ハラメノウミミズ

　フトミミズ科 ＞ ハラメノウミミズ属 ＞ ハラメノウミミズ Perionyx excavatus Perrier, 1872

Perionyx excavatus Perrier, 1872: 126; Rosa, 1889: 134, 1890: 2, 1891: 404, 1892a: 2; Beddard, 1891: 275, 1893: 689, 1895: 437; Michaelsen, 1900: 208, 1922: 22; Cognetti, 1906f: 1, 1908e: 309; Stephenson, 1914: 386, 1916: 317, 1917: 375, 1921: 760, 1922: 435, 1923: 329, 1924b: 340, 1926: 258, 1929: 238, 1931c: 275, 1932: 50, 1933: 930; Gates, 1926b: 472, 1926c: 162, 1926d: 182, 1926f: 213, 1927c: 352, 1932a: 546, 1941a: 161, 1955c: 79; Kobayashi, 1938c: 201; 小林, 1938e: 779; Kale & Krishnamoorthy, 1978: 24; Julka, 1981: 21; Nakamura, 1990: 102; Zicsi, 1993: 193; 中村 & Zicsi, 1999: 89; Shih et al., 1999: 438; Reynolds, 2009b: 57, 2015v: 117; Hong et al., 2001: 188; Paliwal & Julka, 2004: 1976; James, 2004c: 11; Dhiman & Battish, 2006: 2136; Huang et al., 2006: 17; Joshi & Dabral, 2008: 83; Tsai et al., 2009: 38; Csuzdi & Pavlíček, 2009: 10; Sinha et al., 2013: 304.

Megascolex perionyx Vaillant, 1889: 69.

Perionyx gruenewaldi Michaelsen, 1890g: 331.

Perionyx koboensis Stephenson, 1914: 389, 1923: 339.

Perionyx turaensis Stephenson, 1920: 216, 1923: 360.

Perionyx fulvus Gates, 1926b: 472, 1926d: 182, 1926f: 213.

＜外部形態＞

　全長 23-180 mm (23-120 [Stephenson, 1923]、30-180 [Julka, 1982]、40-80 [James, 2004c]、78-139 [Hong et al., 2001b]、120 [Vaillant, 1889]、139-153 [Gates, 1960c], 162 [Kobayashi, 1938c])、体幅 2-7 mm (2-5 [Stephenson, 1923]、2.7-3.5 [Hong et al., 2001b]、3-7 [Julka, 1982]、4 [Kobayashi, 1938c])、体節数 75-178 (75-165 [Stephenson, 1923]、118-171 [Hong et al., 2001b]、123-178 [Julka, 1982]、131 [Kobayashi, 1938c])。第一背孔は第 4/5 体節間溝 (Gates, 1932a)、もしくは第 5/6 体節間溝 (Kobayashi, 1938c)、第 4/5 または 5/6 体節間溝 (Stephenson, 1923; Hong et al., 2001b)、第 2/3-5/6 体節間溝のいずれか (Julka, 1982) に開口する。

　体色は背面は紫赤色で、環帯より前は濃紫色で緑色光沢があり、腹面は背面より淡色で (Kobayashi, 1938c)、黄色味を帯びる (Hong et al., 2001b)。環帯は黄灰色で、背面はやや紫色がかる(Kobayashi, 1938c)。

　剛毛は第 7 体節で 40-49 本 (Hong et al., 2001b)、第 20 体節で 41-52 本 (41-46 [Hong et al., 2001b]、46-52 [Julka, 1982]、52 [Kobayashi, 1938c])。剛毛は腹側で密で、背では疎 (Stephenson, 1923)。

　環帯は第 13-17 体節の 5 体節分を占め、全周がつながり、体節間溝や背孔は目立たない。受精囊孔は 2 対で第 7/8/9 体節間溝にあり、0.5 mmほど離れる (Hong et al., 2001b)。受精囊孔間剛毛は 4-6 本 (Kobayashi, 1938c; Julka, 1982)。雄性孔は第 18 体節で近接しており、円形 (Stephenson, 1923) もしくは卵型 (Hong et al., 2001b) に凹んだ中に開口する (Stephenson, 1923)。この凹みの前縁と後縁ははっきりとしているが、側面ははっきりしない (Stephenson, 1923)。雄性孔間剛毛は 0 本 (Hong et al., 2001b)。

＜内部形態＞

　隔膜は、特別に厚いものはなく、全てほぼ同一の厚さ (Stephenson, 1923)。砂嚢は痕跡的で、第 6 体節にあるか、もしくは確認できない (Stephenson, 1923)。石灰腺を欠く (Stephenson, 1923)。食道は第 13 体節で広がる (Stephenson, 1923)。腸管は第 15 体節から開始する (Stephenson, 1923)。

　最終心臓は第 12 体節 (Stephenson, 1923)。

　精巣は第 10、11 体節 (Stephenson, 1923)。貯精嚢は第 11-14 体節 (Stephenson, 1923)。摂護腺は小さく、通常第 18 体節のみ (Stephenson, 1923)。導管は短く直線的 (Stephenson, 1923)。受精囊の主嚢は大きく卵型 (Stephenson, 1923)。導管は短く細い。副嚢は1-4 個で、ごく小さく、いぼ状 (Stephenson, 1923)。

　性的剛毛は 0.60-0.69 mm (Julka, 1982)、太さ 15-25 μ (Julka, 1982)。

＜異常個体＞

　インドでは、第 7/8/9/10 体節間溝に受精囊孔を持ち、第 15 体節に雌性孔、第 19 体節に雄性孔を持つ異常個体が発見されている (Julka, 1981)。また、第 6/7 体節間溝にも追加の受精囊孔を持つ個体、受精囊孔や雌性孔、雄性孔が 1 体節分前にずれた個体、同様に 1 体節分後方にずれた個体が採集されている (Julka, 1981)。また、第 8/9/10 体節間溝に受精嚢孔があり、雌性孔が第 15-16 体節にある個体も報告されている (Gates, 1960c)。ただし、雄性孔は通常通り第 19 体節にあった (Gates, 1960c)。

　日本では、東京都の小学校のコンポストの中から採集されたのみ (中村・Zicsi, 1999)。

　国外では、インド (Stephenson, 1914, 1916, 1917, 1921, 1922b, 1923, 1924b, 1926; Gates, 1960c; Kale & Krishnamoorthy, 1978; Julka, 1981; Paliwal & Julka, 2004; Dhiman & Battish, 2006; Sinha et al., 2013)、スリランカ (Stephenson, 1923)、アンダマン諸島 (Stephenson, 1923; Julka, 1982)、ミャンマー (Rosa, 1890; Stephenson, 1923, 1924b, 1929; Gates, 1926b-d, f, 1927c, 1932a, 1955c; Stephenson, 1933; Reynolds, 2009b)、タイ (Rosa, 1891)、マレーシア (Stephenson, 1931, 1932)、ベトナム (Vaillant, 1889)、台湾 (Kobayashi, 1938c; 小林, 1938e; Shih et al., 1999; Huang et al., 2006; Tsai et al., 2009)、インドネシア (Rosa, 1889; Michaelsen, 1890g, 1922)、フィリピン (Gates, 1960c)、フィジー (Gates, 1960c)、サモア (James, 2004c)、ハワイ (Gates, 1960c; Nakamura, 1990; Reynolds, 2015v)、グアデルーペ (Csuzdi & Pavlíček, 2009)、ドミニカ (Zicsi, 1993)、マダガスカル (Cognetti, 1908e)、レユニオン島 (Cognetti, 1906f) に分布する。

　倒木の下や樹皮の下、腐朽した材の中、水草の葉の中、貯水槽の脇の石の下や泥の中に生息する (Stephenson, 1923)。枯れ葉の積もった樹洞の中にも生息する (Stephenson, 1923)。

　インドでは、バナナの葉の下の腐植の中や、小川の石の下、小川の土手の砂地などから採集されている (Julka, 1981)。

　驚かせた場合、逃げる時にトビムシのようにジャンプする (Hong et al., 2001b)。

　研究室内では、１日に１頭が1.1 ± 0.5 卵を産卵した (Joshi & Dabral, 2008)。卵胞の重さは0.29 ± 0.02 mg (Joshi & Dabral, 2008)。卵胞は18.7 ± 1.8 日で孵化した (Joshi & Dabral, 2008)。環帯が形成されるまでに、孵化後18 ± 3.4 日を要した(Joshi & Dabral, 2008)。

　和名は中村・Zicsi (1999) に基づく。

Perionyx excavatus Perrier, 1872: 126.

Perionyx excavatus Rosa, 1889: 134.

Megascolex excavatus Vaillant, 1889: 69. [同定形質のみ] (?, 1889)

Perionyx excavatus Rosa, 1890: 2.

Perionyx gruenewaldi Michaelsen, 1890g: 331.

Perionyx excavatus Rosa, 1891: 404.

Perionyx excavatus (? incl. P. m'intoshii) Beddard, 1891: 275.

Perionyx excavatus Rosa, 1892a: 2.

Perionyx excavatus Beddard, 1893: 689.

Perionyx excavatus Beddard, 1895: 437.

Perionyx excavatus Michaelsen, 1900: 208 (syn. gruenewaldi).

Perionyx excavatus Cognetti, 1906f: 1. [記述なし] (Oct 20, 1906)

Perionyx excavatus Cognetti, 1908e: 309. [記述なし]

Perionyx excavatus Stephenson, 1914: 386.

Perionyx koboensis Stephenson, 1914: 389.

Perionyx excavatus Stephenson, 1916: 317.

Perionyx excavatus Stephenson, 1917: 375.

Perionyx turaensis Stephenson, 1920: 216.

Perionyx excavatus Stephenson, 1921: 760.

Perionyx excavatus Stephenson, 1922b: 435.

Perionyx excavatus Michaelsen, 1922: 22.

Perionyx excavatus Stephenson, 1923: 329 (syn. intermedius, gruenewaldi, parvulus).

Perionyx koboensis Stephenson, 1923: 339.

Perionyx turaensis Stephenson, 1923: 360.

Perionyx excavatus Stephenson, 1924b: 340.

Perionyx excavatus Gates, 1926b: 472.

Perionyx excavatus Gates, 1926c: 162.

Perionyx fulvus Gates, 1926b: 472.

Perionyx excavatus Gates, 1926d: 182.

Perionyx fulvus Gates, 1926d: 182.

Perionyx excavatus Gates, 1926f: 213.

Perionyx fulvus Gates, 1926f: 213.

Perionyx excavatus Stephenson, 1926: 258.

Perionyx excavatus Gates, 1927c: 352.

Perionyx excavatus Stephenson, 1929: 238.

Perionyx excavatus Stephenson, 1931c: 275.

Perionyx excavatus Stephenson, 1932: 50.

Perionxy excavatus Gates, 1932a: 546. [記述のみ]

Perionyx excavatus Stephenson, 1933: 930.

Perionyx excavatus Kobayashi, 1938c: 201. [形態記載]

Perionyx excavatus 小林, 1938e: 779. [同定形質のみ]

Perionyx excavatus Gates, 1941a: 161.

Perionyx excavatus Gates, 1955c: 79.

Perionyx excavatus Gates, 1960c: 224. [形態記載]

Perionyx excavatus Kale & Krishnamoorthy, 1978: 24. [記述のみ] (Mar ??, 1978)

Perionyx excavatus Julka, 1981: 21. [記述のみ]

Perionyx excavatus Julka, 1982: 146. [形態記載]

Perionyx excavatus Nakamura, 1990: 102.

Perionyx excavatus Zicsi, 1993: 139.

Perionyx excavatus Shih et al., 1999: 438. [記述のみ]

Perionyx excavatus Paliwal & Julka, 2004: 1976.

Perionyx excavatus 中村・Zicsi, 1999: 89.

Perionyx excavatus Hong et al., 2001: 188. [形態記載]

Perionyx excavatus Dhiman & Battish, 2006: 2136.

Perionyx excavatus Huang et al., 2006: 17. [記述のみ]

Perionyx excavatus Joshi & Dabral, 2008: 83.

Perionyx excavatus Reynolds, 2009b: 57. [記述のみ] (Aug, 2009)

Perinoyx excavatus Tsai et al., 2009: 38.

Perionyx excavatus Csuzdi & Pavlíček, 2009: 10. (syn. gruenewaldi, koboensis, fulvus, turaensis) [記述のみ]

Perionyx excavatus Sinha et al., 2013: 304.

Perionyx excavatus Reynolds, 2015v: 117. [記述のみ] (Dec, 2015)

[要検討]

excavatus Perrier, 1872: 126, Beddard, 1886: 308, Rosa, 1896: 511, Beddard, 1893: 689, Gates, 1933: 549, 1936: 466.

Megascolex excavatus Vaillant, 1889: 69

gruenewaldi, Michaelsen, 1891: 33, Horst, 1893: 75, Beddard, 1893: 685.

turaensis

koboensis

fulvus

Beddard, F.E., 1891. The classification and distribution of earthworms. Proceedings of the Royal Physical Society of Edinburgh 10: 235-290.

Beddard, F.E., 1893. On some new species of earthworms from various parts of the world. Proceedings of the Zoological Society of London 1892: 666-706.

Beddard, F.E., 1895. A Monograph of the order of Oligochaeta. Oxford at the Clarendon Press. pp. 769.

Cognetti, L., 1906f. Lombrichi di Madagascar e dell'isola Riunione. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Università di Torino 21(537): 1-9.

Cognetti, L., 1908e. Di alcuni Oligocheti esotici. Annalen des Naturhistorischen Museums in Wien 22(4): 308-321.

Csuzdi C, Pavleíček T, 2009. New records of earthworms from Guadeloupe with description of a new species (Oligochaeta: Glossoscolecidae, Acanthodrilidae, Megascolecidae and Eudrilidae). Opuscula Zoologica Budapest 40(1): 9-15.

Dhiman, N., Battish, S.K., 2006. Earthworms from northern Indian states with Ocnerodrilus occidentalis Eisen, 1878 as a new report from Punjab. Zoos' Print Journal 21(1): 2135-2137.

Gates, G.E., 1926b. Notes on Rangoon earthworms: The peregrine species. Annals and Magazine of Natural History Ser. 9. 17: 439-473.

Gates, G.E., 1926c. Notes on earthworms from various places in the provinces of Burma, with descriptions of two new species. Records of the Indian Museum 28: 141-170.

Gates, G.E., 1926d. Notes on the seasonal occurrence of Rangoon earthworms. J. Bombay Nat. Hist. Soc. 31:180-185.

Gates, G.E., 1926f. The earthworms of Rangoon. Journal of the Burma Research Society 15: 196-221.

Gates, G.E., 1927c. Regeneration in a tropical earthworm Perionyx excavatus (E. Perr.). Biological Bulletin 53: 351-364.

Gates, G.E., 1932a. The earthworms of Burma. III. The Megascolecinae. Records of the Indian Museum 34: 357-549.

Gates, G.E., 1941a. Further notes on regeneration in a tropical earthworm, Perionyx excavatus E. Perrier, 1872. The Journal of Experimental Zoology 88(2): 161-185.

Gates, G.E., 1955c. The earthworms of Burma VI. Records of the Indian Museum 52: 55-93.

Gates, G.E., 1960c. On Burmese earthworms of the family Megascolecidae. Bulletin of the Museum of Comparative Zoology 123: 203-282.

Hong, Y., Kim, T.H., Na, Y.E., 2001b. Identity of two earthworms used in vermiculture and vermicomposting in Korea: Eisenia andrei and Perionyx excavatus. The Korean Journal of Systematic Zoology 17(2): 185-190.

Huang, J., Qin, X., Zheng, J.S., Chong, W., Dong, M.Z., 2006. Research on earthworm resources of China: I. Check-list and distribution. Journal of China Agricultural University 11(3): 9-20.

James, S.W., 2004c. An illustrated key to the earthworms of the Samoan Archipelago (Oligochaeta" Glossoscolecidae, Megascolecidae, Moniligastridae). Micronesica 37(1): 1-13.

Joshi, N., Dabral, M., 2008. Life cycle of earthworms Drawida nepalensis, Metaphire houlleti and Perionyx excavatus under laboratory controlled conditions. Life Science Journal 5(4): 83-86.

Julka, J.M., 1981. Taxonomic studies on the earthworms collected during the Subansiri expedition in Arunachal Pradesh, India. Records of the Zoological Survey of India, Miscellaneous Publication, Occasional Paper 26: 1-53.

Julka, J.M., 1982. Earthworm fauna of the Andaman and Nicobar Islands, India. Records of the Zoological Survey of India 80: 127-155.

Kale, R.D., Krishnamoorthy, R.V., 1978. Distribution and abundance of earthworms in Bangalore. Proceedings of the Indian Academy of Sciences, Section B, Animal Sciences 87(3): 23-25.

Kobayashi, S., 1938c. Occurrence of Perionyx excavatus E. Perrier in north Formsa. The Science Reports of the Tohoku Imperial University, 4th series (Biology) 13(2): 201-203.

小林新二郎, 1938e. 新竹及びその附近の蚯蚓. II. 動物学雑誌 51: 777-779.

Michaelsen W., 1890g. Oligochaeten des Naturhistorischen Museums in Hamburg. IV. Jahrbuch der Hamburgischen Wissenschaftillchen Anstalten 8: 299-340.

Michaelsen, W., 1900. Oligochaeta. Das Tierrich 10: 1-575.

Michaelsen, W., 1922. Oligochäeten aus dem Rijks museum van natuurlijke historie zu Leiden. Capita Zoologica 1: 1-72.

Nakamura M, 1990. How to identify Hawaiian earthworms. Chuo Daigaku Ronshu 11: 101-110.

中村好男, Zicsi, A., 1999. ハラメノウミミズの日本からの初記録. Edaphologia 63: 89-90.

Paliwal, R., Julka, J.M., 2004. Checklist of earthworms of western Himalaya, India. Zoos' Print Journal 20(9): 1972-1976.

Perrier, E., 1872. Recherches pour servia a I’histoire des Lombriciens terrestres. Noubelles archives du Muséum d’histoire naturelle 8: 5-198.

Reynolds, J.W., 2009b. Species distribution maps for Gates' Burmese Earthworms and current nomenclatural usage. Megadrilogica 13(6): 53-83.

Reynolds, J.W., 2015v. A checklist by counties of earthworms (Oligochaeta: Acanthodrilidae, Glossoscolecidae, Lumbricidae, Megascolecidae, Ocnerodrilidae, Octochaetidae) in Hawaii, USA. Megadrilogica 19: 123-145.

Rosa, D., 1889. I lombrichi raccolti nell'isola Nias dal Signor E. Modigliani e descritti dal Dott. Daniele Rosa, Assistente al R. Museo Zoologico di Torino. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a 7: 125-136.

Rosa, D., 1890. Terrucikas ex Birmania et ex austral America. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Università di Torino 5(93): 1-3.

Rosa, D., 1891. Die exotischen terricolen des k. k. naturhistorischen hofmuseums. Annalen des k. k. Naturhistorischen Hofmuseums, Wien 6: 379-406.

Rosa, D., 1892a. I Terricoli Esotici. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Università di Torino 7(114): 1-2.

Shih, H.T., Chang, H.W., Chen, J.H., 1999. A review of the earthworms (Annelida: Oligochaeta) of Taiwan. Zoological Studies 38(4): 435-442.

Sinha, M.P., Srivastava, R., Gupta, D.K., 2013. Earthworm biodiversity of Jharkhand: taxonomic description. The Bioscan 8(1): 293-310.

Stephenson, J., 1914. Zoological results of the Abor Expedition, 1911-1912, XXIX. Oligochaeta. Records of the Indian Museum 8: 365-410.

Stephenson, J., 1916. On a collection of Oligochaeta belonging to the Indian Museum.Records of the Indian Museum 12: 299-354.

Stephenson, J., 1917. On a collection of Oligochaeta from various parts of India and further India. Records of the Indian Museum 13: 353-416.

Stephenson, J., 1921. Oligochaeta from Manipur, the Laccadive Islands, Mysore, and other parts of India. Records of the Indian Museum 22: 745-768.

Stephenson, J., 1922b. Some earthworms from Kashmir. Bombay and other parts of India. Records of the Indian Museum 24: 427-443.

Stephenson, J., 1924b. On some Indian Oligochaeta with a description of two new genera of Ocnerodrilinae. Records of the Indian Museum 26: 317-365.

Stephenson, J., 1926. Descriptions of Indian Oligochaeta. I. Records of the Indian Museum 28: 249-267.

Stephenson, J., 1929. The Oligochaeta of the Indawgyi Lake (Upper Burma). Records of the Indian Museum 31: 225-239.

Stephenson, J., 1931. Oligochaeta from the Malay peninsula. Journal of Federated Malay Museum 16: 261-285.

Stephenson, J., 1932. Oligochaeta from the Malay Peninsula. II. Bulletin of the Raffles Museum 7: 42-51.

Stephenson, J., 1933. Oligochaeta from Australia, North Carolina and other parts of the world. Proceedings of the Zoological Society of London 1932: 899-941.

Tsai, C.F., Shen, H.P., Tsai, S.C., Lin, K.J., Hsieh, H.L., Yo, S.P., 2009. A checklist of oligochaetes (Annelida) from Taiwan and its adjacent islands. Zootaxa 2133: 33-48.

Vaillant, L., 1889. Histoire naturelle des Anneles marins et d'eau douce. 3. Lombriciniens, Hirudiniens, Bdellomophes, Teretulariens et Planariens. 766 pp.

Zicsi, A., 1993. Some Megadrili Oligochaeta of the Caribbean region. Revue Suisse de Zoologie 100(1): 137-142.