Scolopendra japonica Koch, 1878

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 : 34-36

Taxon classification Animalia Scolopendromorpha Scolopendridae

Scolopendra japonica Koch, 1878 Figs 7D, 9C, 29, 36, 37, 38, 39, 40, 41

Scolopendra japonica Koch, 1878: 790. Daday 1889: 149. Haase 1887: 48, pl. 3, fig. 48. Takakuwa 1942: 41, 1943: 171, 1947: 938. Kronmüller 2012: 24, figs 3D, 4B.

Scolopendra subspinipes japonica - Kraepelin 1903: 263. Attems 1909: 10, 13, 1914: 107, 1930b: 30. Shinohara 1949: 81, 1961: 75. Takashima 1949: 11, 1952: 4. Chamberlin and Wang 1952: 180. Miyosi 1955: 151. Wang 1955: 16. Takano 2001: 211. Lewis 2010b: 114.

Otostigmus politoides Attems, 1953: 147. Lewis 2004: 32, Figs 14-17. Kronmüller 2012: 25.

Otostigmus puncticeps Attems, 1953: 146, figs 16-17. Lewis 2004: 30, figs 8-13. Kronmüller 2012: 25.

Type locality.

Japan.

Material.

Syntype: NHMW Inv. No. 5368, one female from Japan, leg. Roletz, don. Latzel, 1919, det. Attems C (Figs 38, 39). Probable syntypes: NHMW Inv. No. 762, seven spms., Japan, leg. Roletz, red (type) label in bottle.

Additional material.

Laos - CUMZ 00297.1-3, two spms., Phu Fah Mountain, Phongsaly, Laos (21°41'19.6"N, 102°06'30.4"E). CUMZ 00298.1-5, five spms., Plain of Jar, Xiang Khouang, Laos (19°25'51.5"N, 103°09'10.4"E).

Japan - CUMZ 00319, one spm., Shinshu University, Matsumoto, Japan (36°13'22.4"N, 137°54'35.0"E). NHMUK 1893.1.15.3, one spm., Tokyo. NHMUK 1912.12.12.914, two spms., Izu Peninsula. NHMUK 1937.9.9.59, one spm., Japan, det. K.W. Verhoeff. NHMW Inv. No. 758, six spms., Yokohama. NHMW Inv. No. 755, seven spms., Japan. NHMW Inv. No. 757, one spm., Okayama, leg. H. Sauler. NHMW Inv. No. 759, one spm., Kioto [Kyoto] with anther label “344”. NHMW Inv. No. 760, two spms., Kioto [Kyoto], leg. H. Sauler. NHMW Inv. No. 756, two spms., Kanagava [Kanagawa], leg. H. Sauler, 17/11/1905, bottle with labels, "Hans Lauter 4122, Kanagawa 17/11/1905 Jan Haus" and "Hans Lauter 3236, Kanagawa 25/6/1905 Sichen".

Indonesia - NHMUK 1882.62, one spm., Sumatra.

China - NHMUK, one spm., Loc. 273, Peak of Flat No. 2, Hong Kong. NHMUK 1904.7.23.5-8, two spms., Yunnan-Fu [Kunming], South China.

Undetermined - NHMW Inv. No. 761, 17 spms., Kuile? [Possibly referring to Korea?].

Diagnosis. 17-19 antennal articles, 6 basal articles glabrous dorsally. Each tooth-plate with 4-6 teeth. Tergites (3)4-20 with paramedian sutures. Complete margination from TT(10)12-21. Tergite of ultimate leg-bearing segment without depression or suture. Complete paramedian sutures on sternites 2-20. Coxopleural process with 3 apical spines. Ultimate leg prefemora with 2-3 VL, 1-2 M, 1-3 DM and prefemoral process with 0-5 spines. One tarsal spur on legs 1-19(20).

Composite description.

Body length up to 12.9 cm. Two colour morphs; morph 1 with antenna and legs 1-20 yellowish, morph 2 with antenna and legs 1-20 reddish. All tergites greenish brown. Cephalic plate with median sulcus. Paramedian sulci or sutures absent on posterior part.

Antenna usually with 18 articles (atypically with 17 or 19), basal 6 subcylindrical and glabrous dorsally (Fig. 40A), the rest spherical. Antennae reach to tergite 2 (Fig. 36A). Forcipular trochanteroprefemoral process bearing denticles in two groups, one apical and 2-3 inner (Fig. 36 B–C). Tooth-plates wider than long, with 4-6 teeth (Figs 36B, 40 D–E). Tooth-plate with straight, transverse basal suture. Coxosternite smooth without median suture (Figs 36C, 40C). Article 2 of second maxillary telopodite with spur.

Anterior margin of T1 underlying cephalic plate (Fig. 36A). Complete paramedian sutures from TT3-4; margination typically starting on T10. Tergite surface (Figs 37A, 40B and 41A) with median posterior sulci in TT10-20. Tergite of ultimate leg-bearing segment (Figs 37D, 41B) curved posteriorly, without median furrow or depression; ratio of width: length of tergite of ultimate leg-bearing segment 0.82:1. Sternites (Figs 37B, 41C) with complete paramedian sutures, without depression or pit on surface. Sternite of ultimate leg-bearing segment (Fig. 37C) with sides converging posteriorly, surface with obscure depression on median part. Lateral part of coxopleuron with pore-field terminating beneath margin of tergite of ultimate leg-bearing segment.

Coxopleural process moderately long, usually with three apical spines (Fig. 39A); pore-free area extending 70-90% length from distal part of coxopleural process to margin of sternite of ultimate leg-bearing segment.

All legs without setae and tibial spur. One tarsal spur on legs 1-19 (on leg 20 in one spm.). Ultimate legs: moderately thick and long (Figs 37G, 41E), with ratios of lengths of prefemur and femur 1.1:1, femur and tibia 1.3:1, tibia and tarsus 2 1.4:1; tarsus 1 and tarsus 2 1.8:1. Prefemoral spines (Figs 37 E–F, 41 D–E): 2-3(4) VL, 1-2 VM, 1-3 DM, prefemoral process with 0-5 spines.

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 (Fig. 7D). Tergites of genital segment without small setae. Gonopod present in male.

Discussion.

The validity of Scolopendra japonica at the species level was defended by Kronmüller (2012) from morphological surveys of former subspecies of Scolopendra subspinipes . Diagnostic characters of this species by comparison to the other former subspecies are the number of apical spines on the coxopleural process ( Scolopendra subspinipes s.str. having two apical spines versus Scolopendra japonica having three spines, including a subapical spine), and the number of ventral spines on the prefemur of the ultimate legs. With respect to the latter, Scolopendra japonica has three spines whereas Scolopendra subspinipes s.str. has only two, although in this study an asymmetrical number of spines in Scolopendra japonica was also found in syntypes (Fig. 39 C–F). Four specimens among the japonica syntypes have been recorded as presenting only two or three ventral spines on the prefemur of the ultimate leg either on only one side or on both (six spines is the observed maximum). This variability might represent ontogenetic variation more so than geographical variation. Chao (2008) cited the lack of a tarsal spur on leg 20 as an additional diagnostic character for this species but from our study one specimen of Scolopendra japonica from the Izu Peninsula, Japan (NHMUK 1912.12.12.914), also exhibits a tarsal spur on leg 20. As such, the occurrence of a spur on leg 20 and number of ventral spines on the ultimate leg prefemur are not completely reliable for diagnosing Scolopendra japonica .

The sympatric distribution of this species and former subspecies of Scolopendra subspinipes as well as other Asian temperate Scolopendra complicates morphological delimitation of species boundaries except using the three phenotypic characters discussed above. In this paper, we compared taxonomic characters based on collections in the NHMUK and NHMW (Table 8). Additional characters of these two species that might be useful for species identification are the proportions of the ultimate leg podomeres and the number of spines on the prefemoral process on the ultimate leg. In Scolopendra subspinipes s.str., two spines are usually present on the prefemoral process whereas in Scolopendra japonica there are typically three. The length of the antenna also permits a distinction berween these two species; the antenna extends backwards only as far as TT2-3 in Scolopendra japonica whereas it can reach to TT4-5 in Scolopendra subspinipes (Figs 10A, 14B and 36A). Moreover, molecular analysis of three combined genes (COI, 16S and 28S) indicated a genetic distinction between Scolopendra japonica and Scolopendra subspinipes (Fig. 1), and the validity of these species has been corroborated herein. Ratios of ultimate leg podomeres have been used as diagnostic characters of some putative species of Scolopendra in Asia, such as Scolopendra negrocapitis Zhang and Wang, 1999 from Jingshan (northeast coastal area of China). The authors mentioned the close similarity between that species and Scolopendra japonica but the Chinese species can be distinguished from the latter only by the width:length ratio of the ultimate leg prefemur, which is twice as long as broad. The lack of further information from fresh material from the type locality and molecular data from Scolopendra negrocapitis renders the status of these two closely related species questionable.

In the current phylogenetic framework of Scolopendra , Scolopendra japonica is resolved in the same clade as Scolopendra cingulata Latreille, 1829. The two species are morphologically similar despite their markedly disjunct distributions, i.e., Scolopendra cingulata in the Mediterranean versus Scolopendra japonica in East Asia (Table 8). However, exploration of micro-refugia of populations of Scolopendra cingulata during glacial maxima in Europe ( Simaiakis et al. 2012, Oeyen et al. 2014) and a record of Scolopendra japonica in the northern part of Laos could indicate that these two species may be more widespread than previously recognised. However, distributional data for Scolopendra japonica are patchy due to incomplete faunistic surveys in several parts in Asia. For these reasons, the relationship between these two species warrants further scrutiny in both morphological and molecular studies.

Distribution.

Probably distributed throughout the temperate zone of East Asia including mainland and insular territory (Fig. 29). The distribution range is likely to be sympatric with several Asian species ( Scolopendra subspinipes , Scolopendra multidens , Scolopendra dawydoffi and Scolopendra dehaani ). Previous study on the subspecies complex of Scolopendra subspinipes indicated that Scolopendra japonica might occur in northern Vietnam (Tonkin) and Cambodia, based on the type localities of " Otostigmus politoides Attems, 1953" and " Otostigmus puncticeps Attems, 1938" (= Scolopendra subspinipes fide Lewis 2004), the types of both of which are adolescent stages ( Kronmüller 2012) that are compatible with Scolopendra japonica . The current distribution of this species gathered from previous literature and this study is as follows: Southeast Asia: Vietnam (Chapa, Tonkin [probably referring to Sa Pa, Lao Cai Province, northern Vietnam]), Laos (Phongsaly and Xieng Khuang), Cambodia (Ream, Koh Kong Island and Sre Umbell [Sre Ambel]), Indonesia (Sumatra). East Asia: Japan (Yokohama, Enoshima, Murayama, Tokyo, Matsumoto, Kanakawa, Sendai, Kii, Hachijo island and Izu Peninsula), Taiwan and China (Hong Kong, Yunnan-Fu [Kunming] and Ningbo).