History and Current Status of Systematic Research with Araceae

Copyright © 2000 by Thomas B. Croat
Missouri Botanical Garden
P. O. Box 299
St. Louis, MO 63166

This is the second edition of an article that first appeared in Aroideana, Volume 21, 1998. This document is also available as a PDF file here. Adobe Acrobat Reader™, a free software, is required to view this PDF file.

Present Focus on Aroid Research

The current focus of research with Araceae is to a great extent covered by the past history since many of the participants are doing similar research and continue to be active. Simon Mayo, Josef Bogner, and Peter Boyce have released their long awaited book, The Genera of Araceae (1997) published by Kew. Simon Mayo is back at Kew and he continues his efforts in Brazil along with several Brazilian collaborators. Mayo is working on a checklist for the Araceae of Brazil. Jim French has discontinued his molecular work with Araceae and has spent his sabbatical year in Costa Rica studying pollination biology. Tom Croat is pursuing a revision of Dieffenbachia of Central America and is completing a revision of Rhodospatha, and will soon embark on a revision of Anthurium sect. Porphyrochitonium. With Richard Mansell, University of South Florida, Tampa, he is working on a revision of Anthurium sect. Semaeophyllium. He also continues to work with floristic projects in South America, especially with Dorothy Bay on the Araceae of Bajo Calima and the Araceae of the Guianas. Croat organized a three day International Aroid Conference that followed the XVI International Botanical Congress in St. Louis in 1999.

Alistair Hay, Peter Boyce, Wilbert Hetterscheid, and others on the Flora Malesiana team continue to work toward finishing that major Asian project, now due to be completed in the year 2000. Hay is also responsible for organizing an aroid conference in Sydney in 1989. Hetterscheid works independently on his revision of Asiatic Amorphophallus, with S. Ittenbach on the African Amorphophallus species, and with Ittenbach and Bogner on the Amorphophallus species from Madagascar. Jin Murata spends a lot of time in China working on a revision of Arisaema for that region as well as for the Flora Malesiana region. He is also undertaking molecular studies on all genera he can acquire. Li Heng is working on a revision of the Araceae treatment for the Flora of China to be included in the English version of the flora, a project being done in part with collaboration by the Missouri Botanical Garden. Li also was responsible for organizing the VI International Aroid Conference held in Kunming, China in late June 1995. The field of Araceae research is attracting new researchers in both Latin America and in Asia.

A large group of researchers, though perhaps having a long-standing interest in Araceae, began publishing articles regarding Araceae in the present decade and some are only beginning their studies.Matyas Buzgó, from the Botanical Garden and Institute for Systematic Botany at the University of Zürich has studied floral development in Araceae, especially Pistia (Buzgó, 1994) and Lagenandra. A recent paper deals with odor differentiation in Lagenandra (Buzgó, 1998).

A small group of researchers in the Department of Horticulture at the University of Hawaii continues work begun by H. Kamemoto with Anthurium. Most of the work, carried out under the supervision of Adelheid R. Kuehnle, deals with aspects of plant breeding, morphology and embryology. Tracie K. Matsumoto, a student of Kuehnle, did her thesis on the embryology of Anthurium (Matsumoto, 1994) and has subsequently published other papers dealing with the origin of somatic embryos (Matsumoto et al., 1996), the improvements of observing plant structures with light microscopy (Matsumoto et al., 1995), and on micropropagation of anthuriums (Matsumoto & Kuehnle, 1996). Nuttha Kuanprasert works on fragrance of Anthurium species and hybrids (Kuanprasert & Kuehnle, 1995, 1999).

Others who have been working with Araceae are Marcus Nadruz (Rio de Janiero Botanical Garden), Shrirang Ramchadra Yadav, from Shivaji University, Gladys Benevides, and Jimena Rodríguez de Salvador. Benevides did her thesis on a study of the Araceae of the `La Favorita' Biological Reserve in Pichincha Province (Benevides & Ordoñez, 1993). She is continuing her studies with Araceae in other parts of Ecuador. Ileana Arias Grande, working at the botanical garden in Havana, Cuba has a strong interest in Cuban Araceae (Arias Grande, 1992, 1994). Shrirang Ramchandra Yadav, formerly of Goa University in India and now at Shivaji University in Kolapur, India, works on the Araceae of western Ghats (Yadav et al., 1993). He presented a paper at the VI International Aroid Conference in Kunming (Yadav, 1998).

Two Brazilian botanists, E.L.M. Catharino and A.R.R. Olaio (Sao Paulo) have published jointly, describing a new species of Anthurium (Catharino & Olaio, 1990) and in the preparation of the Araceae treatment for Caroza Island in the Atlantic Ocean off the coast of Sao Paulo State (Olaio & Catharino, 1991).

Elke Seubert conducted a thorough survey of fruits and seeds of the Araceae and proposed a novel system of classification based on that information (Seubert, 1993). Her book, "Die Samen der Araceen" has a wealth of information about fruits and especially seed and contains excellent line drawings. To summarize her work, her observations are divided into five categories: (1) flower characteristics; (2) vegetative characteristics; (3) seed characteristics: seedcoat; (4) seed characteristics: endosperm and embryo; and (5) location of crystals. Each category is further subdivided into four or five features. Diagrams are presented with the use of colored lines denoting tribes and subfamilies possessing each feature. A review is planned by Josef Bogner. Seubert also studied the distribution and frequency of sclereids within aerial-roots, leaf sheaths, petioles, blades, spadices, and flowers in Araceae (Seubert, 1997).

Elizabeth Widjaja, working at the Bogor Botanical Garden herbarium in Indonesia did her graduate work in England on the genus Amorphophallus. Sunu [monomial], a student of Elizabeth A. Widjaja in Bogor, Indonesia, is working on a revision of Anadendrum.

Mikhail Serebryanyi, Moscow Main Botanical Gardens has a principal interest in the Araceae of Vietnam, especially Pothos and Pseudodracontium. He has prepared a revision of the latter (Serebryanyi, 1995). His first paper dealing with Araceae dealt with pigmentation in new leaves of Anubias (Serebryanyi & Filimonove, 1990). During his field work in Vietnam he discovered new species, some of which have been published (Serebryanyi, 1991; Hetterscheid & Serebryanyi, 1994). Serebryanyi, in collaboration with other computer staff at the Moscow Main Botanical Garden, has developed a computerized database for Araceae nomenclature. He organized the very successful IV International Botanical Congress in Moscow in August 1992.

Wilbert Hetterscheid began his career at the University of Utrecht and now works for Vaste Keurings Commissie in Alsmeer. His major botanical connection is at the Leiden Botanical Garden where his living collection is housed. He is working on a revision of the Asian species of the large and complex genus Amorphophallus of Asia, and he has been successful in bringing many of the species into cultivation, an essential task since herbarium material of these huge plants is notoriously poorly prepared. With S. Ittenbach (Hetterscheid & Ittenbach, 1996), many species of Amorphophallus from Asia and Africa were described and illustrated. Hetterscheid estimates that there are a total of 200 species in the genus. He has already published a considerable number of new species, 36 to date (Hetterscheid, 1991, 1992, 1994a, 1994b; Hetterscheid & Sarker, 1996; Hetterscheid & Serebryanyi, 1994; Hetterscheid et al., 1994, 1996, 1999). With the help of Ching-I Peng from the Academica Sinica in Taipei, he revised the Amorphophallus of Taiwan (Hetterscheid & Peng, 1995), with S. R. Yadav and K. S. Patil (Hetterscheid et al., 1994) he worked on members of Amorphophallus section Raphiophallus, and with D. DeSarker (1997) he looked at the cytological details of Amorphophallus margaritifer. A major participant in the Araceae treatment for the Flora Malesiana, Wilbert Hetterscheid is the European coordinator for the project headquartered in Leiden. He will contribute Amorphophallus to the flora project. As a part of this project he was a coauthor of the checklist and bibliography for the Flora Malesiana region (Hay et al., 1995, 1995a). A recent paper profiled Filarum manserichensis (Hetterscheid & Sizemore, 1997) and another deals with the odor presentation of Amorphophallus and Pseudodracontium (Kite & Hetterscheid, 1997) and with P.C. Boyce (2000) reclassified Sauromatum to Typhonium. Wilbert also made significant contributions to the recently published Amorphophallus titanum monograph (Barthlott & Lobin, 1998).

Larry Klotz, though not otherwise known as an aroid researcher, made an interesting study of Orontium aquaticum (Klotz, 1991, 1992).

A. Lourteig (1990), at the Paris Herbarium, has attempted to typify some of the Araceae depicted in the illustrations done by Charles Plumier (1755-1760).

Duangchai Sookchaloem (nee Sriboonma), who completed her graduate work under the direction of Jin Murata in Tokyo, works at the Forestry Herbarium at the Royal Forest Department in Bangkok, Thailand. She has revised Typhonium with Jin Murata and K. Iwatsuki (Sriboonma et al., 1993; Sookchaloem, 1994). Her work involved molecular studies with restriction site analysis of chloroplast DNA (Sriboonma et al., 1993).

Guanghua Zhu, a student of Tom Croat, completed a revision of Dracontium for his Ph.D. study (1994b, 1995b). He has published a new species (Zhu, 1995a) and several papers on the nomenclature of Dracontium (Zhu, 1994a, 1996; Zhu & Grayum, 1995) as a step toward the publication of his monograph (Zhu, 1997). His interests continue with the New World Lasioideae, especially Urospatha Schott and Montrichardia Crueg. Zhu has been instrumental in designing and establishing the International Aroid Society web site which is associated with the Missouri Botanical Garden's Web site.

Nguyen Van Dzu (Institute of Ecology and Biological Resources, Hanoi, Vietnam) is working on the Araceae of Vietnam (Nguyen, 1994). Several of his recent papers report new records for Vietnam (Nguyen, 1998a, 1998b, 1999, 2000). With Peter Boyce he published a paper on Pothos grandis (Boyce & Nguyen, 1995) and a new revision of Amydrium which includes two new species (Nguyen & Boyce, 1999b). He has also worked with Tom Croat describing a new species of Typhonium (Nguyen & Croat, 1997).

Dorothy Bay (Missouri Southern State College), a former student of Tom Croat, prepared a floristic survey of a species-rich site along the coast of western Colombia at Bajo Calima (see above). This massive work, with complete descriptions of over 100 species (a large percentage of them being new to science), will be published in the Annals of the Missouri Botanical Garden.

Jenn-Che Wang from National Taiwan Normal University in Taipei has completed an excellent study of the Taiwanese Arisaema (Wang, 1992, 1996). Also at National Taiwan Normal University, T.C. Huang has described a new species of Arisaema (Huang & Wu, 1997).

Jimena Rodríguez de Salvador has worked in Ecuador on the Araceae of the ENDESA Biological Reserve (Pichincha Province). (Rodríguez, 1987, 1989; Croat & Rodríguez, 1995). Despite being a region frequented by Sodiro, a high percentage of the flora proved to be new to science.

Frieda Billiet, of the National Botanical Gardens in Brussels and in charge of the living collections there, has collected in French Guiana and elsewhere. She has long been devoted to the Araceae and made her publication debut with Araceae in Curtis's Botanical Magazine (Billiet, 1996) with a discussion of Philodendron and a redescription of P. billietiae Croat (see Croat, 1995a).

Stephan Ittenbach from the University of Bonn in Germany, working under the guidance of Wolfram Lobin, did his Ph.D. dissertation on African Amorphophallus. Ittenbach published new species and subspecies of African Amorphophallus with Lobin (Ittenbach & Lobin, 1997) and contributed to the Amorphophallus titanum monograph mentioned below. Lobin has published a new species of Eminium in the Near East with P. Boyce (Lobin & Boyce, 1991) and recently edited an extensive monograph of Amorphophallus titanum with W. Barthlott (Barthlott & Lobin, 1998).

Bruce Hoffman studied aerial root fiber products in Guayana made from Heteropsis flexuosa for his M.S. Thesis at Florida International University (Hoffman, 1997).

Brett E. Serviss, with the assistance of Sidney T. McDaniel and Charles T. Bryson, has studied Alocasia, Colocais, and Xanthosoma in the southeastern part of the United States (Serviss et al., in press).

In Brazil, a number of aroid researchers have established themselves and this bodes well for the future of Araceae studies in that country. Simon Mayo has played a critical role in coordinating and promoting these studies (Mayo & Nadruz, 1992).

Marcus Nadruz Coelho began working on aroids in 1986 at the suggestion of Dr. Graziela Barroso, who worked on Brazilian aroid taxonomy in the 1950s and 1960s. Marcus, based at the Jardim Botânico in Rio de Janiero has worked with various aroid projects (Nadruz-Coêlho & Mayo, 1998) and has played an important role in stimulating and supporting other workers in Brazil. His Masters thesis, supervised by Dr. barroso, was undertaken at the Universidade Federal do Rio de Janeiro--Museu Nacional and was a study of the Philodendron species of the montane Atlantic Forest of Macaé de Cima in Rio de Janeiro state (Nadruz-Coêlho, 1995). This resulted in the recognition of 5 new species Nadruz Coelho & Mayo, 1999). He is now working on his Ph.D. thesis (Universidade Federal do Rio Grande do Sul, Porto Alegre) which is a revision of Philodendron subgenus Pteromischum of Brazil. Since 1992 he has been organizing the annual Araceae Specialists Workshop at the Brazilian National Botanical Congress and has a focal role in the Brazilian aroider network. Nadruz most recently has discovered a new species from Brazil (Nadruz & Sakuragui, 2000; Nadruz & Mayo, 2000).

Eduardo Gonçalves completed his Masters thesis at the Universidade de Brasilia on the Araceae from the Brazilian Federal District (Gonçalves, 1997) and has worked extensively with the Araceae of Central Brazil where he has discovered new species of Philodendron (Gonçalves, 1997; Gonçalves, 2000b; Gonçalves & Mayo, 2000). He has conducted a study of the biogeography of the aroids of Central Brazil (Gonçalves, in press) and studied the rare genus Gearum along with Josef Bogner (Bogner & Gonçalves, 1999). For his Ph.D. thesis, being carried out at the Universidade de Sâo Paulo, he is focusing on a molecular systematic and revision of the genus Spathicarpa. For this study, he is working on the whole tribe Spathicarpeae and will also be revising the genus Asterostigma. Independently of these projects, he has studied the petiolar anatomy (patterns of distribution of collenchyma) in the whole family Araceae (Gonçalves et al., in press) and has become very knowledgeable with Xanthosoma of Brazil and the Amazon basin (Gonçalves, 1999; Gonçalves, 2000a). Finally, he is preparing a study of the distribution of aroids in regions of Cerrado vegetation, with the help of Carolyn E.B. Proenca and Luiz Guimaraes, and making use of multivariate analysis to define the patterns of distribution.

Cassia Sakuragui is the first Brazilian botanist of recent times to have been awarded her Ph.D. (University of Sâo Paulo, 1999) on Araceae systematics. She began working on aroids in the early 1990s, and went on to carry out a survey of the aroids on the montane (Cadeira do Espinhaço) vegetation of Minas Gerais State for her Masters thesis (University of Sâo Paulo, Sakuragui, 1994) which resulted in the recognition of several new species of Philodendron and Anthurium (e.g. Sakuragui & Mayo, 1997; Sakuragui & Mayo, 1999). Her Ph.D. thesis was on the taxonomy and phylogeny of Philodendron subgenus Phildoendron sect. Calostigma (Sakuragui, 1998; Sakuragui, in press), during which she carried out a pioneer molecular systematic study on a sample of species of the genus. She continues with her aroid research at the Univesidade de Maringá in the state of Paraná, describing two new species of Philodendron (Kakuragui, 2000). She has interest in other groups of Araceae, such as the Monsteroideae, and along with Peter Boyce and Josef Bogner has made some notes on Alloschemone (Boyce et al., 2000). She has also worked on some small floras (Sakuragui, 2000; Sakuragui, in prep.); and general comments on cultivated aroids (Sakuragui, in press). She is currently supervising a Masters student, Livia Temponi, who is working on a floristic study of the Araceae of "Parque Estadual de Rio Doce" in Minas Gerais State. Temponi is working at the Universidade Federal de Viçosa in Minas Gerais State.

Maria de Lourdes Soares, at the Instituto de Pesquisas da Amazônia (INPA) in Manaus, works on the aroids of northern Brazil. Her Master's thesis (Universidade Federal Rural de Pernambuco, Recife) was a taxonomic survey of the species of Philodendron occurring in the Ducke Reserve north of Manaus (Soares, 1996; Soares & Mayo, in prep.) She also published a general field guide treatment to the aroids of the Ducke Reserve (Soares & Mayo, 1999) and is working on a detailed flora treatment for the same area. She has made a survey of the Araceae of the state of Amazonas based on collections in the INPA herbarium in Manaus (Soares, in press). She currently has begun work on a revision of Heteropsis in Brazil for her Ph.D. thesis, based at INPA.

Ivanilza Moreira de Andrade works on the systematics of the Araceae of Northeast Brazil, especially the state of Ceará. She has a special interest in the ecological morphology and architecture of aroid climbers and studied three such species in Pernambuco for her Master's thesis at the Universidade Federal de Pernambuco in Recife (Andrade, 1996; Andrade & Mayo, 1998; Andrade & Mayo, in prep.). She is now working on the biosystematics of the Araceae of montane forests in Northeast Brazil for her Ph.D. Ivanilza, along with Nadruz, Gonçalves, Sakuragui and Soareas, made their debut to most of the International Aroid Society at the VIII International Aroid Conference in St. Louis (1999) where they all presented their research. 

Alba Lins at the Museo Paraense Emilio Goeldi in Belem studies the anatomy of Amazonian aquatic aroids. Her Masters thesis (Universidade Federal do Rio Grande do Sul) was on root anatomy and morphology in Montrichardia (Lins, 1994; Lins & Oliveira, 1995) and more recently she has been working on Urospatha anatomy. She is now beginning her Ph.D. studies, also at the Universidade Federal do Rio Grande do Sul, Porto Alegre.

Other aroid workers in Brazil include Cicero Barros, from the Instituto de Meio Ambiente de Alagoas in Mutange, Maceió, Alagoas, who works with the aroids of the state of Algoas. He completed his Masters thesis (Universidade Federal de Pernambuco) in 1998 on a survey of aroids from a relict forest reserve in the Atlantic Forest of that state of Northeast Brazil; Jorge Wachter, at the Universidade Federal do Rio Grande do Sul in Porto Alegre, who is interested in the aroids of southern Brazil and who discovered Mangonia tweedieana; Ricardo Lainetti, from the Universidade Federal do Rio de Janeiro, who is interested in the pharmacological effects of Xanthosoma (specifically X. violaceum) and certain members of the Araceae; Luciedi Tostes, from the Universidade Estadual de Sâo Paulo-Botucatu, who is working on the anatomy of the secretory structures and their significance in the biosystematics of the Philodendron/P. selloum complex; Livia G. Temponi, from the Universidade Federal de Viçosa, is working on a floristic inventory of aroids at the Rio Doce Reserve in Minas Gerais State. Finally, Emerson M. Vieira, from the Universidade Estadual de Campinas, and Patricia Izan from the Universidade Sao Paulo, have published a study on the interaction of aroids and arboreal mammals in the Brazilian Atlantic rainforest (Vieira & Izar, 1999).

Two major projects currently under way which combine the efforts of Brazilian aroid systematics are the treatment for the Flora of São Paulo (Sakuragui, Nadruz Coelho, Gonçalves) and the Checklist of the Araceae of Brazil, coordinated by Nadruz Coelho.

In Asia a number of students are working on projects involved with Araceae. Melanie Medecilo is doing a revision of Philippine Epipremnum under the supervision of Domingo Madulid at the Philippine National Herbarium. Lim Sheh Ping, under the supervision of Ruth Kiew at the University Pertanian Malaysia, is working with Araceae (and other families) occurring on the limestone formation in Sabah. Yasamni [monomial], also working at Kebun Raya (Bogor National Herbarium) under the direction of Alistair Hay, is working on terrestrial species of Araceae from Java. Baharuddin Sulaiman is working on the taxonomy of wetland Araceae in North Peninsular Malaysia at the University Sains Malaysia on Penang Island in Malaysia. A Japanese botany student, Yasuko Mori, under the supervision of H. Okado, is working on pollination and population dynamics in Furtadoa in Sumatra and the Malay Peninsula.

In China, a large number of researchers working with Araceae made their international debut by presenting papers at the VI International Aroid Conference in Kunming. Many of these papers have now been presented in a special edition o the Acta Botanica Yunnanica. Some of these researchers are students or former students of Li Heng at the Kunming Institute of Botany, including: Peng Hua (Peng & Li, 1998); Wang Ping-Li (Wang & Li, 1998), and Xiao Tiao-Jiang, Long Chun-Lin, and Xia Li-Fang (Xiao & Li, 1998). Others, such as Zhang Sheng-Lin, Liu Pei-Ying (discussed above and Sun Yuan-Ming from the Southwest Agricultural University in Chongquing, presented papers (Zhang et al., 1998) as did Guo Qiao-Sheng, Zhang Guo-Tai and Wang Kang-Cai from the Nanjing Agricultural University also presented a paper (Guo et al., 1998). Finally, Yang Yong-Kang from the Yunnan Agricultural University in Kunming (Yang, 1998).

Finally, current students of Tom Croat, John Gaskin (Washington University) and Jane Whitehill (University of Missouri-St. Louis), will both be working with Araceae for their Ph.D. dissertations. Whitehill has done studies on reproductive biology in Araceae (Whitehill, 1993) and is conducting molecular studies with members of the former Colocasioideae. Gaskin, currently doing molecular studies to understand the relationships of different sections of Anthurium (especially those with glandular punctations) will do a revision of Anthurium sect. Digitinervium.



Please send your comments to Tom Croat at the address here.

This page was created by Scott E. Hyndman for the IAS on December 9, 2000.