biaceae. Flora of Micronesia. Smithsonian Contr. Bot.
35
1. Estudios embriológicos en la tribu
Spermacoceae (Rubiaceae). Parte 1: Anatomía он).
ра eo eg Megasporogénesis. Bol. Soc. Arg
Bot. 27 (1-2): 7
Howard, R. А. 1989. Rubiaceae. Flora of the Less r An-
968. Morphology Encyclopedia of Pal-
ynology. Univ. Arizona Press, Tucson
Pire, S. M. & E. L. Cabral. 1992. El valor del polen en
Ба A" ción de Galianthe ee p
. Darwiniana 31 (1—4): 1
ы ark, J. А. 1974 маен à oceae, Flor.
i Venn 9 (3): 1834-1978. Instituto Botánico, Ca-
2
Tarak B. 1976. Rubiaceae in R. M. Polhill, Flora of
Tropical East Africa: 333-380. мн {ог Оуегзеа Соу-
ernments and Admin., London
THE TANAKA-KAIYONG Li Xi-wen? and Li Jie?
LINE—AN IMPORTANT
FLORISTIC LINE FOR THE
STUDY OF THE FLORA OF
EAST ASIA!
ABSTRACT
The “Tanaka-Kaiyong Line" is a line separating two floristic subkingdoms of East Asia, the Sino-Japanese to the
east and the Sino-Himalayan to the west. It lies in the provinces of Sichuan and Yunnan in China. It is comprised of
the “Tanaka Line of Citrus Distribution" in Yunnan province and the “Kaiyong Line of Orchid Distribution” in Sichuan
province, which nearly connect at the 757 of NW Yunnan and SW Sichuan. After a survey of the distribution patterns
a, we are sure that it exists. The Tanaka-Kaiyong Line is significant not only for the subdivision of the
mportant floristic regions, the Hengduan Mountain region, Central China
region, and Dian—Qian—Gui region, irk û are noted for species abundance, endemism, and high speciation, are situated
near this Line. Compared with Sino-Himalayan genera, the Sino-Japanese are more completely e by the Tanaka-
Kaiyong Line, possibly due to their different florogenetic backgrounds and different times o
gin. recien. it
seems that the centers of biodiversity (or abundance centers) of Chinese endemic genera on seii Side of the Tan
Kaiyong Line are different. The centers on the west side of the Line are m
mainly paleoendemic. This difference may be caused by geological-geomorphologi
climatic conditions in China. The formation of Hengduan Mountain and Yun
mainly pde: but those on the east are
nnan A which was induced by the
uplifting of the Himalayas, greatly affected the floristic nature of western East Asia ee therefore, Uh floristic distri-
butions that led us to define the Tanaka-Kaiyong Line have probably arisen since that t
Recently in the study on the flora of East Asia
it was noted that an important floristic line exists
in the western part of East Asia. This line is im-
portant in the understanding of the characteristics
and biodiversity of the East Asian flora.
I. WHAT IS THE TANAKA-KAIYONG LINE?
The Tanaka-Kaiyong Line is named for the “Та-
naka Line" in Yunnan province and the “Kaiyong
Line” in Sichuan province. These two lines almost
connect at the border of NW Yunnan and SW Si-
chuan and act as a dividing line between Sino-Jap-
anese and Sino-Himalayan genera.
The Tanaka Line was first suggested by the Jap-
anese scholar Tyózaburó Tanaka (Tanaka, 1954) in
the treatment of the taxonomy and distribution in
Citrus. Tanaka discovered that a line can be drawn
in between the two groups of chains (Metacitrus
predominating and Archicitrus predominating) that
is significant in distinguishing the mode of distri-
bution of citrus fruit. This line might be called the
“Tanaka Line of Citrus Distribution” in connection
with the natural development of the Citrus flora of
the Far East. The approximate position of the Ta-
naka Line can be shown as a straight line starting
at the intersection of 28°N, 98°E southward to ap-
proximately 18°45’ or 19°N, 108°E. Tanaka tested
the existence of this line by plotting the distribution
of the genera in Aurantioideae. Of a total of 22
genera in continental Asia, 10 (45%) do not occur
east of the Tanaka Line, while 1 (Poncirus) stays
inward, totaling 11 genera (50%) that stop at this
line. Likewise, a great number of Indo-Malayan
plant families stop at this line, as does the westward
advance of Sino-Japanese plant families.
The “Kaiyong Line” was first suggested by the
Chinese scholar Lang Kaiyong (Lang, 1994) as a
line separating the Sino-Himalayan and Sino-Jap-
anese subkingdoms in Sichuan Province. It is
based on the distribution patterns of some genera
or subgenera of Orchidaceae typical for these two
' Dedicated to our advisor, Wu Zheng-yi (Wu oe We congratulate him on his 80th birthday and his great
achievements in botanical research, in which he
tireless teaching. This paper is one of the research reports о
has been T for 60 years. We = thank him for his years of
f the project of “The floristic study on the Chinese seeds
plants” (9390010), which was financed by the NSFC. We thank Diana (Nan) ht of the Missouri Sob
Garden for editorial help with the manuscript.
? Kunming Institute of Века Chinese Academy of Sciences, Kunming, Yunnan 650204, People’s Republic of China.
ANN. Missouni Bor. GARD. 84: 888-892. 1997.
Volume 84, Number 4
1997
Li & Li
Tanaka-Kaiyong Floristic Line
889
subkingdoms because the family is more restricted
by environmental conditions than other families.
The Sino-Himalayan genera are Risleya, Diplom-
eris, Diphylax, Platanthera subg. Stigmatosa, and
the Sino-Japanese are Neofinetia, Vexillabium, and
Sedirea. The Kaiyong Line runs north to south as:
Naping (Jiuzhaigou), Songpan (Huanglongzhi),
Maowen, Guanxian (Guangguanshan), Baoxing, Er-
langshan (west of Tianquan), Emei Mountain, Shim-
ian, Xichang, Dechang, Miyi to Panzhihua City.
This line is not named by Lang. We named this line
the “Kaiyong Line of Orchid Distribution” because
Lang was the first person to suggest the line in W
Sichuan in the literature and in relation to the Ta-
naka Line. We have treated the Kaiyong Line as a
northern extension of the Tanaka Line because the
two are nearly connected, and both act as dividing
lines between Sino-Japanese and Sino-Himalayan
genera.
II. SIGNIFICANCE IN FLORISTIC STUDY
The Tanaka-Kaiyong Line as a dividing line in
the flora of East Asia is of great significance in
floristic study. It is characterized by the following:
1. IT ACTS AS A DEMARCATION FOR THE SUBDIVISION OF
THE EAST ASIA KINGDOM.
Recently many Chinese scholars have suggested
East Asia as a floristic Kingdom in the floristic re-
gionalization of the world, because East Asia has
no less than 30 endemic families and 577 endemic
genera (excluding the Pteridophytes) [according to
Wu (1993), but according to Takhtajan (1986), it
has more than 20 endemic families and over 300
endemic genera]. In addition, East Asia has a high
level of endemic species. The area that comprises
Wu's “Sino-Japanese” region is almost the same as
the East Asia Kingdom. East Asia borders on the
Pacific Ocean with a broad east side and a narrow
west side (Fig. 1). It is floristically closely related
to the Holarctic Kingdom in the north and the Pa-
leotropic Kingdom in the south. The East Asia
Kingdom can be divided into two subkingdoms, the
Sino-Japanese in the east and Sino-Himalayan in
the west. The floras of these subkingdoms exhibit
vicariance between eastern and western taxa, wi
the line of demarcation situated in western East
Asia. The Tanaka-Kaiyong Line evidently divides
these two subdivisions on the whole. Because the
Line is based on the distribution patterns of the
Sino-Japanese genera and the Sino-Himalayan gen-
era, it is of great significance for the subdivision of
the East Asia Kingdom floristically, and also for the
floristic regionalization of China. Moreover, three
important floristic regions are situated near this
line: the Hengduan Mountain region on the west
and the Central China and Dian-Qian-Gui regions
on the east. These three regions have a great abun-
dance of species, the highest levels of endemism at
the specific level and speciation in China (Li,
2. IT HELPS THE SEPARATION OF SOME CRITICAL SPECIES
AND THE STUDY OF VICARIANCE.
Because the Tanaka-Kaiyong Line divides the
Sino-Japanese and Sino-Himalayan floras, it is
helpful in the delineation of some critical species
belonging to these two floras. The species endemic
to one flora are quite different from those endemic
to the other, because the floras have different spe-
ciation in time and space. For example, Cycloba-
lanopsis glauca (Thunb.) Oerst. of the Sino-Japa-
nese flora is different from C. glaucoides Schott. of
the Sino-Himalayan flora, although they are very
similar morphologically. The concept of the Tanaka-
Kaiyong Line helps in separating the species ac-
cording to their distribution patterns. Vicariance is
an important problem in phytogeographical study,
and is evident in China between eastern and west-
ern taxa, not only for the oaks, conifers, and other
evergreen or deciduous trees at low altitudes, but
also for hemlocks and firs at high altitudes. For
example, Pinus yunnanensis Franch. of W China is
a vicariant species of P. massoniana Lamb. of E
China, Keteleeria evelyniana Mast. of K. davidiana
Bertr.) Beissn., Castanopsis delavayi Franch. of C.
sclerophylla (Lindl.) Schott., Cyclobalanopsis dela-
vayi (Franch.) Schott. of C. gilva (BL) Oerst., Alnus
nepalensis D. Don of A. crematogyne Burkill, and
so on. The concept of the Line, together with an
analysis of phylogeny, will lead to a determination
of what is true and what is false vicariance. It
seems that the Line is helpful in the study of the
vicariance between the Sino-Himalayan and Sino-
Japanese floras.
3. IT FACILITATES UNDERSTANDING OF THE
SINO-JAPANESE AND THE SINO-HIMALAYAN ELEMENTS.
After surveying distribution patterns of some
genera in both the Sino-Japanese and the Sino-
Himalayan floras (Li & Li, 1992), we are sure that
the Tanaka-Kaiyong Line is real. In comparison
with the Sino-Himalayan genera, the Sino-Japanese
genera follow the line more strictly, perhaps be-
cause of their different florogenetic backgrounds
and different origin times. In general, the Sino-
Himalayan elements are younger and more dynam-
ic, so they do not conform as strictly to the Line,
Missouri Botanical Garden
Annals of the
890
['(0961) ueleryyRy, Jo Yeu иеці JOMOT SI BISY 158] JO лорлод UISYLOU sny :910N] 291099 возио
1ouu[ YINOS 'д оәјиәо ueussueqre], uinog-oeriZuou7 ‘2 чәиәо иецепшиеуу-иецеЗиеп ^9 чәдиәә Sur2ui) e 193099 urejunojy иепрёиәң әүрру ^p *191u99 urejunoyy uenpauoH
ціпос̧ 'e чәшәә тпе)-иет()-иеі `@ 4194029 2213ueX-PIN "Т :еләџәЯ әгшәриә onÁgdojeuueds asautyo jo (влодиоо oouepunqe 10) злојиоо Áirs1oArporg ‘Surg ZuoÁrey + ‘Surg еҹеие,
„у :aur] 2uo&rex-exeue] '2лоџод oruopuo опАудојешлоде asauty;) jo (вләїцәә eouepunqe 10) Апзлолтрота jo вләиәә әц pue әш SuoÁrew-exeue] әці jo dew qojoxs y ст әл
021 su от Set — gi о 901 001 56 06
E * ы BH oe
E £5». ET T I 1 м Jj и ~ жао 08 жр d
«= LE : E а с 7 ], e -ZEHE мит
1E. = : gk ћи. E
$ 4 RY î!
У“ | / "4247 215 UE g TE y
01 7 rg T. an” 7 d -a N
о
S Га om? Сао e» «e o en o "Та. a И. xis а WR, eee ЖОШ Ri
{ aa ——r. T la sf ا THAI! oz
' T ax E ales TOL LAA
ч = T 52 à ~ an C | m YE =- SE
ог E ۰ x $ j 2 y me у da Е ~ A
4 : G "oet R l^ SL wi PA ы
EA ав d 7 E = A e i Very d ај М, „4 da
P a | + Y i ' X tan п т + ^ sz
+ у иф awe °бу,
у 6 ЗЕ q E p у $ ~
"a онај | 71 ~ : х] н (
"E М, йз ~ хр R y IS wo 3
Т F b fcf Je e VES X \ = lag -
9 e m MIT Ew] m. PS >
o fi O "Y 4 т) 9e 3 oe
ME d ЈЕ i а)" СА. Ж = |
р mo "E [3] { Y
gc 5, = бо ~:
e LES vd d »
МА з ~ A, б
Sv 4 f e у 2-7 d
= T
۹ 14 x
-- au, | nm. 0 >
+ Ы A a x ы” HE
~ ~ ЗА ho A Га
fal 4
SPI opt sel сет 1 ozi su от $01 001 56 06 s8 08 СП 02 4
Volume 84, Number 4
1997
Li & Li 891
Tanaka-Kaiyong Floristic Line
and the Sino-Japanese elements are relict, so they
conform more strictly to it. This conclusion is in
accordance with Wu: “From the florogenetic point
of view, the genera of the Sino-Japanese distribu-
tion pattern are usually epibiotic and relict in na-
ture, being the historical elements from an ancient
Laurasian flora, while the Sino-Himalayan are
mixed with several neoendemic elements derived
from both Laurasian, Tethyan and Gondwanic ele-
ments. The Tethyan origin shows especially in such
families as Apiaceae, Boraginaceae, Brassicaceae,
Dipsacaceae, Lamiaceae, etc. Probably it implies
that the Sino-Japanese distribution pattern is older
than the Sino-Himalayan distribution pattern, and
also implies that the degree of diversification 15
stronger and more progressive in Sino-Himalayan
flora.”
4. IT AIDS THE UNDERSTANDING OF BIODIVERSITY
CENTERS (OR ABUNDANCE CENTERS) OF CHINESE
ENDEMIC GENERA.
We noted that the centers of biodiversity (or
abundance centers) of Chinese endemic genera on
each side of the Tanaka-Kaiyong Line are quite dif-
ferent, those west of the Line being mainly neoen-
demic and those east mainly paleoendemic. Ac-
cording to Ying and Zhang (1984) the west side of
the Line has one center of biodiversity, W Sichuan—
NW Yunnan, but according to Wang and Zhang
(1994) there are two centers, the South Hengduan
Mountain center and Middle Hengduan Mountain
center. Ying and Zhang’s concept of the east side
of the Line has two centers, SE Yunnan-W Guangxi
and E Sichuan-W Hubei, while Wang and Zhang's
concept (Fig. 1) has six: Mid-Yangtze, Dian-Qian-
Gui, Qingling, Huangshan-Tianmushan, Zhong-
tiao-South Taihangshan, and South Inner Mongolia.
We think the South Inner Mongolia center, unlike
the others, is mainly dry desert, and that Qingling
is not a center in a strict sense because it has no
endemic genera but supports Chinese endemic gen-
era shared with other regions, especially with Cen-
tral China (Ying, 1994); thus the east side of the
Line has a maximum of four centers in the strict
sense for the East Asia Kingdom. Therefore, the
different natures of the centers on both sides of the
Line are parallel with those of the elements on both
sides of the Line. The cause of the difference be-
tween the centers of biodiversity on either side of
the Line may be directly related to geological-ge-
omorphological patterns and large-scale climatic
conditions in China. The area on the west side of
the Line is situated on the transitional site from the
second to the first terraces geomorphologically and
climatically affected by monsoons from both the In-
dian and Pacific Oceans, while the area on the east
side is on the site from the third terrace, or coast
land, to the second terrace geomorphologically, and
is climatically affected mainly by the monsoons
from the Pacific Ocean.
III. ORIGIN OF THE FLORISTIC LINE
The Tanaka-Kaiyong Line, which divides two
subkingdoms of the East Asia Kingdom, is a result
of the large-scale geological-geomorphological pat-
terns in China, as already mentioned above. Liu
and Ding (1984) summarized the history of the geo-
logical structure of China. The main Chinese plat-
form had risen above the sea by the end of the
Triassic, except that some residual epi-continental
waters remained, including one over present-day
Yunnan. А trough also extended across what is now
the Qinghai-Xizang plateau north of the present
Himalayas. From the end of the Jurassic-Creta-
ceous, a sequence of underthrusts from the south
extended the continental land area of China south-
ward and raised the whole platform, particularly in
the west. By the end of the Paleocene the present
outline of continental China had become estab-
lished, except that the Himalayan region remained
marine until the Eocene. The Himalayas were the
product of northward underthrusting of the Indian
Plate (Chang & Pan, 1981) and only came into be-
ing as a mountain range late in the Cenozoic, or
perhaps in the Miocene-Pliocene. It seems that up-
lift took place at an accelerating rate. The average
for the 3000 m rise during the Quaternary is 1.5
mm per year, but the rise during the Holocene may
have been four or five times greater (Xu, 1984). The
great uplifting of the Himalayas induced the rising
and the climatic fluctuation of Henguan Mountain
and the Yunnan Plateau, and also greatly affected
the floristic nature of the west of East Asia, and the
Tanaka-Kaiyong Line thus appeared since that
time.
Literature Cited
Chang C. & Pan Y. 1981. A brief discussion on the tec-
tonic evolution of Qinghai-Xizang plateau. Pp. 1-18 in
Liu D. S. (editor), Geological and Ecological Studies of
Qinghai-Xizang Plateau. Volume 1. Science Press, Bei-
ng.
tok: Y. 1994. Studies on the distribution patterns of
some significant genera in orchid flora. Acta Phytotax.
Sin. 32: 328-339.
Li X. W. 1996. Floristic statistics and analyses of seed
plants from China. Acta Bot. Yunnan. 18(4): 384.
i X. W. & Li J. On the validity of Tanaka Line
and its significance viewed from the distribution of east-
ern Asiatic genera in Yunnan. Acta Bot. Yunnan. 14(1):
-12.
892
Annals of the
Missouri Botanical Garden
Liu T. S. & Ding M. L. 1984. The characteristics and
evolution of the palaeoenvironment of China since the
late Tertiary. Pp. 11-40 in R. O. Whyte (editor), The
Evolution of the East Asian Environment, Volume I.
Center of Asian Studies, Univ. Hong Kong.
Takhtajan, A. 1986. Pp. 39-78 in Floristic Regions of the
World. Univ. California Press, Berkeley, Los Angeles,
Tanaka, T. 1954. Species problem in Citrus. Pp. 58-69 in
Miro Society for the Promotion of Science, Ueno,
Tokyo
Wang н. S. & Zhang У. L. 1994. The biodiversity and
daga of spermatophytic genera endemic to China.
a Bot. Yunnan. 16: 209-220.
Wu C. Y. - Delineation and unique features of the
ese Floristic region.—A speech at the XV
yo.
Xu R. 1984. Cha anges of the ooo of south-
ern east Asia since the late Tertiary. 25 in R.
O. Whyte (editor), The Evolution of m East Asian En-
vironment, Volume II. Center of Asian Studies, Univ.
Hong Kong.
Ying T. S. 1994. An analysis of the flora of Qingling moun-
tain range: Its nature, ina de d and origin. Acta
Phytotax. Sin. 32(5): 3 Е
Ying Т. 5. & Zhang 2. i we Endemism in the flora of
China—Studies on the endemic genera. Acta Phytotax.
Sin. 22(4): 259-268.
CHROMOSOME NUMBERS IN
COMPOSITAE, XVII:
SENECIONEAE ПГ
Harold Robinson”, Gerald D. Carr’,
Robert M. King?, and A. Michael Powell*
ABSTRACT
matinae are one of the most distinct groups with a base o
x = 9, consi
new subtribe, Abrotanellinae, is established for the southern South American and southwest Pacific genus Abrotanella,
This paper continues a series dealing with chro-.
mosome numbers of Compositae (Raven et al.,
1960; Raven & Kyhos, 1961; Ornduff et al., 1963,
1967; Payne et al., 1964; Solbrig et al., 1964, 1969,
1972; Anderson et al., 1974; Powell et al., 1974,
1975; King et al., 1976; Tomb et al., 1978; Rob-
inson et al., 1981, 1985, 1989). This is the third
of the series ostensibly dealing with the tribe Se-
necioneae. It presents an updated summary of the
chromosome numbers in the tribe (Appendix 1) and
78 new reports of chromosome numbers, including
19 previously uncounted species listed below in the
results (Table 1). The new counts are particularly
important in some of the less well known neotrop-
ical genera such as the Central American Jessea
and the Andean Gynoxys, Lasiocephalus, and Pen-
tacalia. Nevertheless, the new counts are not con-
sidered to be as important in this paper as the over-
view of the chromosome number patterns in the
tribe, to the extent that they are presently known.
The limits of the Senecioneae in the present pa-
per are in the newer, more restricted sense (Bremer,
1994), following more closely the studies of Nor-
denstam (1977), Jeffrey et al. (1977), and Jeffrey
E
thanked for help in numerous other field trips. Liz Zimmer
and B. L. Turner are thanked for careful reviews of the m
_ nat of Botany, National Museum of Natural
.S.A.
3 Department of Botany, University of Hawaii at Manoa, Но
anuscri
Histo:
(1979) and are essentially those of Jeffrey (1992).
Previous studies in the present series (Ornduff et
al., 1963, 1967), treating the Senecioneae, used the
aphyletic traditional concept of the tribe (Bentham
& Hooker, 1873) that included, sometimes with res-
ervations, taxa that we now know were misplaced:
the cichorioid Liabeae and Gongrothamnus (— Dis-
tephanus, Vernonieae) and the asteroid paleaceous
Heliantheae (Dimeresia, Neurolaena, an histo-
carpha), the epaleaceous Heliantheae (Arnica, Mal-
lotopus, Peucephyllum, Psathyrotes), and the Inu-
ean Adenocaulon. For dispositions of these genera,
see the studies of the Liabeae (Robinson, 1983),
Distephanus (Robinson & Kahn, 1986), and the He-
liantheae (Robinson, 1981). Ornduff et al. (1963)
also felt that the Blennospermatinae were not Se-
necioneae, but the subtribe is included in the pres-
ent concept.
The first paper on the Senecioneae in the present
series (Ornduff et al., 1963) had many records and
comments on the genera that remain in the tribe.
Included were a brief mention of Crocidium (of the
Blennospermatinae) with n — 9, some Emilia with
n — 5, examples of species of Senecio in Australia
—
identification of the Australian members of the tribe. The
ed by a National Geographic Society grant. Paul Peterson is thanked
George Proctor, and Robert Garvey are
is thanked for obtaining reference material. Robert Kowal
ip
ry,
nolulu, Hawaii 96822, U.S.A.
2, U.S.A.
t.
Smithsonian Institution, Washington, D.C. 20560,
* Department of Biology, Sul Ross State University, Alpine, Texas 79832, U.
ANN. MISSOURI Bor. Garp. 84: 893—906. 1997.
894 Annals of the
Missouri Botanical Garden
Table 1. Results. The following are newly reported chromosome counts and citations of vouchers for species of
Senecioneae. Species not previously reported are marked with an asterisk (*). K numbers represent King collection
numbers. Counts are reported as 2n (Carr) or n (Powell). Vouchers are in US (duplicates in MO).
Species Chromosome number Voucher
*Arnoglossum ovatum (Walt.) H. Rob. 2n = ca. 28-30 pairs U.S.A. Florida: Liberty Co.,
K10297
Barkleyanthus salicifolius (HBK) Н. Rob. & п = са. 30 Guatemala. Quezaltenango: K7261
B. salicifolius = 30 Guatemala. Totonicapán: K7292
Crassocephalum crepidioides (Benth.) S. Moore te 20 pairs Puerto Rico. Mpio. Bayamén,
K10607
Emilia coccinea (Sims) Sweet п = 8 ог 9 Dominica. St. Patrick, K6366
E. fosbergii Nicolson 2n = 10 pairs Costa Rica. San José: K10023
E. se п = са. 10 Ecuador. edm K6947
E. fosbergi 2n — 10 pairs Puerto Rico. Mpio. Salinas, K10598
Erechtites hieraciifolia (L.) Raf. ex DC. 2n = 20 pairs U.S.A. Virginia: Fairfax Co.,
K10596
Erechtites valerianiaefolia (Wolf) DC. п = 17-20 Dominica. St. George, K6302
*Gynoxys acosta trec. 2n — ca. 40 pairs Ecuador. Pichincha: K10056
*G. buxifolia (HBK) Cass. n — ca. 40 Ecuador. Chimborazo: K6602
*G. fuliginosa e: Cass. 2n — 40 pairs Ecuador. Carchi: 10117
*G. hallii Hiero. very sticky, possibly n = са. 40 Ecuador. Pichincha: K6735
*G. sancti-antonii me e var. brevifolia n = 40 + 4-5 B chromosomes Ecuador. Azuay: K6902
(Hieron.) Cuat
Gynura pn рта (L.) D п = 10 Thailand. Kamphaeng Phet, К5438
* Jessea cooperi (Greenm.) " Rob. & Cuatrec. n — ca. 50 Costa Rica. San José: K6743
* J. cooperi n — ca. 50 Costa Rica. Cartago: K6771
* J. cooperi 2n — ca. 49 foie Costa Rica. San José: 10031
*Lasiocephalus involucratus (HBK) Cuatrec. 2n = 20 p Ecuador. Carchi: K10098
*L. involucratus 2n = 20 a Ecuador. Chimborazo: K10174
*L, patens (HBK) Cuatrec. 2n — 20 pairs Ecuador. Carchi: K10083
Pentacalia andicola (Turcz.) Cuatrec. 2n — 20 pairs Ecuador. Carchi: K10099
andicola var. trm tidus Cuatrec. n — 20 Ecuador. Azuay: K6643
+P firmipes Green 2n — 20 pairs Costa Rica. Cartago: K10022
*P. phanerandra (Cufod.) Н. Rob. & Cuatrec. n = 17 or 20 Costa Rica. Cartago: K6436
P. reflexa (HBK) Cuatrec. 2n — ca. 50 pairs Venezuela. Mérida: K10496
2n — ca. 50 pairs Venezuela. Mérida: K10590
*P. sotarensis (Hieron.) Cuatrec. 2n = ca. 50 pairs Ecuador. Carchi: K10078
P. vaccinioides (HBK) Cuatrec. n = 20 Ecuador. Azuay: K6662
P. vaccinioides 2n — ca. 20 pairs . . Ecuador. Carchi: K10088
Psacalium sinuatum (Cerv.) H. Rob. & Brettell 2n = ca. 30 pairs Mexico. ine K9916
*Pseudogynoxys sonchoides (HBK) Cuatrec. n — ca. 46 Ecuador. Azuay: K6688
Roldana petasioides (Greenm.) H. Rob. & n — 30 Guatemala. cM EVI K7328
Brettell
R. pe n — ca. 30 Guatemala. Nat K7286
* Senecio canchahuinganquensis Cabrera 2n — ca. 20 pairs Argentina. Neuquén: K9408
5. costaricensis R. M. King n Costa Rica. pn K5392, Type
5. costaricensis n — ca. 17 Costa Rica. San José: K6759
S. coahuilensis Greenm. 2n — 23 pairs Mexico. Coahuila: K10356
5. culcitioides Wedd. 2n — 20 pairs Ecuador. Pichincha: K10051
S. dej Griseb. 2n — 20 pairs Argentina. Catamarca: K9465
S. ow DC. 2n — 20 pairs exico. Sonora: K9806
S. elegans L. 2n — ca. 10 pairs Australia. Victoria: K10268
S. = (Респе.) Sch. Bip. 2n = 10 pairs Quarzazate: K10653
S. glomeratus Desf. ex Poir. 2n = 30 pairs :
8. gunnii (Hook. f.) Belcher
5. gunnii 2n = 20 pairs Australia. Victoria: K9773
Volume 84, Number 4
Robinson et al.
Chromosome Numbers in Compositae
895
Table 1. Continued.
Species
Chromosome number
Voucher
*S. hieronymi Griseb.
S. jacobaea L.
S. lautus G. Forst. ex Willd.
S. lautus
S. lautus
S. lautus
2n = 10 pairs
2n = 20 pairs
2n = ca. 20 pairs
2n = 10 pairs
2n = 20 pairs
2n = ca. 20 pairs
2n = 30 pairs
2n = 30 pairs
2n = ca. 30 pairs
2n = ca. 30 pairs
S. lucidus (Sw.) DC. n = ca. 50
S. lucidus 2n = 50-52 pairs
5. madagascariensis Poir. 2n = 10 pairs
5. minimus Poir. 2n = 30 pairs
S. nivalis (HBK) Cuatrec. 2n = 20 pairs
S. odoratum Hornem 2n = 30 pairs
S. pterophorus DC 2n = 10 pairs
S. pteropho 2n = 10 pairs
5. purpureus L 2n = 10 pairs
S. aff. purpureus 2n = 10 pairs
*S. rigidus 2n = 10 pairs
S. runcinatus Less. n = 17 or 20
S. rudbeckiaefolius Meyen & Walp. 2n = 20 pairs
*S. subulatus G. Don ex Hook & Arn. 2n = 20 pairs
*S. subumbellatus Phil. 2n = ca. 20 pairs
*S. tephrosioides Turcz. 2n = 20 pairs
S. vagans F. Muell. 2n = ca. 49 pairs
5. veilleiodes A. Cunn. ex DC.
S. veilleiodes 2n = 19 pairs
S. viravira Hieron. 2n = 20 pairs
5. viravira 2n = 20 pairs
S. warszewiczii A. Br. & Bouché п = 20
S. warszewiczii n — 20
and the genus Ligularia and its relatives with n —
30, and a discussion of the Senecio aureus group
with n = 22-24. The paper ends with, "The oc-
currence of species of Senecio with n — 10 in por-
tions of Europe and Africa, and the concentration
of Senecioninae and Othonninae with n —
cylindrical versus balust
Argentina. Buenos Aires: K10289
: K9785
Australia. Victoria:
Australia. Western Australia:
Australia. Western Australia:
K9597
Australia. Victoria: K9776
New Zealand. North Island:
K10266
Australia. Victoria: K9739
Australia. Victoria: K9752
Australia. Victoria: K9786
Australia. New South Wales: K9975
Dominica. St. George: K6379
France. Martinique: K10647
Australia. New South Wales: K9972
Argentina. La R
Guatemala. Chimaltenango: K7200
Guatemala. Quezaltenango: K7029
erform (with a zone of en-
larged cells below, Robinson & Brettell, 1973b;
Nordenstam, 1978; Vincent, 1996), and (2) the stig-
matic surface covering the whole inside of the style
branch versus forming two separated lines on each
= 10 in branch. These characters were observed and illus-
trated by Cassini (1818), but they were cited only
sparingly afterwards until they were used to delimit
the generic concepts of Senecio sensu lato and the
Tussilaginae (as cacalioids) by Robinson and Bret-
tell (1973b) and Nordenstam (1978). This narrower
concept of Senecio has been resisted by some au-
thors, and the value of the microcharacters has
Africa strongly suggest an Old World origin for the
tribe, with subsequent wide migration and diversi-
fication nearly throughout the world." These points
are enlarged upon below. The present paper adds
and discusses a few chromosome records of note
from the Andean genera that were previously re-
o 1 r et al. (1967 pis
ee и s been questioned due to some variability (Wetter,
Within the Senecioneae, at the subtribal and ge- 1983); however, the value of the microcharacters is
neric levels, two particular microcharacters have now generally recognized (Vincent, 1996) em the
come into taxonomic use: (1) the anther collar being narrower generic concepts are now generally ac-
896
Annals of the
Missouri Botanical Garden
cepted (Jeffrey, 1992; Nordenstam, 1996b; Barkley
t al., 1996; Vision & Dillon, 1996).
MATERIALS AND METHODS
The new reports in this paper are based on ma-
terial collected by R. M. King and counted by G.
D. Carr (reported as 2n) and А. М. Powell (reported
as n). The chromosome counts have been made
from aceto-carmine or aceto-orcein squashes of mi-
crosporocytes in meiosis. Voucher specimens are in
US and a second set is at MO.
DISCUSSION
Blennospermatinae, x = 9
Ornduff et al. (1963) strongly suggested that Cro-
cidium did not belong to the Senecioneae. However,
the TEM study of pollen (Skvarla & Turner, 1966)
and flavonoid data (Ornduff et al., 1973) support
placement of both Crocidium and Blennosperma in
the tribe. Inclusion of Crocidium in the Seneci-
oneae, in a subtribe Crocidiinae (= Blennosper-
matinae), was accepted by Robinson and Brettell
(1973a), with the addition of Ischnea. The latter
addition was supported by the TEM study by Gadek
et al. (1989). The subtribe is distinguished within
the tribe by small pollen, 23-25 ym diam., mea-
sured in fluid, as in the Astereae. Cytologically,
Blennosperma has n = 7, 9; Crocidium n = 9 (Jef-
frey, 1992), and Ischnea n = 9 (Borgmann, 1964).
The subtribe is evidently a basal offshoot of the
Senecioneae with an aneuploid reduction series of
chromosome numbers from x = 10.
Abrotanellinae H. Rob., G. D. Carr, R. M. King
& A. М. Powell, subtribus nov., х = 9. TYPE:
Abrotanella Cass.
Plantae herbaceae perennes caespitosae ad 12 al-
brevibus, stigmatibus marginalibus; е
culini vel hermaphroditi; corollae pallidae, tubis interne
ad apicem non annulate ornatis, filamenta in partibus su-
perioribus cylindracea; rami stylorum breves vel nulli,
stigmatibus marginalibus vel nullis; achaenia glabra vel
raro setulifera; grana pollinis 27—40 jum diam.; chromo-
somatum numerus 2n —
Abrotanella of South America and the Austral-
asian species that have been placed in that genus
were included in the Blennospermatinae by Nor-
denstam (1977), Bremer (1994), and Swenson
(1995a, b), but are excluded from the subtribe here.
The base chromosome number of x — 9 seems to
isolate the group from all members of the Seneci-
oneae except the Blennospermatinae, and Abrota-
nella has glandular-punctate leaves, disciform
heads, more incrassate corolla tissue, and no an-
nulus inside at the base of the corolla limb (Rob-
inson & Brettell, 1973a), all of which distinguish
it from the Blennospermatinae. The hairs of the
achenes, when present, are never the mucilage
hairs seen in members of all three genera of the
Blennospermatinae. The somewhat larger pollen
grains and the texture of the corollas in the Abro-
tanellinae are more like other Senecioneae than
like the Blennospermatinae. The habit of Abrota-
nella is similar to that of Werneria and Xenophyl-
lum, but the latter clearly belong to the Senecion-
inae with balusterform anther collars and
chromosome numbers of n = ca. 50 or higher in
most species.
Tussilaginae, x = 30
One group mentioned by Jeffrey (1992) with n =
30, the tussilaginoid group or Tussilagininae, con-
tains many mostly Northern Hemisphere genera,
some of which were included within Senecio in the
traditional classification of the tribe (Bentham &
Hooker, 1873). The group is characterized by cy-
lindrical anther collars, stigmatic papillae contin-
uous over the inner surface of the style branches,
and a strong tendency for palmately veined or pel-
tate leaves. Included are many species outside of
Europe that were placed in Cacalia, but excluded
is Hasteola (Anderson, 1994). Sixteen species from
various genera with counts of n = 25, 26, 28, 29,
and 30 were reported by Ornduff et al. (1963,
1967). Counts of n = 30 were reported for 15 er-
adiate, white-flowered Mexican species of the group
by Pippen (1968). Counts for a few more Mexican
members of the group were reported by Powell et
al. (1974). Robinson and Brettell (1974) first gen-
erally recognized the Tussilagininae (as the caca-
lioid group) and correlated it with the chromosome
number of n = 30, listing the 27 Mexican species
then known to have this number. The correlation
was also noted by Nordenstam (1977). The Central
American genera were reviewed by Barkley et al.
(1996) with some recent additions such as Robin-
sonecio, with n = 30 (Barkley & Janovec, 1996).
The summary of the 35 European, Asiatic, Austra-
lian, North American, and Mexican genera of the
group by Jeffrey (1992) shows n = 30 as the pri-
mary or only chromosome number in 25 of the 30
genera that have been sampled cytologically. Some
genera have variations from n = 30 (Jeffrey, 1992),
especially aneuploid reductions: to n = 25 or ca.
25 in Arnoglossum and Psacalium: to n = 26 in
Parasenecio, Miricacalia, and Syneilesis: and to n
Volume 84, Number 4
1997
Robinson et al.
Chromosome Numbers in Compositae
897
= 28 in Rugelia and Endocellion. Lower numbers
occur in some genera, within which n = 30 pre-
dominates: n = 16 and 24 in Ligularia, n = 15
and 20 in Doronicum, and n = 14 and 16 in Pe-
tasites.
Although a simple point, it should be noted that
n = 30 in the Senecioneae is not a simple polyploid
derivative of n = 10 like n = 20, 40, or 80, but
requires two steps: first a tetraploid (n = 20) cross-
ing with a diploid (n = 10) to give a triploid (2n
= 30), which then is stabilized by a second dou-
bling to 2n = 60 giving n = 30. In the Tussilagi-
ninae with lower chromosome numbers, only Do-
ronicum has n = 15 and 20. Ligularia has n = 16,
with the next lowest number at 24, and Petasites
has n = 14 and 15, with the next lowest number
at 26. These lower numbers in the group, of mostly
European origin, cannot be ancestral if n = 30 is
basic to the Tussilagininae. At present, these lower
numbers are considered extreme reductions, far ex-
ceeding the modest aneuploid reductions seen in
other members of the group such as Arnoglossum,
Rugelia, and the Asiatic genera Parasenecio, Mir-
icacalia, and Syneilesis.
While n = 30 seems to be basic in the Tussi-
lagininae, that number also occurs in some mem-
bers of the Senecioninae, mentioned below. Also,
the aneuploid reductions in the Tussilagininae ap-
proach or duplicate the aneuploid increases in the
Senecioninae, e.g., the aureoid group with n = 22,
23.
Gynoxoid group, n = 40
Jeffrey (1992) listed a few, mostly American,
genera as gynoxoid, including Gynoxys, Paragy-
noxys, Paracalia, and Aequatorium. For information
on the most recent additions to Gynoxys, Aequato-
rium, and Paragynoxys, see Robinson and Cuatre-
casas (1992). These genera have a single continu-
ous stigmatic surface on the inside of the style
branches and cylindrical anther collars like the
Tussilagininae, and Gynoxys falls generally with the
tussilaginoid genera in the cpDNA study of Kad-
ereit and Jeffrey (1996). The one feature that might
hold the group apart from the Tussilagininae is the
chromosome number of n = 40 or ca. 40 in Par-
agynoxys (n = 3840 pairs, Powell & Cuatrecasas,
1970) and Gynoxys (Turner et al., 1967; this paper).
Counts for two species of Gynoxys have been re-
ported previously: G. hutchisonii H. Rob. & Cua-
trec. (as G. parvifolia Cuatrec.) and G. tomentosis-
sima Cuatrec. (Turner et al., 1967), with n — ca.
40 or 40 + B's. Counts for five additional Ecua-
dorian species are reported in the present study, all
with n — 40 or ca. 40, one of these with 4—5 B
chromosomes.
The dioecious genus Chersodoma (Dillon & Sa-
gástegui-Alva, 1996) of southern South America
has 2n — 20 (Hunziker et al., 1989) based on one
species, C. argentina Cabrera, and was placed in
the Senecioninae near Senecio by Jeffrey (1992).
However, the anther collars are cylindrical and the
stigmatic surfaces cover the inside of at least the
distal halves of the style branches (Robinson &
Brettell, 1973b), for which reasons we provisionally
place the genus near the Tussilaginae.
Senecioninae
(1) Some elements with n — 30 and n — 50
The chromosome number n — 30 is inevitable in
any large group that has a base of x — 10 and many
multiplies of that number. Of special interest are
groups in the Senecioneae that have stabilized at n
— 30. Jeffrey (1992) mentioned a number of such
ups. One in which n — 30 seems to be basic is
the Tussilagininae. Others appear to be derived
separately within the Senecioninae (Jeffrey, 1992),
such as Pericallis and a group of Australian Senecio
that was sampled by Ornduff et al. (1963) and stud-
ied extensively by Lawrence (1980, 1985а, b).
In the Macaronesian Pericallis with 15 species,
including the common *Cineraria" of horticulture,
n = 30 occurs in all eight species that have been
counted (Jeffrey, 1992).
A group of Australian species of Senecio with n
— 30 includes one radiate species, five discoid, and
four classified as erechtitoid (Lawrence, 1980). Six
of the Australian species of Senecio with n — 30
are self-incompatible perennial herbs, and six oth-
ers, including S. biserratus Belcher, which have n
= 50, are self-compatible annuals (Lawrence,
1985a, b). Unlike the Tussilagininae with n = 30,
this group of unusually abundant Australian hex-
aploids was considered morphologically typical of
the genus Senecio Lawrence (1985a). Lawrence fur-
ther suggested that strict autoploidy in the Austra-
lian species was unlikely since no known native
diploids appeared to exist. However, both diploid
and tetraploid counts are reported here for the com-
mon Australian species S. lautus. As this is the first
report of n = 10 for an Australian Senecio, addi-
tional populations of S. lautus should be surveyed
in order to determine the frequency and extent of
this chromosome condition. Regular meiosis with-
out multivalents was observed in the Lawrence
study, even in the highest, decaploid, levels.
In the Senecioneae, n = 50 has an origin anal-
ogous to that of n = 30: a cross of n = 20 with n
898
Annals of the
Missouri Botanical Garden
= 30 and a doubling to give 2n = 100. The number
is apparently basic to the Costa Rican/Panamanian
species of Jessea (Robinson & Cuatrecasas, 1994;
Nordenstam, 1996a), included in Jacmaia by Jef-
frey (1992). Two species were known to have n =
ca. 50 (Jeffrey, 1992), and our three counts of J.
cooperi are the same
The African idad is reported to have
chromosome numbers of n = 10, ca. 40, and 50
(Jeffrey, 1992); however, Knox and Palmer (1995)
and Knox (1996) accepted the conclusions of Knox
and Kowal (1993) that the genus consistently has
n = 50 and the other numbers are in error. Knox
and Palmer (1995) stated, “The giant senecios, ap-
parently decaploid (n = 50), show virtually no mei-
otic irregularities that might suggest a recent origin
from the predominantly tetraploid (n = 20) puta-
tively close relatives suggested in the literature.”
Knox and Kowal (1993) mentioned many compar-
atively unrelated elements of the Senecioneae that
share the number n = 50, including the Australian
Arrhenechthites, African Kleinia, neotropical Den-
drophorbium, Graphistylis, Jessea (as Jacmaia),
Pentacalia, and Xenophyllum (as Werneria). Study
of cpDNA indicated that the closest relatives of
Dendrosenecio are the African genera Cineraria (n
= 10) and Euryops (mostly n = 10, seldom n =
20, 30) (Knox € Palmer, 1995; Knox, 1996). A
separate cpDNA study (Kadereit & Jeffrey, 1996)
placed Dendrosenecio as the most divergent mem-
ber of a diverse group including the neotropical
Pseudogynoxys and Dorobaea and the Macarone-
sian Pericallis, but unfortunately neither Cineraria
nor Euryops were included in the study.
(2) The aureoid senecios, n = 22, 23
The aureoid senecios, often treated as a separate
genus, Packera, have the appearance and almost
all the technical characteristics of Senecio sensu
stricto. The aureoid species have been treated in
Senecio as subgroups Aurei, Bolanderiani, Lobati,
Sanguisorboidei, and Tomentosi in Ornduff et al.
(1963, 1967), with reports of n = 22, 23, or 24
from 22 species. Additional records were provided
by Kowal (1975) and Bain (1985), and the group
was discussed by Barkley (1988), who stated that
chromosome numbers had been reported for 48 of
the 59 species. Our count of n = 23 for the Mex-
ican Senecio coahuilensis agrees with previous re-
ports for the species and is the commonest number
in the group. Some 15 or 16 species occur in Mex-
ico (Freeman & Barkley, 1995; Barkley et al.,
1996). Counts of n = 20 for Senecio flettii Wiegand
and S. bellidifolius HBK are anomalous in the au-
reoid group (Barkley, 1988), but S. bellidifolius is
well nested within the aureoid group on the basis
of ITS sequence data and pollen structure (Bain &
Jansen, 1995). Kowal (1994) cited ca. 65 species
in the group, all in North America from Mexico to
the Arctic, with one Arctic-alpine species extend-
ing into Asia. Most species have n = 23, and n =
22 occurred only in seven species in the area of
most diversity, including four species restricted to
eastern North America (Kowal, 1994). Counts of n
= 20 and 40 in Senecio pseudaureus were probably
wrong (Kowal, 1994).
Packera, with Senecio aureus L. as type, was ac-
cepted as a genus by Jeffrey (1992) using such
structural characters as lack of interfascicular cam-
bium, branching fibrous roots, apparently obscure
carpopodia, and specific forms of apically flagellate
hairs, which are neither exclusive to the aureoid
group nor very convincing. The group lacks the flu-
ked pappus of typical Senecio (Drury & Watson,
1966), but so do many other elements of Senecio.
A more substantial difference is the helianthoid ul-
trastructure of the pollen wall (exine with internal
foramina), which occurred in all 19 aureoid species
examined in sections Aurei, Tomentosi, and Lobati
(Bain & Walker, 1995), and the latter authors ques-
tioned the value of continuing to recognize the sec-
tions. Non-aureoid North American species exam-
ined, including some tephroseroid Senecioneae
(Tephroseris, Tussilagininae, x = 24) and the Se-
necio sections Lugentes, Integerrimi (n = 20), and
Triangulares (n = 10, 18, 20, 40), had a senecioid
exine wall pattern (lacking internal foramina).
A few features of the aureoid senecios are clear.
It is the only significant element of the Senecioneae
in which the chromosome number does not readily
fit the general pattern derived from x =
multiples, although it may have been derived by an
aneuploid increase from groups with n = 20, to
which it seems otherwise closely related. The num-
ber n = 20 has been reported in the group (Barkley,
1988), but Kowal (1994) considered this an error.
Also, the number n = ca. 30 has been reported for
S. sanguisorbae DC. (Turner et al., 1961), for which
all other counts are n = 23. Chromosome numbers
of this group might be more unstable than in taxa
with x = 10, 20, 30, or 50.
Unfortunately, the best characters differentiating
the aureoid senecios have limited value for ordinary
taxonomic identification. Neither chromosome num-
ber nor ultrastructure of the pollen exine is readily
observable. The lineage is distinct, but the phyletic
distance from typical Senecio may still be compar-
atively slight. Senecio aureus falls into a group of
intermixed Senecio-Kleinia-Gynura species in the
cpDNA study of Knox and Palmer (1995). The ev-
idence could be used to justify many further seg-
Volume 84, Number 4
1997
Robinson et al 899
h
Chromosome Numbers in Compositae
regates from Senecio, but the necessary detailed
studies are still to be done. Recognition of Packera
as a distinct genus seems premature at this time.
(3) Mulgedifolii group, n = 20, 40
The Mulgedifolii group of Mexico lacks ray flo-
rets and has red or white, but never yellow, corollas.
The combination of characters led some early au-
thors to place some of the species in the broad
concept of Cacalia, and they appear among the ex-
cluded taxa listed by Pippen (1968). The Mulge-
difolii have the characteristic balusterform anther
collars and separate stigmatic lines of the Sene-
cioninae. group was 7
Barkley et al. (1996). There is a report of n = 22
for S. runcinatus Less. (Keil & Stuessy, 1975), and
the same species is reported here as n = 17 or 20.
Kowal (pers. comm.) has indicated that the n = 22
was probably wrong. The unpublished summaries
of the Mulgedifolii chromosome counts by Villaseñ-
or list 10 counts for 6 species (1986) and 33 counts
for 15 of the 17 species (1991). All listed counts,
except the dubious n = 22, are п = 20 with one
tetraploid n = 40.
(4) Emilia, n = 5
Emilia continues to show chromosome numbers
of n = 5 and 10 (Baldwin, 1946). The related Af-
rican species of Senecio that were reported with n
= 5 by Turner and Lewis (1965) and Lawrence
(1985a, b) have all been transferred to Emilia by
Jeffrey (1986). The lower numbers correlate with
the short-lived, weedy habit of the genus. This sit-
uation parallels that of Fleischmannia microstemon
(Cass.) K. M. King & H. Rob. (Baker, 1967) and
many other short-lived species with lower DNA
content (Bennett, 1972). The low number is con-
sidered here a derived condition in Emilia, a re-
organization of chromatin on fewer centromeres as
in Fleischmannia microstemon (Baker, 1967) and
Crepis (Tobgy, 1943). Such a reorganization, or “ge-
nome congealing” (Wagner et al., 1993: 422-423)
has been supported by a measurement of the nu-
clear DNA content of E. discifolia (Oliv.) C. Jeffrey
(as Senecio discifolius Oliv.) (Lawrence, 1985b).
In structural features, Emilia is a member of the
Senecioninae, with paired stigmatic lines and bal-
usterform anther collars, and comes out in the gen-
eral subgroup with Othonna and Dendrosenecio in
the cpDNA study of Kadereit and Jeffrey (1996).
(5) Senecioninae with n = 10, 20
In the Senecioneae, n = 10 is largely restricted
to Europe and Africa in the Eastern Hemisphere
(Ornduff et al., 1963), and the pattern is reinforced
by new reports in the present study. A few inter-
esting exceptions occur. Emilia, with n = 5 and
10, is only a recent introduction into the New World
from the Old World. The one old citation of n =
10 in the neotropical Pseudogynoxys, questioned by
Jeffrey (1992), must be incorrect. All other reports
for the genus are n = 45-48 including P. cheno-
podioides (HBK) Cabrera with n = 45 + 5 (Turner
et al., 1962, as S. confusus Britton) and our report
of P. sonchoides (HBK) Cuatrec., representing two
morphological extremes of the genus. Jeffrey's
(1992) summary also gives both n — 10 and n —
20 for S. flaccidus Less. of Mexico and S. bras-
iliensis (Spreng.) Less. of South America. These iso-
lated occurrences of n — 10 in basically n — 20
groups seem unlikely to be survivors of an ancestral
number, but are more likely reductions resulting
from polyhaploidy or aneuploid reduction as in Em-
ilia and some Tussilagininae such as Doronicum,
Ligularia, and Petasites.
An interesting close relationship between an Af-
rican species with n — 10 and an American species
with n = 20, both autogamous, annual, desert spe-
cies, is suggested by isozyme analysis (Liston et al.,
1989) and cpDNA evidence (Liston & Kadereit,
1995). In these papers, the North American Senecio
mohavensis A. Gray was considered a closely re-
lated derivative, through long-distance dispersal, of
S. flavus (Decne.) Sch. Bip. subsp. breviflorus Kad-
ereit of the Saharo-Arabian deserts.
Radford et al. (1995) indicated that the aggres-
sively weedy taxon commonly identified as Senecio
ascariensis Poir. was consistently 2n — 20
based on material from Madagascar, South Africa,
Australia, and Argentina specimens, and our count
is the same. In contrast, Radford et al. (1995) re-
rted the common Australian species $. lautus G.
Forst. ex Willd. as consistently 2n = 40 (see also
the present study; Turner, 1970; Lawrence, 1980;
Webb, 1988). However, see the report here of 2n
= 10 pairs for material determined by Lander,
Short, and Wilson as 5. lautus from Western Aus-
tralia. Sindel (1996) summarized the literature on
the four subspecies of S. lautus.
(6) Andean genera Lasiocephalus and Pentacalia,
n = 20, 30, 45-52
Our counts confirm the few previous counts for
two Andean genera. Further work on chromosome
numbers of Andean Senecioneae is needed. Lasi-
ocephalus (including Aetheolaena) has previously
been known cytologically from one species, L. loe-
seneri (Hieron.) Cuatrec. (Turner et al, 1967).
Counts for additional species, L. involucratus
(HBK) Cuatrec. and L. patens (HBK) Cuatrec., are
reported in the present study. All three species
show n = 20.
Robinson and Cuatrecasas (1978) resurrected
Pentacalia from synonymy within Senecio and in-
900
Annals of the
Missouri Botanical Garden
cluded 13 Central American species. Cuatrecasas
(1981) transferred 177 South American species into
Pentacalia. He accepted two subgenera, the often
scandent subgenus Pentacalia and the often heath-
like subgenus Microchaete. Jeffrey (1992) split Pen-
tacalia into three groups: typical Pentacalia; a new
genus Monticalia Jeffrey, roughly equivalent to the
subgenus Microchaete; and a broadly delimited ge-
nus Dendrophorbium. The broader view of Penta-
calia is retained here because of some unresolved
problems of intergradation. In personal discussion,
Cuatrecasas pointed out that, while most members
of subgenus Pentacalia are scandent, the type spe-
cies, Р. arborea (НВК) H. Rob. & Cuatrec., and its
closest relatives are trees with hanging branches.
The isolated branches can be mistaken for vines.
Thus, the nature of the type species complicates
any tendency to base generic distinctions on habit.
The genus Monticalia is not considered distinct
from Pentacalia here. In any case, priority would
go to Scrobicaria, which was imperfectly distin-
guished from Monticalia only by its opposite leaves
and which was recently reduced to synonymy under
Pentacalia (Cuatrecasas, 1994). Dendrophorbium is
also interpreted more narrowly here to include only
its more typical element that usually lacks tails on
the bases of the anthers. As a result, the following
chromosome counts cited by Jeffrey (1992) all fall
within Pentacalia sensu lato: Pentacalia n = 20 (1
sp.), ca. 40 (1 sp.), 45-50 (2 spp.), 50 (1 sp.), and
ca. 51 (1 sp.); Dendrophorbium n = ca. 40, 49-50
based on 1 sp.; and Monticalia n = 20 (9 spp,) and
40 + 1-5 (1 sp.). The reports in the present study,
of which three are new, show the same range of
numbers, n = 20, n = 17 or 20, and n = ca. 50.
The higher numbers in Pentacalia have undoubt-
edly arisen many separate times in higher elevation
members of the genus.
Senecio lucidus (Sw.) DC., from the Lesser An-
tilles, is reported here with n = ca. 50 and 2n =
50-52 pairs. The species evidently belongs to Pen-
tacalia, but dispositions of West Indian species of
the group are very incomplete (Robinson, 1982).
(7) Culcitium and Dorobaea, n = 20, 50
Culcitium sensu stricto (n = 20) and Dorobaea
(n = ca. 50), from the northern Andes, have fistu-
lose receptacles and a rosulate habit with decres-
cent or progressively smaller upper leaves. For
these reasons, Jeffrey (1992) and others, including
Robinson, have considered both genera as part of
Senecio, in spite of their individually distinctive ap-
pearances. Recent cpDNA evidence placed Doro-
baea in a lineage with Dendrosenecio and Pseudo-
gynoxys, less close to Emilia and Othonna, and
isolated from Senecio sensu stricto (Kadereit & Jef-
frey, 1996). Both Andean genera have stigmatic
lines and anther collars of the Senecioninae. On
the basis of the cpDNA study, Dorobaea (3 sp., Nor-
denstam & Pruski, 1995) and the probably closely
related Culcitium sensu stricto are recognized here
as distinct genera.
(8) Werneria and segregates, n = ca. 50, 100
Werneria has been treated very broadly in the
past to include many densely leaved, rosulate or
strobiform species in tropical America specialized
for high elevations. Genera recently segregated by
Funk (1997a, b, c) are Xenophyllum, including two
species with 2n = ca. 104 + 4 and 2n = ca. 108
+ 4 (Diers, 1961), and the monospecific Misbroo-
kea, with n = ca. 106 + 4 (Diers, 1961). The type
of Werneria, W. nubigena HBK, has been reported
as 2n = ca. 212 + 8 from Peru (Diers, 1961) and
as n = 50 from Guatemala (Beaman & Turner,
1962). Other Andean species that presently remain
in Werneria, for which Diers’s (1961) counts are
available, include W. apiculata Sch. Bip. vien 2n
= ca. 206 + 8, W. orbignyana Wedd. with 2n =
ca. 100 + 4, W. caespitosa Wedd. with 2n = ca.
154 = 6, W. pygmaea Gillies ex Hook. & Arn. with
2n = ca. 212 + 8, W. solivifolia Sch. Bip. with 2n
= ca. 42 + 2, W. spathulata Wedd. with 2n = ca.
106 = 4, and W. villosa A. Gray with 2n = ca. 150
* 6. The high chromosome numbers that predom-
inate in the group are evidently associated with the
harsh environment of the high-elevation Andes.
Adenostylinae Benth. & Hook. f., x = 20
Jeffrey (1992) recognized, within his Senecioni-
nae, a Eurasian group of perennial rhizomatous
herbs with 4-lobed corollas. The group includes the
genus Cacalia (n — 19), in its Linnaean sense
(Robinson & Brettell, 1973b), which is the same as
the later-named Adenostyles, and the subtribal
name Adenostylinae is applied here. Other genera
included in the group are Dolichorrhiza (n = 15-
16, 20, 22), Iranecio (n = 12, 20), and Pojarkovia
(n — 20) (Jeffrey, 1992). A fifth genus, Caucasalia
(n — 19), has been described by Nordenstam
(1997), who has suggested base numbers of n — 20
for Dolichorrhiza, Iranecio, and Pojarkovia. The
distribution of the numbers suggests more than one
aneuploid reduction series from x — 20
CONCLUSIONS
Comparisons between recent taxonomy in the Se-
necioneae and available chromosome counts have
demonstrated again the important point already
suggested in the study of the Heliantheae (Robin-
son et al., 1981): that a phyletic taxonomy can rare-
Volume 84, Number 4
1997
Robinson et al. 901
Chromosome Numbers in Compositae
ly be constructed primarily on the basis of chro-
mosome counts, but chromosome counts usually
make sense at some level when compared with a
phyletic taxonomy. The positions of the most diver-
gent elements of the tribe, Blennospermatinae and
Abrotanellinae, with x = 9, could have been fore-
seen using either taxonomic or cytological ap-
proaches. In the remainder of the Senecioneae, sig-
nificant chromosome number patterns usually recur
frequently, such as x = 30 in the distinctive Tus-
silaginae but also in a number of separate elements
of the Senecioninae. Some of the more unique chro-
mosome number trends in the tribe, such as n = 5
in Emilia and n = 22, 23 in the aureoid group of
Senecio, are taxonomically comparatively close to
typical Senecio. Some genera such as Dendrosenecio
of Africa, the Macaronesian Pericallis, the tropical
American Jessia and Pentacalia, and an Australian
group still placed in Senecio, seem to stabilize at
counts of n = 30 and n = 50. Still other members
of the tribe, including some related pairs of species
in Senecio, show simple polyploid variations be-
tween n = 10 and n = 20
Some taxonomically distinctive groups of the Se-
necioneae show distinctive chromosome counts.
Nevertheless, separate elements of the tribe with
the same counts and evidence of different numbers
in closely related groups indicate the peril of trying
to justify generic or subtribal distinctions in the
tribe solely on the basis of chromosome counts.
Literature Cited
Anderson, L. C. 1994. A revision of pe (Asteraceae)
in the New pu Syst. Bot. 19: 211—
‚ D. W. Kyhos, T. Mosquin, A. M. ical & P. H.
Raven. 1974. roms osome numbere in Compositae, IX:
Haplopappus and other Astereae. Amer. J. Bot. 61: 665—
67
Bain, J. F. 1985. mosome numbers in the aureoid
Senecio complex genome Canad. J. Bot. 63: 539–
542.
. Jansen. 1995. A phylogenetic analysis of
the еса Senecio (Asteraceae) complex sed on ITS
sequence data. ағ Syst. Evol. 195: 209-219.
т. 1995. A comparison of the pollen
wall lid of aureoid and non-aureoid Senecio
species (Asteraceae) in North America. Pl. Syst. Evol.
195: 199-207.
Baker, H. G. 1967. The evolution of weedy taxa in the
Eupatorium microstemon species aggregate. Taxon 16:
293—
Baldwin, J. x 1946. Cytogeography of Emilia Cass. in the
Americas. Bull. Torrey Bot. Club 73: pitt
Barkley, T. M. 1988. даба among the aureoid Senecios
of North America: A geohistorical mt Bot.
ien — 54: 82-106.
& J. P. Janovec. 1996. Robinsonecio €
Senecioneae) a new genus from Mexico and Guatemala.
Sida 17: 77-81.
— ———, B. L. Clark & A. M. Funston. 1996. The segre-
gate genera of Senecio sensu lato and Cacalia sensu lato
(Asteraceae: Senecioneae) in Mexico and Central Amer-
ica. In D. J. N. Hind & H. J. Beentje (editors), Com-
positae: Systematics. Proceedings of the International
Compositae Conference, Kew, 1994. Vol. 1: 613-620.
W.
Beaman, J. H. & B. L. Turner. 1962. Chromosome num-
rs in Mexican and Guatemalan Compositae. Rhodora
64: 271-276.
Bennett, M. D. 1972. Nuclear DNA content and minimum
Ее time in herbaceous plants. Proc. Roy. Soc.
п, Ser. B, Biol. Sci. 181: 109–
егет “e 4 J. D. € 1873. Compoeitae. In Gen-
era plantarum 2(1): 1 |
Borgmann, Е. 1964. 5 der Polyploiden i Flora
des Bismarcksgebirges von Ostneuguinea. . 92
118-172.
sa. Káre. 1994. Asteraceae: зе & Classifica-
tion. Timber Press, Portland, Ore;
Cassini, Н. 1818. Composées. In с ‘Cuvier (editor), Dic-
tionnaire des Sciences Naturelles 10: 131-159, 3 pl.
Paris. [Repri nted in R. M. King & H. W. Dawson (ed-
itors). 1975. Cassini on Compositae. Pp. 110-138, pl.
1-3. Oriole Editions, New Yor!
Cuatrecasas, J. 1981. Studies in neotropical Senecioneae
II. Transfers to том Pentacalia of north Andean spe-
cies. Phytologia 49: 241-260.
. Miscellaneous notes in Neotropical Flora
XXII. Identification of the genus mmn and a
e. Phytologia 76:
few few v
402-405.
Diers, L. 1961. Der Anteil an Polyploiden а са е
tationsgúrteln дег Westkordillere Perus. 49:
437-488.
Dillon, M. O. & A. Sagástegui-Alva. 1996. Revision of the
dioecious Chersodoma Phil. (Senecioneae, Asteraceae),
including a new species and status c change. Brittonia
48: 582
Drury, D. G. & L. Watson. 1966. A bizarre pappus form
in Senecio. Taxon 15: 309-3
Freeman, C. C. & T. M. Barkley. 1 1995. A synopsis of -
genus Packera (Asteraceae: Senecioneae) in Mex
Sida 16: 699-709.
sons i A. 1997a. Misbrookea, a new monotypic genus
ved from Werneria s.l. (Compositae: Senecioneae).
Brittonia 49: 110-117.
————. 1997b. Werneria s.l. (Compositae: Senecioneae)
in Ecuador. Pp. 25-35 in R. Valencia & H. Balslev
(editors), Estudios sobre diversidad y ћи де рјап-
tas. Memorias del II Congresso Ecuatoriano de Botanica
realizado en la Pontificia Universidad Católica del Ес-
uador, н 16-20 Octubre 1995.
зае a new Andean genus е
iracted [ox Werneria s.l. (Compositae: adas
Novon 7: 2
Gadek, P. A. : TB C. J. Quinn. 1989. Exine struc-
ture in the ка чя сетете Asteraceae). Grana
28: 163-178.
Hunziker, J. H., A. Wulff, С. С. Xifeda & A. Escobar.
1989. Estudios cariológicos en Compositae V. Darwin-
9.
iana 29: 25-3
Jeffrey, C. 1979. Generic and sectional limits in Senecio
cu m "e Evaluation of some recent studies.
Kew B gom
pos енны in east tropical Africa.
3-943.
Notes on lides IV. Kew Bull. 41: 87
___- 1992. The tribe Senecioneae (Compositae) in the
902
Annals of the
Missouri Botanical Garden
Mascarene Islands with an annotated World check-list
of the genera of the tribe. Notes on Compositae VI. Kew
Bull. 47: 49-109.
‚ P. Halliday, M. Wilmot-Dear & S. У. Jones.
1977. Generic and sectional limits in — ciam ай
I. Progress report. Kew Bull. 32: 4
al analysis
о оп pos-
itae). In D. J. N. Hind & H. J. Beentje (editors), Com-
positae: Systematics. Proceedings of the International
Compositae Conference, in 1994. Vol. 1: 349—360.
Royal Botanic Gardens,
Keil, D. J. & T. F. eines, 1975. Chromosome counts of
Compositae from the United States, Mexico, and Gua-
temala. Rhodora 77: 171-195.
King, R. M., D. W. Kyhos, A. M. Powell, P. H. Raven €
H. Robinson. 1976. Chromosome numbers in Compos-
itae, XIII: Eupatorieae. Ann. Missouri Bot. Gard. 63
—888.
Knox, E. B. 1996. What is the origin of the giant senecios
in eastern Africa? /n D. J. N. Hind & H. J. Beentje
(editors), Compositae: Systematics. Proceedings of the
— Compositae Conference, Kew, 1994. Vol
1: 691—703. ime Botanic Gardens, Kew.
owal. 1993. Chromosome numbers of
the East intr bind Senecios and giant Lobelias and
their evolutionary significance. Amer. J. Bot. 80: 847-
853
& J. D. Palmer. 1995. The origin of Dendrosenecio
within the Senecioneae (Asteraceae) based on chloro-
plast DNA evidence. Amer. J. Bot. 82: 1567-1573.
Kowal, R. R. 1975. Systematics of Senecio aureus and
allied species on the ie Peninsula, Quebec. Mem.
Torrey Bot. Club 23: 1-1
. Chromosome PES of the aureoid se-
necios (Packera) and their evolutionary significance
[Abstract.] P. 22 in D. J. N. Hind (coordinator), Com-
positae: Systematics Biology Utilization, Paper and
Poster
Lawrence, M. E. 1980. Senecio L. (Asteraceae) in Austra-
lia: Chromosome numbers and the occurrence of poly-
ploidy. Austral. J. Bot. 28: 151—165.
85a. Senecio L. (Asteraceae) in Australia: Re-
combination systems of a polyploid series. Austral. J.
Bot. 33: 209-219.
. 1985b. Senecio L. (Asteraceae) in Australia: Nu-
clear DNA amounts. Austral. J. Bot. 33: 221-232.
. W. Kadereit. 1995. prie an ev-
idence for introgression and long dista:
the desert annual Senecio flavus Тачке Ёл n.
Evol. 197: 3.
„Н. Rieseberg & T. S. Elias. 1989. Genetic
similarity i is high between intercontinental disjunct spe-
cies of Senecio (Asteraceae). A J. Bot. 76:
Nordenstam, B. 1977. Senecioneae о. arg ге!
чт review. Рр. O in . Heywood, J. В
e € В. L. Turner (editors) The Biology and E
ps of the Compositae. vols. Academic Press,
London, New York, San Раш
9
т, + == ~.
8. T AGAUIUIHIU
(Composite): Opera Bot. 44: 1-83.
———.. 1996a. Jessea gunillae B. Nord. San ч
necioneae), a new species from Costa Rica. Bot. Jah
Syst. 118: 147-152.
ES ecent revision of Senecioneae and Cal-
— таванице In D. J. ind & H. J. Beentje
(editors), Compositae: Systemintion: Proceedings of the
International Compositae Conference, f 1994. Vol.
1: 591-596. Royal Botanic Gardens
. Caucasalia, a new genus of t ^it Asteraceae,
Senlécionéne. Pl. Syst. Evol. 206: 19-32.
. Pruski. 1995. Additions to Dorobaea and
falamancalia (Compositae-Senecioneae). Comp.
Newsl. 27: ы.
Ornduff, R., . M. Saleh & B. A. Bohm. 1973. The
Ravens and phos: of Blennosperma and Crocidium
dus roam Taxon 22: 407-412.
Mosquin, D. W. Kyhos & P. H. Raven. 1967.
hromosome n numbers i in гранате VI: Senecioneae
A. R. Kruckeberg.
Payne, W. W., P. H. Raven & D. E “6б . Chro-
mosome numbers in Compositae, IV: Ambrosieae.
Amer. J. Bot. 51: 419-424.
ee R. W. 1968. Mexican “cacalioid” genera allied to
Senecio (Compositae). Contr. U.S. Natl. Herb. 34(6):
363-447.
Powell, A. M. & J. Cuatrecasas. 1970. С
bers in Compositae: Colombian and — spe-
cies. Ann. Missouri Bot. Gard. 57: 374—379.
, D. W. Kyhos & Р. Н. Raven. 1974. Chromosome
numbers in Compositae, X. Amer. J. Bot. 61: 909-913.
& . 1975. Chromosome numbers
in ف XI: Heleniete. Ли. Ј. Вог. 62: 1100–
1103.
Radford, I. J., Q. Liu & Р. = wm У пар Chromosome
counts for the Australian weed as Senecio mad-
agascariensis revu ra ar vn Bot. 8: 1029—
1033.
Raven, P. H. & D. W. Kyhos. 1961. Chromosome numbers
in Compositae, II: Helenieae. Amer. J. Bot. 48: 842-
, O. T. Solbrig, D. W. Kyhos € R. Snow. 1960.
side numbers in Compositae, I: Astereae.
Amer. J. Bot. 47: 124—132.
Robinson, H. 1981. A revision of the tribal and subtribal
limits of the Heliantheae (Asteraceae). Smithsonian
Contr. Bot. 51: i-iv, 1– 102.
ul . 309-313 in С. R. Proctor, More ad-
ditione to the Flora of [rie J. Arnold Arbor. 63.
983. A generic review of the wm Liabeae M
ненне), Smithsonian Contr. Bot. 54: i
———— & R. D. Brettell. 1973a. Tribal re revisions in the
Asteraceae. IX. The relationship of Ischnea. Phytologia
158.
1973b. Studies in ре Senecioneae
— The genera Mesadenia, Syneilesis, Mir-
acalia, Koyamacalia and Sihacalio.- er 27:
265 276,
. 1974. Studies i in the Senecioneae (As-
Am an apecios of Pentacalia (Asteraceae: Seneci-
ones) й 40: 37-50.
———— & —. 1992. Additions to Aequatorium and
pia (Asteraceae: O in Bolivia, Ecuador,
and Peru. Novon 2: 411-416.
ea and Talamancalia, two
new genera of the Oe (Asteraceae) from Costa
Rica and Panama. Моуоп 4: 48-52.
& B. Kahn. 1986. Trinervate leaves, yellow flow-
Volume 84, Number 4
1997
Robinson et al.
Chromosome Numbers in Compositae
ers, tailed anthers, and pollen variation in Distephanus
Cassini (Vernonieae: Asteraceae). Proc. Biol. Soc. Wash.
99: 493-501.
‚ A. M. Powell, R. M. King & J. F. Weedin. 1981.
Chromosome numbers in Compositae, XII: Heliantheae.
Smithsonian Contr. Bot. 52: 1-28.
——, &
1985. Chromosome
ы їп Compositae, ХУ: Lieben. Ann. Missouri
Bot. Gard. 72: 469-479.
" ‚ С. D. Carr, В. M. King & J. F. Weedin.
1989. Chromosome numbers in Compositae, XVI: Eu-
patorieae II. Ann. Missouri Bot. Gard. 76: 1004—1011.
Sindel, B. M. 1996. Impact, ecology and control of the
io madagascariensis in Australia. п P. D. S.
Caligari & D. J. N. Hind (editors), Compositae: Biology
& Utilization. Proceedings of the International Compos-
itae Conference, Kew, 1994. Vol. 2: 339-349. Royal
Botanic Gardens, Kew.
Skvarla, J. J. & B. L. ње 1966. Pollen wall ultrastruc-
ture and its bearing on the systematic position of Blen-
idium (Compositae). Amer. J. Bot.
Solbrig, O. T., a С. Anderson, D. W. Kyhos & P. H. Ra-
ven. 1969. Chromosome numbers in Compositae, VII:
Astereae Ш. Amer. J. Bot. 56: 348-353
. W. Kyhos, А. M. Powell & P. H. Raven. 1972.
Chromosome numbers in Compositae, VIII: Helian-
theae. Amer. J. Bot. 59: 869-878.
L. C. Anderson, D. W. Kyhos, P. H. Raven & L.
tae, V: Astereae III. Am
Swenson, U. 1995a. پا of the Blenriospermatinse
(Asteraceae, Senecioneae). Acta Univ. Ups., Comp
hensive Summaries of Uppsala Dissertations from се
Faculty of Science and Technology 162: 1—34. Up
995b. Systematics of A ich plo:
pacific genus of Asteraceae (Senecioneae). Pl. Syst.
Evol. 197: 149-193
Tobgy, H. A. ems A cytological study of Crepis fuligi-
a, C. neglecta, and their F1 hybrids and its bearing
on the рае ома of its етж рамы in chro-
mosome nu . Genet. 45: 67-1
Tomb, A. S., K. L ре. р. W. ад А. М. Powell
ЈЕ 2 Raven. 1978. Chromosome numbers i
itae,
Turner, B. L. 1970. Chrom
positae. XII. Australian species. Amer. 1 Bot. 57: 382-
– 8: W. Н. Lewis. 1965. Chromosome numbers in
the Compositae. IX. African species. J. S. African Bot.
31: 207— Е.
Н. F. L. Rock. 1961. Chromo-
ѕоте а in the Compositae. У. Rhodora 63: 121-
129.
„ А. M. Powell & J. Cuatrecasas. 1967. Chromo-
some numbers in Compositae. XL Shee species.
Ann. Missouri Bot. Gard. 54
&
54: 172-1
. King. 1962. > mosome num-
bers in the Compositae. VI. Additional Mexican and
64: 251-271.
gedifolit (Composita ecionea
cias (Biologia) Tesis, foreare ees
cd
—. e systematics of Senecio section Mul-
ыз bite raceae: Senecioneae). Ph.D. apes
Claremont Graduate Schoo i
7 РЗ доби Systematics, Proceed
ternational Com ше Сайенс, Kew, 1994. Vol. 1:
597-611. Royal cusan Gardens, Kew.
Vision, T. J. & M. O. Dillon. 1996. Sinopsis de Senecio
L. (Senecioneae, Asteraceae) para e el Perá. Arnaldea 4:
mmm vete dol
904 Annals of the
Missouri Botanical Garden
Appendix 1. General distributions, habit, and apparent basic chromosome numbers, geographical ranges, and habits
of genera of the Senecioneae. Arrangement follows the genera and some of the groupings of Jeffrey (1992) with numerous
modifications to coincide with discussions in the text.
Genus General distribution Habit Chromosome number
Blennospermatinae
Blennosperma Less. W U.S., Chile, annual herbs n=7,9
Crocidium Hook W U.S annual herbs n=9
Ischnea F. Muell. New Guinea annual herbs n=9
Abrotanellinae
Abrotanella (Gaudich.) Cass. S South America, SW perennial herbs n = 9, 18
Pacific
New Guinea Montane Endemic Group
Papuacalia Veldkamp New Guinea woody n = 36
Tussilagininae
Traversia Hook. f. New Zealand shrubs n = 30
Bedfordia DC. Australia small trees n = 30
Brachyglottis J. R. Forst. & G. Forst. New Zealand & Tas- shrubs n = 30
ania
Telanthophora H. Rob. & Brettell Mexico, Central weak shrubs n = 30
America
Pittocaulon H. Rob. & Brettell Mexico, Central seasonally deciduous n = 30
America shrubs
Barkleyanthus H. Rob. & Brettell U.S., Mexico shrubs n = 30
Roldana La Llave SW U.S., Mexico, subshrubs, shrubs n = 30
Digitacalia Pippen
Central America
Mexico, Guatemala
U.S., Mexico
perennial herbs
rosulate perennial herbs
Pippenalia McVaugh Mexico rosulate perennial herbs 30
Robinsonecio T. M. Barkley & Janovec Mexico, Guatemala perennial herbs 30
ina Benth. i subshrubs = 30
Cacaliopsis A. Gray perennial herbs = 30
Rugelia Shuttlew. ex Chapm. E 05. perennial herbs = 28
Arnoglossum Raf. EUS perennial herbs = 93 26,27. 55
Tetradymia W U.S., Mexico shrubs = 30
Lepidospartum A. Gray W U.S., Mexico shrubs = 30, ca. 45
Dolichoglott rd. New Zealand perennial herbs = 30
ndrocacalia (Nakai) Nakai ex Tuyama Bonin Island weak shrubs = 30
Farfugium Lind i perennial herbs = 30
Cremanthodium Benth.
Sinacalia H. Rob. & Brettell
Parasenecio W. W. Sm. & Small
Miricacalia Kitam.
m
China
= 24, 27, 29, 30
= 29
= 30
= 26, 29, 30, 60
Syneilesis Maxi E Asia perennial herbs = 26, 39
Doronicum L Euro perennial herbs = 15, 20, 30, 60
Homogyne Cass. Eurasia perennial herbs = 30, 60, 70, 80
Tussilago L. Eurasia perennial herbs =
Endocellion Turcz. & E Asi perennial herbs = 28, 29, 30
Petasites Mill.
Sinosenecio B. Nord.
Eurasia, North
merica
5 & SE Asia, W
perennial herbs
S 5353553533535353353535335355353353353353353з3з35
| | | ||
= 14, 16, 26, 28, 30
perennial herbs n = 23, 24
North America
Nemosenecio (Kitam.) В. Nord. E Asia perennial herbs n = 24
Tephroseris (Rchb.) Rchb. Central Europe, perennial herbs n — 24, 25
rctic
Volume 84, Number 4 Robinson et al.
1997
905
Chromosome Numbers in Compositae
Appendix 1. Continued.
Genus General distribution Habit Chromosome number
Gynoxoid
Paragynoxys (Cuatrec.) Cuatrec. N Andes shrubs, trees n = ca. 40
Gynoxys Cass. N Andes hrubs n = 40, ca. 40
Chersodoma
Chersodoma Phil. S South America perennial herbs, sub- п = 10
hrub
Senecioninae, numbered groups discussed in text. им
(1) Some generic groups with n = 30 or 50, some with penicillate styles.
Jessea H. Rob. & Cuatr. Central America perennial herbs n = ca. 50
Robinsonia DC. Juan Fernandez weak shrubs n = 20
Lordhowea SW Pacific shrubs п = 19
Arrhenechthites Matt New Guinea shrubs n = ca. 50
Dendrosenecio (Hauman ex Hedb.) B. Central Africa rosette-trees n = 50
Nord.
Pericallis D. Don Macaronesia rennial herbs n = 30
Graphistylis B. Nord. Brazil perennial herbs n = 50
locenes B. Nord. Argentina, Chile perennial herbs n — 20
E € S Africa, Arabia herbs n — 10
Cineraria L.
(2, 3) Senecio including aureoid group and Mulgedifolii.
Senecio L. (incl. Packera A. Lóve € D. America, Eurasia, annual & perennial
Lóve) Africa, Australia herbs
(4) Emilia, n = 5.
Emilia (Cass.) Cass. paleotropics, wide ad- annual herbs
ventive
(5) Genus discussed with Senecio under Senecioninae with n — 10, 20.
Pseudogynoxys (Greenm.) Cabrera tropical America vines
(6) Andean genera Lasiocephalus and Pentacalia, n = 20, 30, 45-52.
N Andes spreading subshrubs
Central America, shrubs & vines
West Indies, South
America
Odontocline B. Nord. West Indies
(7) Culcitium and Dorobaea, n — 20, 50.
Culcitium Humb. & Bonpl. Andes
Dorobaea Cass. Andes
Lasiocephalus Schltdl.
Pentacalia Cass. (incl. Scorbicaria Cass.
& Monticalia C. Jeffrey)
scandent shrubs
perennial herbs
perennial herbs
(8) Werneria and segregates, n = ca. 50, 100.
Werneria HBK Andes perennial herbs
Misbrookea V. A. Funk Andes perennial herbs
Xenophyllum V. A. Funk Andes perennial herbs
Other Senecioninae not discussed in text, giving Jeffrey (1992) subgroups.
(Senecionoid)
Hasteola Raf. (incl. Synosma Raf. ex
Britton . Br.)
Erechtites Raf.
E US. perennial herbs
North & South perennial herbs
Ameri
rica
Crassocephalum Moench tropical Africa, Asia, perennial herbs
n = 10, 18, 19, 20, 22,
23, 30, 40, 46, 50, etc.
n ="5; 8, 10,15
n = 45, ca. 46
n = 20
n
= 20, ca. 40, 45-50,
50
n = 30
n = 20
n = са. 50
п = са. 48, 50, 52, са.
75, са. 77, са. 103, са.
106
п = са. 53
п = са. 50, 54
п = 18,20
п = 20
һ = 10, 20
906
Annals of the
Missouri Botanical Garden
Appendix 1. Continued.
Genus ` General distribution Habit Chromosome number
(Synotoid)
Faujasiopsis C. Jeffre Mauritius shrubs п = 10
Synotis (С.В. it G: Jeffrey & У. L. Chen E Asia subshrubs n = 10, 18, 20
Delairea Le South Africa scandent п = 10
нае
Solanecio (Sch. Bip.) Walp. tropical Africa herbs, weak shrubs n = ca. 90
inia Mill. Macaronesia, Africa, fleshy herbs, shrubs = 9, 10
Asia
Gynura Cass. paleotropics scandent herbs п = 10
(Othonnoid)
Steirodiscus Less. South Africa annual herbs n = 8, 10
Othonna L South Africa, SW succulent perennial = 10, 20
Asia, Australia herbs, subshrubs
us Less. South Africa annual herbs 9
Euryops (Cass.) Cass. tropical & South annual & perennial n = 10, 20
Africa, Arabia herbs, subshrubs
посева nie group)
Iranecio B. N SW rhizomatous herbs n — 12, 20
Dolichorrhiza ds Galushko Caucasus, Iran rhizomatous herbs n = 15-16, 20, 22
Pojarkovia Askerova Caucasus rhizomatous herbs п = 20
асаћа L. Europe izomatous herbs п = 19
навр В. Nord. Caucasus rhizomatous herbs п = 19
BOOK REVIEW
Mori, S. A., G. Cremers, C. Gracie, J.-J. de Gran-
ville, M. Hoff & J. D. Mitchell. 1997. Guide to
the Vascular Plants of Central French Guiana.
Part 1. Pteridophytes, Gymnosperms, and Mono-
cotyledons. Hardcover. ISBN 0-89327-398-8.
Memoirs of the New York Botanical Garden
76(1): 1-422. Retail price: $50 U.S.
In a time when many country-wide or regional
floras are under way in tropical America, yet few are
close to completion, it is refreshing to see that a new
round of smaller neotropical floras is being com-
pleted. The Guide to the Vascular Plants of Central
French Guiana follows the recently published Flora
of St. John by Pedro Acevedo and will soon be suc-
ceeded by a florula of several biological reserves
around Iquitos, Peru, by Rodolfo Vasquez and a flo-
rula of Amacayacu National Park in Amazonian Co-
lombia by Agustin Rudas. Within the next year or
two we should also be regaled by an illustrated field
guide and a separate flora of the Reserva Ducke
outside Manaus, Brazil. Each of these floras covers
between 1000 and 3000 taxa and will provide a solid
baseline of plant data that will facilitate more in-
depth studies at these sites in the future.
This first of two volumes of the Central French
Guiana flora covers the pteridophytes (194 spp.),
gymnosperms (1 sp.), and monocotyledons (426
spp.), as well as a brief introduction, an extensive
“Aids to Identification” section, a key to the major
groups of plants, and a glossary of botanical terms
at the end. Informative and easy-to-use keys are
also provided for the families of monocotyledons
and pteridophytes. Volume 2, which is due out in
late 1998, will cover the estimated 1435 species of
dicotyledons in the study area, for a total of just
over 2150 species of vascular plants in the whole
flora. The area covered by this flora is 1400 km’,
although the level of knowledge and collecting is
clearly concentrated on the immediate surround-
ings of the village of Saiil. I was at first confused
when looking at the map of the flora area (fig. 2),
because the scale indicates a much larger area. To
correlate with the given coordinates and the stated
size of the flora area, the scale bar should read “7
km” instead of “20 km.”
The beauty of this book is that it is so lavishly
illustrated. There are 240 excellent color photo-
graphs by Carol Gracie of plant species and their
distinguishing characters, interspersed in small fas-
cicles throughout the text. There are also 165 line
drawings, most of them illustrating individual spe-
cies, but 8 of these (figs. 4-11) are exquisite full-
page composite drawings by Bobbi Angell that
show features such as different fruit types, leaf
glands, and adaptations for climbing. Two more
full-page figures in the glossary pack in a multitude
of useful characters as well.
I have just a few quibbles with the style and
layout of the volume. The size of the type is too
small, at least where there are pages of uninter-
rupted text. I would like to see more precise or
informative headers used, for example, family
names rather than the few high level groups used
on recto pages or the same journal name repeated
throughout on the verso pages. The “Aids to Iden-
tification” section provides a myriad of valuable
field characters, but the lists go on for a full 36
pages, and a single list such as “Trees, shrubs, and
lianas with glands on the leaves” covers 2% pages
with 57 bulleted entries. There are clearly three
very different styles of illustrations combined in the
volume. Bobbi Angell’s drawings of the monocots
(except palms) are typically elegant and delicate.
The palms, however, are excessively dark and too
highly contrasted. Last, the pteridophyte figures use
dashed lines to separate different species on the
same page and then label them with overly bold-
faced, Leroy-drawn species names. The photo-
graphic inserts are treated as “plates” to distin-
guish them from the black-and-white “figures,” but
then they are numbered with Roman numerals,
which presents a minor challenge when the text
refers the reader to “Plate LXXII.” Toward the end
of the volume there is a nine-page “Index to species
illustrated in Part 1,” but since the general index
already does a fine job of referencing all the figures
and plates, I would recommend not repeating this
kind of index in the next volume.
The taxonomic treatments were prepared mostly
by specialists in the particular families, and the
careful degree of editing is apparent throughout.
Specialized families such as the orchids, grasses,
and sedges have separate figures that effectively
illustrate the main descriptive characters used in
the treatments. The scope of the descriptions,
whether family, genus, or species, is not universal
but rather is restricted to the flora area itself. Con-
sequently, the claim that the flora will help identify
families and genera from other lowland areas in
northern South America will not always hold up.
Ann. Missouri Bor. GARD. 84: 907-908. 1997.
908
Annals of the
Missouri Botanical Garden
Still, botanists from as far away as Manaus have
found a great deal of overlap with this area and
their own local flora. One helpful addition would
be to include some information about each species’
overall distribution, for instance, if it is a narrow
endemic or a widespread American weed.
Altogether, this volume ranks right at the top of
its class. It follows the tradition of extremely infor-
mative local floras like Tom Croat’s Flora of Barro
Colorado Island and then provides the kind of vi-
sual aids that will entice even casual aficionados
to explore the flora of lowland South America, be
it vicariously, browsing through this book, or by
getting their feet dirty and visiting a now well-doc-
umented site such as the region surrounding the
French Guiana village of Saiil—Paul E. Berry,
Missouri Botanical Garden, Р.О. Box 299, St. Louis,
Missouri 63166, U.S.A.
NOTICE
THE 1997 Jesse M. GREENMAN AWARD
The 1997 Jesse M. Greenman Award has been
won by Elena Conti for the publication “Circum-
scription of Myrtales and their relationships to oth-
er rosids: Evidence from rbcL sequence data,” co-
authored by E. Conti, A. Litt, and K. J. Sytsma,
and published in American Journal of Botany
83(2): 221-233 (1996). This study is based on a
Ph.D. dissertation from the University of Wisconsin
under the direction of Dr. Kenneth J. Sytsma.
The Greenman Award, a certificate and a cash
prize of $1000, is presented each year by the Mis-
souri Botanical Garden. It recognizes the paper
judged best in vascular plant or bryophyte system-
atics based on a doctoral dissertation published
during the previous year. Papers published during
1997 are now being accepted for the 30th annual
award, which will be presented in the summer of
1998. Reprints of such papers should be sent to
Dr. P. Mick Richardson, Greenman Award Com-
mittee, Missouri Botanical Garden, P. O. Box 299,
St. Louis, Missouri 63166-0299, U.S.A. In order to
be considered for the 1997 award, reprints must be
received by 1 June 1998.
ANN. Missouni Bor. GARD. 84: 909, 1997.
ANNALS OF THE MISSOURI BOTANICAL GARDEN: CHECKLIST FOR AUTHORS
1. General Instructions
LJ Text is in English or Spanish on numbered pages.
[] Manuscript is typed on one side of nonglossy 8% X
1l in. paper.
LJ At least 1 in. is left as margin all around, except on
the first page, which has 3 in. left blank at the top.
Three copies of double- or triple-spaced printed manu-
script, including abstract, legends, tables, specimen
lists, Literature Cited, and footnotes, are enclosed.
L] Manuscript is also submitted on MS-DOS/WINDOWS
3% in. diskette, as WordPerfect® [preferable] file.
(О Right-hand margin is not justified, and words are not
broken there.
[0 Special typefaces are not used. Common Latin words
or phrases are not italicized (e.g., et al., i.e., sensu,
etc.).
[0 Only names at the rank of genus and below are itali-
cized.
[Г] Correct accents, umlauts, and other diacritical marks
have been included.
Û All figures and tables are cited in the text and аге
numbered in the order in which they are to appear.
[0 Photocopies of the figures are included with each copy
of the manuscript.
2. Style
O Recent issue of the Annals is used as a model
L] Chicago Manual of Style, latest edition, is used as a
reference.
3. First Page
L] Footnotes are typed as double-spaced paragraphs on
the first page. The first footnote contains acknowledg-
ments, including information on granting agencies,
herbaria that loaned specimens, and the name of the
4. Abstract
A one-paragraph abstract precedes text. Papers in
Spanish have an English abstract in addition to a
nish resumen.
O The abstract is concise (1 paragraph) and includes
brief statements about the paper's intent, materials and
methods, results, and significance of findings.
9. Tables
O Tables are neat, double-spaced, and easily understood
rather than long and complex.
(0 Tables do not contain vertical or horizontal lines. [Ed-
itor and/or printer will add them as needed.
ÛJ Captions are typed double-spaced as paragraphs at the
tops of the tables.
O Each table starts on a separate sheet and is double-
paced.
6. Abbreviations
L] Periods are used after all abbreviations (which are
minimized) except metric measures, compass direc-
tions, and herbarium designations.
L] When dates are given as part of collection information,
three-letter month abbreviations are used, except for
months with four letters, which are spelled out in full.
States are not abbreviated, and cities are spelled out.
bbreviated according to Brummit
& Powell's Authors of Plant Names.
О Book titles are abbreviated according to Taxonomic Lit-
erature, edition 2, but with initial letters capitalized.
itles are spelled out in the Literature Cited.
L] Herbaria are abbreviated according to Index Herba-
riorum, edition 8.
7. Taxonomic Treatment
author, literature citation, type citation, e.g., Pleurothy-
rium amplifolium (Mez) Rohwer, Mitt. Inst. Allg. Bot.
Hamburg 20: 43. 1986. Nectandra amplifolia Mez, Ar-
beiten Kónigl. Bot. Gart. Breslau 1: 131. 1892. TYPE:
Brazil. Rio de Janeiro: Alto Macahé, Glaziou 17731
(holotype, B; isotypes, B, G, K, NY, P).
[0 Lectotype designations are included together with an in-
dication of where they were designated, the year, and the
author. This reference is listed in the Literature Cited. If
d
and types not seen are indicated as such (e.g., MO!,
US no
ri
for parenthetical extremes: “peduncle (8.2—)14.3—
28.0(-31.9) cm long," unless intermediate values are
not expected: ovary with (2)4(6) locules. Length X
width are given in the following manner: lamina 36.4—
82.8 X 9.1–16.8 ст
When relevant, nomina nuda, misapplied names and
excluded names are included in the discussion follow-
ing the description, or at the end of the paper, but are
not part of the formal synonymy.
О
8. Specimens Examined
O If many specimens were examined, those cited in the
text are limited to ca. 1% manuscript pages.
i i ed is placed at the end
of the paper, following the Literature Cited. It is ar-
ranged alphabetically by collector, followed by collec-
tion number, followed by the number of the taxon in
the text. Names (including initial(s) of first and second
ing are provided, “et al." if three or more.
ns are cited in the text as follows: Additional
12737"N, 85714", 950-1100 m, 3 Feb. 1987 (fl), Jer-
gensen 865 (BM, G, K, US). [Dates and reproductive
status are optional but are omitted from longer lists.]
Countries are run together in the same paragraph, e.g.,
COUNTRY А. Major pues division: . OUN-
TRY B. Major political division: . . oem par-
agraphs are used for major Ишен АР regions within
major political divisions.
9. Vouchers and Genetic Sequences
[0 If paper presents original data, it includes the oon
of herbarium vouchers, as well as vouchers for
collections, etc. очи on the type of paper, pat
erence to the original wild source may also
quired.] oran are also cited from common names
and uses taken from specimen labels
Herbarium vouchers state the collector and number,
herbarium in which the voucher is located, and a clear
annotation that the material represents the voucher for
the study in question.
O a ne have been banked, and accession
mbers ar ided.
O pem ign esponsibility 1 for establishing the ac-
curacy of information provi
О
10. Keys
O Keys are clear and have been checked carefully for
consistency with the descriptions. Leads of each cou-
plet are parallel.
O Dichotomous keys are indented.
O oo taxa are keyed separately, not in species
key:
11. Literature Cited
[0 bae аар Cited contains all references cited in
the
O All pss in the Literature Cited are cited in the text.
O Spelling of author(s) ме and years of publication
have been double-checked
[0 All entries have been verified against original sources,
especially journal titles, accents, diacritical marks,
ows: ашћогз lunt name,
initial(s). Year. Full title of — Journal abbreviated
as in B-P-H/S. Volume: o parenthetical part
numbers after volume Mision are given unless each
part is paginated separa
C] For more than one author, this style is followed: au-
thors last name, initial(s), second author's initial(s),
last name & third Uum initial(s), last name.
L] Books appear as follows: author's last name, initial(s).
Year. Full Unabbreviated Title (edited br Editor), 3rd
is Vol..2. incor City of Publicatio:
O Сион of work “in prep., {зшен theses and
dissertations, and iura references to inaccessible
sources have been eliminated or kept to a minimum
They are not included in the Literature Cited.
–
o
—
<
12. Illustrations
O Ê name(s) and figure number(s) are written in
pencil on the back o figure or plate, on both
originals d review co
O Scale bars appear on al photographs, and
O Magnifications/reductions are not indicated in cap-
tions.
O All illustrative materials are mounted on stiff card-
board no thicker than % in
O Illustrations are presized to fit either column width (2%
in. or ca. 68 mm) or full page width (5% in. ог са. 140
mm), or illustrations no larger than 11 X 16 in. (= 29
х 40 cm). [Maximum size for printed а is
5% X 8% in.] Oversized or delicate figures are su
mitted as photostats.
O Figures are numbered in Arabic numerals іп the order
of their citation in the text. Parts of figures are labeled
with capital letters
O Photographs are crisp black-and-white prints.
O Figures are grouped into composite plates when pos-
sible; edges of photographs are abutt
O No stripping is inserted between бие or figure seg-
ments. [Printer will insert stripping.]
O суза of figures are squared.
O Maps include reference to latitude and longitude and
are bounded by a fine border.
O Seanning electron E инно are free of conspicuous
charging.
O Axes on graphs are all labeled.
O Казр provide all explanatory text. No text appears
n the figures. Captions are separate from other text,
one paragraph for each group of figures, and following
the style in current issues of the Anna
O Symbols on maps are legible and reduction has been
taken into consideration.
Electronic Artwork—Printer's Specifications
1. We accept line art, halftones and oe figures on disk
or via electronic delivery. nclude a printout or
email of the file directory that س the file name(s),
size and kind of file.
~
. You should discuss figure sizing specifications with the
journal prior to disk submission, and the figures on
disk should be the final size desired.
Graphic files should be submitted in TIFF or EPS file
format. Other applications that are acceptable are as
follows: Adobe Photoshop; Corel Draw; Adobe Illustra-
tor; Adobe Freehand.
. All line art should be scanned at 1200 d.p.i. All gray-
scale and color images should be scanned at 350 d.p.i.
Color images should be submitted in CMYK mode, not
RGB or Indexed Color mod
. Include the screen and printer fonts used within the
graphic file. Use only Adobe Type 1 postscript fonts.
True Type fonts and System *bitmap" fonts should not
be used.
. If the files have been compressed, please indicate what
compression program has been use
The disk submission should be accompanied by a hard
copy of the figure that exactly matches what is on the
disk in the event that the disk is unuseable.
P
+
e
е
а
Acceptable Electronic Artwork Media
3%" floppy disks; Syquest cartridges; EZ Flyer; ZIP
disks; JAZ disks; Magneto Optical disks; CD ROM;
Artwork may also be submitted via email attachment
or FTP. Please contact Tara Schuley, e-mail address:
tschuley@allenpress.com for more information.
13. What to Submit
O Full mailing and email ee tah and fax
numbers are submitted in cover let
ÛJ Three copies of manuscript are ара НА
O —— of original figures are enclosed
L] An MS-DOS/WINDOWS 3% in. diskette, as Word-
Perfect? [preferable] file, is submitted in addition to
the printed documents.
SCOPE OF THE ANNALS
The Annals publishes original articles in systematic bot-
any and related fields. Papers whose purpose is the estab-
lishment of new nomenclature entities in vascular plants
and bryophytes are not accepted. Rather, they should be
submitted to Novon for consideration. (Novon instructions
are available from the managing editor.
CONDITIONS FOR PUBLICATION
Authors are urged to contact the managing editor prior to
submitting manuscripts. Manuscripts sent already re-
viewed and revised are generally subject to the full normal
review process initiated here. Manuscripts not properly
prepared may be returned for revision prior to review. It
is the authors responsibility to supply missing biblio-
graphic information; the editor does not perform library
ior arrangement with authors not
having access to crucial reference materials. Communi-
cation with the editor at all points of the publication pro-
cess is encouraged.
PAGE CHARGES
Page costs are $80 per page, although charges may be
reduced under certain circumstances. Corrections in proof
will be billed nonnegotiably to the author at $3 per line
changed.
CORRESPONDENCE
Amy McPherson,
Managing Editor
Scientific Publications
Missouri Botanical Garden
P.O. Box 299
St. Louis, MO 63166-0299,
USA.
Tel: (314) 577-5112
Fax: (314) 577-9594
Electronic mail:
amcpher@
admin.mobot.org
Updated 9/97
Volume 84, Number 4, pp. 705-912 of the ANNALS OF THE MissoURI BOTANICAL GARDEN
was published on January 15, 1998.
Experimental and Molecular Approaches to Plant Biosystematics
The proceedings of the Fifth International Symposium of the International Organization of Plant
Biosystematists ПОРВ)
Edited by Peter C. Hoch and A. G. Stephenson
Twenty-three original contributions that span the breadth of biosystematics, a dynamic field of study
that bridges the realms of systematics and population biology. The papers are arranged in four groups,
reflecting the original four symposia of the 1992 meeting. DNA and Plant Biosystematics presents
innovative work that uses the rapidly developing nucleic acid methods adapted from molecular biology.
Plant Growth Patterns and Biosystematics includes comparative and developmental analyses of plant
architecture and branching patterns. Plant Reproductive Strategies surveys new approaches in the anal-
ysis of plan} reproductive biology, an area central to both systematic and population-level studies.
Phylogenetic Analysis and Population Biology emphasizes the application of the powerful new methods
of phylogenetic analysis to problems at the species and population levels. Monographs in Systematic
Botany from the Missouri Botanical Garden, Volume 53. ISBN: 0-915279-30-4. 416 pp. Illustrated.
1995. $60.00 U.S. $62.00 Non-U.S. |
Annals of the Missouri Botanical Garden, Volume 82, Number 2: Alternative Genes
for Phylogenetic Reconstruction in Plants | :
А symposium cosponsored by the American Society of Plant Taxonomists and the Botanical Society of
America, organized by Pamela S. Soltis and Douglas E. Soltis, and presented at the 1993 AIBS meetings.
Although the chloroplast gene rbcL has been successfully used to reconstruct plant phylogeny, many
important questions of plant phylogeny and evolution. cannot be addressed using it. The contributors to
this issue of the Annals explore the potential of eight alternative genes or DNA regions for phylogenetic
reconstruction at a variety of hierarchical levels. Both nuclear and chloroplast genes are evaluated. Three
ored: the 185 gene, the internal transcribed spacers
(ITS), and the 26S gene. Small multigene families from the nuclear genome may also carry phylogenetic
signal: the phytochrome gene family and the small heat shock gene family. Thre chlo-
roplast genome are also considered: atpB, ndhF, and тик. Each paper describes с e
structure, and rate of evolution of the chosen gene and discusses its potential for phylogenetic study. This
issue also contains: “The Comparative Pollination and Floral Biology of Baobabs (Adansonia-Bombec
. ceae)" by David А. Baum and “In Memoriam: Peter G. Martin.” Annals 82(2) 1995. 174 pages. $27.
U.S. $28.00 Non-U.S. ; | ато | avit
Я В Y 4 - - | : а | 4 " а | : 1 e |
To order, please indicate method of payment below. Checks or money orders should be in na lli $
berths through a U.S. bank, to Missouri Botanical бира Or must be Medos No Mapa
ee sndling fee will be to orders requiri و y
се on all orders. Additional $2.00 handling fee wil 314) 577-9534, fax: (314) 577-9594, аай;
are made until payment is received. Phone: (
` deptllGmobot.org. | | 4
Send order to: , A EE ` Please send — copy(ies) of Monograph No. 53
Department Eleven, Missouri Botanical Garden Please send ——- copylies) of Annals 82(2)
P.O. Box 299 | |
St. Louis, MO 63166-0299, U.S.A. AS >
О Check/money ondex enclosed: · Send books to: - (
O Send invoice ($2.00 fee will be added to total) - ze
O Charge card number (MasterCard/Visa) Р A а LL
| . · Name Ж egre нам
Expiration LPS аа qn od a Om : Add EB e dec Eu тхл r r i 15
Мате as it appears on card | | ress 455 e |
Telephone number (daytime) AENEA Eee уя РЕ О
Postal Code
| 8414)
س ن
- Prices ARE Suner то CHANGE Wrriour NONE
СОМТЕМТ5
A Revision of Styrax (Styracaceae) for Western Texas, Mexico, and Mesoamerica __
Peter W. Fritsch
Systematics of Kalimeris (Asteraceae: Astereae) ____ Hong-ya Gu & Peter C. Hoch
A Review of the Genus Cydista (Bignoniaceae) Warren D. Hauk
Systematics of Eleusine Gaertn. (Poaceae: Chloridoideae): Chloroplast DNA and Total
Evidence __. Khidir W. Hilu & John L. Johnson
The Fruits of Jasminum mesnyi (Oleaceae), and the Distinction Between Jasminum
and Menodora _ Jens С. Rohwer
Revisión del Generi Galianthe subg. Ebelia stat. nov. (Rubiaceae: Spermacoceae) _
— Elsa L. Cabral y Nélida M. Bacigalupo
Género Galianthe subg. Ebelia (Ка. рсе: Estudio Palinológico __
Stella Maris Pire
The Tanaka-Kaiyong Line—An Important Floristic Line for the Study of the Flora of
t Asia .. Li Xi-wen & Li Jie
3 Chromosome Numbers in Compositae, XVII: Senecioneae III
eflet o ‘Harold Robinson, Gerald D. Carr, Robert M. King & A. Michael Powell
Book Review. Guide to the Vascular Plants of Central French Guiana. Part 1 by S.
А. Mori et al., reviewed by Paul E. Berry __
Notice
Checklist for Authors