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Revision of Dracontium (Araceae)

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Revision of Dracontium (Araceae) REVISION OF DRACONTIUM Guanghua Zhu2 and Thomas B. Croat2 (ARACEAE)1 ABSTRACT A taxonomic revision of the genus Dracontium L. (Araceae) is presented predominantly from morphological, anatom- ical, and phytogeographic data. Dracontium is treated as comprising 23 species, including 10 new species: D. ama- zonense G. Zhu & Croat, D. angustispathum G. Zhu & Croat, D. asperispathum G. Zhu & Croat, D. bogneri G. Zhu & Croat, D. grandispathum G. Zhu & Croat, D. grayumianum G. Zhu & Croat, D. guianense G. Zhu & Croat, D. peruvianum G. Zhu & Croat, D. plowmanii G. Zhu & Croat, and D. prancei G. Zhu & Croat. Five names are newly lectotypi®ed herein: Amorphophallus nivosus Lem., A. papillosus hort. ex Rafarin, Dracontium elatum Masters, D. pittieri Engl., and D. soconuscum Matuda. The following species are newly synonymized: D. carderi Hook. f., D. costaricense Engl., D. dressleri Croat, D. lineare G. S. Bunting & Tillett, D. loretense K. Krause, D. ornatum K. Krause, D. trianae Engl., and Echidnium regelianum Engl. All taxa are described and illustrated. Key words: Araceae, Dracontium, monograph, taxonomy. The Araceae are a family of herbaceous mono- sist of a single leaf arising from an underground cots with 106 genera and approximately 3500 spe- tuber with a more or less mottled, reptilian-pat- cies, the vast majority occurring in the New World terned petiole up to several meters long with a tropics (Croat, 1988, 1992). The World Checklist highly divided, compound, spreading blade reach- and Bibliography of Araceae (and Acoraceae) lists ing as much as three meters or more in diameter. a total of 2824 recorded species (Govaerts & Fro- In¯orescences of both genera are notorious for their din, 2002). Members of the family are highly di- fetid scents at anthesis. However, Dracontium dif- verse in life form, leaf morphology, and in¯ores- fers sharply from Amorphophallus in having bisex- cence characteristics. Life forms range from ual ¯owers on a uniform spadix. submerged or free-¯oating aquatics to terrestrial Accurate identi®cation of Dracontium species (sometimes tuberous), epiphytic or hemiepiphytic has been dif®cult because of the paucity and in- plants, to climbers. Leaves range from simple and adequacy of herbarium material and their sporadic entire to compound and highly divided, and may phenology. Herbarium specimens, though compar- be basal or produced from an aerial stem. The fam- atively abundant for some species such as Dracon- ily is best characterized by its distinctive in¯ores- tium spruceanum (Schott) G. Zhu, are frequently cence, a spadix with bisexual or unisexual ¯owers sterile and often incomplete since mature adult (sometimes with a sterile region) subtended by a leaves are almost always too large to be mounted solitary pedunculate spathe. on a single herbarium sheet. Frequently, a single The present study of Dracontium L. comprises a leaf is mounted on several to many herbarium taxonomic revision of one of the most poorly known sheets. Hence, intensive study of living material in tuberous genera in the Araceae. Dracontium, as the ®eld or under cultivation is essential. Though here recognized with 23 species, represents the local populations may contain many plants, popu- most species-rich tuberous genus of the Araceae in lations of Dracontium are often widely scattered the New World. Vegetatively, it shares a great sim- and single individuals not frequently encountered. ilarity with the Old World genus Amorphophallus Under unfavorable growing conditions, plants of Blume ex Decne. Plants in both genera usually con- Dracontium may remain vegetative for many years, 1 This revision is based on a study completed as a partial ful®llment of a Ph.D. degree in biology for the ®rst author at the University of Missouri±St. Louis. Many individuals have contributed to the study. We thank John Banta, Josef Bogner, Julius Boos, David Boufford, Dewey Fisk, Mike Grayum, Wilbert Hetterscheid, Victoria Hollowell, Richard Keating, Petra Schmidt, Emily Yates, Dan Nicolson, P. Mick Richardson, Hong Song, Hans Vissers, Henk van der Werff, and Scott Zona for providing plant materials and helpful communications. We also thank the curators of the following herbaria for providing loans of specimens: AAU, B, BBS, BM, BR, CAY, CM, COL, CR, DUKE, EAP, ECON, F, FTG, G, GH, IAN, IBE, INB, INPA, JAUM, K, M, MEXU, MG, MOL, NY, P, PMA, PORT, QAME, QCA, QCNE, R, RB, RSA, S, SEL, SJ, SP, TRIN, TULV, U, UCLA, US, USJ, VEN, and WIS. This research was supported by NSF grant DEB-9310171, an award from the International Center for Tropical Ecology at UM±St. Louis, and an OTS 92-3 Fellowship. A teaching assistantship from the University of Missouri±St. Louis and an Andrew W. Mellon Foundation Fellowship provided partial ®nancial support. 2 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ANN.MISSOURI BOT.GARD. 91: 593±667. 2004. 594 Annals of the Missouri Botanical Garden and often go dormant for two to four months in the gram (Dallwitz et al., 1995). The program TAXA- dry season. In¯orescences of Dracontium often SOFT (Gouda, 1994) was used for data mainte- arise from the apex of the tuber before or after leaf nance of DELTA. The exsiccatal information was development. Therefore, matching fertile with ster- prepared by using TROPICOS. Ecological zones ile herbarium material of the same Dracontium spe- are estimated by using the Holdridge Life Zone sys- cies is often problematic. tem (Holdridge et al., 1971). Live plants of Dracontium are not uncommon in Material for chromosomal studies was obtained cultivation in botanical gardens and aroid nurser- from plants cultivated at the Missouri Botanical ies, as they share an ornamental appeal with Amor- Garden. Growing root tips were treated in 0.002M phophallus species and are easy to propagate and 8-hydroxyquinoline for 4 hr. before being ®xed in maintain. Unfortunately, cultivated material of Dra- ethanol-glacial acetic acid-chloroform (6:3:1) at contium is limited to only a few species, such as 88C overnight. Root tips were then rotated through D. gigas (Seem.) Engl., D. polyphyllum L., and D. a hydrated 70%±50%±25%±10% ethanol series, spruceanum. Locating these plants in the ®eld is were macerated with 1N HCl (room temperature) often not dif®cult since populations of Dracontium for 10 minutes, 1N HCl (608C) for 8 min., and 1N are usually well known to native people due to the HCl (room temperature) for another 10 min., and distinctive, reptilian-patterned petioles and medic- rinsed with deionized H2O for 30 min. before being inal uses. stained in 0.5% Feulgen for 1.5 hr. (Masahiro Tak- ei, pers. comm.). Finally, the plant material was MATERIALS AND METHODS rinsed with deionized water, and slides with the chromosome squashes were prepared. This revision was conducted mainly at the Mis- souri Botanical Garden, where the largest living TAXONOMIC HISTORY and herbarium collections of Dracontium in the world are housed. Field research for this study was The ®rst mention of a species referable to Dra- conducted in Costa Rica, Panama, and Brazil be- contium appeared in Hermann's Paradisi batavi tween 1992 and 1995. Trips were made to several prodromus (1689: 315) as ``Arum polyphyllum, dic- botanical gardens to study cultivated plants, in- tum Dracontium, caule scabro punicante, surina- cluding the Marie Selby Botanical Gardens and The mense'' and ``Dracontium americanum caule scabro New York Botanical Garden in the Unites States, puniceo radice Cyclaminis.'' The plant was intro- and the SaÄo Paulo Botanical Garden and Botanical duced into the gardens of Holland by the early Garden of Rio de Janeiro in Brazil, as well as to Dutch plant-hunters from Surinam in the second herbaria CR, F, GH, HB, IAN, IBGE, ISC, MG, NY, half of the 17th century. A corresponding plate was PMA, R, RB, SCZ, SEL, SP, SPSF, UB, UEC, US, later published by Plukenet (1696, t. 149, ®g. 1) in and WSU. Over 670 collections comprising about his Almagestum under the name ``Arum polyphyl- 3000 herbarium sheets, including loans from 49 lum surinamense caule atrorubente glabro et ele- major institutions worldwide, were studied (see Ap- ganter variegato.'' This same species was rede- pendices 1, 2). Data from all cited specimens are scribed by Hermann (1698: 93±94) in his recorded in the Missouri Botanical Garden data- Paradisus batavus as ``Dracontium americanum base TROPICOS. scabro puniceo caule radice Cyclaminis,'' along with In this study, we have relied heavily upon live a ®gure (t. 93) under the name ``Arum polyphyllum collections, especially the second author's gatherings caule scabro punicante.'' Linnaeus (1737) published during his more than 35 years of ®eldwork in the the ®rst comprehensive description of this species Neotropics. Some species, such as Dracontium gui- after he personally studied a ¯owering specimen in anense G. Zhu & Croat, are known only from a few the Clifford Garden at Hartecamp, Holland. Later, herbarium collections. In such cases, descriptions he validly published the genus Dracontium with are from herbarium specimens only. Due to the rarity ®ve species in Species Plantarum and established of many Dracontium species, all specimens with the binominal Dracontium polyphyllum (Linnaeus, proper collecting information are cited, including 1753), which is the only species still recognized those prepared from cultivated plants. For species within Dracontium. Under D. polyphyllum, Linnae- determinations, we have also relied on photographs us (1753: 967) cited both Plukenet's and Her- prepared by Croat and many other individuals to mann's ®gures. Hermann's ®gure was later selected reconstruct the natural habits for these plants. as the lectotype of D. polyphyllum (Hay, 1992). Descriptions of species were prepared using the Since D. polyphyllum was referred to as ``typical'' DELTA (DEscriptive Language for TAxonomy) pro- by Britton and Wilson (1923: 130), it was accepted Volume 91, Number 4 Zhu & Croat 595 2004 Revision of Dracontium as the generic type of Dracontium (Jarvis et al., portance of the ¯oral characters, Engler (1889) 1993: 43).
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