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.

Renewed Revisionary Efforts After 1950

In addition to the floristic efforts which began in the early 1950's considerable new revisionary activity began as well with a number of new aroid researchers. One of first of these was H. C. D. de Wit who worked exclusively with the limited number of aquatic aroids that can be grown in fish tanks. These plants have a good commerical value and have spawned a number of research projects, most of them centered at the University of Wageningen in Holland and in other places in Europe. The most active research on these aquatic aroids began with de Wit and Karel Rataj and progressed until the time that Niels Jacobsen published several important papers on the genus Cryptocoryne [see below]. New species continue to be discovered in the genus.

De Wit published his first papers, all dealing with Cryptocoryne in 1953 (de Wit, 1953a-c). They were published in popular aquarium magazines Fishkeeping and Waterlife and Het Aquarium. Other species of Cryptocoryne were treated in papers published in succeeding years (de Wit, 1954, 1956, 1957, 1958a-g), and it was not until 1958 that de Wit described his first new species (de Wit, 1958b). He then published many short papers (de Wit, 1959a-i; 1960a-e, 1961a-e, 1963a-d, 1971a-c, 1975a, 1975b, 1976, 1979) in Het Aquarium a Dutch magazine popular with the aquarium enthusiasts. Written in Dutch and of one to four pages in length, each article deals with a single species of Cryptocoryne. Each fascicle is illustrated with photographs or detailed drawings and sometimes with colored paintings or photographs. One additional species was described in Artedi (de Wit, 1975b). The first widely useful paper by de Wit presented a key to all the known species of Cryptocoryne along with detailed photographs (de Wit, 1969).

De Wit's other publications include a short article on pollination in Cryptocoryne (de Wit, 1978a), chromosome numbers (de Wit & Jacobsen, 1982), and a complete revision of another aguatic genus, Lagenandra Dalzell (de Wit, 1978b). The publication is written in Dutch and contains excellent illustrations and keys as well as details on the biology. De Wit has summarized his work nicely in a series of books, all well illustrated with detailed drawings and colored plates. These books deal with numerous aquatics but the Araceae constitute the largest share (de Wit, 1983). The first version was printed in Dutch (de Wit, 1966), reprinted in 1982 (de Wit, 1982), and an essentially identical version was published in 1990 in German (de Wit, 1990).

Simultaneously, the Czech botanist, Karel Rataj was doing revisionary work on Cryptocoryne. His book on Cryptocoryne (Rataj, 1975) divided the genus into 4 subgenera and 16 sections and described three of the subgenera and all of the sections as new. Rataj recognized 52 species of Cryptocoryne while describing seven new species and six new varieties. Another paper described new cultivated species of the genus (Rataj, 1974). Rataj also published a book Aquarium Plants, which he co-authored with T. Horeman (Rataj & Horeman, 1977) as well as a paper dealing with Typhonium flagelliforme (Rataj, 1982).

In the 1950s, Haruyuki Kamemoto, working at the University of Hawaii, carried out an extensive breeding program with Anthurium in order to provide new and beautiful stock for the Hawaiian cut flower industry. Much of his original stock of wild collected material came from a field trip to Panama with Yoneo Sagawa in the early 1960s. Kamemoto's successes were many and most of his publications dealt with the development of new cultivars (Kamemoto & Nakasone, 1955, 1963; Kamemoto & Sheffer, 1978, 1982; Kamemoto et al., 1986, 1993). He also did genetic research with Araceae, especially the inheritance of color in the spathe, in collaboration with R. Y. Iwata, C. S. Tang, S. Wannakrairoj and M. Marutani (Iwata et al., 1985; Marutani et al., 1987; Kamemoto et al., 1988; Wannakrairoj & Kamemoto, 1990a, 1990b). Other technical research done by Kamemoto and his students include the use of gel electrophoresis for the identification of Anthurium cultivars (Kobayashi et al., 1987) and an extensive use of cytology, especially by R. Sheffer, S. Wannakrairoj, K. Kaneko (Kaneko & Kamemoto, 1978), and M. Marutani (Marutani et al., 1988, 1993) [see also Sheffer below].

Kamemoto's many years of research with aroids are summarized in a book entitled Breeding Anthurium in Hawaii (Kamemoto & Kuehnle, 1996), co-authored by Adelheid R. Kuehnle [see Kuehnle below] who took Kamemoto's position at the University of Hawaii upon his retirement.

A small group of researchers in the Department of Horticulture at the University of Hawaii continues the Anthurium studies begun by H. Kamemoto. Most of the work, carried out with students under the supervision of Adelheid R. Kuehnle or in collaboration with her colleagues, Tessie Amore and Nellie Sugii (the latter two, who deal with classical plant breeding and histology), histologist David Webb and biochemist T. S. Tang, deals with aspects of plant cultivation (Kuehnle et al., 1996), breeding, morphology, embryology (Kuehnle et al., 1996), novel methods of regeneration (Kuehnle et al., 1992; Kuehnle & Sugii, 1991a, 1991b) and gene transfer (Kuehnle & Chen, 1994; Chen & Kuehnle, 1996; Kuehnle & Nan, 1991). Tracie K. Matsumoto, a student of Kuehnle, did her thesis on the embryology of Anthurium (Matsumoto, 1994) and has subsequently published other papers on the origin of somatic embryos (Matsumoto et al., 1996) and on improvements of observing plant structures with light microscopy (Matsumoto et al., 1995), and on micropropagation of anthuriums (Matsumoto & Kuehnle, 1966). Nuttha Kuanprasert, another student, has begun a study of Anthurium fragrances (Kuanprasert & Kuehnle, 1995).

Monroe Birdsey, who did his graduate work at the University of California then taught at Miami-Dade Community College in Miami, was active in the Araceae with research during the 1950s. His unpublished thesis, entitled "The morphology and taxonomy of the genus Syngonium" Schott (Birdsey, 1955a), was a thorough study with emphasis on anatomy. Always interested in cultivated plants, he published The Cultivated Aroids, one of the earliest popular books of its kind devoted to Araceae (Birdsey, 1951). This work illustrated and described 70 species of Araceae. Shorter works include articles placing Pseudohomalomena pastoensis into synonymy with Zantedeschia aethiopica (Birdsey, 1955b), taxonomic problems with the confusing cultivar "golden pothos" (Epipremnum pinnatum cv. aureum) (Birdsey, 1962b), and the reintroduction of Homalomena roezlii (Birdsey, 1962a) into cultivation. His collection of living plants at his estate in Miami was until his death one of the finest of its kind in the world, possessing many fully grown species of Araceae.

The late Donald G. Huttleston, who revised Arisaema of North America, published a discussion of three subspecies of Arisaema (Huttleston, 1949) even before he completed his thesis. His thesis involved a taxonomic study of the Araceae of North America (Huttleston, 1953). Other papers dealt with the nomenclature of Lysichiton (Huttleston, 1955) and further reports on Arisaema (Huttleston, 1981, 1984).

B. Bergdolt (1955), working at Freiburg University in Germany, worked with anatomical and embryological research on leaf mottling and other leaf types.

Finally, in the late 1950s and early 1960s, the Dutch botanist, R. C. Bakhuizen v.d. Brink, though never deemed to be a major player in Araceae, did make significant contributions with his studies of the Araceae of Java (Bakhuizen, 1957; Bakhuizen & Koster, 1963). He also published a paper dealing with the differences in certain members of the Monsteroideae (Bakhuizen, 1958).

Despite the scattered and localized research mentioned earlier it was not until the beginning of the 1960s that much serious research with Araceae re-occurred since the time of Engler and Krause. The early 1960s saw the active publication by taxonomists George S. Bunting, Graziela Maciel Barroso, Dan Nicolson, Mitsuru Hotta, Cecil T. Prime, Harald Riedl, and Hiroyoshi Ohashi. Though Bunting's publication career was strongest during the 1960s, his first paper, a key to the genera of Araceae in Venezuela written in Spanish, was published in the mid-1950s (Bunting, 1956). Articles regarding cultivated aroids came next (Bunting, 1955, 1956b, 1959, 1961b) followed by the publication of his Ph.D. thesis, a revision of the genus Spathiphyllum Schott (Bunting, 1960a) and a continued flurry of publications throughout the 1960s, many of which dealt with mostly Venezuelan genera (Bunting, 1960b; Bunting & Steyermark, 1969) or floristic regions in Venezuela such as Chimantá (Bunting, 1963a) or Auyán-tepuí (Bunting, 1967) and the Sierra de Lema (Bunting, 1963b), the description of new species (Bunting, 1963c) or taxonomic problems with cultivated plants including Spathiphyllum (Bunting, 1961a), Dieffenbachia (Bunting, 1962a, 1963d, 1966a, 1988c), Alocasia (Bunting & Nicolson, 1963), Anthurium (Bunting, 1963d), Monstera (Bunting, 1962d, 1966a, 1966b), Syngonium (Bunting, 1966b), and Philodendron (Bunting, 1966d, 1966e; Moore, 1974). Other papers dealt with the differences between Schizocasia and Alocasia (Bunting, 1962b), the delimitation of genera of the Monsteroideae (Bunting, 1962c), and a discussion of Philodendron hederaceum-scandens complex (Bunting, 1963f). Of his papers treating species of areas other than Venezuela, the most useful are his commentary on the Araceae of Mexico (Bunting, 1965) and a discussion of the anatomy and taxonomy of the Philodendron scandens complex (Bunting, 1968).

Part of Bunting's early work was begun at the Missouri Botanical Gardens, then continued at the Bailey Hortorium at Cornell University. From Cornell he moved to Venezuela where his extensive field studies and collections of Araceae formed the basis for a treatment of the Araceae for the Flora of Venezuela. Toward this end he has published many new species for Venezuela and adjacent countries (Bunting, 1975, 1986, 1987, 1988a, 1989a) and a new genus (Bunting, 1988b, 1989b) for Venezuela and adjacent countries. Though the Araceae treatment for the Flora of Venezuela has not been finished, a major synopsis of the flora has been published (Bunting, 1979). It contains most of the species and taxonomic keys but no descriptions or illustrations. Bunting has also published a treatment of the Araceae for the flora of the Cerro Aracamuni in Venezuela (Bunting, 1989c) and the Flora of the Venezuelan Guayana (Bunting, 1995). Aside from his work with the flora of Venezuela, his principal interest for many years was a revision of the genus Philodendron. One of his major contributions has been with cultivated plants, a major emphasis in his earlier years. He contributed all the Araceae for Hortus Third (Bunting, 1976) and the Philodendron for the European Garden Flora (Bunting, 1984). Bunting has also described the genus Jasarum as well as many other species, 195 in all, mostly from Venezuela.

Graziela Maciel Barroso, working on Brazilian Araceae, made the description of Philodendron camposportoanum G. M. Barroso in her first publication (Barroso, 1956). This was followed by a series of papers describing other new species (Barroso, 1957, 1959, 1965, 1970). All but one, Anthurium wendlingeri G. M. Barroso from Costa Rica, represented Brazilian species. Barroso published 15 species in all.

Dan H. Nicolson, working at Cornell University, published extensively throughout the 1960s. His first publications on Araceae were a review of the classification of the Araceae (Nicolson, 1960a) and a paper describing the occurrence of trichosclereids in the Monsteroideae (Nicolson, 1960b). Several other papers, including one on Gorgonidium (Nicolson, 1963) and

revisions of small groups, e.g. Filarum (Nicolson, 1966); Xenophya (Nicolson, 1968a); Asian Spathiphyllum (Nicolson, 1968b, 1992a); and Amydrium (Nicolson, 1968c) were published in part even before his Ph.D. thesis, a revision of the genus Aglaonema (Nicolson, 1967b, 1969). The work with Aglaonema involved extensive fieldwork in the Asian tropics which led to a continued interest in Asian floristic accounts including floristic accounts of the Araceae of the Hassan District (Nicolson, 1976a) and the Tamilnadu Carnatic regions (Sivadasan & Nicolson, 1983) of India, as well as for Fiji (Nicolson, 1978, 1979), and Sri Lanka (Ceylon) (Nicolson, 1988). Nicolson (1984f) also treated the Aglaonema for the European Garden Flora. With C. R. Suresh and K. S. Manilal in India he discussed H. A. van Rheede's Hortus Indicus Malabaricus [SW India] (Nicolson et al., 1988). His interest in fieldwork also resulted in papers on collecting Araceae (Nicolson, 1965, 1976b).

Along with Josef Bogner, Nicolson has been involved with studies and revisions of the suprageneric system of classification (Bogner & Nicolson, 1991) resulting in one of the competing systems of classification of the Araceae. This was among the first to question the classification of Engler.

Early in his career Nicolson became interested in the legal matters of nomenclature (Nicolson, 1963b, 1964, 1967a, 1968d, 1975a, 1975b, 1977, 1981b, 1984b-d, 1987a; Nicolson & Bogner, 1977, 1981; Nicolson & Mayo, 1984a, 1984b; Nicolson et al., 1984), an interest that continues to this day. He has spent much of his career dealing with nomenclature of Araceae and suprageneric systems of classifications (Bogner & Nicolson, 1991). Other work includes a survey of floral anatomy of Araceae carried out in conjunction with R. H. Eyde and P. Sherwin (Eyde et al., 1967). In a collaborative efforts he published papers on Alocasia (Bunting & Nicolson, 1963), a revision of Gorgonidium (Bogner & Nicolson, 1988), new species of Arisaema (Sivadasan & Nicolson, 1983) and Theriophonum (Sivadasan & Nicolson, 1981), a revision of Typhonium with M. Sivadasan (Nicolson & Sivadasan, 1981) as well as one on the taxonomy of Theriophonum (Nicolson & Sivadasan, 1982). One of his major accomplishments is his study of the complex publication history of Luis Sodiro (Nicolson, 1984a), who had the practice of publishing each new species several times. Until Nicolson's enlightening work, many of the earliest valid publications were overlooked. He also alphabetized and indexed Schott's Icones Aroideae and Reliquae (Nicolson, 1984e). Nicolson's translation of Engler's classification of the Araceae including the key to genera made understanding and ultimate revision of that system possible (Nicolson, 1982a). Nicolson is a member of the editorial board for both Taxon and Aroideana and has an interest in aroid literature (Nicolson, 1989, 1992b). His understanding of classical languages as well as German, the history of early aroid taxonomy (Nicolson, 1982a, 1987a), the rules of nomenclature, and his role as Senior Curator at the Smithsonian Institution where he has worked since he left Cornell make Nicolson the person to whom many of us turn for advice. His contributions to the Araceae are unique.

Harald Riedl, working at the Naturhistorisches Museum in Schott's hometown of Vienna, published his first paper on the Araceae in the Flora Iranica (Riedl, 1963). He is also a specialist on the genus Eminium (Riedl, 1969) and the flora of Middle Europe and the Middle East, and has published on Arum (Riedl, 1967), and Stylochiton (Riedl, 1990) [the latter from Africa] as well as having written the Araceae treatment in Hegi's Illustrierte Flora von Mitteleuropa (Riedl, 1979) and the Araceae treatment in the Flora of Iraq (Riedl, 1985). Riedl is also an expert on the history of H. W. Schott and has published several papers dealing with Schott's work and collections (Riedl, 1965a-c, 1966, 1978; Riedl & Riedl-Dorn, 1988). He published several papers in the earliest issues of Aroideana, including a discussion of the aroids described in Russel's Natural History of Alepo in 1794 (Riedl, 1980a), a partial treatment of Biarum Schott (Riedl, 1980b), and a paper stressing the importance of ecology in defining genera (Riedl, 1980c).

The earliest of these was Mitsuru Hotta, working at the Kyoto University in Japan, who began publishing papers on Araceae in 1963 (Hotta, 1963a, 1963b). Some of his first papers dealt with Arisaema of Japan (Hotta, 1963a, 1963b, 1964, 1966a, 1970a-d) but he soon began working in more tropical parts of Asia, including Borneo, where he studied the Schismatoglottidinae (Hotta, 1965, 1966b, 1987) and made phytogeographic and floristic surveys (Hotta, 1966c, 1967). His work in Borneo resulted in the description of two new genera, Pedicellarum, Phymatarum, and Heteroaridarum (Hotta, 1976). His work in Sumatra has resulted in floristic surveys (Hotta, 1984), another new genus, Furtadoa M. Hotta (Hotta, 1981), other miscellaneous new species (Hotta, 1985, 1993), a survey of Homalomena and Anadendrum of Sumatra (Hotta, 1986a, 1986b), and a paper on taro uses (Hotta, 1962, 1983). Hotta (1982) also made detailed comparisons of the Homalomeninae and the Schismatoglottidinae in Malesia.

One of Hotta's major accomplishments was a system of classification in which he proposed major first-time changes in the suprageneric system of classification of the Araceae (Hotta, 1970a). His system of classification is discussed in detail and is compared with other major systems (Croat, 1990). Another major publication (Hotta, 1971) provided a detailed discussion of the relationship of the Araceae to other families and discusses the morphological, anatomical, and cytological characteristics of different aroid subfamilies. In recent years, Hotta has been involved (sometimes with Hiroshi Okada and Motomi Ito) in ecological studies with Araceae in West Sumatra (Okada, 1986; Hotta et al., 1985; Okada & Hotta, 1987). His skills in cytology, broad ranging ecological interests and astute observations have made Hotta's contributions to Southeast Asian studies of Araceae diverse, unique, and important.

Hiroyoshi Ohashi, at the University of Tokyo and Tohoku University, began publishing on Japanese Arisaema in the early 1960s. Miscellaneous notes on Arisaema (Ohashi, 1963, 1964) were followed by a complete revision of the genus for Japan (Ohashi & J. Murata, 1980), and the Araceae treatment for the Wildflowers of Japan (Ohashi, 1982). Continuing the work of Hara on the Flora of Eastern Himalaya, Ohashi published a third report of that work which included additional Arisaema (Ohashi, 1975). Ohashi also compiled a list of types of Arisaema in Japanese herbaria (Ohashi, 1981a, 1981b) and studied pollen morphology of Japanese Arisaema (Ohashi et al., 1983). A recent paper describes a new species of Piptospatha that regularly produces adventitious bulbils along the midrib on the lower blade surface (Ohashi et al., in press).

A European specialist in Arum, Cecil T. Prime, was also active during the 1960s. His major work on the biology of Arum maculatum, Lords and Ladies (Prime, 1960), is one of the most detailed and interesting books ever written on the biology of an aroid. Prime also contributed the Arum section for the Flora Europaea (Amaral Franco et al., 1980).

Although the taxonomists noted above were the main players within Araceae research, several post-Englerian non-specialists made significant or unique studies in specific areas of aroid research. Three individuals working with Zantedeschia are worthy of mention. Hamilton Traub produced a single work on Zantedeschia that has been one of the most useful ever done for that genus (Traub, 1949). The second work, published in the same year was written by L. Mirzwick (Mirzwick, 1949). Most recently, Cynthia Letty (1973) also published a revision on Zantedeschia. Another work carried out by non-specialists was a minor revision of Arisaema by Walter Robyns and R. Tournay (Robyns & Tournay, 1955). This work treated five species of Arisaema from tropical Africa. Another useful work on Arisaema deals with the plants of the Himalayas (Pradhan, 1986, 1990).

With the advent of Bunting and Nicolson, interest in Araceae accelerated with several new workers, including: Josef Bogner, Tom Croat, Dorothy Shaw, Jorge Crisci, Mike Madison, Li Heng, Marija Bedalov, Richard Sheffer, Simon Mayo, and Jin Murata beginning their publishing careers with Araceae, in the late 1960s and 1970s.

Probably no one in the history of work with the family has had a greater focus on research with Araceae than Josef Bogner. His unprecedented interest in the family is so universal that it is difficult to define. Beginning with a paper dealing with Theriophonum (Bogner, 1968), Bogner has been involved with the Araceae in nearly all parts of the world, including a major study of the Araceae of Madagascar, the Comoros Islands, and with the Arophyteae (Bogner, 1972a, 1972b, 1973a, 1973b, 1975), as well as the Seychelles with Protarum (Bogner, 1973a, 1973e); and in Africa with Amorphophallus (Barthlott & Bogner, 1981; Bogner & Hetterscheid, 1992); Andromycia (Bogner, 1969b); Callopsis (Bogner, 1969a [= Nephthytis hallaei (Bogner) Bogner, see Bogner, 1980b]); Cercestis (Bogner & Knecht, 1994); Culcasia (Bogner, 1980a); Gonatopus (Obermeyer & Bogner, 1979); Nephthytis (Bogner, 1980b; de Namur & Bogner, 1994); Pseudohydrosme (Bogner, 1981a); Stylochiton (Bogner, 1984f); in Asia with Amorphophallus (Bogner, 1976d, 1981a, 1981b, 1989a, 1995; Bogner et al., 1985; Bogner & Hetterscheid, 1992); Aridarum (Bogner, 1979, 1981c, 1983a); Cryptocoryne (Bogner, 1974, 1984a-c, 1984i, 1985a, 1989c; Bogner & Jacobsen, 1985, 1986; Ehrenberg & Bogner, 1992); Pycnospatha (Bogner, 1973b); Lagenandra (Bogner, 1974, 1978; Bogner & Jacobsen, 1987); Homalomena (Bogner, 1976a); Thomsonia (Bogner, 1976b); Bucephalandra (Bogner, 1980c, 1984g); Plesmonium Schott (Bogner, 1980d), Hottarum (Bogner, 1983b, 1984g; Bogner & Hotta, 1983a); Schismatoglottis (Bogner & Hotta, 1983b; Bogner, 1988); Hapaline and Phymatarum (Bogner, 1984e); Scindapsus (Bogner & Boyce, 1994); and Typhonium (Bogner, 1987a), as well as in the Americas with Xanthosoma (Bogner, 1986a); Mangonia (Bogner, 1973d, 2000); Scaphispatha (Bogner, 1980e); Jasarum (Bogner, 1977, 1984d, 1985d), Dracontium (Bogner, 1981d), Caladium (Bogner, 1980f, 1984h); Chlorospatha (Bogner, 1985b, 1985e);Gearum (1999); Homalomena (Bogner & Moffler, 1985a, 1985b); Taccarum (1989b); Philodendron (Bogner & Bunting, 1983); and Gorgonidium (Bogner & Nicolson, 1988) and in Turkey with Biarum (Bogner & Boyce, 1989). With James French he described the tribe Anadendreae (Bogner & French, 1984). His most recent paper described species from both the Old and New World [Spathantheum intermedium, Asterostigma cryptostylum, Zomicarpella amazonica, Ulearum sagittatum var. viridispadix and Nephthytis afzelii var. graboensis]. He has described a total of 54 new species.

Bogner, because of his broad interests and deep understanding of all matters regarding Araceae, has long been principally interested at the subfamilial and tribal levels. He has been responsible for a rethinking of the system of classification of the Araceae beginning with his "critical list" of aroid genera (Bogner, 1978), the reduction of genera (Bogner, 1985c), new name and combinations (Bogner, 1986b), the placement of Jasarum (Bogner, 1980e), and his revised classification of the family (Bogner & Nicolson, 1991). A recent paper (Mayo et al., in press) defining the relationship of the Araceae to other closely related families reflects his deep understanding of the family. He is a coauthor of The Genera of Araceae (Mayo et al., 1997) that describes and illustrates all the genera of Araceae. Another summary paper by Bogner deals with the wide variation in morphology of Araceae (Bogner, 1987b), and another is on new taxa of Araceae (Bogner, 1997). Bogner is also one of the few aroid researchers who has dealt with fossil Araceae (Bogner, 1976c; Gregor & Bogner, 1984, 1989). He is a member of the team doing the Flora Malesiana and is a coauthor of a checklist and bibliography of the region (Hay et al., 1995a, 1995b) and has written about a collecting trip to Sarawak (Bogner & Boyce, 1995).

Last but not least are Bogner's capabilities as a grower, where few are his equal. His collection of living aroid genera, housed at the Munich Botanical Garden, is unparalleled. His many field trips to three continents where he successfully sleuths yet another poorly known genus or species are largely financed with his personal funds and his accumulated vacation time. This largely unrewarded effort on the part of Josef Bogner is one of the greatest contributions ever to the field of aroid research.

One of the few South American botanists who played a role in research with Araceae is Jorge Crisci from the Museo de la Plata in La Plata, Argentina. His first paper dealing with Araceae was a treatment of the Araceae for the Flora of Buenos Aires Province (Crisci, 1968a). He then described new additions to the flora (Crisci, 1968b; Crisci et al., 1991) or new species (Crisci, 1970). Most of his work involves floristic accounts of Argentina (Crisci, 1971; Crisci & Katinas, 1999). Finally, he wrote a systematic and ethnobotanical study of Philodendron bipinnatifidum (Schott) Schott (Crisci & Gancedo, 1971).

Tom Croat, of the Missouri Botanical Garden, became interested in the Araceae in 1967 when, in conjunction with his work on the Flora of Barro Colorado Island (Croat, 1978a), he found the family to be the most difficult in the flora. Perplexed by the immense variation in species and the confusion of juvenile, preadult, and adult forms, he collected elsewhere in Panama where the wet forests were much richer and replete with undescribed species. Plants were collected and grown at Summit Gardens in the Canal Area, near where he lived, and later were transported to the Missouri Botanical Garden in St. Louis. His earliest paper on Araceae described a Dracontium from Barro Colorado Island (Croat, 1975a), followed later that year by a discussion of the Anthurium gracile (Rudge) Schott-A. friedrichsthalii Schott complex of Central and South America (Croat, 1975b).

Croat's earliest interest was involved with the Araceae of Panama, the richest part of Central America (Croat, 1985a), and this led to floristic work with Anthurium in Central America sponsored by the National Geographic Society (Croat, 1977). This was followed by descriptions of new species of Anthurium (Croat, 1978c, 1979a, 1981a, 1983a), a concern for the standardization of species descriptions of Anthurium (Croat & Bunting, 1979), a revision of Central American Anthurium sect. Polyphyllium Engl. (Croat & Baker, 1978), a treatment of Anthurium for Costa Rica (Croat & Baker, 1979), a revision of the Araceae of the La Selva Reserve in Costa Rica (Croat & Grayum, in prep.), a study of the flowering behavior of Anthurium (Croat, 1980), a study of the sectional classification of Anthurium (Croat & Scheffer, 1983), and a review and analysis of chromosome information for Anthurium (Sheffer & Croat, 1983b). Additional publications during this time were a revision of Syngonium (Croat, 1981b) and a review of the distribution of Araceae worldwide (Croat, 1979b).

A National Science Foundation supported revision of Anthurium of Central America began in 1977 (Croat, 1983a, 1986a, 1986b) and was followed, between 1980 and 1986, by a revision of Anthurium sect. Pachyneurium Schott for the neotropics (Croat, 1991a). The resulting field work in South America brought other involvements including papers on the Araceae of Venezuela (Croat & Lambert, 1987), a treatment of the Araceae for the Flora de Paraguay (Croat & Mount, 1988), a checklist for the Flora of the Guianas (Croat, 1992c, 1997a), the treatment for the Flora of Central French Guiana (Croat, 1997c) and for the flora of Nicaragua (Croat & Stiebel, in press), the checklist for the flora of Peru (Croat, 1993), and for Ecuador (Croat, in prep.) as well as more specific floristic studies of Colombia and Ecuador. In Colombia, a National Geographic grant sponsored study involved a comparative study of lowland pluvial forest-wet forest transition at Bajo Calima (Valle) and an upland premontane wet forest at La Planada (Nariño) (Croat, 1992). This in turn has led to student involvement and (as yet unpublished) treatments of the aroid floras of both areas, for La Planada with Jeff Lake (Croat & Lake, in prep.) and for Bajo Calima with Dorothy Bay (Croat & Bay, in prep.). Similar floristic studies in Ecuador resulted in a comparison of six biological reserves in Ecuador (Croat, 1995b) and a study of Anthurium in the Reserva ENDESA (Croat & Rodríguez, 1995).

Other efforts dealing with Araceae include review papers on the use of neotropical Araceae as medicinal plants (Croat, 1994a), the locality of neotropical aroid collections (Croat, 1988a), the ecology and life forms of Araceae (Croat, 1988b, 1989), a comparative survey of three modern systems of suprageneric systems of classification (Croat, 1990), and a history and status of systematic aroid research (1998). Smaller revisionary studies include the Anthurium bredemeyeri Schott complex in Venezuela (Croat, 1985e), a treatment of the Araceae entitled In Gardens of Hawaii (Croat, in press), a treatment of Syngonium (Croat, 1984a) and Alocasia for the European Garden Flora (Croat et al., 1984).

Miscellaneous papers dealing with Araceae include historical briefs on aroid specialists including Matuda (Croat, 1978b) and Engler (Croat, 1983c), as well as reports on aroid conferences (Croat, 1985f, 1992b, 1994b; Croat & Cosgriff, 2000), discussions of collecting and preparation procedures for Araceae (Croat, 1985b, 1969), special drying facilities needed for Araceae collections (Croat, 1979d), labeling living collections (Croat, 1984d), germination of seeds (Croat, 1979c), and propagation of cuttings (Croat, 1981c). Other miscellaneous papers deal with a discussion of Anthurium andraeanum Linden (Sheffer et al., 1980), A. leuconeurum Lem. (Croat, 1983d), the description of new South American species Croat (Croat, 1987; [Peru] in press), reports on living collections of Araceae (Croat, 1979d, 1988c) or of field trips (Croat, 1982a, 1982b, 1991b), reports on rare (Croat, 1983b, 1984b, 1985c, 1995a) or poorly known species (Croat, in prep.), new combinations (Croat & Grayum, 1987, 1994), new records (Croat & Pérez-Farrera, 2000), and illustrative profiles of aroids, including Philodendron rugosum Bogner & G. S. Bunting (Croat, 1984c), Taccarum weddellianum Brongn. ex Schott (Croat, 1985d), and Syngonium steyermarkii Croat (Croat & Bogner, 1987). With Kay Rossmann, he produced an index for the first ten volumes of Aroideana (Croat & Rossmann, 1991).

Other recent NSF sponsored projects have been completed or are being carried out, including a revision of Philodendron subg. Philodendron of Central America (Croat, 1997b), a revision of Rhodospatha (Croat, in prep.), and a revision of Dieffenbachia of Central America (Croat, in prep.). In addition Anthurium sect. Semaeophyllium is being revised with the help of R. L. Mansell at the University of South Florida (Croat & Mansell, in prep.) and Anthurium sect. Calomystrium ser. Rupicola ser. nov. is being revised with the assistance of Jane Whitehill (Croat & Whitehill, in prep). These five as yet unpublished revisions will result in a total of 96, 67, 30, 22 and 8 species respectively with a total of 139 taxa (65, 52, 22, 6 and 3 species respectively) new to science. Additional new, as yet unpublished taxa have resulted from floristic studies. For example, 20 of the 50 species for Reserva La Planada in Colombia are believed to be new; 84 of 150 for the Flora of La Planada (Nariño); and 11 of the 31 taxa at the Reserva ENDESA in Ecuador. To this date, Croat has published 352 taxa. The resources which have been built up for aroid research at the Missouri Botanical Garden include one of the largest living collections of aroids and the largest collection of herbarium specimens of neotropical aroids. The living and dried collections include a large percentage of Croat's more than 80,000 personal collections.

Dorothy Shaw has published a series of mostly technical, experimental or ecological papers concerning the Araceae of Australia and Papua New Guinea. These include observations on the behavior of Colocasia esculenta (Shaw, 1975, 1982; Shaw et al., 1979), pollination in Alocasia macrorrhizos (L.) G. Don [A. brisbanensis] (Shaw et al., 1982; Shaw & Cantrell, 1983a, 1983b), fruit dispersal in Alocasia macrorrhizos (Shaw et al., 1985), plant damage and fruit ingestion of seeds of Alocasia brisbanensis by birds (Shaw, 1998a) and lizards (Sha2, 2998b), stomata of Monstera deliciosa Liebm. (Shaw, 1992c), aroids of botanical gardens in Brisbane (Shaw, 1987), germination of Typhonodorum seeds in cultivation (Shaw, 1990), the occurrence of the fungus Puccinia on Monstera (Shaw, 1991, 1992a, 1992b, 1993a, 1994, 1995a, 1995b), habitat of Gymnostachys anceps (Shaw et al., 1997) and of fruit dispersal (Shaw et al., 1997), and postage stamps that exhibit plants of the family (Shaw, 1993, 1999). With R. Greber she reported on the dasheen mosaic virus in Queensland (Greber & Shaw, 1986).

S. Serizawa published on Japanese Arisaema during the late 1970s and mid-1980s (Serizawa, 1975, 1980a, 1980b, 1981a, 1981b, 1982a, 1982b, 1986). These works, published only in Japanese, meant that he was not widely recognized internationally.

A brief research effort was made by Richard Baker at the Field Museum in Chicago. His efforts, before embarking on a new career in the early 1980's, were entirely in Costa Rica. With W. C. Burger, in charge of the Flora Costaricensis project at the Field Museum, Baker revised Spathiphyllum for Costa Rica (Baker & Burger, 1976). A few years later he collaborated with Tom Croat in the revision of Anthurium for Costa Rica (Croat & Baker, 1979).

Mike Madison played an important role in the late 1970s and early 1980s before changing careers. He began with a flourish, publishing five papers the first year; two (Madison, 1976b, 1976c), dealing with new species (Rhodospatha and Asterostigma respectively), another comparing Alloschemone and Scindapsus (Madison, 1976a), and another comparing Caladium and Xanthosoma (Madison, 1976d), and finally a paper dealing with the seeds of Monstereae (Madison & Tiffney, 1976). His Ph.D. thesis, a revision of Monstera, was published the following year (Madison, 1977a). Though Madison did make an expedition to Brazil (Madison, 1979a), his principal fieldwork was in Ecuador where he collected widely, describing species in Caladium (Madison, 1981a), Philodendron (Madison, 1977b), Stenospermation (Madison, 1977c) and Xanthosoma (Madison, 1978e). In addition, he described a plant from Brazil as a new Ulearum (Madison, 1980). This later proved to be the new genus Bognera. Aside from his revision of Monstera, other major papers included a revision of the palmately-lobed Anthurium species (Madison, 1978g), a major paper discussing the ecology of the genera of Araceae of the northern Andes (Madison, 1978f), and a partial revision of the Caladieae (Madison, 1981a).

Madison published many miscellaneous short papers, especially while he was editor of Aroideana. These include reports on nomenclature (Madison, 1978a, 1978d), plant culture (Madison, 1978h), the living collections at Selby (Madison, 1978i), packing and shipping aroids (Madison, 1981b), Monstera seeds and the fossil record (Madison & Tiffney, 1976), the rediscovery of Philodendron frits-wentii (Madison, 1978b), and a synopsis of Caladiopsis (Madison, 1978j). Another includes the protection of developing seeds in Araceae (Madison, 1979b). He also wrote illustrative profiles of Monstera deliciosa (Madison, 1978c), Xenophya [= Alocasia] lauterbachiana (Madison, 1979c), Anthurium lilacinum (Madison, 1979d), A. punctatum (Madison, 1979f), and A. superbum (Madison, 1979e).

One of Madison's major accomplishments was the founding of the journal Aroideana with the International Aroid Society [see discussion below] in 1977. Madison began publishing the journal and was its editor for several years when he changed careers. Many of his earlier papers were written specifically for the journal. Madison was also responsible for organizing the first International Aroid Conference at Selby Gardens in Sarasota, Florida on March 28-29, 1980. These conferences have been continued, albeit, irregularly, and they have contributed greatly to the dissemination of knowledge about aroids. With the completion of the conference in St. Louis, Missouri in August 1999, seven such conferences have been held, three of them in conjunction with the International Botanical Congresses. Aroid research was dealt a severe blow with Madison's retirement. This brilliant Harvard-trained researcher left a significant mark in his five short years of work with Araceae.

Wim Crusio, one of H. C. D. de Wit's students, completed a revision of the genus Anubias, and this excellent work was published twice, once in English (Crusio, 1979a) and once in German (Crusio, 1987). Another description of the genus and a discussion of its taxonomy also appeared in German (Crusio, 1980). Crusio has also published short papers on Cryptocoryne (Crusio, 1979b, 1979c). Along with Arie de Graaf he describes a new species of Lagenandra, L. dewitii (Crusio & de Graaf, 1986) and in another redescribes L. ovata Thwaites (Crusio & de Graaf, 1987).

Taking up where de Wit left off, Niels Jacobsen, from the Royal Veterinary and Agricultural University in Denmark, did additional work on Cryptocoryne, including extensive fieldwork in Southeast Asia. In a series of general papers he described the biology and ecology of Cryptocoryne. His first paper dealing with Araceae is about the ecology of Cryptocoryne (Jacobsen, 1976) while others deal with its pollination (Jacobsen, 1977a), chromosomes (Jacobsen, 1977b, 1977c; Arends et al., 1982), and flowering behavior (Jacobsen, 1980a), vegetative morphology (Jacobsen et al., 1989a-c) as well as with the description of new species (Jacobsen, 1977d, 1979a, 1980b, 1980e, 1981a, 1982, 1985a), a discussion of C. undulata (Jacobsen, 1981b), C. ferruginea (Jacobsen, 1980d), and a revision of the Cryptocoryne albida complex (Jacobsen, 1980c). A 1991 paper (Jacobsen, 1991) treated the small-leaved Cryptocoryne species. A paper co-authored with Marian Ørgaard involved a SEM study of surface features in the spathes of Cryptocoryne and Lagenandra (Ørgaard & Jacobsen, 1998). In a series of papers with Josef Bogner, he revised the Cryptocoryne of the Malay Peninsula (Jacobsen & Bogner, 1986, 1987a-c) then published a complete revision for Borneo (Jacobsen, 1982, 1984, 1985b) and later for Ceylon (Jacobsen, 1988), and Tasek Bera (Jacobsen, 1986). These publications were precursors to his full revision of the genus. The complete revision of Cryptocoryne was published in two versions, one in Danish (Jacobsen, 1979b) and one in German (Jacobsen, 1979c). He will contribute Cryptocoryne for the Flora Malesiana and is a coauthor of a checklist and bibliography for the region (Hay et al., 1995a, 1995b). Finally, Jacobsen published the treatment of the Arales in R. M. J. Dahlgren, H. J. Clifford & P. F. Yeo's, The Families of Monocotyledons: Structure, Evolution and Taxonomy (Jacobsen, 1985c).

Li Heng, working for the Chinese Academy of Sciences at the Kunming Institute of Botany and doing research on Chinese Araceae even before China's opening to the West, has become the authority on the family in China. Her earliest publication deals with the medicinal value of certain Arisaema (Li, 1976) and another, (Li et al., 1977) "Claves diagnosticae et taxa nova Aracearum Sinicarum", provides a key to the genera of Araceae of China and describes 30 new taxa. Her principle interest is in Arisaema, including its phytogeography (Li, 1980a, 1980b, 1981) and taxonomy, describing many new species (Li, 1985, 1988a, 1992a; Li et al., 1999) as well as Amorphophallus (Li, 1988b-d; Li et al., 1989, 1990; Li & Long, 1989; Long et al., 1989), Typhonium (Li & Zgeb-quian, 1983), Remusatia Schott (Li, 1987a, 1991, 1992b; Li & Hay, 1992b; Long et al., 1989b), Rhaphidophora (Li, 1992b), Gonatanthus [later reduced to Remusatia] (Li, 1987b; Li & Hay, 1992b), and Colocasia (Li & Wei, 1993). Her papers frequently deal with cytological details of the species described (Gu et al., 1992; Li et al., 1989). Still others deal with floristics such as that of the Dulongjian Region (Li, 1993b), Xizang area (Li, 1987c), the Gaoligong Mountains (Li et al., 1999), or plants of medicinal value (Li, 1988). A paper dealing with the phytogeography of the Araceae (Li, 1986), divides the family into 12 distribution patterns and 29 subpatterns; another deals with the origin and phylogeny of Araceae (Li, 1983). Li presented a paper at the XVI International Botanical Congress in Yokohama dealing with the species diversity of Chinese Araceae (Li, 1993a). Perhaps her major accomplishment is the treatment of the Araceae of China written with C. Y. Wu (Li, 1979a, 1979b) that deals with 34 genera and 191 native species. The largest aroid genus in China, Arisaema, is reported with 82 species. Li Heng is also chiefly responsible for the treatment of the Araceae in the "Iconographia cormophytorum sinicorum (Anonymous, 1976) published by an editorial committee of that publication. This work treats 26 genera and 51 species, all illustrated with line drawings. Li is currently working on the English version of the Flora of China and presented information about that project at the VI and VIII International Aroid Conferences (Li & Long, 1998a). Another recent paper (in Chinese) (Li & Long, 1998b) deals with the taxonomy of Amorphophallus and includes a key to the Chinese species.

Also in China in the same year, Kao Pao-Chung [Gao Baochum], working with the Academica Sinica in Chengdu, Sichuan, did a revision of the Araceae for the Flora Sichuanica (Kao, 1989a) and described new species of Araceae (Kao, 1989b). That flora, not as tropical as Yunnan, treated 13 native genera and 62 native species.

In southwestern China, Liu Pei-Ying at the Research Center of Konjac at the Southwest Agricultural University in Chonqing, has been working on Amorphophallus breeding. She presented a paper at the VI International Aroid Conference in Kunming entitled "Research and Utilization of Amorphophallus (Liu, Zhang & Zhang, 1998).

Simon Mayo of the Royal Botanic Gardens, Kew is today one of the foremost aroid researchers. His first paper dealing with Araceae was a report on his early fieldwork in Brazil (Mayo, 1978a), making it one of the first contributions in the new journal Aroideana. His next two papers (Mayo, 1978b; Mayo & Barroso, 1979) dealt with Brazilian species, and his interest in Brazilian Araceae continues to this day (Mayo, 1983a, 1986e, 1987c, 1988b, 1989b, 1995; Mayo & Barbosa, 1996; Mayo & Féliz, 2000; Mayo & Fevereiro, 1982; Mayo & Zappi, 1993; Fevereiro & Mayo, 1982; Mayo et al., 1995; Sampaio et al., 1996). Mayo has in recent years lived and worked in Brazil where he has taught and organized the research of several Brazilian students who were interested in Araceae (Mayo & Nadruz, 1992; Ramalho, 1995; Sakuragui, 1994; Nadruz, 1995; Andrade, 1996; Soares, 1996; Sakuragui & Mayo, 1997). Together with Brazilian colleagues, he has prepared a checklist for all of Brazil (Mayo et al., in prep.) and has been especially interested in the Atlantic coastal regions (Mayo, 1990b). He has published papers on the phytogeography (Mayo, 1984b) and taxonomy of Bahía (Harley & Mayo, 1980; Mayo, 1984b) and has recently prepared a revision of the Araceae of Bahía (Mayo, manuscript). On a broader topic Mayo discussed aroid phytogeography in Africa and South America (Mayo, 1993). Other Mayo papers dealing with New World aroid species are those describing a new Caladium (Mayo & Bogner, 1988) and rediscovering Gearum N. E. Br. (Mayo et al., 1994).

Among Mayo's earliest efforts were his work with the flora of Trinidad (Mayo, 1981, 1986a) and with taxonomic problems in the West Indies, such as a resolution of the poorly understood Anthurium acaule and its relationship to the A. sect. Pachyneurium (Mayo, 1982a). Mayo has also been heavily involved with African Araceae, and he has produced the treatment of the Araceae for the Flora of the Mascarene Islands (Mayo, 1983b, 1984c) and the Flora of Tropical East Africa (Mayo, 1985a). This in turn has led to considerable involvement with the taxonomy of African Amorphophallus (Mayo et al., 1982; Bogner et al., 1985), Araceae in the Flora of Cyprus 2 (Mayo & Meikle, 1985), and Arisaema (Mayo, 1982b, 1984a, 1985b, 1986b, 1987a, 1987b; Mayo & Gilbert, 1986). A short paper deals with Biarum (Mayo, 1980a) while others focus on aroids at Kew (Mayo, 1979) and an aroid symposium at Selby Gardens (Mayo, 1980b). Still another paper discusses the presence of anthocyanins and flavonoids in the Araceae (Williams et al., 1981). Mayo's participation in a special volume of Curtis's Botanical Magazine resulted in articles dealing with the "Genera of Araceae" project (Mayo et al., 1995a, 1995b, 1995e), and Roberto Burle Marx (Mayo, 1978c, 1982c, 1982d; Mayo et al., 1995c). He has also reviewed various books on Araceae (Mayo, 1980d, 1982d, 1983c, 1986c, 1991b).

For his Ph.D. work Mayo chose to do a revision of Philodendron subg. Meconostigma (Schott) Engl. (Mayo, 1986d) but his work went well beyond Meconostigma, leading him to conduct research in various aspects of the whole Philodendroideae. In a series of papers he discusses the evolution (Mayo, 1988a), the gynoecial structure (Mayo, 1989a) and the taxonomy of P. subg. Meconostigma (Mayo, 1991a), and the history and infrageneric nomenclature of Philodendron (Mayo, 1990). He was the first to formally recognize tribe Pteromischum as a subgenus. Mayo has subsequently put all of his information on this group and other genera together in a massive computer-driven cladistic study to reappraise the suprageneric classification of the Araceae. This system is presented in The Genera of the Araceae (Mayo et al., 1997). Using the same cladistic information, the authors (including J. Bogner & P. Boyce) present the case for the inclusion of the Lemnaceae into the Araceae as a subclade of a monophyletic Araceae (Mayo et al., 1995). They have also completed a treatment of the Araceae in K. Kubitzki's The Families and Genera of Vascular Plants (Mayo et al., in press), and done an article on the acolytes of the Araceae (Mayo et al., 1995d).

Simon Mayo's decision in 1973 to leave the Horticulture Department at Kew Gardens and to become involved with the taxonomy of the Araceae was an important event for research with the Araceae. His research, especially his broad general studies in the evolution of the Philodendroideae and his cladistic studies which have reclassified the genera of Araceae, are on the cutting edge.

Richard Sheffer did important breeding studies and cytological work with Anthurium during his Ph.D. dissertation (Sheffer, 1974, 1977) at the University of Hawaii and later at Indiana University Northwest. The work was carried out in part with his major professor, cytologist and Anthurium breeder H. Kamemoto (Sheffer & Kamemoto, 1976a, 1978; Sheffer et al., 1980). Sheffer published a review of chromosome numbers for Anthurium (Sheffer & Kamemoto, 1976b; Sheffer & Croat, 1983), and he conducted a cytotaxonomic study of the Anthurium scandens complex (Sheffer et al., 1980). Another major accomplishment was breeding studies carried out with Anthurium sect. Pachyneurium (Croat, 1991a) [see also Kamemoto above]. Sheffer has a new greenhouse facility filled with Araceae which he uses in his cytological research.

Another important plant breeder dealing primarily with Araceae is R. J. (Jake) Henny from the Central Florida Research and Education Center, University of Florida in Apopka. His work has been largely experimental involving culture techniques of Aglaonema, Anthurium, Dieffenbachia, and Spathiphyllum (Henny, 1980a, 1989a, 1989b; Henny & Fooshee, 1990a, 1990b; Henny et al. 1980a, 1994, 1995), the use of growth regulators to induce flowering (Henny, 1980b, 1981, 1983a, 1983c, 1988c, 1989b, 1991, 1992; Henny & Fooshee, 1983, 1989b, 1990b, 1990c, 1991; Henny & Rasmussen, 1980b), breeding (Henny, 1982a-c, 1983b, 1984, 1988a, 1989a; Henny & Rasmussen, 1980a, 1980c, 1980d), and aroid introductions (Henny, 1988b, 1995a, 1995b; Henny et al., 1987a, 1987b). Ann Chase, also from the C.F.R.E.C.-Apopka, works on aroid research and has published results on various cultural aspects (Chase, 1989; Chase & Henny, 1990; Chase & Poole, 1991) as have C.A. Conover (Conover & Henny, 1995), R.W. Henley (Henley, 1992), and D. Norman (Norman, 1996).

Marija Bedalov, working at the University of Zagreb in Croatia, has worked many Araceae of the Balkan region. Her Ph.D. thesis, written in Croatian dealt with the cytotaxonomy of the Araceae of Yugoslovia (Bedalov, 1973a, 1976b). Since then she has worked with several genera including Arisarum (Bedalov & Broni?, 1999), Biarum (Bedalov, 1969b), Calla (Bedalov, 1983b, 1994), and Dracunculus (Bedalov, 1972, 1976b, 1994; Bedalov & Hesse, 1999), but most of her work has been with Arum, especially dealing with aspects of cytology, phytogeography and palynology (Bedalov,1975a-c, 1976a, 1977, 1978, 1980, 1981a, 1981b, 1982, 1983a, 1984, 1985, 1999; Bedalov & Guttermann, 1982; Bedalov & Bronic, 1989; Bedalov & Hesse, 1989; Bedalov et al., 1991; Bedalov et al., 1993a, 1993b, 1999a, 1999b; Bianco et al., 1994; Bedalov & Fischer, 1995; Bedalov & Drenkovski, 1997; Bedalov & Bronic (1998a, 1998b; Bedalov et al., 1998). She also has been very active publishing chromosome reports (Bedalov, 1973; Favarger & Bedalov, 1998) and especially in IOPB Chromosome Data 10 [see list of literature]. With M. Hesse she has studied pollen types within Dracunculus (Bedalov & Hesse, in prep). Other papers in preparation include a cytotaxonomical study of Arisarum vulgare (Bedalov & Bronic, in prep.), a paper on the artificial hybridization in Arum (Bedalov et al., in prep), on observations in meiosis in Arum (Bedalov et al., in prep), and on studies with Arum in Denmark (Bedalov et al., in prep).

Bedalov has also dealt with the cytology and phytogeography of Calla and Acorus (Bedalov, 1983b). Her work has largely been concentrated in the Balkan region, especially in the former Yugoslavia. A participant in the first International Aroid Conference as well as third, fourth and sixth, she is an active and productive researcher. In addition to her work in Zagreb, she carries out independent investigations with a colleague in Switzerland.

Jin Murata, an expert on Asian Arisaema, published his first paper on Arisaema late in the decade (J. Murata, 1978). Several other papers describing new species followed (J. Murata, 1956, 1983a, 1985a; J. Murata & Ohashi, 1980; J. Murata & Ohno, 1989; J. Murata et al., 1994), one on a new combination in Typhonium (J. Murata & Mayo, 1991) and another describing the first leaves of a species (J. Murata, 1986a). In a recent paper in Aroideana, Murata provides keys, photos and a discussion of the Japanese species of Arisaema (J. Murata, 1990a). Other papers include information on chromosomes of Arisaema (J. Murata, 1983b, 1990b; J. Murata & Iijima, 1983), a study of the stem morphology (J. Murata, 1988), a study of shoot organization recognizing four types of stems (J. Murata, 1990c), a study of developmental pattern of pedate leaves (J. Murata, 1990d) and allozyme differentiation in Arisaema (J. Murata & Kawahara, 1994a-c). Two papers deal with attempts of infrageneric classification (J. Murata, 1984, 1990f) and others deal with revisions of minor groups or species complexes of Arisaema (J. Murata, 1962, 1985b, 1986b, 1986c, 1990d, 1990g, 1991, 1995; J. Murata & Ohno, 1991). Murata also wrote a memoriam for Hiroshi Hara (J. Murata, 1987).

Among Murata's major publications are a complete revision of Arisaema in Japan (Ohashi & J. Murata, 1980), complete with keys and illustrations [see Ohashi above] and a proposed infrageneric classification of Arisaema (J. Murata, 1984). Murata also participated in the research of his student, Duangchai Sriboonma in a molecular study of the genus Typhonium (Sriboonma et al., 1993) and a revision of the genus (Sriboonma et al., 1994). Murata has recently published an extensive review of the cytology of Arisaema with the help of senior author Kuniaki Watanabe and Tomiki Kobayashi (K. Watanabe et al., 1998).

Murata is a member of the team of taxonomists working on the Araceae treatment for the Flora Malesiana and is a coauthor of a checklist and bibliography for this region (Hay et al., 1995, 1995a). He organized the VI International Aroid Conference as a part of the XI International Botanical Congress in Yokohama in 1993.

Another Japanese botanist, sometimes publishing with Hotta, is H. Okada. His work has been largely experimental and behavioral, including cytotaxonomical studies of populations of Araceae in West Sumatra (Okada, 1984, 1986), studies of population dynamics of Schismatoglottis in Sumatra (Okada, 1989; Okada & Hotta, 1987; Okada & Mori, in press) as well as on chromosome behavior in Colocasia (Okada & Hambali, 1989) and Schismatoglottis (Okada, 1992a). Two other papers discuss cytotaxonomic studies of rheophytic aroids (Okada, 1992b, 1993).

Working in Hungary on Arum, A. Terpo has made studies on the distribution and taxonomy of Arum species (Terpo, 1971, 1973) in Pannonian territories (now mostly Hungary and Yugoslavia).

Toward the end of the 1970s, P. Blanc in France carried out important studies on the growth behavior of the Araceae. These studies (Blanc, 1977a, 1977b, 1978, 1980) were a precurser to the more extensive growth behavior studies carried out by Tom Ray (see below).

As mentioned earlier, the late 1970s also saw the creation of the International Aroid Society, known initially as the American Aroid Society. This organization, founded in Miami, Florida on June 18, 1977, was started through the inspiration of Bette Waterbury, Allen Fernández, John Faust, Shirley Crete, Marilyn Turner, Peggy Fischer (all constituting the first officers with Waterbury as president), Tom Fennell, Monroe Birdsey, Ron Weeks, De Hull, Gary Antosh, Joan Hackler, and others. In the words of Michael Madison, in the leading article of Aroideana, the International Aroid Society was intended to "promote the study of the aroid family in all of its aspects." Madison started publishing the society's journal, naming it Aroideana at the suggestion of Dan Nicolson. The society and certainly the journal have had their share of difficulties, mostly the result of a volunteer-driven and sometimes inadequately prepared staff, but also because of editors who gave up, and even one (Mark Moffler) who died suddenly, along with the many difficulties in finding the right publisher. The journal has by all accounts been immensely valuable in dealing with Araceae. Many of the papers published throughout the years simply would not have been written at all had it not been for the need to "feed the presses." Major contributors in the first few years were Madison himself, Simon Mayo, Josef Bogner, Tom Croat, and Harald Riedl. Other aroid researchers who contributed were George Bunting, Niels Jacobsen, Tom Ray, Dan Nicolson, Alistair Hay, Peter Boyce, M. Sivadasan, Richard Sheffer, Dorothy Shaw, Mike Grayum, Richard Henny, H. Kamemoto, Marianne Knecht, Farah Ghani, Larry Klotz, and Mark Moffler.

From the onset, the journal has encouraged and received articles from horticulturists and plant collectors since the aroid society is primarily supported by plant lovers of all kinds. Many of them have contributed greatly, such as Bette Waterbury, John Banta, Frank Brown, an expert on the genus Aglaonema (Brown, 1980, 1982, 1984), Fred Dortort, Amy Donovan, Lawrence Garner, dealing with hybridizing Alocasia (Garner, 1983), James Watson, Luis Bueno, Julius Boos, David Leedy, Linda Theus, David Prudhomme, John Johnston, Joe Wright, Mike Bush, William Drysdale, Marcel Lecoufle, Stu Cramer, and Arnold Melim. Naturalist Julius Boos, a recent contributer, is particularly knowledgeable about the aroids of Trinidad and some members of the Lasioideae (Boos, 1997), especially Urospatha (Boos, 1993; Boos & Boos, 1993). Fanny Phillips made an important contribution to the understanding of Amorphophallus (Phillips, 1988). Libbe Besse wrote a paper on the native south Florida aroids (Besse, 1980). She has also played an important role in the development of Selby Gardens and has sponsored and participated in several important expeditions to Ecuador with Mike Madison and others at Selby Gardens. These expeditions were among the most productive, ever, in the procurement of living Araceae and her assistance is to be commended. David Burnett made a major contribution where he contributed an unpublished table of contents to earlier Aroideana volumes that was useful in preparing the published indices (Croat & Rossman, 1991; Donovan & Malesevich, 1994), and he published an illustrated introduction to the cultivated Alocasia (Burnett, 1984). Since there is no modern revision of this genus, the work by Burnett has been immensely useful. Another paper discusses a proposal for hybrid and cultivar names (Burnett, 1982).

In addressing the success of the journal and the International Aroid Society itself, both of which have been instrumental in promoting activity with Araceae, a major tribute must go to a small band of enthusiasts in Miami who have provided the impetus to keep things running. The faces have changed over the years although many devoted members are worthy of mention, only a few can be mentioned here. Foremost is the late Bette Waterbury, founder and first president (who also served subsequent terms) and president emeritus, who did so much to keep the society alive. Other productive members such as Linda Theus, Allen Fernandez, and Maree Winter are no longer with us. Special thanks must go to people like Amy Donovan, present editor of the journal and one of the most dedicated society members; Dewey Fisk who served as board member, President, Corresponding Secretary, journal editor, plant sale promotor and auctioneer; David Burnett, who served as a board member, recruited many Australian members and ran Australian membership affairs; David Leedy who served as newsletter editor; Bruce McManus who served as newsletter editor, membership chairman and especially as Show Chairman for the annual meeting and show; Denis Rotalante, Ron Weeks and others who could always be counted on to bring big plants for the Annual Show and Sale; Tricia Frank, Susan Staiger, Jerry Bengis, Donna Rich, and many others who have served as officers or who were heavily involved in the work of putting together the Annual Meeting and Show are all to be commended for their efforts. Petra Schmidt Malesevich, who has been my faithful assistant for many years, first as aroid greenhouse manager and later as research assistant, is one of the unsung heroes of the aroid community. She has served as a board member, membership chairperson, assistant editor, book sales and compiler of membership lists and Aroideana indices in addition to carrying out the multitude of tasks necessary in my own research program. Finally, the many others who were there making the society work; they are the ones who sold the plants, the T-shirts and books to make the profits which kept the journal in publication during the lean years. Certainly all of us owe them a debt of gratitude.

The 1980s were, in many respects, some of the most important years for aroid research. This period saw the greatest increase of knowledge since the time of Engler, Krause and Sodiro, around the turn of the century. Most researchers who had begun their work in the 1960-1970 period were still active. It was also a time of real ferment with a number of excellent, new, well-trained researchers beginning their careers with Araceae. Peter Boyce at Kew began work with Mediterranean Araceae. The focus of research on Araceae also became increasingly diverse; no longer mostly systematic, but instead focus widened to a number of behavioral and experimental approaches. Hegnauer reviewed the chemical attributes of the Araceae (Hegnauer, 1963, 1986, 1987). Tom Ray began working with a wide variety of aspects of growth behavior (See, Ray below). H. J. Tillich reported on seedling development (1985). Jim French conducted an extensive and comprehensive review of technical aspects of the Araceae, including a wide spectrum of anatomical features and a broad molecular survey before embarking on studies with pollination biology of the Araceae (See French below). Mike Grayum startled the aroid world with his astounding new suprageneric classification that followed his thorough SEM review of pollen and a review of virtually all character states in the Araceae (Grayum, 1984). Marianne Knecht published her biosystematic study of the Araceae of the Ivory Coast. Denis Barabé began studies of floral anatomy. William Carvell followed with studies on the Pothoideae and Monsteroideae (Carvell, 1989a; 1989b). Robin Scribailo at Purdue North Central in Westville, Indiana, is now doing similar studies on floral anatomy. He published work on the developmental anatomy of Peltandra (Scribailo & Lloyd, 1993) and on shoot and floral development in Calla (Scribailo & Tomlinson, 1992). Gitte Petersen began her work with the cytology of the Araceae (see "Miscellaneous Disciplines" below). Helen Young (currently at Barnard College in New York), working at La Selva in Costa Rica, observed Philodendron rothschuhianum (Engl. & K. Krause) Croat & Grayum (Young, 1987), and often with the assistance of George Schatz (Missouri Botanical Garden) studied reproductive biology of Dieffenbachia (Young, 1986, 1988a, 1988b) thereby discovering many interesting features that gave insight into all other beetle pollinated genera. Lloyd Goldwasser (University of California) worked on similar pollination projects. Long Chun-lin, working with Li Heng at the Kunming Institute of Botany, began working with Chinese Araceae. Finally, this decade saw one of the first and certainly the best book devoted to Araceae written for the general public, Aroids, written by Deni Bown, an amateur aroider and a highly regarded professional writer. Her book has gone a long way toward introducing Araceae to the general public. It is not only highly informative, but it is easy reading and is filled with excellent pictures (Bown, 1988). She also contributed papers for Aroideana on naturalized English aroids (Bown, 1985) and on the history of Acorus calamus L. (Bown, 1987) to Aroideana.

The published aroid research of James C. French, first at the University of Mississippi and later at Rutgers, began in the early 1980s. Perhaps no aroid worker was ever as prolific over a single decade. His first paper (French, 1977) dealt with growth relationships of leaves and internodes in vining angiosperms with different modes of attachment. His first paper dealing exclusively with Araceae was a collaborative survey of the vascular system in Araceae (French & Tomlinson, 1980). A series of papers followed which revealed the vascular anatomy of all the subfamilies: Pothoideae (French & Tomlinson, 1981a), Philodendroideae (French & Tomlinson, 1981b, 1984), Calloideae and Lasioideae (French & Tomlinson, 1981c), Monsteroideae (French & Tomlinson, 1981d), and Colocasioideae, Aroideae and Pistoideae (French & Tomlinson, 1983). Another work dealt with a much larger survey of Philodendron, a genus of especially variable vascular anatomy (French & Tomlinson, 1981b) while still another dealt with stems in general (French, 1983). French also embarked on an independent survey of a variety of anatomical features with the hope of understanding their taxonomic significance. These included meristems (Fisher & French, 1976, 1978), endothecial thickenings in stamens (French, 1985a, 1985b, 1986c), ovular vasculature (French, 1986a), sclerotic hypodermis in roots (French, 1987a), anastomosing laticifers (French, 1988), and patterns of staminal vasculature (French, 1986b). With M. G. Fox he studied the systematic occurrence of sterols in the latex of Colocasioideae (Fox & French, 1988). These broad surveys contributed greatly to a better understanding of the evolution of the Araceae and the attempt, so intense at that time, to resolve the differences in competing systems of classification (Grayum, 1984; Bogner & Nicolson, 1991).

In a thorough survey of the chloroplast DNA of Araceae (French et al., 1995), French and his colleagues contributed greatly to the most recent revision of the suprageneric classification of the Araceae (Mayo et al., 1997). Other molecular studies dealt with Acorus and Gymnostachys (French & Kessler, 1989) and the Colocasioideae (Kessler & French, 1989). Jim French has covered a variety of distinct research topics in Araceae and has done them all well. His latest approach is with pollination biology which he is conducting during his sabbatical while living with his family in Costa Rica.

Alistair Hay began his career in New Guinea and published a treatment of the Araceae of Papua New Guinea (Hay, 1981). For his D.Phil. dissertation he revised Cyrtosperma (Hay, 1986, 1988a). With D. J. Mabberley he published a paper (Hay & Mabberley, 1991) on a controversial theory of evolution in Araceae and discussed its implication for the evolution of other angiosperms. Hay, now working at the Royal Botanic Gardens, Sydney in Australia, had a major interest in Australasia and published a treatment of Alocasia for Australasia with R. Wise (Hay & Wise, 1991), discussed collecting Alocasia in New Guinea (Hay, 1990a), did a revision of Typhonium (Hay, 1993a), and Amorphophallus (Hay, 1988b) for Australasia, described the new genus, Lazarum, discovered in Australia (Hay, 1992a), published the Araceae of New South Wales (Hay, 1993b), and new taxa of Alocasia (Hay, 1989, 1994; Hay et al., 1997 in press), Colocasia (Hay, 1996b), Nephthytis in Borneo (Hay et al., 1994), Rhaphidophora (Hay, 1993c), and aroids of Papua New Guinea (Hay, 1990b). Recently he completed a revision of the genus Pothos for New Guinea, Solomon Islands, and Australia (Hay, 1995), introduced a new species of Typhonium (Hay & Taylor, 1996), and a revision of Schismatoglottis for Peninsular Malaysia and Singapore (Hay, 1996a). He is also publishing a revision of Homalomena in New Guinea, the Bismarck Archipelago and Solomon Islands (Hay, 1997a), an article on Alocasia melo (Hay et al., 1997), and on new species of Typhonium (Hay, 1997b). Hay has recently revised Alocasia in the Philippines (Hay, in press), reviewed Schismatoglottis in the Philippines (Hay, in press) and is working on a treatment of the Flora of Australia (Hay, in prep.), a revision of Alocasia in West Malaysia (Hay, in prep.), and a popular account of the Araceae of Sabah and Sarawak (Hay, in prep.). Other recently completed papers involve studies with shoot architecture in Pothos (Hay, in press) and a discussion of the value of living collections for taxonomic studies and for conservation (Hay, in press).

Hay's interest has continued with the subfamily Lasieae worldwide. He described the neotropical genus Anaphyllopsis A. Hay (Hay, 1988c) and discussed the tribal and subtribal distribution and circumscription of the Lasieae (Hay, 1992b) as well as the proper circumscription of Lasia concinna Alderw. (Hay, 1988d). Currently Hay is the coordinator of the Araceae treatment for the Flora Malesiana (Hay, 1994b), a project involving contributions from about a half dozen aroid taxonomists from all over the world, including, in addition to Hay, Josef Bogner, Peter Boyce, Wilbert Hetterscheid, Niels Jacobsen, Jin Murata, and Elizabeth Widjaja. A recent joint effort by several of these contributors has resulted in a checklist (Hay et al., 1995a) and a bibliography (Hay et al., 1955b). As the prime mover on the Flora Malesiana project and a member of the Flora Malesiana Foundation Board, his contribution to aroid research is certain to continue to be profitable.

Tom Ray, carried out studies on growth and heterophylly on Syngonium for his Ph.D. thesis at Harvard (Ray, 1981). While working at the University of Delaware, he played an important role in understanding growth of Araceae and his first paper in Araceae dealt with skototropism (Strong & Ray, 1975). Other early papers describe the physical aspects and variability of growth behavior in general terms (Ray, 1979) and use of specific cases to demonstrate growth behavior (Ray, 1983a, 1983b; Oberbauer et al., 1980). In another paper (Ray, 1986) he began to define terms and discuss the universality of the stem segment regardless of its age. In another he discusses cyclic heterophylly of plants displaced from trees (Ray, 1987a). In a paper entitled "Leaf types in the Araceae" (Ray, 1987b) he begins to define the complex terminology that will become a part of his work. In the next two papers (Ray, 1987c, 1988) the system is further defined and a schematic formulation is provided for each type of shoot organization. Ray also describes metamorphosis, i.e. the abrupt change from one growth form to another, and discusses how this varies in different genera (Ray, 1990, 1992a). Another paper discusses a novel method to measure and record leaf shape using Syngonium podophyllum Schott as a test case (Ray, 1992b). Though not a systematist, Ray did describe a new species of Syngonium (Ray, 1980). Ray's comparative surveys of most genera proved very useful in helping to sort out the generic relationships. With the assistance of Susanne Renner (Ray & Renner, 1990) he translated Part 2 of Engler's (1877) "Comparative Studies on the Morphology of the Araceae." This information was yet another important element in the body of knowledge accumulated during this productive period of Araceae research. Unfortunately for aroid research, Ray has embarked on another field of learning. His high intellect and imagination will be sorely missed in Araceae research.

Hiromichi Yoshino from Kyoto University in Japan explored the Himalayas of Nepal and Bhutan (and wrote two general interest books on the subject) before he began his work with Araceae at the Kihara Institute for Biological Research in 1975, continuing his studies after he moved to Okayama University, publishing molecular studies on Colocasia and Alocasia in Japan and China (Yoshino, 1975, 1994). Another paper describes the morphological characteristics of the wild species of Colocasia (Yoshino, 1984). His Ph.D. dissertation dealt with a phylogenetic differentiation in taro, Colocasia esculenta (Yoshino, 1995). A recent paper deals with phosphate and nitrate absorption ability of wild species of taro (Yoshino, 1995).

Michael H. Grayum, while still a student at the University of Massachusetts, spent a summer in Costa Rica working at La Selva. His first paper dealing with Araceae described the characteristics of that flora (Grayum, 1982). His Ph.D. thesis, written at the University of Massachusetts, involved an SEM study of pollen, but in characteristic thoroughness, he studied every other known character state as well, and on this basis he developed a new system of classification of the genera of Araceae (Grayum, 1984). The system, quite at odds with that of Engler, has for the most part proven over time to better fit our modern state of knowledge of the family. Other papers dealing directly with the information assembled for his thesis include: one on the evolution and ecological significance of starch in pollen of Araceae (Grayum, 1985), the phylogenetic implications of pollen nuclear number in the Araceae (Grayum, 1986a, 1986b), correlation between pollination biology and pollen morphology with some implications for angiosperm evolution (Grayum, 1986b), and the systematic embryology of the Araceae (Grayum, 1991). The bulk of his thesis was published under the titles "Evolution and Phylogeny of the Araceae" (Grayum, 1990) and "Comparative External Pollen Ultrastructure of the Araceae and Putatively Related Taxa" (Grayum, 1992a). Another major contribution was his investigations supporting the removal of Acorus from the Araceae, listing 13 unique characteristics of Acorus not shared with other Araceae (Grayum, 1987a).

Grayum's principal work with Araceae, since his thesis, has involved Costa Rica where he lived for seven years (Grayum, in press), and with a revision of Philodendron subg. Pteromischum (Grayum, 1996). Some new species of P. subg. Pteromischum have been published (Grayum, 1992b, 1996) as well as a new Anthurium (Grayum, 1993). He has also had an interest in Caladium and Chlorospatha (Grayum, 1987b, 1991b). Grayum now serves as Editor of the Annals of the Missouri Botanical Garden and co-investigator of the Costa Rican Manual project. His latest efforts in this project involved a revision of the Araceae of Costa Rica. This has resulted in his current paper (Grayum, in press).

Marianne Knecht, from Switzerland, did a biosytematic study of the Araceae of Côte d'Ivoire (Ivory Coast) in tropical West Africa. This was a thorough study of every aspect of each species occurring there from morphology to phytogeography, cytology, anatomy, palynology and flowering behavior. The results are published in a book in French (Knecht, 1983). Another paper deals with African traditional medicine (Knecht, 1980).

M. Sivadasan (Das), from the University of Calicut in Kerala, India, has worked with the Araceae extensively since the mid-1970s. His unpublished doctoral thesis was a taxonomic study of the Araceae of South India (Sivadasan, 1982). His first published paper described a new species of Typhonium (Sivadasan & Nicolson, 1981), and his first major production was a revision of Theriophonum (Sivadasan & Nicolson, 1982). Other publications include popular articles published on rare Indian aquatic plants, including Cryptocoryne (Sivadasan, 1985a, 1989b) and Lagenandra (Sivadasan, 1990; Sivadasan & Babu, 1995; Sivadasan & Bogner, 1986), and he co-produced the Araceae for the flora of Tamilnadu Carnatic (Sivadasan & Nicolson, 1983). Sivadasan (1983) also wrote on threatened species of Indian aroids as well as the description of new species or new names of Arisaema (Sivadasan, 1985b; Sivadasan & Kumar, 1987; Sivadasan & Nicolson, 1981, 1983a), Lasia (Sivadasan & Sajeev, 1996), Amorphophallus (Sivadasan, 1986a, 1986b, 1989a), Pothos (Sivadasan et al., 1989; 1994), and Theriophonum (Sivadasan & Wilson, 1997, in press). Another paper discusses the pollination biology of Amorphophallus (Sivadasan & Sabu, 1989). In collaboration with C. R. Suresh and K. S. Manilal, he discussed the taxonomy of aroid species in H. A. van Rheede tot Draakestein's Hortus Indicus Malabaricus. A recent addition to the study of the Indian flora is his study of the Araceae of the Silent Valley area (Sivadasan, 1999). This treats 9 genera and 21 species with 17 color figures.

Peter Boyce's interest in Araceae began about 1980. He was involved in the cultivation of aroids and exploration, made a collecting trip to Crete as early as 1986 (Boyce, 1986). In his official capacity at the Royal Botanic Gardens at Kew, he began his work with Araceae as Simon Mayo's assistant. He properly chose to work in a different part of the world than Mayo, initially doing revisionary work with the Mediterranean genera, especially Arum, about which he has published a book (Boyce, 1993a). This work, complete with color paintings, covers all aspects of the taxonomy and biology of the genus. A similar book, this one dealing with Biarum, is to be published. Other publications on Mediterranean plants deal with Arisarum (Boyce, 1989, 1990), Biarum (Boyce, 1987b, 1995h, 1999; Boyce & Athanasiou, 1991), Arum (Boyce, 1987a, 1988, 1989, 1994a, 1995j), and include descriptions of new species. Boyce has also published a treatment of both Dracunculus and Helicodiceros (Boyce, 1994b).

As noted above, Boyce is a member of the team working on the Araceae for the Flora Malesiana and has co-authored a checklist and bibliography of the region (Hay et al., 1995, 1995a) and written about collecting in Peninsular Malaysia (Boyce, 1994). He is responsible for Epipremnum (Boyce, 1998), Pothos (Boyce, 2000), Rhaphidophora (Boyce, 1999), and Scindapsus. He is also doing the Araceae for the Flora of Brunei (Boyce, 1994c, 1997). In addition, he is coordinator of the Araceae project for the Flora of Thailand, contributing the accounts for the Pothoideae and Monsteroideae. Most of his recent publications deal with the biogeography (Boyce, 1996c), and architecture and growth patterns of Pothos (Boyce & Poulsen, 1994) and P. grandis (Boyce & Nguyen, 1995, 1996), and miscellaneous papers as well as other genera from southeast Asia including Amydrium (Boyce, 1995a), Bucephalandra (Boyce, 1995b; Boyce et al., 1995a), Eminium (Lobin & Boyce, 1991), Hapaline (Boyce, 1996), Homalomena (Boyce, 1994b), Pinellia Ten. (Boyce, 1988), Rhapidophora (Boyce, 1996b), Schismatoglottis and Pycnospatha (Boyce, 1993b), Scindapsus (Boyce, 1993c), and Steudnera (Boyce, 1995b). Besides working with Asian genera his studies have also included African genera, namely Culcasia (1995g) and the American genera Anthurium (Boyce, 1995e) and Ulearum (Boyce, 1995f). Boyce was largely responsible for compiling a special issue of Curtis's Botanical Magazine that was devoted entirely to Araceae and included genera from Asia, Africa, and America (Boyce, 1995d-i). Many of his individual contributions to this are cited above but he also wrote for this work an introduction to the family Araceae (Boyce, 1995d), Araceae at the herbarium of Kew (Boyce, 1996b), and an article on aroid conservation (Boyce, 1995i) for the special issue. Boyce co-authored an article on aroid cultivation with P. Brewster and R. Wilford (Boyce et al., 1995) and along with Mayo and Bogner he contributed an article on the history of Araceae research (Mayo et al., 1995). Boyce (1996d) also contributed Arisarum, Arum, Biarum, and Pothos to The World of Plants series. Finally, Boyce is a member of a team (including Mayo and Bogner) who are working on a new understanding of the family at both the supra-family level (Mayo et al., 1995b) and at the suprageneric level (Mayo et al., in press). His intimate knowledge of the Asian Araceae is critical to that effort. Along with Mayo and Bogner, he is an author of The Genera of Araceae (Mayo et al., 1997) and he lectured on this subject at the VI International Aroid Conference in Kunming, China (Mayo et al., 1998).

Miklos Treiber, working at the University of North Carolina at Chapel Hill, did his Ph.D. thesis on a biosystematic study of the Arisaema triphyllum complex (Treiber, 1980).

Collette Ntépé-Nyame of the University of Cameroun has described a new species of Rhektophyllum (R. camerunense Ntépé-Nyame [now Cercestis] (Ntépé, 1981). She also wrote a paper, with R. Letouzey in Paris, on the nomenclatural and taxonomic problems with Culcasia scandens (Letouzey & Ntépé, 1981). Her most recent contribution is the treatment of the Araceae for the Flore du Cameroun (Ntépé-Nyame, 1988). The treatment, written in French, has 56 plates each with one or more detailed line drawings.

Sue Thompson of the Carnegie Museum published her first paper on the distribution and ecology of Cyrtosperma chamissonis Merr. (Thompson, 1982). She has begun a revision of Xanthosoma (Thompson, 1984, 1985, 1989) and did her Ph.D. thesis on the systematics and biology of Araceae and Acoraceae of temperate North America (Thompson, 1990). She contributed the treatment of the Araceae for the Flora of North America (Thompson, 2000) and another paper detailing the biology of North American Araceae (Thompson, in prep). Thompson is a member of the Honorary Board of Directors for the International Aroid Society and has also been editor of the I. A. S. Newsletter.

Chinese botanists, Wen-yen Lien and Ru-zhi Feng, published a survey of Arisaema and Pinellia in China (Lien & Feng, 1982). Z. Xie et al. (1996) reported on morphological variation within P. ternata populations, and Z.-L. Wang introduced a new species of Sauromatum from Gaoligong Mountains (Z.-L. Wang & H. Li, 1999).

Denis Barabé, at the Montreal Botanical Garden and working with different collaborators, made extensive studies of the floral anatomy (Barabé, 1982; Barabé & Chrétien, 1985, 1986; Barabé & Forget, 1988a-b, 1992; Barabé & Labrecque, 1983, 1984, 1985; Barabé et al., 1984, 1985, 1986a, 1987a) and development (Barabé, 1993, 1994, 1995; Barabé & Bertrand, 1996; Barabé & Jean, 1996; Boubes & Barabé, 1996, 1997; Barabé et al., 1986b, 1987b, 1996). These studies include a discussion of neotany in the Araceae (Barabé, 1987) and a cladistic analysis of the Calloideae (Barabé & Forget, 1987a).

The late Mark Moffler, former editor of Aroideana and student of Homalomena, was working on a revision of the genus Homalomena at the time of his death. His published papers deal with the anatomy of Homalomena (Moffler, 1983), a description of a new species with Josef Bogner (Moffler & Bogner, 1984), the cold tolerance of Araceae (Moffler, 1980), and on Anthurium araliifolium (Moffler, 1981). His partially finished thesis on the genus is being revised by Richard Wunderlin, Tom Croat, and Richard Mansell; they are making final revisions on the manuscript before publishing it (Moffler et al., in prep.).

Farah D. Ghani from Malaysia published a paper on useful Araceae of Peninsular Asia (Ghani, 1983) and one on edible aroids (Ghani, 1984a) that includes keys for the identification of cultivars of Colocasia esculenta in Malaysia (Ghani, 1984b).

George R. Haager, currently Director of the Prague Botanical Garden in the Czech Republic has had a long interest in Araceae, collecting in Mexico, Venezuela and Ecuador. He was responsible for collecting Anthurium sarukhanianum in Mexico (Croat, 1991a). He has also described other new species from Ecuador (Haager & Jenik, 1984) and Mexico (Haager, 1991).

W. Greuter (Greuter, 1984) produced a revision of Arum for the island of Crete in the Mediterranean.

Arie de Graaf, sometimes with J. C. Arends and J. D. Bastmeijer, has published papers on Cryptocoryne. Graaf & Arends published on the occurrence of Cryptocoryne and Lagenandra in Sri Lanka (Graaf, 1987a, 1987b, 1988, 1991; Graaf & Arends, 1986). J. D. Bastmeijer published a series of short papers, each featuring a species of Cryptocoryne (Bastmeijer, 1982, 1984, 1986a, 1989, 1991, 1992, 1993; Bastmeijer & Leenen, 1983; Bastmeijer & Kettner, 1991; Bastmeijer et al., 1984) and along with Arends published on the chromosomes of Lagenandra (Arends & Bastmeijer, 1978). He published biographic notes about de Wit and Jacobsen (Bastmeijer, 1986b). Finally Arends and F. M. van der Laan published a paper on the chromosomes of Lagenandra (Arends & van der Laan, 1978).

Richard Keating from Southern Illinois University has spent part of the past 15 years working on the vegetative anatomy of the Araceae for Metcalfe & Chalk's Anatomy of Monotocotyledons (Keating, in prep.). He has presented papers at major meetings that dealt with the anatomical distinction between the Pothoideae and Monsteroideae (Keating, 1980) and with other relationships within the family (Keating, 1982). Other general papers entitled "Techniques for studying aroid anatomy", "Vegetative anatomical features in the Araceae," and "Structural trends in lamina histology in the Araceae" are expected to be published in upcoming issues of Aroideana.

William N. Carvel has completed an as yet unpublished thesis dealing with the floral anatomy of the Pothoideae and Monsteroideae (Carvell, 1989a, 1989b). Another work worthy of mention is a completed revision of Stenospermation for Central America by Alcira Pérez de Gómez (1983), a Venezuelan student from Barquisimeto, working under the direction of Tom Croat.

Kerim Alpinar, at the Department of Pharmacy of the University of Istanbul in Turkey and has published a key (with illustrations) to the Turkish species and recognized new taxa as well as reported on the starch and protein content of the Turkish species (Alpinar, 1985). In addition, he has published chromosome information on Arum in Turkey (Alpinar, 1987) as well as phytochemistry of Dracunculus vulgaris Schott (Alpinar & Meridi, 1987).

Surawit Wannakrairoj at the Department of Horticulture at Kasetsart University in Bangkok, Thailand, a former student of H. Kamemoto at the University of Hawaii, works on a breeding program with Aglaonema. His thesis involved studies of Anthurium spathes and the inheritance of color, a feature so important to the cut flower industry (Marutani et al., 1988; Wannakrairoj & Kamemoto, 1990a, 1990b).

Long Chun-lin from the Kunming Institute of Botany in China published his first papers on Amorphophallus in China in collaboration with Li Heng (Li & Long, 1989a, 1989b). Working with Li Heng, Gu Zhijian and Liu Xianzhang, he produced a cytogeographic study of Remusatia (Long et al., 1989) and a study of the karyotypes of Amorphophallus from China. A recent paper dealt with ethnobotanical uses of Amorphophallus (Long, 1992, 1998). He presented a paper on Amorphophallus of China at the VII International Aroid Conference (Long et al., in press). He has been involved with many papers with Li Heng, including the recent study of aroids of the Gaoligong Mountains (Li et al., 1999) and a new species of Amorphophallus (Long & Li, 2000). See also papers discussed under Li Heng.

Greg Ruckert, founder of the Australian Area collection and the journal Area which features information about tuberous aroids, especially Arisaema, has published articles on Arisaema and recently participated in the work on the Gaoligong Mountains (Li et al., 1999; Li & Ruckert, 1998).

Z. Y. Zhu (1985) redescribed Alocasia cucullata (Lour.) G. Don in Sichuan, China as the "new" genus Panzhuyuia with a single species, P. omeiensis.

P. J. Matthews published several papers on taro, including their cultivation (Matthews, 1987), the origins, dispersal and domestication (Matthews, 1990, 1995; Matthews et al., 1992), and on ribosomal and mitochondria DNA variation (Matthews et al., 1992). S. Chandra (1984) published a work on the taro and other edible aroids.

Yashica Singh, from the National Botanic Institute in Durban, South Africa in collaboration with A. E. van Wyk and H. Baijnath, has published a guide to identifying members of Zantedeschia (Singh et al., 1995). They have also worked on the floral biology of Z. aethiopica (Singh et al., 1996a) and on taxonomic notes of the genus (Singh et al., 1996b). These publications form part of an M.Sc. study on the systematics of Zantedeschia (Singh, 1996) undertaken by Singh at the University of Pretoria.

Jenn-Che Wang from Taiwan Normal University in Taipei, Taiwan has completed a systematic revision of Taiwanese Arisaema (Wang, 1996). The work contains detailed drawings, photographs of spathe and spadix, and pollen micrographs.



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.