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.

Regional Studies with Araceae


Among the earliest who described and often illustrated Araceae in Asia were the Dutchmen, H. A. van Rheede tot Drakestein (1688, 1692), who did massive studies of the Malabar coast of India; P. Hermann (1689), who produced the Paradisus Batavus; and G. E. Rumphius (1747), who studied the flora of the island of Amboina [one of the Mollucas Islands, now Maluku in Indonesia]. Another Dutch botanist, N. L. Burmann (1768), published Araceae in his Flora Indica, and C. L. Blume dealt extensively with Malesian Araceae (Blume, 1836-1837). Another early botanist who described and illustrated Araceae in Malesia was the Italian, Odoardo Beccari (Beccari, 1879, 1882, 1889; Engler, 1879b).

Many of the species in Asia were described by regional workers doing floristic studies on particular regions. Except for Japan the earliest workers with aroids in Asia were generally not natives but were primarily botanists from colonial powers working in their own sphere of influence. In the Dutch East Indies it was primarily the Dutch. Other botanists who were describing Araceae from the Dutch East Indies included the Britian, R. Brown (1810), the German, J. C. Hasskarl (Hasskarl, 1842), and Dutch botanists, J. E. Teijsmann (Teijsmann & Binnendijk, 1862), F. A. W. Miquel (Miquel, 1855-1856, 1856, 1860, 1864, 1867), Hans Hallier (1898, 1901, 1915), C. R. W. K. van Alderwerelt van Rosenburgh, thankfully abbreviated "Alderw." (Alderwerelt van Rosenburgh, 1920, 1922a, 1922b), and C. A. Backer (Backer, 1913a, 1913b, 1913c, 1914, 1920, 1928).

In Indochina, it was a Portuguese naturalist Joao Loureiro who published his Flora Cochinchinensis (1790) and the Frenchman, F. R. Gagnepain, who wrote the Araceae treatment for Lecomte's "Flore général de l'Indochine" (Gagnepain, 1942a) and other works describing new genera and species (Gagnepain, 1941a-c).

In the Malay Peninsula it was H. N. Ridley, Director of the Singapore Botanical Garden from 1888 to 1912 (Ridley, 1908, 1916, 1922, 1938), and C. X. Furtado (Furtado, 1930, 1935, 1939, 1941, 1958, 1964a, 1964b), also from Singapore who described Araceae in the area.

In the Philippines it was the Spaniard, F. M. Blanco (1837) and Americans, E. D. Merrill (1912, 1915, 1916a, 1916b, 1917, 1918, 1919, 1921b, 1923, 1924, 1925, 1926, 1932, 1935b, 1937, 1948, 1949, 1952) and A. D. E. Elmer (1919, 1938, 1939). Merrill also described species from Guam (Merrill, 1914), Borneo (Merrill, 1921a, 1922a, 1929); Sarawak (Merrill, 1922b, 1928, 1934a) and Hainan in China (Merrill, 1927, 1930a, 1930b; Merrill & Metcalf, 1945), Sumatra (Merrill, 1933) and Vietnam (Merrill, 1942).

In Malesia, much of the recent work with the flora was organized by C. G. G. J. van Steenis who began the Flora Malesiana project as a resident in Asia for much of his lifetime. Most of his studies were carried out on Java (van Steenis, 1948a, 1948c, 1949, 1965a, 1965b, 1972, 1975). Despite the fact that some of the earlier European workers, including Ridley and van Steenis, spent considerable portions of their lives in Asia the majority of the Asian species were described in the European centers of botany, in Paris, Brussels, Leiden, Berlin, and Kew.

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).

Many other floristic studies, both regional and general, throughout many years have contributed to the generally high levels of knowledge about Asian plants. Owing to the early English involvement, many of the early floristic studies were made on the Indian subcontinent, including those in W. Roxburgh's Flora Indica (Roxburgh, 1820b, 1832) and in Hortus Bengalensis (Roxburgh, 1814) which cataloged the holdings of the East India Company's living collection in Calcutta. Others early works include those by the German A. W. Roth (1821) and the Dane N. Wallich (1830, 1831). Still other general works on India include those by Burkill (1925), Suresh et al. (1983) and Zhongguo (1994). Works dealing with specific parts of India include that for the Coromandel Coast [SE India] (Roxburgh, 1820a); Assam, South India (Rao & Verma, 1968, 1976; Barnes & Fischer, 1936a); the Calicut area (western sectors of Calicut and Malappuram Districts) (Manilal & Sivarajan, 1982); the Presidency of Madras (Fischer, 1931); the Howrah District (Bennet, 1979); the Presidency of Bombay (Cooke, 1906; Blatter & McCann, 1931); Nainital in Uttar Pradesh (Gupta, 1968), the Bashahr Himalayas (Nair, 1977); Manipur State [NE India] (H. Singh, 1993), the Chikihagular District, Karnataka, India (Bhat, 1993; Yoganarasimhan et al., 1981); as well as of Majuli (Islam, 1990); Punjab (Sharma, 1990), Andaman and Nicobar Islands (Kurz, 1893; Srivastava & Kumar, 1993) and Barren Island (Prain, 1893). C. Fischer published a series of papers during the 1930s dealing with Indian Araceae (Fischer, 1931, 1933, 1934, 1936a, 1936b, 1939) and F. Baius (1936) wrote on the medicinal and poisonous aroids of India. New species of Arisaema (Rao & Srivastava, 1991; Yadav et al. 1993) have been recently described from India (see also Sivadasan below).

Studies of Araceae in non-Indian areas on the Indian subcontinent include those in West Pakistan (Nasir, 1978); Bangladesh (Khan & Halim, 1987); Bhutan (Noltie, 1994); Nepal (Wallich, 1824); Hara, 1978); Himalayas (Polunin & Stainton, 1984) and Eastern Himalaya (Hara, 1966). Araceae studies in areas adjacent to India include those in Ceylon (Thwaites, 1864; Trimen, 1898; Alston, 1931) and Burma (Kurz, 1873).

Elsewhere in Southeast Asia studies were made in Thailand (Ridley, 1911a; Craib, 1912, 1913; Hu, 1968; Suvatti, 1978) and the Malay Peninsula including West Malesia (Jack, 1820; Burkill & Holttum, 1923; Ridley, 1885, 1893, 1902, 1904, 1907a, 1907b, 1909, 1910; 1911a, 1911b, 1912, 1925a; Rendle, 1924-1925; Henderson, 1926, 1927, 1928, 1939, 1954; Merrill, 1952; Chin, 1982; van Steenis-Kruseman, 1963, 1966, 1975); Malaya [Perak] (Hemsley, 1887); Malaya (Johore) & Singapore (Corner, 1978); [Pahang, Gunung Ulu Kali] (Stone, 1981); Singapore (Ridley, 1900). Studies in Indochina were those by Gagnepain (1942a) and by Merrill (1935a); others were in Vietnam (Pham Hoàng Hô, 1960; Thin, 1997). Other studies were made in Borneo (Rendle, 1901; Ridley, 1905, 1913, 1914; Gibbs, 1914; Masamune, 1942); Sabah (Stapf, 1894); and Sarawak and Brunei (Anderson, 1963; Wong, 1990).

Additional works from the Dutch East Indies and vicinity included works by the following: de Vriese (1851); Miquel (1855-1856, 1956a, 1956b, 1860, 1864, 1867); Zollinger (1845, 1854, 1857); Soepadmo (1977); van Steenis (1949); other areas mostly Indonesia (Java, Sumatra, Celebes, Borneo, New Guinea and rarely the Philippines) by Alderwerelt van Rosenburgh (1920, 1922a, 1922b); Java (Thunberg, 1825; Hasskarl, 1842a, 1842b, 1844, 1848; Koorders, 1901a, 1901b, 1911, 1918a, 1918b, 1923; Backer, 1913a-c, 1914, 1920, 1928; Backer & Bakhuizen, 1968; Backer et al., 1950; Bakhuizen v.d. Brink, 1957, 1963); Sumatra (Hotta, 1984; Ridley, 1917, 1923, 1925a, 1925b); Celebes (Koorders, 1898, 1922b; Kawakami, 1912); Tjibodas [Indonesia] (Koorders, 1922); Amboina [Indonesia] (Rumphius, 1747, 1750); Bangka (Kurz, 1864); Talaud Islands [Indonesia] (Holthuis & Lam, 1942) and Mentawi Islands [Indonesia] (Ridley, 1926), and finally Christmas Island [a British island south and west end of Java] (Rendle, 1900; Ridley, 1906).

Works in New Guinea included: (Ridley, 1886; Schumann & Hollrung, 1889; Schumann & Lauterbach, 1905; Engler, 1907, 1911; Engler & Krause, 1911; Rechinger, 1913; Ridley, 1916; Rendle, 1923; Borrell, 1989); Papua New Guinea (Gilli, 1980; Lane-Poole, 1925; Mueller, 1876b; Schumann, 1887; Warburg, 1891; White, 1922; White & Francis, 1927); Dutch New Guinea (Gibbs, 1917); Bismarck Archipelago [Papua New Guinea] (Peekel, 1984) including New Ireland (Lauterbach, 1911) and New Britain (Schumann, 1898), and Kairiru Island [New Guinea] (Borrell, 1989).

In the Philippines works included are those by: Usteri (1905); Brown (1919); Quisumbing & Merrill (1928); Pancho (1959); and Hatusima (1966). [See also papers by E. D. Merrill and A. Elmer cited above].

In northern Asia studies were made in China (Li et al., 1977); southwestern China (Handel-Mazzetti, 1936; Wu & Li, 1979); Hainan Province (Chun et al., 1977), the Dulongjian region (Li, 1993b); Taiwan (Hayata, 1915, 1916, 1919, 1920; Liu & Chen, 1984; Liu & Huang, 1963, 1977; Huang, 1960, 1979, 1982; Masamune, 1943); and Korea (Lee, 1976; Lee, 1985). [See also the contributions by Merrill elsewhere.] In Japan most species of Araceae were described by T. Makino (1892, 1893, 1901a, 1901b, 1910a, 1910b, 1911, 1913, 1918a, 1918b, 1928, 1931, 1932, 1960, 1961) and T. Nakai (1917, 1918, 1928, 1929, 1931, 1934, 1935a-d, 1937a, 1937b, 1938, 1939a-c, 1940a-d, 1943). Other works from Japan included those by M. Honda (1939), S. Kitamura et al. (1977); G. Koidzumi (1928), T. Koyama (1965), J. Ohwi (1953, 1965), and J. Ohashi (1982). Another dealt with Okinawa and the southern Ryukyu Islands (Walker et al., 1976).

Floristic works and miscellaneous papers on Araceae in Australia include: general areas (Brown, 1810; Mueller, 1858, 1874, 1876a, Bentham, 1878; Maiden, 1889, 1905; Domin, 1911, 1915; Green et al., 1994; Jones et al., 1977; Pate & Dixon, 1982; Elliot & Jones, 1982, 1984, 1990; Morley, 1983; Jones et al., 1977; Jones & Gray, 1988, Briggs & Leigh, 1988; Hnatiuk, 1990; Hay, 1989, 1992a, 1993a, 1993b, 1995); Queensland (Bailey, 1883, 1891, 1897, 1902, 1913, 1914; Orsino & Dameri, 1992; Williams, 1979); New South Wales (Moore & Betchie, 1893; Dixon, 1908; Evans, 1961, 1962; Hay, 1993c), Lord Howe Island (Oliver, 1916), Sydney region of New South Wales (Beadle et al., 1963, 1982; Beadle, 1987); Carolin & Tindale, (1993); Victoria (Ewart, 1930; Willis, 1962); Central Australia (Jessop, 1981); South Australia (Black, 1909, 1943, 1978, 1986; Eichler, 1965; Jessop, 1986); Western Australia (Gardner, 1931); Kimberley region [Western Australia] (Rye, 1992); Northern Territory (Ewart & Davies, 1917; Blake, 1954; Lazarides et al., 1988); Cocos Islands [north of Australia] (Prain, 1891); and Norfolk Island (Maiden, 1903).

The region of Oceania, though not rich in Araceae, has been well studied from the standpoint of aroid floristics. Among the earliest studies made in the region was that by J. G. A. Forster (1786) who made a floristic study of the so called Australian Islands (Southern Islands, i.e. New Zealand, New Caledonia, and many of the smaller Oceanic islands. Still another early study was that by B. C. Seemann (1868, 1869a, 1869b) on Fiji. Other studies in the Oceanic region included that on the Samoan Islands (Christophersen, 1935; Reinecke, 1898); Rarotonga [Cook Islands] (Wilder, 1931); the southeastern Moluccas (Hemsley, 1885b); Ryukyu Islands (Hatusima, 1962); Micronesian Islands (Koidzumi, 1916; Hosokawa, 1937; Hatusima, 1939; Kanehira, 1933, 1935); Makatea (Wilder, 1934); Guam (Safford, 1905; Stone, 1964); Tonga Islands (Hemsley, 1893; Yuncker, 1959; Hotta, 1963a; Whistler, 1991); Christmas Island (Rendle, 1900; Ridley, 1906); Niue [New Zealand] (Yuncker, 1943; Sykes, 1970); Kermadec [New Zealand] (Sykes, 1977; Parham, 1972), Ponape [Caroline Islands] (Glassman, 1952), New Hebrides (Guillaumin, 1932, 1938), and New Caledonia (Guillaumin, 1937, 1943, 1947, 1948, 1962; Rendle, 1921).

In contrast to much of Asia the Japanese have largely studied their own flora. Local Japanese botanists who made great contributions to the understanding of the Araceae include Tomitaro Makino (1862-1957) and Takenoshin Nakai (1882-1952) from the University of Tokyo and Director of the National Science Museum. Nakai described most species, mostly species of Arisaema and he also published a new system of classification (Nakai, 1943) which excluded a few long-standing genera of Araceae, incorporating them into their own families, Pistiaceae, Cryptocorynaceae, and Acoraceae (only the latter is still excluded from the Araceae). Nakai's students, Fumio Maekawa and Hiroshi Hara, were also very much interested in Araceae, especially Arisaema. Maekawa (1924, 1932, 1934, 1936, 1937) described new species from Japan. Hara described new species of Arisaema (Hara, 1935a, 1935b, 1961, 1965; Hara & Ohashi, 1973) and made critical revisions for the Flora of the Eastern Himalaya (Hara, 1966, 1971a). Later he proposed an infrageneric system of classification for the genus Arisaema (Hara, 1971b), a system since adopted by Wu Cheng Yih and Li Heng (1979) and recently revised by Jin Murata (1984). A later paper described additional species from Himalaya (Hara, 1973).

Another Japanese botanist, Shiro Kitamura, working about the same time as Hara, published several significant papers on Araceae of Japan (Kitamura, 1941, 1949; and compiled Coloured Illustrations of Herbaceous Plants of Japan (Kitamura et al., 1977) that contains line drawings and paintings of Araceae.


Most floristic projects on the continent were initiated by the colonial powers, including Belgium and Germany, and especially Britian and France. In general, the Araceae treatments were completed by botanists who were competent researchers but not formally trained aroid specialists. One such botanist was F. N. Hepper, who contributed the Araceae treatment for the Flora of West Tropical Africa (Hepper, 1968a-c). Hepper's treatment of the Araceae for that flora [a revision of an earlier flora by the same name (Hutchinson & Dalziel, 1936)] is a remarkably good one. His understanding of the genus Culcasia (Hepper, 1965, 1967), probably the most complex genus in Africa, was particularly good. An exception to the rule that most flora writers were non-aroid specialists might be N. E. Brown who prepared the Araceae treatment for the Flora of Tropical Africa (Brown, 1901). Other publications that contributed to the generally high level of knowledge about African flora include some generic studies for the entire continent (Thonner, 1915), South Africa (Dyer, 1976), and Central Africa (Bamps, 1982; Malaisse & Bamps, 1993). Other publications include general floristic studies in the Mascarene Islands (Mayo, 1983b), German East Africa (or Tanganyika and now mostly Tanzania) (Mildbraed, 1936; Peter, 1929), Belgian Congo (Katanga) [until recently Zaire and now Congo] (Wildeman, 1921, 1922), Senegal (Lykke, 1994), as well as Egypt and western Asia (Boisser, 1884) and Ghana (Beath, 1993). Papers focusing mainly on African species include those on Zantedeschia (Letty, 1973; Perry, 1989), Zamioculcas (Obermeyer & Strey, 1969); Stylochaeton (Malaisse & Bamps, 1994); Remusatia (Robyns, 1931), and Gonatopus (Obermeyer 1977; Obermeyer & Bogner, 1979). A recent paper ennumerates species of Araceae in 30 genera (including introduced genera) from tropical West Africa (Lebrun & Stork, 1995). [See also Ntépe-Nyame and Knecht, below.]

Madagascar was a special interest of Samuel Buchet (Bogner, 1980g), a French botanist who described Arisaema and Pothos species from Asia (Buchet, 1911a, 1911b) and studied the Arophyteae (Buchet, 1939a, 1942), the dominant element of the Malagasy flora. Josef Bogner (see below) has had the greatest impact on the study of Araceae in Madagascar. The Seychelles near Madagascar also have a flora and the island is important as having the endemic genus Protarum (Robertson, 1989).


In contrast to Africa and Asia where there were strong botanical interests within the colonial powers, the neotropics had little early exploration and very few floristic projects. Neither Spain nor Portugal made much headway towards the production of floras nor did they even launch major collecting programs. Exceptions were the expeditions of Sesse & Moçino, Ruiz & Pavon, and Triana & Planchon, but they collected few specimens of Araceae.

Regardless of their origin, most 18th and 19th Century botanists collected and described relatively few Araceae, though some are worthy of mention. E. F. Poeppig collected and described a number of important species of Araceae from Peru and Brazil (Poeppig, 1845). Also important was A. F. M. Glaziou and H. W. Schott, both working in Brazil. While Glaziou collected many species described by Schott and others he did not publish any species himself. F. A. W. Miquel collected important Araceae in the Guianas as did R. Spruce in the Amazon basin (though the latter did not publish any new species). Aside from Luis Sodiro who collected Araceae in Ecuador [see Sodiro above] most of the other important collectors of Araceae in the 19th Century did not publish on Araceae. These included F. Lehmann, collecting in the western Andes of Colombia and Ecuador, E. Ule in the upper Amazon basin, as well as August Fendler and H. Pittier collecting in Venezuela.

With the notable exception of J. M. da Conceiçao Vellozo (1742-1811) who published Araceae in his Flora Fluminensis (Vellozo, 1825(1829), 1831a, 1831b) few early Portuguese or Spanish described Araceae [see Stellfeld, 1950 for an account of the work of Vellozo]. With the minor exceptions of M. S. Bertoni (Paraguay), G. M. Barroso, specializing on Araceae at the Rio de Janiero Botanical Garden and P. R. Reitz, also from Brazil and working on the Aráceas Catarinensis (Reitz, 1957), few Araceae were described by native-born Latin Americans. In Latin America as in Asia, the new species were in part described by the flora writers from other countries or by plant explorers such as Europeans N. J. Jacquin, F. A. W. Miquel, and others. The works of Jacquin (1760, 1763, 1772, 1790a, 1790b, 1797) based on his travels in the West Indies are important because he was one of the first to describe Araceae from the New World. His colored paintings in Icones Plantarum Rariorum (Jacquin, 1790a) are particularly noteworthy. John G. Baker described and illustrated a number of Araceae, especially Anthurium in Saunder's Refugium Botanicum (Baker, 1871). S. L. Moore included Araceae in his studies of the plants of Mato Grosso in Brazil (Moore, 1895).

During the 20th Century, American institutions began serious studies in Latin America and authors such as P. C. Standley, who wrote a series of floras in Central America, and J. F. Macbride, who worked on the Flora of Peru, described a number of new species. In addition, small but important collections were described from Colombia by R. E. Schultes during his ethnobotanical studies of the neotropics and L. Diels (1937a, 1937b), working at the Berlin Botanical Garden, described collections made in Ecuador. The Dutch under Pulle also started a flora in Suriname, then a Dutch territory. Floristic studies did not play as great a role in the Americas as they did in Asia and Africa; however, a number were done. For South America these include: Argentina (Crisci, 1971; Crisci & Katinas, 1999; Hauman & Vanderveken, 1917); Buenos Aires Province (Crisci, 1968a); Bolivia (Rusby, 1910, 1927); Brazil (Stellfeld, 1950), Amazonas (Smith, 1939), Bahía (Harley & Mayo, 1980); Rio Grande do Sul State (Rambo, 1950), Rio de Janiero State (Casiri, 1982); Cardoso Island (Olaio & Catharino, 1991); Colombia (Garcia-Barriga, 1974; Forero & Gentry, 1989; Gines et al., 1953; Galeano & Bernal, 1993); Ecuador (Dodson & Gentry, 1978; Dodson et al., 1985); French Guiana (Aublet, 1775; Lemée, 1955; Croat, 1995b); Guyana (British Guiana) (Gleason, 1929; Graham, 1934); Paraguay (Bertoni, 1916; Chodat & Hassler, 1903; Chodat & Vischer, 1919; Croat & Mount, 1988); Peru (Macbride, 1936); Surinam [see Jonker-Verhoef & Jonker below]; Uruguay (Herter, 1943; Marchesi, 1984); and Venezuela (Pittier et al., 1945; Maguire, 1948; Steyermark, 1951; Steyermark & Huber, 1978; Bunting, 1995; Gines et al., 1953).

The earliest effort to produce a flora from Central America was that by William Botting Hemsley (Hemsley, 1885a). Other floristic works done later include those by: L. O. Williams (1981) in Central America; I. Johnston (1949), F. Liebmann (1849), C. L. Lundell (1937, 1939, 1941), R. McVaugh (1993), M. Martínez & E. Matuda (1979), and Espejo Serna & Lopez Ferrari (1993) in Mexico; McVaugh (1993) and Vásquez et al. (1995) in western Mexico; H. H. Bartlett (1937) in Petén Province, Guatemala; T. K. Yuncker (1940) and A. Molina (1975) in Honduras; D. L. Spellman et al., (1975) in Belize; Engler (1900) in Costa Rica, and Paul Standley (see Standley below) in various other countries.

In the West Indies, floristic studies have been numerous and this area was one of the first to be explored by collectors such as C. Plumier, H. Sloane, N. J. Jacquin, and others. In fact, many of the earliest names of New World plants are based on West Indian types. General floristic accounts include: the Dutch Antilles (Arnoldo, 1971; Boldingh, 1913); Jamaica (Adams, 1972; Proctor, 1982); Cayman Islands (Proctor, 1984); Cuba (Grisebach, 1864, 1866; Sauvelle, 1868; Leon, 1946); Haiti (Barker & Dardeau, 1930; Liogier, 1981); Dominican Republic (Moscoso, 1943; Hodge, 1954a); Puerto Rico (Liogier, 1965; Liogier & Martorell, 1982); Puerto Rico and the Virgin Islands (Britton & Wilson, 1923, 1926; Acevedo-Rodríguez, 1966); the Windward and the Leeward Islands (Beard, 1949); Barbados (Gooding et al., 1965); Guadeloupe and Martinique (Heckel, 1897); and Lesser Antilles (Howard, 1979).


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