Cryptocoryne Excerpts from:
HISTORY AND CURRENT STATUS OF SYSTEMATIC RESEARCH WITH ARACEAE

http://www.aroidsociety.org/literature/croat/croat_araceae_history04.pdf
Thomas B. Croat
Missouri Botanical Garden


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At Kew, the role of describing Araceae was first played by William J. Hooker and his son Joseph D. Hooker (J. Hooker, 1883, 1904), the first two directors, and later by N. E. Brown (Brown, 1901) and to a lesser extent by M. T. Masters (1873, 1876, 1878, 1884, 1893a, 1893b, 1898). The elder Hooker played only a minor role dealing directly with Araceae but did publish a number of short papers in Curtis's Botanical Magazine. These dealt with Alocasia (W. Hooker, 1860b, 1863), Amorphophallus (W. Hooker, 1860a), Arum (W. Hooker, 1828), and Pistia (W. Hooker, 1851). J. D. Hooker dealt more extensively with Araceae. He was responsible for the treatment of the Araceae in Bentham & Hooker's Genera Plantarum (J. Hooker, 1883c), a treatment largely based on that of Schott. He also described many species of Araceae as well as the genus Gonatopus. Many of his new species were described in his Flora of British India (J. Hooker, 1893). J. D. Hooker also treated many species in great detail in Curtis's Botanical Magazine, including Aglaonema (J. Hooker, 1865b), Alocasia (J. Hooker, 1865a, 1896), Amorphophallus (J. Hooker, 1888, 1891a, 1893b), Arisaema (J. Hooker, 1890a, 1890b, 1891b), Colocasia (J. Hooker, 1894), Cryptocoryne (J. Hooker, 1900), Culcasia (J. Hooker, 1869 [described as Aglaonema]), Hapaline (J. Hooker, 1893a), Lysichiton (J. Hooker, 1904), Piptospatha (J. Hooker, 1881b, 1895), Schismatoglottis (J. Hooker, 1881a), and Typhonium (J. Hooker, 1875).


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The French botanist, Samuel Buchet, published several papers dealing with the systematics of the Araceae between 1920 and 1939 (Buchet, 1939a, 1939b, 1942; Buchet & Guillaumin, 1939). He published new plant species descriptions from Asia and especially from Madagascar. Another Frenchman, H. Jumelle, also worked on the plants of Madagascar (Jumelle, 1919, 1928). Still another French botanist, A. Chevalier, published a few papers on Araceae during the same era. These dealt with aquarium plants, Cryptocoryne (Chevalier, 1934a, 1934b), and Cercestis in West Africa (Chevalier, 1920). In the late 1940s and early 1950s the Indian botanist, D. Chatterjee, published new species of Arisaema from Burma, India, and Sikkim (Chatterjee, 1949, 1955).


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RENEWED REVISIONARY EFFORTS AFTER 1950

In addition to the floristic efforts that began in the early 1950’s, considerable new revisionary activity began 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 coauthored with T. Horeman (Rataj & Horeman, 1977) as well as a paper dealing with Typhonium flagelliforme (Rataj, 1982).


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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; Bogner, 2003); 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, 2004; Bogner & Jacobsen, 1985, 1986, Ehrenberg & Bogner, 1992); Jacobsen, 2002; Jacobsen, Bastmeijer & Sasaki, 2002; Sasaki, 2002; 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; Bogner & Hay, 2000); 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), Gorgonidium (Bogner & Nicolson, 1988, Gorgonidium and Anthurium (Bogner & Gonçalves, 2002) and in Turkey with Biarum (Bogner & Boyce, 1989). With James French he described the tribe Anadendreae (Bogner & French, 1984). A recent paper (Bogner, 1997) 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 56 new species. Josef Bogner has 6 specific epithets of Araceae named in his honor, and one genus, Bognera. He has been author or co-author of 2 subfamilies, one subtribe, two tribes, and 94 species of Araceae.

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) and, along with Michael Hesse and other collaborators, has also published a review of the palynology of the perigonate members of the Aroideae (Hesse et al., 2001). 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. In 1999 at the VIII International Aroid Conference in St. Louis he was awarded the first Bette Waterbury Award for Excellence in Horticulture.


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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), C. cordata (Jacobsen, 2002) 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 an 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). A paper in German in Aqua-Planta deals with the Cryptocorye complex and includes two new combinations (Jacobsen, 2002) while another in the same issue coauthored with J. D. Bastmeijer and Y. Sasaki describes a new hybrid from Kalimantan (Jacobsen et al. 2002). 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).


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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. Currently, Sivadasan has an article in press with the journal Aroideana concerning flowering phenology and beetle pollination in Theriophonum infaustum N. E. Br.


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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, 2002; 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).


MISCELLANEOUS PAPERS MOSTLY DEALING WITH SPECIFIC GENERA

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In any review of this kind there are always some research publications or individual horticultural treatments, that though individually worthy of comment, are not cited in any of the above commentary. Generally these are contributions by persons not considered to have played an important role in the history of aroid systematics but whose contributions may be of special interest to those seeking information about a particular genus. These publications will be cited here and will be referenced by genus in alphabetical order.

Cryptocoryne: (Bastmeijer, 2000; Benl, 1960; Bouwmeester & Hoogendoorn, 1966; Davis & Kane, 1995; Dötsch, 1985, 1986, 2001; de Graf, 1980; Eichner, 2001; Hendriks, 1981; Hertel, 1985; Hertel & Mühlberg, 1994; Jones et al., 1980; Kane et al., 1990; Kasselmann, 1986; Koorders, 1981a; Korthaus, 1980; Machlin, 1997; Mansor, 1991; Möhlmann, 1989; Mühlberg, 1980 [mostly Cryptocoryne]; Ørgaard & Jacobsen, 1998 [also Lagenandra]; Reumer, 1984; Schulze, 1971a-d; Watts, 1995, Wendt, 1954.











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Richard J. Sexton