Croat, 2015 A Review of Studies of Neotropical A Review of Studies of Neotropical Araceae

Thomas B. Croat P.A. Schulze Curator of Botany Missouri Botanical Garden P.O. Box 299, St. Louis, MO 63166 [email protected]

ABSTRACT especially section Porphyrochitonium, are The last decade has seen many changes in studied. A Lucid multichotomous key for research with neotropical Araceae. There Anthurium was created by staff at Kew have been a lot of revisionary efforts Gardens and has been used to key out especially with smaller genera but relatively potential new . The diagnosis of the few changes in the largest and most putative species can then be written to show complex genera. Several key floristic studies differences from species already published. have been completed and several more are It has been continuously upgraded during underway. Increasingly phylogenetic studies the last five years by adding new species and are being carried out among genera of all by making modifications in the key to make sizes. This review will provide a history of it more useful. The Lucid Anthurium key past monographic and floristic efforts and now contains about twice as many species will summarize by suggesting the areas most as were present when initially made in need of further work. available.

A recent paper by Croat & Carlsen (2013) REVISIONARY STUDIES realigns the palmately lobed Anthurium The largest , Anthurium, remains still species, merging Schott’s section poorly known overall with only three Schizoplacium to section Dactylophyllium and sections, namely Pachyneurium (Croat, 1991), excluding three Mexican species and a single Polyphyllium (Croat, 1978) and Semaeophyllium Colombian species. (Croat & Carlsen, 2004) that have been revised. With the discovery of many new Marcus Nadruz Coelho published a species, especially in the of Western revision of Anthurium section Urospadix South America, the number of named and subsection Flavescentiaviridia (Coelho et al., described (albeit not yet published) species 2009) and Livia Temponi (Temponi, 2006) exceeds 1690 and the upper limit will did a molecular analysis on section certainly exceed 2000 when more sections, Urospadix resulting in considerable

Aroideana VOL 38E NO 1, 2015 44 Croat, 2015 A Review of Studies of Neotropical Araceae realignment of the section and the sinking expected to be submitted for publication of section Chamaerepium, a previously sometime in 2015. endemic section to Brazil. Revisionary work is being carried out on Philodendron is perhaps more fully known, Monstera for the Flora of Mesoamerica with subgenus Meconostigma (Mayo, 1991; (Croat, in prep.) and Alejandro Zuluaga, Gonçalves & Salviani, 2002), the subgenus now a Ph.D. student at the University of Pteromischum from Central America, West Wisconsin, Madison, is concentrating on a Indies and Pacific Tropical South America molecular phylogeny of the genus. The very and Central America having been fully useful revision by Madison (Madison, 1977) revised (Grayum, 1992; 1996). Some is now somewhat out of date, especially published work also has occurred in the with many new species in Central America Guianas (Croat & Grayum, 1994). In (Grayum, 1997). addition Philodendron subgenus Philodendron section Macrobelium of Brazil has been The genus Spathiphyllum with an estimated revised (Sakuragui, 1998; Sakuragui & D. C. 63 species was last revised by Bunting Zappi, 2005). A Lucid multichotomous key (Bunting, 1960) where 36 species were for Philodendron prepared by Marcela Mora covered. Spathiphyllum is now being studied contains most of the species with the by Felipe Cardona at the Universidad de exception of some new species from Brazil. Antioquia (Cardona, 2000; 2001; 2004). In The inital key with 465 species resides on Central America there are about 25 species the CATE website and Thomas Croat and of Spathiphyllum. his staff of volunteers are continuing to populate it with new species. It now Stenospermation, with an estimated 250 contains 529 species for an increase of 12% mostly undescribed species is one of the during the past 6 years. most poorly know of all aroid genera. No complete revision has been made since that The genus Dracontium with 23 species has of Engler in Das Pflanzenreich (Engler, been revised (Zhu & Croat, 2004) and a 1908) which covered only 20 species. A Lucid key for Dracontium is being prepared revision of Stenospermation for Central by Croat. The Chlorospatha with 68 species America (A. P. Gómez, 1983) comprised (Croat & L. P. Hannon, 2015; in press) have only 9 species but since that time many now been revised and a Lucid key is species have been found, especially in contemplated as well. Panama and Costa Rica with as many as 30 species now reported (Croat, in prep.). Adelonema with 16 species has recently Numbers of new species are even larger in been revised (Croat et al., in prep) and is South America. A recent survey of Stenospermation in Carchi Province in

Aroideana VOL 38E NO 1, 2015 45 Croat, 2015 A Review of Studies of Neotropical Araceae

Ecuador yielded 26 new species. Natalia The study compared subtribe Spathicarpeae Castaño Rubiano from Colombia is working with the related subtribe Bognerineae and on a revision of Colombian species of subtribe Dieffenbachineae. Stenospermation (Castaño Rubiano, 2013) and is also working on the creation of a Lucid Rhodospatha with about 79 species has been key for the genus. at least partially revised and a Lucid key to the genus is anticipated. with an estimated 140 species has been revised for Central America (Croat, 2004) with 26 species represented FLORISTIC STUDIES but most of the species in South America Much of the neotropics is undergoing a are still poorly known. An unpublished key floristic review of the Araceae. In Central to South America with 104 species has been America completed floristic studies have prepared (Croat, in prep.) and a Lucid key been made for Costa Rica (Grayum, 1982; to Dieffenbachia is being planned. Grayum et al. 1983; 2003, 2003a), Guatemala (Croat & Vannini, in prep.) and Xanthosoma with an estimated 90 species Nicaragua (Croat & Stiebel, 2001), Veracruz remains poorly known, certainly one of the (Croat & Acebey, 2015). The Flora of most poorly known genera in the Mesoamerica treatment of the Araceae is neotropics. Central America has about 13 expected to be completed by 2016. species but there are many new species in Although the largest genera Anthurium the western Andes (Croat et al., in prep.) (Croat, 1983; 1986) and Philodendron (Croat, and even in the Guianas (Croat et al., in 1997) and well as several smaller genera, press). Eduardo Gonçalves is working on Chlorospatha (Croat & L. P. Hannon; in Xanthosoma and has described 10 new press), Dieffenbachia (Croat, 2004), species (Gonçalves, 1999; 2000; 2011). Dracontium (Zhu & Croat, 2004) have been completed for Central America this Flora of Most of the smaller neotropical genera are Mesoamerica treatment will result in the by now relatively well known, especially the first revision of most of the smaller genera. recently revised Spathicarpeae inhabiting Floristic studies in South America are a eastern South America and the western more daunting task owing to its much Andes (Gonçalves, 2002a). That revision greater complexity. The Andean countries covered initially 8 genera in subtribe have been partly reviewed with modern Spathicarpeae, namely Incarum, Asterostigma, Araceae Checklists having been created for , Synandrospadix, Mangonia Peru (Croat, 1993), Ecuador (Croat, 1999) Gorgonidium, and Gearum. Later and Bolivia (Kessler & Croat, 1999; Croat & the new genera Croatiella (Gonçalves, 2005) Acebey, 2015). Colombia, clearly the largest and Lorenzia (Gonçalves, 2012). were added. floristic region in all of South America does not have a published flora but an effort is

Aroideana VOL 38E NO 1, 2015 46 Croat, 2015 A Review of Studies of Neotropical Araceae underway by a group of Colombian aroiders close relationship between the Spathicarpeae to produce such a checklist. Venezuela is to both and Dieffenbachia.. reasonably well known floristically owing to work by Bunting (1979) as well as a later Alejandro Zuluaga is carrying out study by Croat & Lambert (1986). The molecular studies with Monstera and these Guianas are now reasonably well known studies will be published as a part of his floristically after a National Geographic Ph.D. thesis. Molecular studies carried out Sponsored expedition of four months made by Peter Boyce and Sin Yeng Wong to the region. The flora of the Guianas principally on Old World Homalomena have comprises about 160 species and treatment shown the American element, namely is expected to be completed in four years. Homalomena section Curmeria Linden & Brazilian aroiders are also generating a André to be distinct from Asian species so Checklist of the Araceae and the current that group will adopt the generic name, total is 477 species; has only17 Adelonema (Wong & Croat, in press). species (Crisci, 1971). Paraguay, the only Adelonema has 12 species and ranges from other country with a significant aroid flora Costa Rica to most of northern South has a completed flora with 9 genera and 16 America as well as the Amazon basin. As species of Araceae (Croat & Mount, 1988). yet unpublished molecular studies with 8 species by Merrow & Croat showed Floristic studies are more all encompassing Adelonema to cluster in three well-supported with published or in press floristic studies clades. Two molecular studies (Gauthier et having been carried out in Paraguay and al., 2008; Tam et al., 2004) found Adelonema checklist treatments of Ecuador, Peru and to fall within Philodendron but this conflicts Bolivia have been completed in the past ten with studies by Wong & Croat ( in press) years. Floristic accounts of Venezuela have that show it to be distinct from Philodendron, been completed and a checklist for the though most closely related to Philodendron Flora of Colombia and for is nearing subgenus Pteromischum and completely completion. The Araceae for the Flora of distinct from the Asian Homalomena. Mesoamerica and for the Guianas is expected to be completed in the next five Currently two separate molecular studies years. are being carried out on Philodendron, one by Nils Köster and Dubán Canal working at the Berlin Botanical Garden and one by MOLECULAR STUDIES Santelmo Vasconcellosas from Brazil. Both Molecular studies of neotropical Araceae involve collaboration with Croat. are still in their infancy but significant studies have been made on Tribe Molecular studies of Anthurium (Carlsen, Spathicarpeae by E. G. Gonçalves pers. com.) have successfully separated 18 (Gonçalves, 2002b). His studies showed a

Aroideana VOL 38E NO 1, 2015 47 Croat, 2015 A Review of Studies of Neotropical Araceae clades and have major advances in our Danny Beath, employing techniques understanding of the sectional classification learned in Central West Africa made of Anthurium. Among the findings are a significant observations on pollination of validation of a West Indian clade Anthurium and Dieffenbachia while working in corresponding to Engler’s section Costa Rica (Beath, 1999). Julio Sierra- Episeiostenium, the isolation of two new Giraldo has made observations on distinct Mexican clades, one of which pollinators in the Andes of western South corresponds to Schott’s section Andiphlium America in Risaralda Department and and one which represents species with reported what appears to be a likely cordate-sagittate blades with dark glandular pollination system involving frogs punctations on the lower blade surfaces. pollinating Araceae (Sierra-Giraldo, 2014). Anthurium section Pachyneurium series Multinervium is recognized at the sectional Considering the vast array of syndromes level as section Multinervium Croat and and the incredible richness of the Anthurium at the sectional level are either Neotropical aroid flora pollination, the completed or ongoing. biology of Araceae is likely to be a very lucrative field of study for many years to Molecular studies are also being carried come. out on Philodendron and Monstera. DISCOVERY OF NEW SPECIES POLLINATION STUDIES Perhaps the most significant effect of Pollination biology of Araceae is still much research with Araceae in recent years is the understudied and most serious studies of realization that more than any other tropical pollination are still in their infancy in the family so far studied the Araceae neotropics with only a few total species represents a group that is proving to be having been studied for their pollinators but exceedingly rich in species. Most genera several important studies are ongoing. have grown in size (numbers of species) but percentage-wise the growth has been Nevertheless we have strong intitiatives, greatest in the two largest genera of especially through the efforts of Marc Araceae, namely Anthurium and Philodendron. Gibernau working in French Guiana and Anthurium has grown from about 600 sometimes working with Denis Barabé at species to 1392 species in less than 5 years the Montreal Botanical Garden, Quebec, with most of the growth in species coming Canada, Gerhard Gottsberger working in from Panama, Colombia, Ecuador and Peru. Brazil and Heiko Hentrich from Germany, Philodendron, though not heavily studied is working in Mexico with Pedro Diaz. also growing dramatically from 469 species to 529 species at present. Moreover it looks

Aroideana VOL 38E NO 1, 2015 48 Croat, 2015 A Review of Studies of Neotropical Araceae like this rate of increase will not soon increased by 9 species; Schismatoglottis with diminish with the addition of hundreds of 110 species has increased by 35 species; new collections from areas not previously Schottariella a new monotypic genus is explored. This single attribute of Araceae, described and Schottarum with 2 species is namely the ability to produce many new described. In Homalomena with 93 species species, makes it seemingly unique though I about 40 species have been described by do not know what is happening to other Boyce and or Sin Yeng alone. families. Single site visits often yield an array of unknown species, most of which are At least in the neotropics and Asia where apparently new and the same thing could be vast expanses of forest in poorly explored found true in clearly related sites. Despite areas still exist and especially in light of the repeated visits to the El Queremal- high rate of narrow endemism in the family, Anchicaya region on the western slopes of this process of discovery of new species is the Andes between Cali and Buenaventura, likely to continue for a long time. each visit presents a surely unique set of discoveries of new species for each stop. USE OF LUCID MULTICHOTOMOUS KEY Interestingly, though perhaps less dramatic, the situation in Asia also shows that many What have made determinations of new new species still exist there. Since 1980 species possible are the Lucid keys which Amorphophallus has grown by about 90 have been developed for both Anthurium species and now totals 210 species. About and Philodendron. The Lucid database stores 80% of this growth is due to the efforts of all of the important qualitative and Wilbert Hetterschied. Both Homalomena and quantitative information on all published Schismatoglottis have grown dramatically species for all species in a specific genus. owing to the efforts of Hay & Yuzammi The key enables one to select a series of (2000) and Bogner & Hay (2000) but conservative features which in turn causes especially by Peter Boyce and Sin Yeng the key to reject any species not possessing Wong in Sarawak. In the tribe that feature. After passing through a series Schismatoglottideae the genus Apobalis was of characters the remnant list of species is resurrected; Aridarum has increased from 9 then compared with the plant being to 24 species; Bakoa was described as a new determined to see if there is a match. genus with 4 new species; Bucephalandra increased from 2 to 28 species; Hestia was Lucid keys are powerful tools when fully described as a new genus; Hottarum with 2 developed and they should be developed for species was resurrected from Piptospatha; other genera. Mention has already been Ooia was described as a new genus with 3 made about the development of Lucid keys species; Pichinia is described as a new for Monstera and Stenopermation which have monotypic genus; Piptospatha with 15 species

Aroideana VOL 38E NO 1, 2015 49 Croat, 2015 A Review of Studies of Neotropical Araceae already been developed but others will be Bucephalandra, Phymatarum and developed for Aglaonema, Chlorospatha, Piptospatha. Telopea 9(1): 179 –222. Dieffenbachia, Dracontium and Rhodospatha. Cardona, F. 2000. Patrones de distribución de epífitas vasculares en robledales. CONCLUSIONS Revista de la Facultad Nacional de The results of our collective investigations Agronomía. Medellín, Vol. 53, pp. with Neotropical Araceae have been fruitful 969–983. in recent years with significant monographic, floristic and molecular studies Cardona, F. 2001. Diversidad y produced especially in smaller taxonomic composición de epífitas vasculares en groups. Floristic, studies though by now robledales de Antioquia (Colombia). widespread, are needed for more areas and Actualidades Biológicas. Universidad de they must be continuously reviewed with Antioquia, Medellín, Vol. 23 No. 74: the ability to make updates as new species 25–31. or new country or regional record are added since they otherwise quickly become outdated. Larger groups, especially Cardona, F.. 2004. Synopsis of the Genus Anthurium and Philodendron have proven to Spathiphyllum (Araceae) in Colombia. be so species-rich that any end product Ann. Missouri Bot. Gard. 91(3): 448– remains somewhat elusive though attempts 456. are under way to at least describe species and add them to Lucid keys. Molecular Carlsen, M. 2013. A molecular phylogeny of studies are beginning to produce reliable the species-rich Neotropical genus end products that were previously dubious. Anthurium (Araceae) based on Nevertheless broader and more species-rich combined chloroplast and nuclear surveys are required to better understand DNA. Syst. Bot. 36(3): 576–588. the sectional classification of the larger genera. Studies of pollination must be Castaño Rubiano, N. 2013. "Revisión encouraged at all levels since we have Taxonómica de Stenospermation para scarcely begun this study and the results are Colombia", Master’s Thesis, so promising at both the biological and Universidad Nacional de Colombia, taxonomic level. Bogota.

LITERATURE CITED Coelho, M. A. N., J. L. Waechter & S. J. Mayo. 2009. Revisāo Taxonômica das Bogner, J. & A. Hay. 2000. Schismatoglottideae Espécies de Anthurium (Araceae) (Araceae) in Malesia II – Aridarum, seçāo Urospadix Subseçāo

Aroideana VOL 38E NO 1, 2015 50 Croat, 2015 A Review of Studies of Neotropical Araceae

Flavescentiviridia. Rodriguésia 60(4): Dactylophyllium (Araceae). PhytoKeys 23: 799–864. 41–54.

Crisci, J. V. 1971. Flora Argentina: Araceae. Croat, T. B. & D. Mount. 1988. Araceae. Pp. Revista Mus. La Plata, Secc. Bot. 11: 1–46. In: R. Spichiger & J. M. 193–284. Mascherpa (eds.), Flora del Paraguay. Conservatoire et Jardin Botanique, Croat, T. B. 1991. A revision of Anthurium Geneva & Missouri Bot. Gard. section Pachyneurium (Araceae) Ann. Missouri Bot. Gard. 78: 539–855. Croat, T. B. & M. H. Grayum. 1994. Philodendron subgenus Pteromischum Croat, T. B. 1993. Araceae. Pp. 71–82. In: L. (Araceae) from Saül, French Guiana. Brako & J. L. Zarucchi, Catalogue of Novon 4: 211–219. the flowering and gymnosperms of Peru. Monogr. Syst. Croat, T. B. & L. P. Hannon. 2015. [in Bot. Missouri Bot. Gard. 45. press]. A Revision of Chlorospatha (Araceae). Ann. Missouri Bot. Gard. Croat, T. B. 1997. A revision of Philodendron 20xx. subgenus Philodendron (Araceae) and Central America. Ann. Missouri Bot. Croat, T. B., V. Pelletier, L. Salmon & J. Gard. 84: 311–704. Weigel. 2014. A new species of Xanthosoma from western French Croat, T. B. & A. Acebey. 2015. [in press]. Guiana. Aroideana, in press. Araceae, In: A. Gomez-Pampa et al., Flora de Veracruz, Xalapa, Mexico. Croat, T. B. & T. Stiebel. 2001. Araceae. In: W.D. Stevens, C. Ulloa Ulloa, A. Pool Croat, T. B. & R. Baker. 1978. Studies in & O. M. Montiel (eds.), Flora de Araceae II, Anthurium Section Nicaragua. Monogr. Syst. Bot. Missouri Polyphyllium. Selbyana 2: 230–238. Bot. Gard. 85: 136–188.

Croat, T. B. & M. Carlsen. 2004. New Croat, T. B. & G. Zhu. 2004. A revision of species of Anthurium sect. Dracontium (Araceae). Ann. Missouri Semaeophyllium. Novon 14: 401–414. Bot. Gard. 91(4): 593–667.

Croat, T. B. & M. Carlsen 2013. A Engler, A. & K. Krause. 1908. Araceae- reassessment of Anthurium species Monsteroideae. Pp. 4–138. In: A. with palmately divided leaves, and a Engler (ed.), Das Pflanzenreich IV. reinterpretation of Anthurium 23B(Heft 37). W. Engelmann, Berlin.

Aroideana VOL 38E NO 1, 2015 51 Croat, 2015 A Review of Studies of Neotropical Araceae

Gauthier, M.-P. L., D. Barabé, and A. Gonçalves, E. G. 2012. Lorenzia (Araceae Bruneau. 2008. Molecular phylogeny — Spathicarpeae): A New Genus of the genus Philodendron (Araceae): from Northern Brazil Supported by delimitation and infrageneric matK Sequence Data Systematic classification. Bot. J. Linn. Soc.156: 13– Botany 37(1):48–52. 2012 doi: 27. http://dx.doi.org/10.1600/03636441 2X616620

Gonçalves, E. G. 1999. A New Pedate- Gonçalves, E. G. & E. R. Salviani. 2002 leaved Species of Xanthosoma Schott New species and changing concepts (Araceae: Tribe Caladieae) with Linear of Philodendron subgenus Meconostigma Leaflets, from the Brazilian Pantanal. (Araceae). Aroideana 25: 2–15. Aroideana 22: 3–6 Grayum, M. H. 1997. Nomenclatural and Gonçalves, E. G. 2000. Xanthosoma riparium taxonomic notes on Costa Rican (Araceae), a New Species from Goiás, Araceae. Phytologia 82(1): 30–57. Brazil. Novon 10(1): 26–28. Grayum, M. H. 1982. The aroid flora of Finca La Selva, Costa Rica: A lowland Gonçalves, E. G. 2002a. New Aroid Taxa wet forest locality. Aroideana 5(2): 47– from Brazil. Aroideana 25: 16–35. 62. Gonçalves, E. G. 2002b. Sistemática e evolução da tribo Spathicarpeae Grayum, M. H. 1986. New taxa of (Araceae). Ph.D. thesis, Universidade , Chlorospatha, and de São Paulo, São Paulo, Brazil Xanthosoma (Araceae: Colocasioideae) from southern Central America and Gonçalves, E. G. 2005. Two new Andean northwestern Colombia. Ann. genera for the tribe Spathicarpeae Missouri Bot. Gard. 73: 462–474. (Araceae). Willdenowia 35(2): 319–326. Grayum, M. H. 1987. New taxa of Caladium and Chlorospatha (Araceae– Gonçalves, E. G. 2011. The Commonly Colocasioideae) from Panama and Cultivated Species of Xanthosoma NW Colombia. Ann. Missouri Bot. Schott (Araceae), including Four New Gard. 74: 659–660. Species. Aroideana 34: 3–23. Grayum, M. H. 1992. New species of Philodendron subgenus Pteromischum (Araceae) from Mesoamerica and

Aroideana VOL 38E NO 1, 2015 52 Croat, 2015 A Review of Studies of Neotropical Araceae

Pacific South America. Phytologia 73: Grayum, M. H. 2003a. Lemnaceae. Pp. 30–39. 618–622 in, B. E. Hammel, M. H. Grayum, C. Herrera & N. Zamora Grayum, M. H. 1993[1992] A remarkable (eds.), Manual de plantas de Costa new Anthurium from Costa Rica. Rica. Vol. II. Gimnospermas y Aroideana 15: 40–43. monocotiledóneas (Agavaceae– Musaceae). Monogr. Syst. Bot. Missouri Grayum, M. H. 1996. Revision of Bot. Gard. 92: 1–694. Philodendron subgenus Pteromischum Hay, A. & Yuzammi. 2000. (Araceae) for Pacific and Caribbean Schismatoglottideae in Malaysia I - Tropical America. Monogr. Syst. Bot. Schismatoglottis Telopea 9(1): 1–177. Missouri Bot. Gard. 47: 1–233. Kessler, M. & T. Croat. 1999. State of Grayum, M. H. 1997. Nomenclatural and knowledge of Bolivian Araceae. taxonomic notes on Costa Rican Selbyana 20(2): 224–234. Araceae. Phytologia 82: 30–57. Mayo, S. J. 1991. A revision of Philodendron Grayum, M. H. 1999. Araceae. In: P. subgenus Meconostigma (Araceae). Kew Jørgensen & S. Leon, Catalogue of Bull. 46: 601–681. the vascular plants of Ecuador. Monogr. Syst. Bot. Missouri Bot. Gard. Sakuragui, C. M. 1998. Filogenia e 75: 109–952. Taxonomia de Philodendron seção Calostigma no Brasil. Unpublished Grayum, M. H. 2003. Araceae. Pp. 59–200 Ph.D. Thesis. University of São Paulo, in, B. E. Hammel, M. H. Grayum, C. Brazil. 238 p. Herrera & N. Zamora (eds.), Manual de plantas de Costa Rica. Vol. II. Sakuragui, C. M. & D. C. Zappi. 2005. Gimnospermas y monocotiledóneas Taxonomic revision of Brazilian (Agavaceae–Musaceae). Monogr. Syst. species of Philodendron section Bot. Missouri Bot. Gard. 92: 1–694. Macrobelium. Kew Bull. 60: 465–513.

Sierra-Giraldo, J. A. 2014. First record of Anura (Dendropsophus columbianus -Anura: Hylidae-) as floral visitors of Araceae (Zantedeschia aethiopica (L.) Spreng). Aroideana 37E, No. 1: 4–10.

Aroideana VOL 38E NO 1, 2015 53 Croat, 2015 A Review of Studies of Neotropical Araceae

Tam, S., P. C. Boyce, T. M. Upson, D. Croat, T. B. A revision of Rhodospatha Barabé, A. Bruneau, F. Forest & J. S. (Araceae). Parker. 2004. Intergeneric and infrafamilial phylogeny of subfamily Croat, T. B. A revision of Stenospermation Monsteroideae (Araceae) revealed by (Araceae) for Central America. chloroplast trn L–F intergenic spacer. Amer. J. Bot. 91(3): 490–498. Croat, T. B., L. P. Hannon & X. Delannay. New species of Xanthosoma from Temponi, L. 2006. Systematics of western Andes of South America. Anthurium sect. Urospadix (Araceae) [online]. São Paulo: Instituto de Croat, T. B., M. Moffler & R. Wunderlin. A Biociências, University of São Paulo, Revision of Adelonema (Araceae). 2006. Doctoral Thesis in Botânica. Croat, T. B. & J. Vaninni. New species of IN PREPARATION Araceae from Guatemala. Croat, T. B. A key to South American species of Dieffenbachia (Araceae).

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