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Native and Exotic Species of Terrestrial Earthworms (Oligochaeta)

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Native and Exotic Species of Terrestrial Earthworms (Oligochaeta) Zoological Studies 39(4): 285-294 (2000) Native and Exotic Species of Terrestrial Earthworms (Oligochaeta) in Taiwan with Reference to Northeast Asia Chu-Fa Tsai*, Huei-Ping Shen and Su-Chen Tsai Habitat and Ecosystem Division, Taiwan Endemic Species Research Institute, Chichi, Nantou, Taiwan 552, R.O.C. (Accepted May 15, 2000) Chu-Fa Tsai, Huei-Ping Shen and Su-Chen Tsai (2000) Native and exotic species of terrestrial earthworms (Oligochaeta) in Taiwan with reference to Northeast Asia. Zoological Studies 39(4): 285-294. Of the 35 nominal species of terrestrial earthworms (Oligochaeta) reported from Taiwan, there are 30 species of the Megascole- cidae, two species of the Lumbricidae, and 1 species each for the Moniligastridae, Glossoscolecidae, and Octochaetidae. The genera Amynthas and Metaphire of the Megascolecidae are dominant and include 27 species, of which 12 species are native. Among these 35 species, the 12 native species occur primarily in hills and mountains of the island, where natural environments (forests) still remain, or in areas relatively less dis- turbed by humans. The other 23 species are exotic, including 4 species with presumed origins from America and Europe, four species from Southeast Asia, and 15 species from southern China. They occur in disturbed en- vironments (urban areas and cultivated lands) on the plains and in some hill and mountainous areas. Endemism based on the nominal species in Taiwan is 34.3%, which is much lower than 74.2% for Hainan Island, 78.1% for China, 65.0% for Japan, 60.0% for Korea, and 37.0% for the Ryukyus. The native species of Amynthas and Metaphire in Taiwan are apparently offshoots of the oligochaete fauna of southern China, resulting from disper- sion differentiation. Key words: Endemism, Distribution, Biogeography. Earthworms have feeble powers of travel and 1996), but more commonly characterized as an are easily isolated geographically. However, many anthropochorous species, defined as a species species of earthworms disperse widely with assis- transported by humans, usually unintentionally tance from nature, such as on tree trunks floating (Gates 1972). downstream and in earth between the hooves of In this study we reviewed the literature to deter- animals (Stephenson 1929), and/or as the result of mine the native or exotic status of the terrestrial human activities, such as in earth on roots of intro- earthworms in Taiwan, and their endemism and geo- duced plants for horticulture, agriculture, and botani- graphic affinities, as compared to those in other geo- cal gardens (Beddard 1912, Gates 1972, Lee 1985). graphical areas of Northeast Asia, including main- Based on the distribution and dispersion mecha- land China, Hainan Island, the Ryukyu Islands and nism, terrestrial earthworms in a geographical area other Japanese islands, the Korean Peninsula, and may consist of both native and exotic species. A Manchuria. native species is generally a species that naturally occurs in a restricted geographical area. It is cer- tainly speciated locally and is thus autochthonous RESEARCH IN TAIWAN (= endemic, indigenous) (Gates 1972). In contrast, an exotic species is a foreign species introduced to a Investigation of the earthworm fauna in Taiwan particular geographical area. It is equivalent to a began at the end of the 19th century, when Goto and peregrine species, that is a species of wide distribu- Hatai (1898) described new and imperfectly known tion in Michaelsens term (Edwards and Bohlen species of earthworms in the Japanese Empire. In *To whom correspondence and reprint requests should be addressed. Tel: 886-4-9761-331 ext. 256. Fax: 886-4-9764-600. 285 286 Zoological Studies 39(4): 285-294 (2000) this report they described Pheretima takatorii and P. the plains of northern and western Taiwan. The ex- candida from Taipei. Michaelsen (1922) examined ceptions were Michaelsen (1922) whose materials the specimens of earthworms collected by Hans were collected from a peripheral hill of southwestern Sauter from the locality presently called Chiahsien in Taiwan, and Chen and Shih (1996) who surveyed a Kaohsiung. He described a new species, P. for- botanical garden in the mountains of northeastern mosae, and also reported a nominal species, P. Taiwan. papulosa Rosa. Kobayashi (1938a,b,d) made a gen- Since 1998 we have conducted an earthworm eral survey in Hsinchu and added 10 nominal survey for the island with particular attention to hill species: Drawida japonica Michaelsen, Bimastos and mountainous areas. We have described 7 new parvus Eisen, P. californica Kinberg, P. hawayana species: Amynthas tungpuensis and A. sexpectatus (Rosa) (= Amynthas gracilis (Kinberg)), P. morrisi from Nantou County, A. tayalis from Taipei Co., A. (Beddard), P. rockefelleri Chen (= A. papulosa binoculatus from Taichung Co., Metaphire puyuma (Rosa)), P. diffringens (Baird) (= A. corticus from Taitung Co. (Tsai et al. 1999), M. paiwanna from (Kinberg)), P. hupeiensis (Michaelsen), Perionyx Pingtung and Hualien Cos., and M. bununa from excavatus Perrier, and Dichogaster bolaui Nantou and Taitung Cos. (Tsai et al. 2000b). We (Michaelsen). Kobayashi (1941b) added P. zoysiae also added Pontoscolex corethrurus (Müller) (Tsai et Chen (= A. minimus (Horst)). al. 2000a) and Metaphire hesperidum (Beddard) as After World War II, Gates (1959) examined the new exotic species to the fauna of Taiwan. We found specimens collected by D. E. Beck on the plains of that P. yushi described from Taipei (Tsai 1964) is a Taiwan, including the Taipei City; Chaochow, Ping- junior synonym of A. formosae described by Michael- tung County; and Suao, Ilan Co. He listed 10 nomi- sen (1922). A. lautus (Ude) and A. robustus (Perrier) nal species of Pheretima earthworms, including P. are considered as 2 valid species (Tsai 1964), how- bicincta (Perrier), P. elongata (Perrier), P. incongrua ever, Easton (1980) considered the former to be a Chen, and P. posthuma (Vaillant) as new records. A junior synonym of the latter. few years later Tsai (1964) collected earthworms in To date, there are 35 nominal species of terres- the Taipei area, and recorded the 3 nominal species, trial earthworms in Taiwan, of which 12 species are P. schmardae (Horst), P. lauta (Ude), and P. robusta native (Table 1). They are classified according to the (Perrier), and described 4 new species, namely P. Raynolds and Cook System (Edwards and Bohlen yushi, P. swanus, P. taipeiensis, and P. polyglandu- 1996) for families, and according to Sims and Easton laris. (1972) and Easton (1979) for the genera of the Sims and Easton (1972) considered Amynthas Pheretima species complex of the Megascolecidae. polyglandularis of Taiwan (Tsai 1964) to be a sub- species of A. omeimontis of Mt. Omei in Sichuan, China (Chen 1934), and called it A. omeimontis LIST OF TAIWAN EARTHWORMS polyglandularis. They also included 2 additional subspecies, A. omeimontis kinfumontis of Mt. Kinfu Order Moniligastrida in Sichuan (Chen 1946) and A. omeimontis kinabalu Family Moniligastridae of North Borneo (Sims and Easton 1972). However, 1. Drawida japonica Michaelsen based on figure 3 of Sims and Easton (1972), their Order Haplotaxida locations and patterns of genital papillae differ so Suborder Lumbricina greatly that specific distinction is possibly permis- Superfamily Lumbricoidea sible. Therefore, in this study we retain A. poly- Family Lumbricidae glandularis as a valid species. 2. Allolobophora trapezoides Duges Recently, Kuo (1995) described a new species, 3. Bimastos parvus Eisen P. hsinpuensis, from Hsinchu, Taiwan. Chen and Superfamily Glossoscolecoidea Shih (1996) made an earthworm survey in the Family Glossoscolecidae Fushan Botanical Garden (elevation 1000 m) in Ilan 4. Pontoscolex corethrurus (Müller) of northeastern Taiwan. They reported 6 nominal Superfamily Megascolecidea species: P. taipeiensis, P. diffringens, P. hawayana, Family Megascolecidae P. formosae, P. rockefelleri, and P. polyglandularis. 5. Amynthas aspergillus (Perrier) Shih et al. (1999) reviewed the literature on Taiwans 6. Amynthas binoculatus Tsai, Shen and earthworms and listed 26 nominal species. Tsai For the past 100 yr, faunal surveys of earth- 7. Amynthas candidus (Goto and Hatai) worms in Taiwan have been conducted primarily on 8. Amynthas corticus (Kinberg) Tsai et al. − Endemism of Taiwans Earthworms 287 9. Amynthas formosae (Michaelsen) 20. Amynthas swanus (Tsai) 10. Amynthas gracilis (Kinberg) 21. Amynthas taipeiensis (Tsai) 11. Amynthas hsinpuensis (Kuo) 22. Amynthas tungpuensis Tsai, Shen and 12. Amynthas hupeiensis (Michaelsen) Tsai 13. Amynthas incongruus (Chen) 23. Amynthas tayalis Tsai, Shen and Tsai 14. Amynthas lautus (Ude) 24. Amynthas minimus (Horst) 15. Amynthas morrisi (Beddard) 25. Metaphire bununa Tsai, Tsai and Liaw 16. Amynthas polyglandularis (Tsai) 26. Metaphire californica (Kinberg) 17. Amynthas papulosus (Rosa) 27. Metaphire hesperidum (Beddard) 18. Amynthas robustus (Perrier) 28. Metaphire paiwanna Tsai, Tsai and Liaw 19. Amynthas sexpectatus Tsai, Shen and 39. Metaphire posthuma (Vaillant) Tsai 30. Metaphire puyuma Tsai, Shen and Tsai Table 1. Terrestrial earthworms reported from Taiwan and their occurrence in China, the Ryukyus, Japan, Korea, and Southeast (SE) Asia Species Taiwan China Ryukyus Japan Korea SE Asia Native species Amynthas candidus (Goto and Hatai) + Amynthas formosae (Michaelsen) + Amynthas swanus (Tsai) + Amynthas polyglandularis (Tsai) + Amynthas hsinpuensis (kuo) + Amynthas tungpuensis Tsai, Shen and Tsai + Amynthas tayalis
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