A taxonomic review of the centipede genus Scolopendra Linnaeus, 1758 (Scolopendromorpha, Scolopendridae) in mainland Southeast Asia, with description of a new species from Laos

Warut Siriwut; Gregory D Edgecombe; Chirasak Sutcharit; Piyoros Tongkerd; Somsak Panha

doi:10.3897/zookeys.590.7950

. 2016 May 17;(590):1–124. doi: 10.3897/zookeys.590.7950

A taxonomic review of the centipede genus Scolopendra Linnaeus, 1758 (Scolopendromorpha, Scolopendridae) in mainland Southeast Asia, with description of a new species from Laos

Warut Siriwut ^1,², Gregory D Edgecombe ³, Chirasak Sutcharit ², Piyoros Tongkerd ², Somsak Panha ²

PMCID: PMC4926625 PMID: 27408540

Abstract Abstract

The centipede genus Scolopendra in mainland Southeast Asia is reviewed taxonomically based on morphological characters, informed by a molecular phylogenetic analysis using sequences from three mitochondrial and nuclear genes (COI, 16S rRNA and 28S rRNA). Eight nominal species of Scolopendra, namely Scolopendra morsitans Linnaeus, 1758, Scolopendra subspinipes Leach, 1816, Scolopendra dehaani Brandt, 1840, Scolopendra multidens Newport, 1844, Scolopendra calcarata Porat, 1876, Scolopendra japonica Koch, 1878, Scolopendra pinguis Pocock, 1891, and Scolopendra dawydoffi Kronmüller, 2012, are redescribed together with some revision of type materials. Geographical variation in each species has been compiled with reference to samples that span their distribution ranges in Southeast Asia and some parts of neighbouring areas such as East Asia, the Indian Ocean, and Africa. Comparative study of traditional taxonomic characters from external morphology provides further information to distinguish some closely related species. Scolopendra cataracta Siriwut, Edgecombe & Panha, sp. n., is described from the southern part of Laos, with additional records in Thailand and Vietnam. The phylogenetic framework for Southeast Asian Scolopendra recognizes Scolopendra calcarata + Scolopendra pinguis, Scolopendra morsitans, and a Scolopendra subspinipes group that unites the other six species as the main clades. Within the Scolopendra subspinipes group, two monophyletic groups can be distinguished by having either slender or short, thick ultimate leg prefemora and different numbers of apical spines on the coxopleuron. Scolopendra arborea Lewis, 1982, is placed in subjective synonymy with Scolopendra dehaani. A survey of external morphology of the genital segments confirms its potential for improving species identification in Scolopendra. Some observations on biology and behaviour are recorded based on field surveys in this area.

Keywords: Chilopoda , Scolopendra , systematics, distribution, phylogeny, species diversity

Introduction

The genus Scolopendra Linnaeus, 1758, is among the predominant centipede groups in tropical regions. These animals are generalist feeders that play an important role as one of the top carnivorous invertebrates in soil ecosystems. In several Asian countries, Scolopendra has symbolic status or figures in superstitions, and is used commercially in traditional medicine (Pemberton 1999). A few species have been proposed as model animals for medical and biological subjects (Minelli and Fusco 2004) but comprehensive work on the regional biota has not been consolidated since the last monograph on Scolopendromorpha (Attems 1930b).

The taxonomic study of Scolopendra dates back to the late 19^th to mid-20^th centuries (Kohlrausch 1881, Pocock 1891a, Kraepelin 1903, Attems 1930b, 1953). Several names have fallen into synonymy through the course of taxonomic revisions (Kohlrausch 1881, Kraepelin 1903, Attems 1930b). Conversely, some populations that had been classified in geographically widespread species have recently been identified as distinct species e.g., Scolopendra antananarivoensis Kronmüller, 2010, versus Scolopendra morsitans Linnaeus, 1758, or Scolopendra subcrustalis Kronmüller, 2009, versus Scolopendra subspinipes Leach, 1816. The phylogenetic position of Scolopendra has been investigated in the context of broad-scale phylogeny of Scolopendromorpha (Vahtera et al. 2012, 2013). Combined molecular and morphological data supported a hypothesis that Old World species of Scolopendra can be distinguished from the New World species, and nested most Old World species in a clade with the genus Asanada Meinert, 1886. Morphological discrimination between two regional groups within Scolopendra is made mainly based on a transverse suture on tergite 1 in New World Scolopendra that is absent in nearly all Old World species. Some Old World species are polymorphic with respect to external phenotypic characters. For instance, Scolopendra subspinipes and Scolopendra morsitans are cosmopolitan species worldwide, and both of them include several colour variants (Kohlrausch 1881, Attems 1930a, Shelley et al. 2005). Previous scolopendrid studies proposed that morphological variation within species is influenced by geographic distribution and ontogeny (Lewis 1968, 1972). Ontogenetic variation in colouration patterns has been recorded in Scolopendra dehaani Brandt, 1840 from Southeast Asia, where the species has been investigated using both molecular and morphological data (Siriwut et al. 2015a).

To date, 99 described species of Scolopendra have been recorded (Bonato et al. 2016), of which fourteen species have been found in the Asian tropics (Schileyko 2007, Lewis 2010b, Kronmüller 2012). Taxonomic studies have been undertaken in the following regions of Asia: the Indian Subcontinent (Jangi and Dass 1984), Indochina including Burma (Pocock 1889, 1891b, Attems 1953, Schileyko 2007), the Malay and Philippine Archipelagos (Pocock 1894, Wang 1962, 1965b, 1967a, b), and the East coast of the China Sea (Takakuwa 1942a, Chamberlin and Wang 1952, Wang 1955a, b, 1956, 1957), including Taiwan (Chao 2008). In Southeast Asia, the following Scolopendra species are endemics: Scolopendra pinguis Pocock, 1891, Scolopendra gracillima Attems, 1898, Scolopendra spinosissima Kraepelin, 1903 and Scolopendra arborea Lewis, 1982. There are also species which are widely distributed: Scolopendra subspinipes, Scolopendra morsitans, Scolopendra dehaani and Scolopendra japonica Koch, 1878; each of them extends into neighbouring territories such as the Indian subcontinent and the Asian temperate region (Koch 1878, Jangi and Dass 1984). In mainland Southeast Asia, Scolopendra comprises ten nominal species. Most of them are cosmopolitan species found synanthropically. For several species, geographical variation has not previously been documented, but we now have access to molecular evidence by which such variability can be mapped to genetic structure among and between populations.

The validity of various scolopendrid species has been ambiguous because their distributions have not been comprehensively documented and/or because the diagnostic value of particular taxonomic characters has been unclear (Lewis 1978, 2003, 2010b, Lewis et al. 2006, Edgecombe 2007). Infra-specific variation within Scolopendra has long been noted as a fundamental problem for distinguishing between similar species (Newport 1845, Attems 1930a, Lewis 2010b). Exclusive reliance on the traditional external morphological characters may not be sufficient to resolve some of these questions, not the least those involving problems of colour variation (Koch 1982, 1983a, Shelley 2005). The phylogenetic relationships of Scolopendra to other scolopendrid genera have also been contentious, as is the monophyly of the genus (Vahtera et al. 2013). Recently, evidence has been presented that morphological identification, molecular phylogeny, and geometric morphometric analyses congruently support the traditional delimitation of Scolopendra species in mainland Southeast Asia (Pocock 1891b, Flower 1901, Schileyko 1992, 1995, Kronmüller 2012). Molecular sequence analyses of Southeast Asian species indicated previously unrecognized groups within nominal species (Siriwut et al. 2015a). These results may indicate that even within a morphologically conservative centipede group, cryptic species can potentially be identified.

In this work, we review Scolopendra species in mainland Southeast Asia. The type material of some species has been re-described and, where available, type material is photographed and illustrated. All species are compared with the most closely allied congeners to provide distinguishing taxonomic characters. Variability in morphological characters is recorded in order to document geographical variation. The description of a new species is based on specimens from three SE Asian countries. Molecular phylogeny of three standard genes is analyzed, adding new samples to previous work on Scolopendra, to test the monophyly of each species and to determine the phylogenetic position of new species. An identification key to Scolopendra is presented and distribution ranges for species are updated.

Methodology

Material examined. Specimens were collected mainly throughout mainland Southeast Asia, principally from Thailand, Laos, Cambodia, Myanmar, Singapore and Malaysia and kept at Chulalongkorn University Museum of Zoology, Bangkok, Thailand. Examination of additional Southeast Asian and other Oriental regional collections including available type material was based on both identified and previously undetermined specimens in several museums. All specimens were observed by using either a LEICA MZ 16A, Nikon SMZ25 or Olympus stereo-microscope. Morphological characters were photographed using montaged image stacks. Each morphological feature was serially captured with a Canon 700d linked to an automated calibration program, either Cell’D imaging or Helicon Focus on a desktop PC. In addition, illustration of some morphological variation was made by free-hand drawings.

Institutional abbreviations: (CUMZ), (MSNG), (NHMUK), (NHMW), (NHRS), (NMB) and (ZMUC).

Behaviour, biology and distribution. Feeding behaviour and brooding of eggs and hatchlings was observed and photographed both in the field and the laboratory. Characteristics of habitats and brood chambers are discussed in Lewis (1981), Mitić et al. (2012) and Siriwut et al. (2014).

The distributional ranges of all SE Asian Scolopendra species were reinvestigated based on field sampling, museum collections, and literature records from this region. Localities cited in the descriptions are arranged geographically and are separated in two sections:

I. A determined locality refers to the corrected name of a locality. In cases of inaccurate spelling and outdated names on old labels, we provide the corrected name in square parentheses based on resources from the internet and/or historical notes. Spellings of new collection localities in Thailand were transcribed by the Thai Romanization program (Wirote 2001). Latitude and longitude coordinates are given for each new collecting locality, tracked by a GPS conductor via a Garmin GPS travelling device.

II. An undetermined locality refers to a name that is localized at only a regional scale such as by region or country.

Distribution maps for each species include the records from recent field surveys and specimens from museum collections that provide sufficiently detailed locations. Each of those localities is marked by a filled symbol. Some localities from previous taxonomic work are included using a blank symbol. All undetermined localities and some specimens which were attributed only to a region, island or country have been excluded from the distribution maps.

Species identification and description. Morphological terminology follows the standardized terminology (Bonato et al. 2010). Taxonomic nomenclature and species identification followed Attems (1930b, 1938, 1953), Schileyko (1992, 1995, 2007), Shelley (2005), Shelley et al. (2005), Chao (2008), Lewis (2010b), Kronmüller (2012) and Siriwut et al. (2015a). The description of the genital segments is based on previous surveys of the genital system of Scolopendridae (Demange and Richard 1969, Iorio 2003).

Abbreviations for terminology applied to morphology used in descriptions and some comparison tables are as follow: PS; AP; SAP; LS; DS; V; VL; VM; M; DM; SP; ULBS. Abbreviated terminology for the genital region used in figures is as follows: IM; AV; GP; LA; LS; PN; SGS I; SGS II; TGS.

The list of synonyms for each Scolopendra species follows Chilobase (http://chilobase.biologia.unipd.it/). Diagnoses are revised from Siriwut et al. (2015a), and the range of geographical variation from the type, voucher specimens and previous surveys.

Phylogenetic reconstruction. Southeast Asian and some temperate Asian Scolopendra sequences were obtained from GenBank, based on previous phylogenetic analyses (Joshi and Karanth 2011, Vahtera et al. 2013, Siriwut et al. 2015a, b). We add more Scolopendra sequences from additional specimens collected during 2014 from various parts of the region (Table 1). DNA extraction methods follow Siriwut et al. (2015a). Three standard genes for centipede phylogeny (cytochrome c oxidase subunit I, 16S rRNA and 28S rRNA) were used to reconstruct phylogenetic trees. Maximum likelihood and Bayesian inference approaches were employed, using RAxML (Stamatakis 2006) and MrBayes (Huelsenbeck and Ronquist 2001, Ronquist et al. 2012), respectively. Standard statistical tests were applied to evaluate branch support (bootstrap support and posterior probability). Algorithms and parameter settings for both analyses followed protocols detailed previously by Siriwut et al. (2015a).

Table 1.

Scolopendra sequences used in phylogenetic reconstruction in present study. Abbreviation names of voucher ID codes refer to museum collections as follow: AMNH; CUMZ; MCZ; NHMUK. References: 1 = Siriwut et al. (2015a), 2 = Siriwut et al. (2015b), 3 = Vahtera et al. (2013).

Species	Taxon locality	Voucher ID number	COI	16S	28S	Reference
Scolopendra morsitans Linnaeus, 1758	Khon Kaen, Thailand	CUMZ 00339	KR705662	KR705600	KR705724	1
	Nan, Thailand	CUMZ 00340	KR705661	KR705599	KR705723	1
	Chonburi, Thailand	CUMZ 00341	KR705660	KR705598	KR705722	1
	Surin, Thailand	CUMZ 00342	KR705666	KR705604	KR705728	1
	Chiang Mai, Thailand	CUMZ 00343	KR705665	KR705603	KR705727	1
	Sa Kaeo, Thailand	CUMZ 00344	KR705664	KR705602	KR705726	1
	Sisophon, Cambodia	CUMZ 00345	KR705663	KR705601	KR705725	1
	Singapore	CUMZ 00315	KR705636	KR705574	KR705698	1
Scolopendra subspinipes Leach, 1816	Papua New Guinea	MCZ IZ-130685	KF676528	KF676488	–	3
Scolopendra subspinipes Leach, 1816	Martinique	AMNH LP3879, MCZ IZ-131452	HQ402554	HQ402502	–	3
Scolopendra cingulata Latreille, 1829	Spain	MCZ IZ-131446	HM453310	HM453220	–	3
Scolopendra dehaani Brandt, 1840	Lopburi, Thailand	CUMZ 00282	KR705689	KR705627	KR705751	1
	Ayutthaya, Thailand	CUMZ 00256	KR705688	KR705626	KR705750	1
	Lan Island, Rayong, Thailand	CUMZ 00320	KR705684	KR705622	KR705746	1
	Sa Kaeo, Thailand	CUMZ 00321	KR705682	KR705620	KR705744	1
	Trad, Thailand	CUMZ 00322	KR705681	KR705619	KR705743	1
	Sichang Island, Chonburi, Thailand	CUMZ 00252	KR705683	KR705621	KR705745	1
	Chiang Mai, Thailand	CUMZ 00323	KR705659	KR705597	KR705721	1
	Chiang Mai, Thailand	CUMZ 00346	KR705658	KR705596	KR705720	1
	Maehongson, Thailand	CUMZ 00324	KR705657	KR705595	KR705719	1
	Maehongson, Thailand	CUMZ 00325	KR705656	KR705594	KR705718	1
	Sakon Nakhon, Thailand	CUMZ 00247	KR705655	KR705593	KR705717	1
	Mahasarakarm, Thailand	CUMZ 00275	KR705651	KR705589	KR705713	1
	Loei, Thailand	CUMZ 00277	KR705653	KR705591	KR705715	1
	Ubon Ratchathani, Thailand	CUMZ 00248	KR705652	KR705590	KR705714	1
	Phatthalung, Thailand	CUMZ 00274	KR705641	KR705579	KR705703	1
	Nakhon Si Thammarat, Thailand	CUMZ 00281	KR705639	KR705577	KR705701	1
	Ranong, Thailand	CUMZ 00262	KR705637	KR705575	KR705699	1
	Phang Nga, Thailand	CUMZ 00251	KR705640	KR705578	KR705702	1
	Chumphon, Thailand	CUMZ 00326	KR705638	KR705576	KR705700	1
	Uthai Thani, Thailand	CUMZ 00243	KR705632	KR705570	KR705694	1
	Prachuap Khiri Khan, Thailand	CUMZ 00327	KR705628	KR705566	KR705690	1
	Kanchanaburi, Thailand	CUMZ 00328	KR705631	KR705569	KR705693	1
	Ratchaburi, Thailand	CUMZ 00253	KR705630	KR705568	KR705692	1
	Tak, Thailand	CUMZ 00329	KR705629	KR705567	KR705691	1
	Siem Reap, Cambodia	CUMZ 00330	KR705687	KR705625	KR705749	1
	Srisophon, Cambodia	CUMZ 00331	KR705686	KR705624	KR705748	1
	Attapue, Laos	CUMZ 00332	KR705678	KR705616	KR705740	1
	Champasak, Laos	CUMZ 00333	KR705673	KR705611	KR705735	1
	Luang Prabang, Laos	CUMZ 00334	KR705677	KR705615	KR705739	1
	Phongsaly, Laos	CUMZ 00335	KR705676	KR705614	KR705738	1
	Perak, Malaysia	CUMZ 00336	KR705669	KR705607	KR705731	1
	Kelantan, Malaysia	CUMZ 00337	KR705668	KR705606	KR705730	1
	Perak, Malaysia	CUMZ 00338	KR705667	KR705605	KR705729	1
Scolopendra multidens Newport, 1844	Qiang Binh, Vietnam	NHMUK, MCZ IZ-131459	KF676540	KF676485	–	3
Scolopendra calcarata Porat, 1876	Kanchanaburi, Thailand	CUMZ 00312	KR705650	KR705588	KR705712	1
	Wat Mae Long, Mae Chaem, Chiang Mai, Thailand	CUMZ 00417	KU512629	KU512632	KU512635	This study
	Lan Sang Waterfall, Mueang, Tak, Thailand	CUMZ 00418	KU512630	KU512633	KU512636	This study
Scolopendra japonica Koch, 1878	Matsumoto, Japan	CUMZ 00319	KR705679	KR705617	KR705741	1
	Xieangkhuang, Laos	CUMZ 00298.1-2	KR705671, KR705670	KR705609, KR705608	KR705733, KR705732	1
	Phongsaly, Laos	CUMZ 00297.1-2	KR705675, KR705674	KR705613, KR705612	KR705737, KR705736	1
Scolopendra pinguis Pocock, 1891	Bo Kaeo, Laos	CUMZ 00309	KR705646	KR705584	KR705708	1
	Kanchanaburi, Thailand	CUMZ 00303	KR705646	KR705584	KR705708	1
	Nan, Thailand	CUMZ 00307	KR705644	KR705582	KR705706	1
	Xieangkhuang, Laos	CUMZ 00306	KR705643	KR705581	KR705705	1
	Huaphun, Laos	CUMZ 00304	KR705642	KR705580	KR705704	1
	Chiang Mai, Thailand	CUMZ 00313	KR705649	KR705587	KR705711	1
	Mae Hong Son, Thailand	CUMZ 00314	KR705648	KR705586	KR705710	1
Scolopendra dawydoffi Kronmüller, 2012	Trad, Thailand	CUMZ 00272	KR705680	KR705618	KR705742	1
	Nakhon Ratchasima, Thailand	CUMZ 00290	KR705654	KR705592	KR705716	1
	Nakhon Ratchasima, Thailand	CUMZ 00294.1-2	KR705635, KR705634	KR705573, KR705572	KR705697, KR705696	1
Scolopendra cataracta sp. n.	Tad E-tu Waterfall, Bolaven Plateau, Pakse, Champasak, Laos	Holotype CUMZ 00316	KR705672	KR705610	KR705734	1
	Tad-Yueang Waterfall, Mueang Singh, Luang Namtha, Laos	Paratype CUMZ 00317	KR705633	KR705571	KR705695	1
	Kao Sok National Park, Surat Thani, Thailand	Paratype NHMUK 010305528	KU512631	KU512634	KU512637	This study
Cormocephalus monteithi Koch, 1983	Queensland, Australia	MCZ IZ-130638	HM453309.1	AF370861.1	HM453274	3
Digitipes kalewaensis Siriwut, Edgecombe & Panha, 2015	Kalewa, Sagaing, Burma	CUMZ 00234	KP204116	KP204112	–	2
Otostigmus astenus Kohlrausch, 1878	Fiji / Vanuatu	MCZ IZ-130669/130670	HM453312	HM453221	–	3
Sterropristes violaceus Muadsub & Panha, 2012	Similan, Thailand	MCZ IZ-130610	KF676519	KF676477	–	3

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Results

Phylogenetic relationships of mainland Southeast Asian Scolopendra and the position of Scolopendra cataracta sp. n.

The phylogenetic tree from the updated concatenated DNA dataset aggregates studied specimens into eleven monophyletic groups within Scolopendrinae that are compatible with morphological identification (Fig. 1: Clade A). The phylogeny supports the monophyly of the genus Scolopendra, in contrast to a previous analysis in which a sampled species of Cormocephalus fell within Scolopendra (Siriwut et al. 2015a). The sequence annotation for each partial marker is given in Table 2. Genetic divergence was calculated by pair-wise comparison of k-2 parameter distance under one thousand bootstrap replicates (Table 3). The genetic distance among Scolopendra species ranges from 15.9–24.4% in COI and 8.3–25% in 16S (COI 13.5–16.8% and 16S 19.3–23.0% for European Scolopendra by Oeyen et al. (2014)). Comparing with different genera from the same/another subfamily, the distances are between 21.6–28.9% and 23.1–26.9% in COI, and 22.2–26.1% and 25.9–34.1% in 16S, respectively. Within populations, intraspecific variation is between 8.3–18.4% in COI and 5.2–11.3% in 16S (Table 4).

Figure 1. — Maximum likelihood tree for *Scolopendra* in mainland Southeast Asia: colours for clades correspond to species and outgroups; black and white circles indicate statistical support values in both ML and BI analyses or only ML or BI analysis, respectively. Numbers at nodes are bootstrap support and posterior probability. Specimen codes in parentheses following localities correspond to Siriwut et al. (2015a, b: table 2) and Vahtera et al. (2013: table 1).

Table 2.

Sequence annotation of three partial genes used in this present study.

Molecular marker	Length	Parsimony informative sites	Variable sites	Conserved sites
COI	611	260	302	309
16S	446	214	271	175
28S	683	149	188	450

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Table 3.

Genetic distance between Scolopendra species in mainland Southeast Asia and outgroups; upper right and lower left distance collected from COI and 16S partial gene pairwise comparisons.

Species		16S
Species		1	2	3	4	5	6	7	8	9	10	11	12	13	14
COI	Scolopendra dehaani ¹		0.143	0.159	0.148	0.216	0.216	0.210	0.159	0.142	0.185	0.253	0.259	0.265	0.294
	Scolopendra cataracta ²	0.159		0.164	0.138	0.238	0.213	0.199	0.156	0.158	0.175	0.259	0.262	0.288	0.300
	Scolopendra dawydoffi ³	0.171	0.190		0.164	0.209	0.220	0.217	0.123	0.127	0.083	0.222	0.292	0.285	0.293
	Scolopendra subspinipes ⁴	0.167	0.194	0.177		0.229	0.238	0.212	0.162	0.166	0.174	0.257	0.271	0.303	0.291
	Scolopendra pinguis ⁵	0.219	0.247	0.215	0.224		0.171	0.240	0.246	0.230	0.233	0.234	0.280	0.298	0.322
	Scolopendra calcarata ⁶	0.228	0.240	0.212	0.230	0.219		0.241	0.237	0.223	0.250	0.255	0.270	0.278	0.315
	Scolopendra morsitans ⁷	0.204	0.230	0.204	0.233	0.244	0.242		0.214	0.215	0.231	0.274	0.323	0.341	0.290
	Scolopendra japonica ⁸	0.202	0.227	0.163	0.208	0.218	0.219	0.232		0.123	0.135	0.258	0.287	0.297	0.307
	Scolopendra cingulata ⁹	0.188	0.197	0.177	0.214	0.224	0.228	0.221	0.181		0.131	0.261	0.274	0.291	0.322
	Scolopendra multidens ¹⁰	0.196	0.203	0.107	0.178	0.216	0.214	0.229	0.163	0.188		0.230	0.304	0.326	0.323
	Cormocephalus ¹¹	0.236	0.263	0.234	0.238	0.269	0.239	0.267	0.243	0.231	0.233		0.266	0.296	0.322
	Digitipes ¹²	0.234	0.216	0.227	0.255	0.249	0.252	0.289	0.260	0.237	0.229	0.279		0.213	0.246
	Sterropristes ¹³	0.234	0.246	0.228	0.249	0.233	0.240	0.245	0.217	0.232	0.242	0.244	0.205		0.199
	Otostigmus ¹⁴	0.217	0.237	0.222	0.237	0.250	0.232	0.251	0.248	0.267	0.236	0.261	0.219	0.208

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Table 4.

Genetic distance under pairwise sequence comparison within populations of Scolopendra species in mainland Southeast Asia.

Species	COI	16S
Scolopendra dehaani	0.083	0.052
Scolopendra cataracta	0.165	0.087
Scolopendra dawydoffi	0.019	0.010
Scolopendra subspinipes	0.146	0.109
Scolopendra pinguis	0.184	0.113
Scolopendra calcarata	0.114	0.072
Scolopendra morsitans	0.086	0.068
Scolopendra japonica	0.128	0.061

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Three main clades are identified in mainland Asian Scolopendra (Fig. 1: Clade B), corresponding to pinguis-calcarata, subspinipes and morsitans groups (Fig. 1: Clade C, D and E). The highest intraspecific variation is observed in Scolopendra pinguis, a species native to this region, and the lowest variation is in Scolopendra dehaani, which is widespread and the dominant species in the region. The high measure of genetic divergence among Scolopendra pinguis populations in our previous study prompted a re-examination of those specimens and additional ones that were added in this study. The updated phylogenetic tree revealed that the former Scolopendra pinguis clade (Siriwut et al., 2015a: fig. 1, clade C) can be divided into two species, Scolopendra pinguis and Scolopendra calcarata (Fig. 1: Clade C), and this separation can also be supported by diagnostic morphological characters. These two species are distributed along the montane areas of Burma and Thailand and occur eastward to the Indochina sub-region in mountain ranges between Laos and Vietnam. Synapomorphic characters shared by these two species are the comparatively robust, vaulted shape of their body segments, the tergite of the ultimate leg-bearing segment being acute posteriorly, four glabrous antennal articles, and the dichromatic colouration on the cephalic plate in all Scolopendra calcarata specimens and most Scolopendra pinguis populations. In addition, the phylogeny indicates that even after re-categorising a closely related species (Scolopendra calcarata) that had previously been classified as an aberrant clade within Scolopendra pinguis, the genetic distance within Scolopendra pinguis is still considerable. This distance might suggest cryptic speciation among different geographical populations.

The remaining Scolopendra species may be divided into two groups, one consisting of Scolopendra morsitans and another including former subspecies of Scolopendra subspinipes sensu Kronmüller (2012). In the case of Scolopendra morsitans, monophyly is corroborated with high bootstrap support and posterior probability in ML and BI analyses, respectively (Fig. 1: Clade D). Within the Scolopendra subspinipes group (Fig. 1: Clade E), a clade uniting Scolopendra cingulata Latreille, 1829, Scolopendra japonica, Scolopendra dawydoffi and Scolopendra multidens (Fig. 1: Clade F) differs from a clade composed of Scolopendra subspinipes, Scolopendra cataracta and Scolopendra dehaani (Fig. 1: Clade I). A morphological feature shared by Scolopendra cingulata, Scolopendra japonica, Scolopendra dawydoffi and Scolopendra multidens is the cingulata-like ultimate legs, which have a dorsally flattened prefemur and femur and are much shorter and stouter than in the subspinipes clade (Fig. 1: Clade I). Attems (1938) referred to similar groupings based on form of the ultimate legs when describing Scolopendra dawydoffi. The number of apical spines on the coxopleural process might also be useful for discrimination of these two groups (one or two versus more than two spines in the subspinipes and cingulata groups, respectively). A monophyletic group composed of Scolopendra subspinipes and the two allied species (Scolopendra cataracta and Scolopendra dehaani) received statistical support both in ML and BI (Fig. 1: Clade I).

All samples of Scolopendra cataracta united as a clade with Scolopendra dehaani to the exclusion of Scolopendra subspinipes (Fig. 1: Clade J). This new Scolopendra species shares the following morphological similarities with various species of the Scolopendra subspinipes group: presence of two ventro-lateral spines on the ultimate leg prefemur (as in Scolopendra subspinipes), long and slender ultimate legs (like Scolopendra dehaani), and incomplete paramedian sutures on the sternites (like Scolopendra dawydoffi). However, Scolopendra cataracta is clearly distinguished from all of them by extremely short tergal paramedian sutures. Interspecific variation of DNA sequences ranges between 15.9–19.4% and 13.8–16.4% in COI and 16S, respectively, among these three related species.

Species diversity of Scolopendra in mainland Southeast Asia

In this region, nine species are identified from our survey. The taxomomic boundaries between species were based on information from both morphology and molecular analysis. Two other species of Scolopendra were not included in this paper, namely Scolopendra mirabilis (Porat, 1876), and Scolopendra hardwickei Newport, 1845. In the case of Scolopendra mirabilis, an African-central Asian species, the single known specimen in SE Asia may be introduced, being found on an island in a coastal area of northern Vietnam (Schileyko 1995). Likewise Scolopendra hardwickei was reported from Singapore, the largest port in Southeast Asia (Decker 2013). This species has been documented from India and the Nicobar and Andaman islands, and it probably occurs in Sumatra and Java (Khanna 2001, Lewis 2010b, Decker 2013). Without further material from Singapore or neighbouring areas, the status of this recorded species in the mainland SE Asian fauna is questionable. Scolopendra gracillima sternostriata Schileyko, 1995, from Vietnam (Schileyko 1995, 1997), is similar to Scolopendra pinguis in most respects. We include it in the key below but have no new material of this subspecies, and accordingly have not revised it. The following key to native species of Scolopendra excludes only the two first aforementioned species.

Key to species of Scolopendra in mainland Southeast Asia

1	Tergite of ultimate leg-bearing segment with median suture	Scolopendra morsitans Linnaeus, 1758
–	Tergite of ultimate leg-bearing segment without median suture	2
2	Sternal paramedian sutures complete	3
–	Sternal paramedian sutures incomplete	4
3	Coxopleural process with three or more apical spines, thick prefemur of ultimate leg with at least two ventro-lateral and four spines on prefemoral process; average ratio of width:length of ultimate leg prefemur1:2	Scolopendra japonica Koch, 1878
–	Coxopleural process with 1–2 apical spines, slender prefemur of ultimate leg with two ventro-lateral and two spines on prefemoral process; ratio of width:length of ultimate leg prefemur 1:3	Scolopendra subspinipes Leach, 1816
4	Ultimate leg prefemur with at least one ventro-lateral spine	5
–	Ultimate leg prefemur without ventro-lateral spines	Scolopendra dehaani Brandt, 1840
5	Coxopleural process with one lateral and one dorsal spine; ultimate leg prefemur with numerous small scattered spines	6
–	Coxopleural process without lateral and dorsal spines; ultimate leg prefemur with a few enlarged spines in rows	8
6	Legs 21 with a tarsal spur	Scolopendra calcarata Porat, 1876
–	Leg 21 without tarsal spurs	7
7	Sternites of anterior body segments with complete paramedian sutures	Scolopendra gracillima sternostriata Schileyko, 1995
–	All sternites with incomplete paramedian sutures, reaching only 20–30% on anterior part of sternites	Scolopendra pinguis Pocock, 1891
8	Complete paramedian sutures on tergites	9
–	Short, incomplete paramedian sutures confined to anterior and posterior parts of tergites	Scolopendra cataracta Siriwut, Edgecombe & Panha, sp. n.
9	Cephalic plate and tergite 1 densely punctate; tergites with short median sulcus on posterior part	Scolopendra multidens Newport, 1844
–	Cephalic plate and tergite 1 sparsely punctate; tergites without median sulcus on posterior part	Scolopendra dawydoffi Kronmüller, 2012

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Systematics

Family Scolopendridae Leach, 1816: Subfamily Scolopendrinae Kraepelin, 1903: Genus Scolopendra Linnaeus, 1758

Scolopendra morsitans

Linnaeus, 1758

Figs 1 , 2 , 3 , 4 , 5 , 6 , 7

Figure 2. — Colouration pattern during developmental stages of most *Scolopendra* *morsitans* populations in mainland Southeast Asia: A Juvenile stage B Adult stage (Colour morph 1).

Figure 3. — *Scolopendra* *morsitans* (*CUMZ* 00344): **A–C** Spiracles 3, 5 and 8, respectively D Tooth-plates E Cephalic plate and trunk segments 1–2 F Forcipular segment G Tergites 9–11.

Figure 4. — *Scolopendra* *morsitans* (*CUMZ* 00344, *NHMUK* 1889.7.15.14): A Sternites 9–11 B Coxopleura and ventral view of ultimate leg prefemora C Tergite of ultimate leg-bearing segment D Dorsal view of ultimate legs E Sternite of ultimate leg-bearing segment and coxopleura.

Figure 5. — *Scolopendra* *morsitans* (*CUMZ* 00300, 00344): A Cephalic plate and basal antennal articles B Tergites 9–11 C Teeth on tooth-plates and trochanteroprefemoral process D Forcipular segment E Sternites 9–11.

Figure 6. — *Scolopendra* *morsitans* (*CUMZ* 00300, 00344): A Tergite of ultimate leg-bearing segment B Lateral view of coxopleuron C Sternite of ultimate leg-bearing segment, coxopleura and ultimate legs D Spines on prefemoral process of ultimate leg **E–F** Spines on ultimate leg prefemora and margination on prefemora and femora (dorsal view).

Figure 7. — Genital segments in some live *Scolopendra* specimens: A *Scolopendra* *dehaani* (male) B *Scolopendra* *dawydoffi* (male) C *Scolopendra* *morsitans* (male) D *Scolopendra* *japonica* (female) E *Scolopendra* *pinguis* (male) F *Scolopendra* *pinguis* (female).

Scolopendra morsitans Linnaeus, 1758: 638. Newport 1844: 97, 1845: 378. Koch 1847: 163. Wood 1862: 23. Kohlrausch 1881: 104. Meinert 1886: 200. Haase 1887: 52, pl. 3, figs 52–54. Daday 1889: 150, 1891: 150. Silvestri 1895: 714. Kraepelin 1903: 250. Attems 1907: 80, 1909: 13, 1914a: 106, 1927: 61, 1930b: 23, Figs 38–39, 1930c: 175 1932: 5, 1938: 334. Brölemann 1912: 54. Muralewicz 1913: 200. Takakuwa 1942a: 359, 1942b: 15, 1942c: 41, 1943: 171, 1947: 936. Bücherl 1974: 107. Chamberlin and Wang 1952: 180. Jangi 1955: 597–607, 1959: 253–257. Wang 1955a: 198, 1955b: 16, 1956: 158, 1957: 27, 1962: 101, 1965: 450, 1967b: 391. Würmli 1975: 201–206. Koch 1983a: 79–91. Jangi and Dass 1984: 29, fig. 1. Lewis 2002: 81, 2010b: 107, figs 4, 33, 34. Shelley 2005: 39, figs 57–64, 2006: 5. Shelley et al. 2005: 39–58. Schileyko 2007: 75. Akkari et al. 2008: 83, map. 2. Decker 2013: 19. Tran et al. 2013: 228. Chagas-Júnior et al. 2014: 138.
Scolopendra brandtiana Gervais, 1837: 16. Newport 1845: 379.
Scolopendra bilineata Brandt, 1840: 155. Kohlrausch 1881: 108.
Scolopendra crassipes Brandt, 1840: 153. Kohlrausch 1881: 108.
Scolopendra erythrocephala Brandt, 1840: 155. Kohlrausch 1881: 108.
Scolopendra limbata Brandt, 1840: 154. Kraepelin 1903: 250.
Scolopendra platypus Brandt, 1840: 153. Newport 1844: 97, 1845: 379.
Scolopendra elegans Brandt, 1841: 21. Kohlrausch 1881: 107.
Scolopendra fulvipes Brandt, 1841: 22. Kohlrausch 1881: 108.
Scolopendra morsitans scopoliana C.L. Koch, 1841: 222, pl. 11. Akkari et al. 2008: 83.
Scolopendra angulipes Newport, 1844: 97. Kohlrausch 1881: 108.
Scolopendra leachii Newport, 1844: 97. Kraepelin 1903: 251.
Scolopendra longicornis Newport, 1844: 97. Kohlrausch 1881: 109.
Scolopendra platypoides Newport, 1844: 97. Kohlrausch 1881: 111.
Scolopendra tuberculidens Newport, 1844: 97. Kohlrausch 1881: 108.
Scolopendra algerina Newport, 1845: 387. Akkari et al. 2008: 83.
Scolopendra fabricii Newport, 1845: 384. Kohlrausch 1881: 107.
Scolopendra formosa Newport, 1845: 383. Kohlrausch 1881: 108.
Scolopendra richardsoni Newport, 1845: 385. Kohlrausch 1881: 109.
Scolopendra tigrina Newport, 1845: 381. Kohlrausch 1881: 108.
Scolopendra varia Newport, 1845: 380. Kohlrausch 1881: 111.
Scolopendra tongana Gervais, 1847: 275. Kohlrausch 1881: 110.
Scolopendra infesta Koch, 1847: 169. Kohlrausch 1881: 112.
Scolopendra planipes Koch, 1847: 168. Kohlrausch 1881: 106.
Scolopendra pella Wood, 1861: 13. Kohlrausch 1881: 111.
Scolopendra porphyratainia Wood, 1861: 15. Kohlrausch 1881: 108.
Scolopendra brachypoda Peters, 1862: 529, pl. 33, fig. 2. Kohlrausch 1881: 107.
Scolopendra mossambica Peters, 1862: 527, pl. 33, fig. 4. Kohlrausch 1881: 107.
Scolopendra compressipes Wood, 1862: 31. Haase 1887: 52.
Scolopendra modesta Wood, 1862: 29. Kraepelin 1903: 251.
Scolopendra carinipes Humbert & Saussure, 1870: 204. Kohlrausch 1881: 111.
Scolopendra afzelii Porat, 1871: 1146. Meinert 1886: 200.
Scolopendra attenuata Porat, 1871: 1148. Meinert 1886: 200.
Scolopendra chlorocephala Porat, 1871: 1149. Meinert 1886: 200.
Scolopendra cognata Porat, 1871: 1145 Meinert 1886: 200.
Scolopendra intermedia Porat, 1871: 1145. Meinert 1886: 200.
Scolopendra picturata Porat, 1871: 1144. Meinert 1886: 200.
Scolopendra pilosella Porat, 1871: 1148. Meinert 1886: 200.
Scolopendra saltatoria Porat, 1871: 1151. Meinert 1886: 200.
Scolopendra vaga Porat, 1871: 1151. Kraepelin 1903: 251.
Scolopendra wahlbergi Porat, 1871: 1150. Meinert 1886: 200.
Eurylithobius slateri Butler, 1876: 446. Kraepelin 1903: 250.
Scolopendra impressa Porat, 1876: 12. Meinert 1886: 200.
Scolopendra morsitans procera Haase, 1887: 53, pl. 33, fig. 53. Kraepelin 1903: 250.
Scolopendra morsitans sulcipes Haase, 1887: 54, pl. 33, fig. 54. Kraepelin 1903: 250.
Scolopendra morsitans calcarata Daday, 1891: 150. Kraepelin 1903: 250.
Scolopendra grandidieri Saussure & Zehntner, 1902: 302, pl. 3, fig. 13, pl. 12, fig. 6. Kraepelin 1903: 251.
Scolopendra lineata Saussure & Zehntner, 1902: 308, pl. 15, fig. 19. Kraepelin 1903: 251.
Scolopendra spinosella Saussure & Zehntner, 1902: 308, pl. 2, fig. 11. Kraepelin 1903: 251.
Scolopendra morsitans fasciata Attems, 1930a: 372. Würmli 1975: 205.
Scolopendra morsitans amazonica Bücherl, 1946: 135. Würmli 1975: 205.
Trachycormocephalus jodhpurensis Khanna, 1977: 154, figs 5–8. Jangi and Dass 1980: 67.

Type locality.

India.

Material.

Thailand — CUMZ 00343, one spm., Hui Hong Khrai, Chiang Mai (18°50'59.5"N, 99°13'16.4"E). CUMZ 00340, one spm., Lainan, Weing Sa, Nan (18°34'16.1"N, 100°46'59.7"E). CUMZ 00405, one spm., Wat Khao Isan, Pak Tho, Ratchaburi (13°23'2.458"N, 99°46'16.525"E). CUMZ 00302, two spms., Wat Mahavanh, Buriram (14°41'09.8"N, 102°52'33.8"E). CUMZ 00342, one spm., Ban Khok Pho, Prasat, Surin (14°32'53.4"N, 103°22'19.1"E). CUMZ 00339, one spm., Ban Dan Chang, Ta Kantho, Kalasin (16°50'06.1"N, 103°16'32.0"E). CUMZ 00409, one spm., Nong Bo, Borabue, Maha Sarakham (16°1'35.695"N, 103°7'42.487"E). CUMZ 00410, two spms., Ban Tha Tum, Mueang, Maha Sarakham (16°10'45.231"N, 103°27'4.134"E). CUMZ 00411, one spm., Wat Pa Sai Mun, Sai Mun, Yasothon (15°56'45.092"N, 104°12'1.929"E). CUMZ 00412, two spms., Wat Tham Pha Koeng, Phu Wiang, Khon Kaen (16°42'10.303"N, 102°14'56.901"E). CUMZ 00414, two spms., Wan Tham Chia, Nong Ruea, Khon Kaen (16°32'27.014"N, 102°33'19.454"E). CUMZ 00413, two spms., Phu Wiang National Park, Nong Bua Rawe, Chaiyaphum (16°41'4.089"N, 102°14'38.477"E). CUMZ 00299, one spm., Wang Bua, Kabin Buri, Prachinburi (13°57'16.3"N, 101°36'37.3"E). CUMZ 00406, one spm., Wang Bo Waterfall, Mueang, Prachin Buri (14°10'33.933"N, 101°25'40.163"E). CUMZ 00407, 16 spms., Khram Yai Island, Sattahip, Chon Buri (12°42'18.095"N, 100°50'29.35"E). CUMZ 00341, one spm., Juang Island, Sattahip, Chonburi (12°31'46.4"N, 100°57'18.4"E). CUMZ 00408, three adult and numerous juvenile spms., Ta Phraya, Sa Kaeo (14°5'1.047"N, 102°45'36.389"E). CUMZ 00300, two spms., Mueang, Sa Kaeo (13°49'07.9"N, 102°03'10.5"E). CUMZ 00344, one spm., Tha Kra Bak Reservoir, Sa Kaeo (13°58'13.9"N, 102°15'57.6"E). CUMZ 00301, one spm., Kuiburi, Prachuab Khiri Khan (12°06'32.0"N, 99°45'53.0"E). CUMZ 00403, 12 spms., Hat Wanakon National Park, Tab Sakae, Prachuab Khiri Khan (11°38'6.012"N, 99°42'5.25’’). CUMZ 00404, five spms., Kui Buri National Park, Kui Buri, Prachuab Khiri Khan (12°8'57.096"N, 99°45'34.433"E). CUMZ 00402, two spms., Ban Laem Sai, Chaiya, Surat Thani (9°24’ 9.691"N, 99°17'19.719"E). CUMZ 00400, one spm., Ching Kho, Singhanakhon, Songkhla (7°16'44.261"N, 100°31'36.759"E). CUMZ 00401, eight spms., Nhai Plao Beach, Khanom, Nakhon Si Thammarat (9°7'52.64"N, 99° 52’ 39.415"E). NHMUK 1897.9.7.29, one spm., Betong, Yala, leg. S.S. Flower. NHMUK, one spm., Ko Kraew [Khao Kaeo, Chonburi]. NHMUK, one spm., Ko Kraam [Kram Islands, Chonburi], leg. S.S. Flower, 1897–1898.

Cambodia — CUMZ 00345, one spm., Wat Phanombak, Srisophon (13°36'05.5"N, 102°57'09.3"E). NHMW, two spms., Mount Cardamones [Cardamom Mountain], 500 m above sea level, Mission Dawydoff, April 1893.

Laos — CUMZ 00419, two spms., Ban Na Ka-Som, Attapue (14°48'30.477"N, 106°50'49.948"E), CUMZ 00420, two spms., Wat Kao Kaeo, Pakse, Champasak (14°11'30.572"N, 105°54'30.821"E), CUMZ 00421, two spms., Savannhaket (16°38'25.053"N, 104°50'0.994"E)

Myanmar — CUMZ 00415, two spms., Old Bagan, Bagan (21°10'19.161"N, 94°51'34.61"E). CUMZ 00416, two spms., Kyaing, Pakokku (21°52'37.982"N, 94°37'47.384"E). NHMUK, one spm., Pyrimana [Pyinmana], Upper Burma. NHMUK 1889.7.15.14-18, 21 spms., Teikiyi (Rangoon) [Yangon], leg. E.W. Oates. NHMUK 1889.7.15-16, two spms., Moulmein. NHMUK 1889.7.15.17, five spms., Townwingyi (upper Burma) [Taunggyi, Shan State], leg. E.W. Oates. NHMUK 1889.7.15.15, six spms., Iharrawady [Ayeyarwady Region], leg. E.W. Oates. NHMUK 1889.7.15-20, 20 spms., Mandalay, leg. E.W. Oates (Cap). NHMW Inv. No. 671, one spm., Osl-Indien, Aracan [Rakhine State], leg. Stoliczka, 1873.

Vietnam — NHMUK 1926.9.30.13, one spm., Thai Nien Basin, Heure Range, Tonkin, leg. Sladen-Godman, Trust Expedition. NHMUK, one spm., Annam, with label “?19”.

Brunei — NHMUK 1973.7.659, one spm., Jerudong, with label “CIE Coll. A6527”. NHMW Inv. No. 641, one spm., Brunei, don. Stimdarlmus.

Philippines — NHMUK, one spm., Mactan, Zebu Island. NHMUK 1913.6.18.851-853, one spm., Philippines. NHMW Inv. No. 646, two spms., Manila, Novara Expedition.

Indonesia — NHMW Inv. No. 659, one spm., Padang, Sumatra, leg. Comal Fehiel, 1901. NHMW Inv. No. 12?, one spm., Java, with label “adeusawer 8”. NHMW Inv. No. 636, five spms, Batavia, Java, Novara Expedition, 1857–1859. NHMUK 1913.6.18.849, one spm., Celebes, with label “Spec. No. 28”. NHMW Inv. No. 632, one spm., Celebes, leg. Beruh. Walt, 3 January 1894. NHMUK, one spm., Seram [Seram Island, Maluku Province], leg. Dr. R.F. Ellen, det. D. MacFarlane, 1975. NHMW Inv. No. 635, one spm., Amboina [Ambon], leg. Doleschal, 1859.

China — NHMUK 1928.3.16.41-42, two spms., Amoy, leg. Prof. B. Ping 4/2/1926. NHMUK, two spms., China. NHMW Inv. No. 652, one spm., Hong Kong, April 1901. NHMW Inv. No. 638, two spms., Takao [Kaohsiung, Taiwan], leg. H. Sauler. NHMW Inv. No. 643, one spm., Fumasra [Formosa: Taiwan], leg. Breitenstein, 1884.

Japan — NHMUK 1913.6.18.853, one spm., Japan, leg. Koch. NHMUK 1892.10.10, one spm., Loochoo [Ryukyu Islands], leg. Holst. NHMW Inv. No. 647, two spms., Ishikagi-Jiwa, Liu Kiu Island [Ryukyu Islands], leg. H. Sauler.

Indian and Middle Asian Territory — NHMUK 1930.4.11.14, one spm., Ahmadabad, leg. Capt. J.B.E. Manning I.M.S. NHMUK, one spm., North Behar, Champharam, leg. Mrs. Campbell Martens with note “see letter 24/9/1930”. NHMUK, one spm., Mandras in 1925, leg. F.A. Turk. NHMUK, one spm., on porch at Bombay, leg. N.H. Soc, 1/8/1902. NHMUK 1894.10.24.70-73, five spms., Madras, leg. J.R. Henderson. NHMUK 1910.4.10.31, one spm., Ceylon, Bainbridge Fletcher’s collection. NHMUK 1903.6.18.848, one spm., Bombay. NHMUK 1948.8.6.5, one spm., Kasual [Kansal], Punjab, leg. S.F. Woodward. NHMUK 1975.12, one spm., Ceylon. NHMUK, one spm., Andaman Island, leg. B.B. Osmaston and P.A. Buxton, London School of Hygiene and Tropical Medicine. NHMUK, one spm., Northern Baluchiotan [Balochistan, Pakistan], brought to Indian Museum, leg. Prof. P.A. Buxton, Indian School of Tropical Medicine. NHMUK, one spm., Calcutta, leg. Dr. S.P.R. Chaudhuri, relocated in 25.4.49. NHMUK, one spm., Indian Ocean, with label “Assumption of J.S. Gardiner in 1952.12.17”. NHMUK, 10 spms., Aldaima, Maldives, November, 1908, det. J.S. Gardiner. NHMUK, two spms., Aldaima, Maldives, 17/12/1952. NHMUK 1952.12.17.247, one spm., Astove Island, Indian Ocean, leg. J.S. Gardiner. NHMUK 1948.10.11.8-4, five spms., Delhi, leg. J.H. Graham. NHMUK 1896.10.2.3-4, two spms., Assam, leg. E.W.P. Cambridge. NHMW Inv. No. 665, one spm., Calcutta, leg. Stoliezka, 1865. NHMW Inv. No. 650, one spm., Kamoly?, Ceylon, leg. M. Hoelui, September, 1892. NHMW Inv. No. 673, two spms., Osindien [East India], with label “Parr.”. NHMW Inv. No. 642, one spm., Kagi Island, Maldives, leg. H. Sauler.

Africa — NHMUK, one spm., imported with bananas from West Africa, leg. J. Knight Co. Ltd Bermondsey, Indian S.E. on 17/5/1952. NHMUK, one spm., St. Helena Island, South Atlantic Ocean. NHMUK 1892.5.16.1, one spm., Delagoa Bay [Maputo], leg. J. de Coster. NHMUK 1954.7.5.17, one female, Tanga, Tanganyika territory, Tanzania, leg. R.H.C. Sweeney. NHMUK, two spms., St. Helena, leg. A. Loveridge. NHMUK, one spm., Shinyanga [Shinyanga, Tanzania?], Tanganyika, leg. P. Gettliffe.

Australia — NHMW Inv. No. 637, two spms., Gayndah [Queensland], New-Holland. NHMW Inv. No. 666, one spm., Neu Holland, leg. Dr. Millas, 1884.

South Pacific — NHMUK 1893.11.15.1-2, two spms., Tongatapu. NHMUK 1975.65, one spm., Samoan Island [Samoa]. NHMUK 1966.72, two spms., Samoan Island. NHMUK 1892.12.27.11-13, two spms., Levuka, Fiji, leg. H. Hjorring, det. G.M. Thomson. NHMUK 1913.6.18.850, one spm., Vamma Levu [Vanua Levu, Fiji]. NHMUK 1976.31, two spms., Samoan Island. NHMW Inv. No. 634, one spm., Tahiti. NHMW Inv. No. 649, one spm., Tahiti, Museum Goddefroi, 3/3/1881.

Undetermined locality — NHMW Inv. No. 640, one spm., Yantempo, leg. H. Sauler. NHMUK 1985.29, three spms., with label “Lnas”.

Diagnosis.

17–23 antennal articles, 5–8 basal articles glabrous dorsally. Each tooth-plate with 5–6 teeth. Tergites 7(12)-20 with paramedian sutures. Tergite of ultimate leg-bearing segment with median suture. Complete paramedian sutures on sternites 2–20. Coxopleural process with 3–4 apical and 0–1 lateral spines. Ultimate leg prefemora with three rows of ventral spines (2–6 VL, 3 V, 2–6 VM), 2–6 M, 2–6 DM and 0–8 spines on prefemoral process. One tarsal spur on legs 1–19 (in Southeast Asia).

Composite description.

Body length up to 12.7 mm (In Australian populations according to Koch (1983a)). Reddish-brown or yellowish colour on body segments. Cephalic plate and tergites monochromatic or dichromatic in adult (Fig. 2B). Tergites usually reddish-orange (Thai, Laos and Cambodian populations); dark band on posterior border of tergites. Cephalic plate with or without small punctae, median sulcus present on anterior part. Posterior part of cephalic plate without paramedian sutures.

Antenna usually with 18–20 articles (sometimes 17, 21 or 23 on one side), basal 5–7 glabrous dorsally (Figs 3E, 5A), 5–8 articles glabrous ventrally. Antennae reach to segment 4. Forcipular trochanteroprefemoral process (Figs 3F, 5D) with denticles in two groups, 2–3 apical and one inner. Tooth-plates wider than long or nearly as long as wide, usually 5–6 teeth (Fig. 3D); rarely 3, 4 or 7. Tooth-plate with straight, transverse basal suture (Fig. 5C). Coxosternite without median suture. Article 2 of second maxillary telopodite with spur.

Anterior margin of T1 underlying cephalic plate (Fig. 3E). Complete paramedian sutures from TT4–5; margination typically starting on T14 (one spm., with margination restricted restricted to last two tergites). Tergite surface (Figs 3G, 5B) smooth. Tergite of ultimate leg-bearing segment (Figs 4C, 6A) curved posteriorly, with median suture; ratio of width: length of tergite of ultimate leg-bearing segment 1.34:1. Sternites (Figs 4A, 5E) with complete paramedian sutures. Sternites without depressions. Sternite of ultimate leg-bearing segment (Fig. 4E) with sides converging posteriorly; surface without depression. Pore-field on coxopleuron terminating well beneath margin of tergite of ultimate leg-bearing segment, pore area slightly widened anteriorly.

Coxopleural process moderately long or short, usually with 4–5 apical and 0–1 lateral spines (Fig. 4E); pore-free area extending 40–50% length from distal part of coxopleural process to margin of sternite of ultimate leg-bearing segment (Fig. 6B).

All legs without setae and tibial spurs. One tarsal spur on legs 1–19 (20 in some African and Indian populations). Ultimate legs: thick and moderately long, with ratios of lengths of prefemur and femur 1.2:1, femur and tibia 1.3:1, tibia and tarsus 2 1.7:1.; tarsus 1 and tarsus 2 2.8:1. In male, lateral margin of prefemora, femora and tibia marginated dorsally. Prefemoral spines (Figs 4B, D, 6C–F): 2–4 VL, 3 V, 2–4 VM, 2–3 M, 2–3 DM and prefemoral process usually with 3–5 spines. Posterior margin of prefemur with shallow median groove.

Genital segments well developed, reaching longer than the distance between posterior margin of sternite of ultimate leg-bearing segment and distal part of coxopleural process (Fig. 7C). Sternite of genital segment 1 round and convex posteriorly, with median suture. In male, sternite of genital segment 2 attached to penis. Tergites of genital segments without small setae. Gonopods with small setae in male. Penis with fine posterior seta.

Colouration. Scolopendra morsitans demonstrates colour variation among its populations in SE Asia. Previously, colour variation has been recorded in African, Australian and Taiwanese populations (Lewis 1968, Koch 1983a, Chao 2008), those studies proposing that latitude and habitat composition might affect this variability. Recent molecular analyses of Thai-Cambodian Scolopendra morsitans suggested that some colour morphs may be specific to local populations (Siriwut et al. 2015a), although similar patterns occur in each of three different continental faunas. We have recorded the colouration pattern in juvenile and adult specimens (Fig. 2A–B, respectively):

Colour morph 1. Dichromatic. Cephalic plate, T1 and tergite of ultimate leg-bearing segment orange, the remaining tergites brownish. Posterior borders and lateral margins of tergites dark. Antenna bright orange. Pleuron of leg-bearing segments with pale grey integument, pleurites orange. Legs 1–21 orangish or yellow. Ultimate legs orangish or light brown.

Colour morph 2. Dichromatic. Cephalic plate, T1 and tergite of ultimate leg-bearing segment dark brown or blackish, the remaining tergites brownish. Posterior borders and lateral margins of tergites dark. Antenna dark blue. Pleuron of leg-bearing segments with pale grey integument, pleurites orange or brown. Legs 1–20 yellowish or pale. Ultimate legs blackish or brown.

Discussion.

Scolopendra morsitans is morphologically varied and subsumes many synonyms that are now attributed to geographical and/or ontogenetic variation. Intraspecific variation has been studied in Africa (Lewis 1969), India (Jangi 1955, 1959), and Australia (Koch 1983a), revealing that some diagnostic characters are inconsistent within its populations. These include: number of glabrous antennal articles, number of teeth on the forcipular tooth-plates, number of tergites that are marginated, and the number of legs with tarsal spurs. This species also demonstrates differences in colour patterns that might be correlated with its geographical distribution. Lewis (1969) noted that a population of Scolopendra morsitans from Bihe, Angola, demonstrated a dark body with red legs whereas specimens from Sudan were straw-coloured. Here we record two colouration patterns in Thai populations that do not occur sympatrically. In addition, some morphological characters might be restricted to certain geographical populations, such as a tarsal spur on leg 20, which has been reported from India and in some African populations. For this reason, the utility of this character for defining boundaries between Scolopendra morsitans and other Scolopendra species that share some morphological characters with it, such as Scolopendra laeta Haase, 1887 and Scolopendra antananarivoensis, is not absolutely clear. Our survey of geographic variation in Scolopendra morsitans is presented in Table 5.

Table 5.

Geographical variation in several populations of Scolopendra morsitans in Old World territory including Australia. ? insufficient data.

Character	India³	Burma¹	Indochina^1,2	Malay Archipelago^1,2	Philippines¹	East Asia^1,6	Australia⁵	Africa^2,4
Number of antennal articles	19–20	18–22	20–21	18–20	17–19	18–20	17–23	17–21
Number of glabrous articles	6–9	6–7	5	6–7	6–7	5–7?	3–8	5–7
Teeth on tooth-plate	5	4–5	3–7	4–5	4–5	5	3–6	5
First tergite with complete paramedian sutures	2	3	4–5	2–3	2–3	3	2–4	2–4
First tergite with margination	7–17	6–17	5–14	6–13	12–13	10–14	5	2–7 (15)
Tergite surface	smooth	smooth	smooth	smooth	smooth	smooth	smooth	smooth
Median furrow on tergite of ULBS	present	present	present	present	present	present	present	present
Paramedian sutures on sternites	incomplete	incomplete	incomplete	incomplete	incomplete	incomplete	incomplete	incomplete
Sternite of ULBS	?	without depression	without depression	without depression	without depression	without depression	without depression	?
Spines on coxopleural process	AP: 3–5 LS: 0–1	AP: 3–5 LS: 1	AP: 1–5 SAP: 0–1 LS: 0–1	AP: 3–5 LS: 0–1	AP: 4–5 LS: 0–1	AP: 4 LS: 0–1	AP: 2–6	AP: 2–6 LS: 0–1
Spine formula on prefemora of ultimate legs	VS: 3 rows DS: 4–5 2 rows (4–5) SP: 4–10	V: 5–10 (3 rows) M: 2–4 DM: 2–4 SP: 3–7	V: 5–10 (3 rows) M: 0–6 DM: 0–5 SP: 2–6	V: 5–10 (3 rows) M: 2–4 DM: 2 SP: 3–7	V: 9–10 (3 rows) M: 2–4 DM: 2 SP: 3–7	V: 7–9 (3 rows) M: 0–3 DM: 2–6 SP: 0–4	VL: 2–6 VM: 2–8 M: 2–6 DM: 2–6 SP: 4–8	VL: 6–12 M: 2–6 DM: 3–6 SP: 3–8
Legs with one tarsal spur	1–19(20)	1–19	1–19	1–19	1–19	1–19	1–19	1–19(20)

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Note: each superscript number refers to description in previous and present studies as follow; ¹ = this study, ² = Attems (1930b, 1953), ³ = Jangi and Dass (1955), ⁴ = Lewis (1967, 1969), ⁵ = Koch (1983), ⁶ = Chao (2003, 2008).

Our phylogenetic analysis corroborates the monophyly of SE Asian populations of this species (Fig. 1). Previous molecular phylogenetic analyses of the Scolopendra morsitans complex in India suggested that Scolopendra morsitans was paraphyetic with respect to specimens that were determined as Scolopendra amazonica, the latter name being used for specimens with tarsal spurs on leg 20 (Joshi and Karanth 2011). From these results, it seems that molecular phylogenetics of this species complex throughout its geographic range may be necessary to clarify the taxonomic value of some variable morphological characters and to more confidently determine the taxonomic status of some phenotypically similar species. It is likely that some names currently treated as junior subjective synonyms of Scolopendra morsitans may be found to be applicable to cryptic species.

Distribution.

This is one of the oldest described centipede species and it is distributed worldwide in the tropics.The native distribution is difficult to determine because of assumed introduction in several areas. Shelley et al. (2005) provided full distribution records. Here we provide a distribution map (Fig. 8) and summarise the occurrence of Scolopendra morsitans in Southeast Asia and some parts of East Asia as follows: Southeast Asia: Thailand (entirely), Laos (southern part; Khammouane, Champasak), Cambodia (probably entirely), Vietnam (fide Schileyko 2007: Bai Tu Long Archipelago, Nghe An (Vinh), Thua Thien Hue (Hai Van Pass), Dak Lak, Khanh Hoa, Ninh Thuan (Phan Rang), Lam Dong (Da Lat), Ba Ria, Tay Ninh and Ca Mau), Spratly Archipelago, Myanmar (probably entirely), Malaysia, Singapore, Indonesia, Philippines (Manilla and Zebu Island), and Brunei (Jerudong). East Asia: China (Amoy, Hong Kong, Taiwan) and Japan (Ryukyu Islands).

Figure 8. — Distribution map of *Scolopendra* *morsitans* in Southeast Asia: Filled triangles indicate data from material examined herein; blank triangles indicate localities in the literature (Shelley et al. 2005, Schileyko 2007).

Scolopendra subspinipes

Leach, 1816

Figs 9A , 10 , 11 , 12 , 13 , 14 , 15 , 16 , 17 , 18

Figure 9. — Habitus photographs of *Scolopendra* species: A *Scolopendra* *subspinipes* (Singapore, *CUMZ* 00315) B *Scolopendra* *calcarata* (Thailand, *CUMZ* 00418) C *Scolopendra* *japonica* (Colour morph 2: Laos, *CUMZ* 00298).

Figure 10. — *Scolopendra* *subspinipes* (*NHMUK*): A Cephalic plate and trunk segments 1–2 B Tergites 9–12 C Forcipular segment D Tooth-plates E–G Spiracles 3, 5 and 8, respectively.

Figure 11. — *Scolopendra* *subspinipes* (*NHMUK*): A Sternites 8–11 B Sternite of ultimate leg-bearing segment and coxopleura C Lateral view of coxopleuron D Ventral view of ultimate leg-bearing segment and ultimate leg prefemora E Dorsal view of ultimate leg-bearing segment and prefemora.

Figure 12. — *Scolopendra* *subspinipes* (Syntype *NHMW* 751 of *Scolopendra* *mutilans* Koch, 1878): A Tooth-plates B Cephalic plate and trunk segments 1–2 C Forcipular segment D Tergites 9–11 **E–G** Spiracles 3, 5 and 8, respectively.

Figure 13. — *Scolopendra* *subspinipes* (Syntype *NHMW* 751 of *Scolopendra* *mutilans* Koch, 1878): A Sternite of ultimate leg-bearing segment and coxopleura B Sternites 10–12 C Sternite of ultimate leg-bearing segment, coxopleura and ventral view of ultimate leg prefemur D Tergite of ultimate leg-bearing segment E Ventral view of ultimate leg prefemur F Dorsal view of ultimate leg prefemur G Ventro-lateral view of ultimate leg.

Figure 14. — *Scolopendra* *subspinipes* (Syntypes *NMB* 391Va of “*piceoflava* Attems, 1934”): A Tooth-plates B Cephalic plate and trunk segments 1–3 C Forcipular segment D Tergites 9–12 E Sternites 8–10.

Figure 15. — *Scolopendra* *subspinipes* (Syntypes *NMB* 391Va of “*piceoflava* Attems, 1934”): A Lateral view of coxopleuron B Tergite of ultimate leg-bearing segment and dorsal view of legs 20 and ultimate leg C Ventral view of Leg 20, coxopleura and ultimate leg D Sternite of ultimate leg-bearing segment, coxopleura and ultimate leg **E–F** Ventro-lateral view of ultimate legs.

Figure 16. — *Scolopendra* *subspinipes* (*CUMZ* 00315): A Cephalic plate and basal antennal articles B Tooth-plates and trochanteroprefemoral process C Tergites 9–11 D Forcipular segment.

Figure 17. — *Scolopendra* *subspinipes* (*CUMZ* 00315): A Sternites 9–11 B Lateral view of coxopleuron C Tergite of ultimate leg-bearing segment D Sternite of ultimate leg-bearing segment, coxopleura, female genital segment and ultimate leg prefemora E Dorsal view of ultimate leg prefemora.

Figure 18. — Distribution map of *Scolopendra* *dehaani* and *Scolopendra* *subspinipes* in Southeast Asia and China-Japan Sea (small map): Filled and blank colours refer to localities from the present study and in the literature, respectively.

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.