Scolopendra subspinipes Leach, 1816

Siriwut, Warut, Edgecombe, Gregory D., Sutcharit, Chirasak, Tongkerd, Piyoros & Panha, Somsak, 2016, A taxonomic review of the centipede genus Scolopendra Linnaeus, 1758 (Scolopendromorpha, Scolopendridae) in mainland Southeast Asia, with description of a new species from Laos, ZooKeys 590, pp. 1-124 : 15-20

Taxon classification Animalia Scolopendromorpha Scolopendridae

Scolopendra subspinipes Leach, 1816 Figs 9A, 10, 11, 12, 13, 14, 15, 16, 17, 18

Scolopendra subspinipes Leach, 1816: 383. Newport 1844: 96. Koch 1847: 163. Kohlrausch 1881: 96. Meinert 1886: 202. Haase 1887: 44, pl. 3, Figs 43-45. Daday 1889: 150. Latzel 1892: 185. Pocock 1894: 312. Silvestri 1894: 624, 1895: 714. Attems 1897: 477, 1903: 81, 1914a: 106, 1914b: 568, 1914c: 380, 1915: 2, 1927: 1, 1930b: 29, fig. 43, 1930c: 175, 1932: 5, 1938: 334; 1953: 145. Flower 1901: 21. Ribaut 1912: 248. Chamberlin 1918: 158, 1920a: 30, 1920b: 391. Muralewicz 1913: 201. Chamberlin and Wang 1952: 190. Takakuwa 1942a: 1, 1942b: 15, 1942c: 41, 1943: 171. Wang 1955b: 16, 1962: 101, 1965a: 449, 1967b: 391. Würmli 1972: 91, Fig. 1. Shelley 2000: 42. Lewis 2002: 83, 2007: 10, 2010a: 129, 2010b: 111, 2010c: 380, figs 1-3. Schileyko 1995: 77, 2007: 75. Chao and Chang 2003: 4, fig. 9, tables 1, 2. Chao 2008: 35, figs 37-39, table 2. Kronmüller 2012: 20, table 1, figs 3-5. Chagas-Júnior et al. 2014: 139.

Scolopendra audax Gevais, 1837: 50. Kohlrausch 1881: 99.

Scolopendra septemspinosa Brandt, 1840: 152. Kraepelin 1903; 256.

Scolopendra borbonica Blanchard, 1829: 7, pl. 1. Kohlrausch 1881: 98.

Scolopendra sexspinosa Newport, 1844: 96, 1845: 391. Kohlrausch 1881: 100. Daday 1891: 149.

Rhombocephalus gambiae Newport, 1845: 392. Kraepelin 1903: 256.

Scolopendra ceylonensis Newport, 1845: 391. Kohlrausch 1881: 98.

Scolopendra flava Newport, 1845: 392. Kohlrausch 1881: 98.

Scolopendra gervaisii Newport, 1845: 390. Kohlrausch 1881: 100.

Scolopendra lutea Newport, 1845: 392. Kohlrausch 1881: 99.

Scolopendra ornata Newport, 1845: 392. Koch 1863: 10, pl. 66, fig. 134. Kohlrausch 1881: 100.

Scolopendra placeae Newport, 1845: 390. Kohlrausch 1881: 100.

Scolopendra planiceps Newport, 1845: 391. Kohlrausch 1881: 99.

Scolopendra rarispina Gervais, 1847: 270. Kohlrausch 1881: 97.

Scolopendra sandwichiana Gervais, 1847: 276. Kohlrausch 1881: 99.

Scolopendra mactans Koch, 1847: 16, 1863: pl. 41, fig. 79. Kohlrausch 1881: 98.

Scolopendra sulphurea Koch, 1847: 156, 1863: 24, table. 11, fig. 21. Kohlrausch 1881: 98.

Scolopendra byssina Wood, 1861: 10. Kohlrausch 1881: 99.

Scolopendra cephalica Wood, 1861: 12. Kraepelin 1903: 256.

Scolopendra cephalica gracilis Wood, 1861: 13. Kraepelin 1903: 256.

Scolopendra dinodon Wood, 1861: 12. Kohlrausch 1881: 98.

Scolopendra gracilipes Wood, 1861: 12. Kraepelin 1903: 256.

Scolopendra parvidens Wood, 1861: 13. Kohlrausch 1881: 98.

Scolopendra plumbeolata Wood, 1861: 14. Kohlrausch 1881: 97.

Scolopendra bispinipes Wood, 1862: 28. Brölemann 1909: 25.

Scolopendra nesuphila Wood, 1862: 31. Kraepelin 1903: 256.

Scolopendra repens Wood, 1862: 31. Kraepelin 1903: 256.

Scolopendra elongata Porat, 1871: 1143. Meinert 1886: 202.

Rhombocephalus smaragdinus Butler, 1876: 446. Kraepelin 1903: 256.

Scolopendra damnosa Koch, 1878: 789. Kraepelin 1903: 256.

Scolopendra mutilans Koch, 1878: 791. Haase 1881: 47, pl. 3, fig. 47. Takakuwa 1947: 938.

Scolopendra aurantiipes Tömösváry, 1885: 67. Haase 1887: 44. Takakuwa 1936: 152.

Scolopendra variispinosa Tömösváry, 1885: 67. Haase 1887: 44. Takakuwa 1936: 152.

Scolopendra rugosa Meinert, 1886: 202. Kraepelin 1903: 257.

Scolopendra meyeri Haase, 1887: 49, pl. 3, fig. 50. Kraepelin 1903: 257.

Scolopendra macracanthus Bollman, 1889: 213. Kraepelin 1903: 257.

Scolopendra flavicornis Tömösváry, 1885: 67. Kraepelin 1903: 256. Takakuwa 1936: 152.

Scolopendra subspinipes gracilipes Daday, 1891: 149. Kraepelin 1903: 256.

Scolopendra subspinipes molleri Verhoeff, 1892: 199. Kraepelin 1903: 256.

Scolopendra polyodonta Daday, 1893: 5. Kraepelin 1903: 257.

Scolopendra machaeropus Attems, 1901: 136. Kraepelin 1903: 257.

Scolopendra aringensis Sinclair, 1901: 529, pl. 31, fig. 46, pl. 32, Figs 67, 85, 86, 93. Kraepelin 1903: 257.

Scolopendra subspinipes mutilans Kraepelin, 1903: 263. Attems 1938: 334, 1953: 138. Takakuwa 1943: 171. Takashima 1952: 4. Shinohara 1961: 75. Wang 1993: 850, fig. 5. Schileyko 1998: 268, 2007: 75. Chao and Chang 2003: 8, table 1-2, figs 6-7. Chao 2008: tab. 2. Lewis 2010b: 111. Kronmüller 2012: 20, table 1.

Scolopendra subspinipes gastroforeata Muralewicz, 1913: 201. Lewis 2010b: 114. Kronmüller 2012: table 1.

Scolopendra subspinipes piceoflava Attems, 1934: 51. Lewis 2010b: 113. Kronmüller 2012: 21, table 1.

Scolopendra subspinipes fulgurans Bücherl, 1946: 148, 1974: 107. Kronmüller 2012: 21, table 1.

Type locality.

Not designed. The whereabouts of the holotype are unknown.

Material.

Specimens referred to Scolopendra subspinipes Leach, 1816: Malaysia - NHMUK 1897.1.25.12, one spm., Penang, Malaysia, leg. S.S. Flower, 26/11/1896, with label "PENANG 226". NHMUK, two spms., Penang, Malay Peninsula, leg. H.N. Ridley. NHMUK (E): 2000-110, one spm., C89, caught in base camp, Mulu, Sarawak, 5/8/1978, leg. J.G.E. Lewis. NHMUK (E): 2000-110, one spm., Mulu, Sarawak, 21/8/1978, leg. Ian Baillie. NHMUK 1952.9.8.576, one spm., Sarawak, Borneo, with label “F.42.24.8.1932”, Oxford University Sarawak Expedition. NHMUK 1906.2.18.3, one spm., Malay Peninsula, leg. Annandale and Robinson, with label "No. 45".

Singapore - CUMZ 00315, one spm., Kentridge Road, Singapore (1°17'08.9"N, 103°47'09.8"E). NHMUK 1886.115, one spm., Singapore, leg. Dr. Invine Russell, don. E.W. Holmes Zgar. NHMUK 1897.12.22.63-64, two spms., Singapore, leg. S.S. Flower.

Indonesia - NHMUK 1893.5.13.30, two spms., east coast Sumatra, leg. Mrs. Findlay. NHMW, 12 spms., Singkarak, Klakah [Singkarak Lake, Sumatra, Indonesia]. NHMUK, one spm., Java, with label "No. 46/108", 2/3/1885. NHMUK 1874.57, two spms., Java, leg. G. Lyon Esq. NHMW Inv. No. 726, six spms., Java, Indonesia, 1884, det. Attems C. NHMUK 1896.6.20.33, one spm., Surabaya, Java, leg. S.S. Flower, April 1896. NHMW Inv. No. 8596, two spms., Klakah, Lumajang Regency, East Java, Indonesia, leg. Thienemanm, October/November 1928, det. Attems C. NHMW Inv. No. 965, two spms., Java, 30/6/1857-1859, Novara expedition. NHMUK, one spm., found between base camp and corner camp near Utakawa River, Expedition to Dutch New Guinea [Irian Jaya]. NHMUK 1911.12.23.63-64, two spms., Muisika River, South Dutch New Guinea, B.O.U. expedition, leg. Mr. Wollaston.

Philippines - NHMUK 1896.3.8.87, one spm., Philippines, leg. Pascal. NHMUK 1913.6.18.897, one spm., Philippines, with label "Typical form" and "Spec. 13". NHMUK 1883.33, two spms., Manila, leg. S.W. Taylor.

China - NHMUK 1928.3.16.64-68, one spm., Amoy, China, leg. Prof. C. Ping, 4/2/1926, with label “No.CAT.3”. NHMUK 1894.12.20.1, one spm., Central Formosa [Taiwan], leg. Holst. NHMUK, one spm., Changsha, Siang River (28°12'N, 112°59'E), leg. L.T. Loomer and R.H.S. Rodger R.N.

Japan - NHMUK, one spm., Japan, leg. Koch, with label "Number 2?". NHMUK 1907.6.18.1-2, two spms., Goto Island, Japan, leg. R. Gordon Smith.

South Asia - NHMUK, one spm., India, with label "No. 45/29". NHMUK, two spms., Ceylon, with label “No.46/104”.

Africa - NHMUK 1881.99, one spm., Kee Road, South Africa

Jamaica - NHMUK, two spms., in bananas from Jamaica, det. P.C. Jerrard. NHMUK, seven spms., Jamaica. NHMUK Entomology: 2000-110, one spm., in case of bananas from Jamaica, 1987. NHMUK, one spm., in bananas from Jamaica, Longford, Kent, leg. Miss S. Truman, 1/11/1950.

Madagascar - NHMUK, one spm., West Africa, with label “n/a”. NHMUK 1989.3.12.3-7, five spms., Madagascant [Madagascar], det. Lewis and Ransome (cap.). NHMUK 1878.30, one spm., Madagascar.

Rodrigues - NHMUK Entomology: 2000-110, six spms., under stones, heavily grazed grassland hill, west of port Mathunn, Rodrigues, 9/4/1995 (63°25'E, 9°41'S). NHMUK Entomology: 2000-110, three spms., Solitude expedition to Rodrigues, 10.11.1995. NHMUK Entomology: 2000-110, two spms., under forest cavern, Patate, Rodrigues (63°23.5'E, 19°45.5'S). NHMUK, six spms., Rodrigues, leg. Slater, October 1876, with label "refer to Scolopendra mossambica (Peter)". NHMUK, one spm., probably from Rodriguez, with label "Rodriguez: Anse aux anglaise". April 1983 CIE A14995". NHMUK 1924.2.9.6-11, six adult spms. and brood, Rodrigues, leg. G.C. Addison-Williamson.

Mauritius - NHMUK Entomology: 2000-110, two spms., Raphael island, St. Brandon, Mauritius, January 1996.

Seychelles - NHMUK, one spm., with label "No. 13216?", Praslin, January 1953, det. E.S. Brown. NHMUK 1952.12.17.248-249, two spms., Silhouette Island, leg. J.S. Gardiner. NHMUK, one spm., Mahe, leg. J.M and R.D. Pope, August 1976, det. MacFarlane. NHMUK 1867.76, two spms., Seychelles, leg. A. Newton Esq. NHMW Inv. No. 8597, two spms., Mahe, leg. Brauer, det. Attems C.

Comoros - NHMUK Entomology: 2000-110, one spm., Comores, leg. Helen Read, May 1995.

Pacific Islands - NHMUK, one spm., South Pacific Islands, leg. J.M. Selfridge Mhl., 2/3/1885. NHMUK 1950.4.19.13, one spm., Nukualofa, Tonga, 22/2/1925, det. Brolemann. NHMUK 1950.4.19.6 and 11, two spms., Apia, Upolu, Samoa, leg. Buxton and Hopkins, 7/5/1924. NHMUK, one spm., Ahui, Tautira, Tahiti, 9/8/1925, leg. Cheesman. NHMUK, one spm., Pahenoo, Tahiti, South Pacific, 3/1925, leg. Col. S.Y. Sr. George P.H. Johnson S.Z.R.Q. NHMUK 1882.60, one spm., Ravatonga, leg. Sir J. Fulbock. NHMUK 1911.12.4.14, one spm., Savau, Friendly Island. NHMUK, one spm., Hawaiian Islands, leg. Henry Edward, 30/12/1875. NHMUK 1882.60, one spm., Rarotonga, Sir J. Lubbock’s collection. NHMUK 1926.1.24.465, one spm., on mango trunk, Fatu Hiva, Marquesas Island, leg. P.H. Johnson S.Z.R.A, 6/1/1925. NHMUK 1926.1.24.466-470, five spms., Hiva Oa, Marquesas Island, January 1924, leg. P.H. Johnson.

United Kingdom - NHMUK, one spm., in flat over banana-ripening store, Poplar, London, leg. E.Z.H.O.H., 9/10/1956. NHMUK 1894.12.23.1, one spm., Kew Gardens. NHMUK, one spm., imported with bananas, Windsor, 14/6/1965, det. P.C. Jerrard, 1965. NHMUK, one spm., West Indies dock, London. NHMUK, one spm., imported with bananas, Boston, Lincs [Lincolnshire, U.K.], det. P.C. Jerrard, 24/10/1960.

Caribbean Sea - NHMUK, two spms., Barbados, det. J. Locke Esq. NHMUK 1886.113-116, one spm., Montserrat. NHMUK 1899.6.1-3, three adults, numerous juvenile spms., St. John (West Indies), leg. J.W. Gregory. NHMUK, one spm., leg. Morne Fortune, R.F.S., det. D.J. Clark. NHMUK, one spm., Bermuda, April, 1873. NHMUK 1896.3.17.21.22, two spms., Antigua, leg. W.R. Forrest. NHMUK, one spm., Bermuda, April, 1873.

Central and South America - NHMUK 1913.6.13.398-399, two spms., Bogotia [ Bogotá, Colombia]. NHMUK 1898.2.12.15, one spm., Rio Jurua, Amazons, leg. Dr. Bach. NHMUK 1913.6.18.900, one spm., Cayenne. NHMUK, one spm., Rio Sofiars, northwest Ecuador, 450 ft., leg. Rosenberg. NHMUK 1905.7.13.1, one spm., Isthmus of Panama, leg. Mr. H. Robert.

Undetermined locality - NHMUK, one spm., unknown locality, with label “No.47/21”. NHMUK 1813.6.18.201-202, one spm., with label " Scolopendra Div.I seet. B.6 Cintillus". NHMUK 1813.6.18.904, one spm., with label " Scolopendra subspinipes I (typical form)". NHMUK 1916.10.4.4-8, one spm., unknown locality. NHMUK, one spm., Rei Islands, leg. Cahl. Lingen, with label “88-100”. NHMW Inv. No. 703, one spm., Ostindien [East Indies].

Specimens referred to Scolopendra mutilans Koch, 1878: Japan - Syntype NHMW Inv. No. 751 of Scolopendra mutilans Koch, 1878, one spm., Japan, with label “Syntype”, leg. Roletz, don. Latzel, 1919 (Figs 12-13). NHMUK 1911.12.12.915-916, two spms., Izu, Japan, leg. S. Akiyama. NHMUK, one spm., Yokohama, Japan, HMS Challenger Expedition, May 1875. NHMUK, two spms., Kole [Kobe], Japan, June 1875. NHMW Inv. No. 746, one spm., Nagasaki, Japan, leg. Rausonel, 1871. NHMW Inv. No. 738, seven spms., Japan, don. Roretz, 1/3/1881. NHMW Inv. No. 740, six spms., Japan.

China - NHMUK 1886.120, one spm., Snowy valley, Ningbo. NHMUK 1892.12.6.1, one spm., Chusan Island [Zhoushan], leg. J.J. Walkes.

Korea - NHMUK 1882.14, two spms., Southeast Korea. NHMUK, two spms., Kang-hwa [Khangwhado], Korea, leg. Miss Scarlett.

Undetermined locality - NHMUK 1888.50, one spm., Seimer Island (Pabva), leg. H.O.F. NHMUK, two spms., Tsur Island, leg. Holst, July-August 1891. NHMW, nine spms., unknown locality determined as " Scolopendra mutilans ".

Diagnosis.

17-19 antennal articles, 6 basal articles glabrous dorsally. Each tooth-plate with (4)5-7 teeth. Tergites 3(4)-20 with paramedian sutures. Complete tergite margination on TT14 (17)-21. Tergite of ultimate leg-bearing segment without depression or median suture. Complete paramedian sutures on sternites 2(3)-20. Coxopleural process with 2 apical spines, without lateral and dorsal spine. Ultimate leg prefemora with 2 VL, 1-2 M, 0-3 DM and prefemoral process with 1-6 spines. One tarsal spur on legs 1-19 or 20.

Composite description.

Body length up to 16 cm. Reddish brown colouration on entire body. Cephalic plate and segments monochromatic or dichromatic. Tergites reddish brown; dark band on posterior border of tergites. Cephalic plate with small punctae on anterior part; median sulcus present. Posterior part of cephalic plate without paramedian sulci.

Antenna usually with 18-19 articles (16-17 articles on one side in some specimens), basal 6 subcylindrical and glabrous dorsally (Fig. 16A), 6 articles glabrous ventrally. Antennae reach segment 3-4. Forcipular trochanteroprefemoral process with denticles in two groups, 1-3 apical and 1-2 inner (Fig. 16B, D). Tooth-plates wider than long or nearly as long as wide, 5-7 teeth (Figs 10D, 16B, D); atypically with 10 on one side (NHMUK specimen from western New Guinea). Tooth-plate with straight, transverse basal suture. Coxosternite without median suture (Figs 10C, 16D). Article 2 of second maxillary telopodite with spur.

Anterior margin of T1 underlying cephalic plate (Figs 10A, 16A). Complete paramedian sutures on TT3-4; margination typically starting on TT5-10 (atypically from TT12-13 in some specimens). Tergite surface (Figs 10B, 16C) smooth. Tergite of ultimate leg-bearing segment (Fig. 17C) curved posteriorly, without median suture or depression; ratio of width: length of tergite of ultimate leg-bearing segment 0.8:1. Sternites (Figs 11A, 17A) with complete paramedian sutures. Surface of sternites smooth, without depression. Sternite of ultimate leg-bearing segment (Fig. 11B) with sides converging posteriorly; surface without depression. Pore-field on coxopleuron terminating far beneath margin of tergite of ultimate leg-bearing segment, dorsal margin of pore area gently sinuous (Figs 11C, 17B).

Coxopleural process moderately long, with two apical and 0-1 subapical spines; pore-free area extending 30-70% length from distal part of coxopleural process to margin of sternite of ultimate leg-bearing segment (Figs 11B, 17D).

All legs without setae and tibial spur. One tarsal spur on legs 1-19 or more commonly 1-20. Ultimate legs: moderately long and slender, with ratios of lengths of prefemur and femur 1.4:1, femur and tibia 1.2:1, tibia and tarsus 2 1.4:1.; tarsus 1 and tarsus 2 2:1. Prefemoral spines: 2 VL, 1-2 M, 0-3 DM and prefemoral process with 1-6 spines (Figs 11 D–E, 17 D–E). Posterior margin of prefemur with short median groove.

Genital segments well developed, reaching longer than distance between posterior margin of sternite of ultimate leg-bearing segment and distal part of coxopleural process. Sternite of genital segment 1 round and convex posteriorly, with median suture. In male, sternite of genital segment 2 attached to penis. Tergite of genital segment without small setae. Gonopods with small setae in male. Penis with apical bristle.

Discussion.

Recently, the taxonomic validity of Scolopendra subspinipes and its former subspecies has been evaluated both by morphology ( Kronmüller 2012) and molecular methods ( Chao et al. 2011, Siriwut et al. 2015a). Three former subspecies of Scolopendra subspinipes , namely Scolopendra subspinipes japonica , Scolopendra subspinipes dehaani and Scolopendra subspinipes cingulatoides (= Scolopendra dawydoffi ), have been raised to species rank ( Kronmüller 2012), whereas the remaining four subspecies (in the classification of Attems (1930b)) have been synonymized with the nominotypical subspecies. However, some subspecies still remain of ambiguous status. Notably, Scolopendra subspinipes mutilans Koch, 1878, a nominal subspecies occurring in East Asia, corresponds to Scolopendra subspinipes in all respects apart from the cephalic plate and T1 showing reddish colouration. Recent morphological revisions have regarded this subspecies to be a synonym of Scolopendra subspinipes ( Schileyko 2007, Kronmüller 2012), whereas molecular analyses based on four loci found it to either resolve as sister taxon to Scolopendra subspinipes s.str. or to group more closely with other species ( Vahtera et al. 2013). In this study, we document a syntype of Scolopendra mutilans Koch, 1878 in the NHMW collection (Figs 12-13) and reconfirmed its taxonomic status by using molecular analysis from the concatenated DNA dataset of Scolopendra subspinipes s.str. and Scolopendra mutilans Koch, 1878. The phylogenetic tree supports the proposition that this subspecies cannot be distinguished taxonomically from Scolopendra subspinipes . According to genetic divergence among examined populations, Scolopendra mutilans Koch, 1878 should be regarded as a geographical variant of Scolopendra subspinipes , as was suggested in other recent taxonomic studies ( Schileyko 1995, 2007). Some morphological comparisons of several populations from Southeast and East Asia are provided in Table 6.

Scolopendra subspinipes piceoflava , another former subspecies of Scolopendra subspinipes from Sulawesi, Indonesia, is currently treated as a synonym of Scolopendra subspinipes (see Kronmüller 2012), but may upon further study prove to be a valid species. Attems (1934) stated that it could be distinguished from other forms of Scolopendra subspinipes by yellowish colouration on the posterior part of its tergites. He also referred to the weakness or near absence of tergal paramedian sutures, which also occurs in some other SE Asian Scolopendra species (see Kronmüller 2012 and discussion of Scolopendra cataracta in this study). Re-examining the syntypes of Scolopendra subspinipes piceoflava leads us to dispute the taxonomic validity of this character because paramedian sutures are visible on the tergites in all syntypes. In order to provide a more complete evaluation of its taxonomic status, a redescription of its syntypes is as follows: