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. 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(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. 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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. 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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. 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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. 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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. 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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. 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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 € —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. 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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 | | | | . 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. 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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. 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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