A taxonomic review of the centipede genus Scolopendra Linnaeus, 1758 (Scolopendromorpha, Scolopendridae) in mainland Southeast Asia, with description of a new species from Laos Author Siriwut, Warut Author Edgecombe, Gregory D. Author Sutcharit, Chirasak Author Tongkerd, Piyoros Author Panha, Somsak text ZooKeys 2016 590 1 124 http://dx.doi.org/10.3897/zookeys.590.7950 journal article http://dx.doi.org/10.3897/zookeys.590.7950 1313-2970-590-1 BE34EA62E27346BB9FE64660953EDFE8 BE34EA62E27346BB9FE64660953EDFE8 Taxon classification Animalia Scolopendromorpha Scolopendridae Scolopendra dawydoffi Kronmueller , 2012 Figs 7B, 29, 42C, 48, 49, 50, 51, 52, 53 Scolopendra subspinipes cingulatoides Attems, 1938: 335, fig. 307, 1953 : 138. Schileyko 1998 : 268, 2001 : 434, 2007 : 76. Lewis 2010b : 112, fig. 24. Tran et al. 2013 : 229. Scolopendra dawydoffi Kronmueller , 2012: 22, table 1, fig. 4E [new replacement name]. Tran et al. 2013 : 229. Siriwut et al. 2015a : 1. Type locality. Two localities were reported in the original description, Hagiang, Haut Tonkin [ Ha Giang Province, northern Vietnam], and Thakek, Laos [Thakhek, Khammouane Province, Laos]. Material. Syntypes NHMW Inv. No. 8234, two females labeled " Scolopendra subspinipes cingulatoides Attems, 1934 typus by Attems", Thakek, Laos (Figs 50, 51). Figure 48. Scolopendra dawydoffi (CUMZ 00290, 00291): A Tooth-plates B Forcipular segment C Cephalic plate and trunk segments 1-3 D Sternites 9-11 E-G Spiracles 3, 5 and 8, respectively. Figure 49. Scolopendra dawydoffi (CUMZ 00290, 00291, 00272): A Tergites 9-11 B Tergite of ultimate leg-bearing segment C Lateral view of coxopleuron, leg 20 and ultimate leg D Sternite of ultimate leg-bearing segment and coxopleura E and F Variation in ventral spines on ultimate leg prefemora G Dorsal view of ultimate leg prefemora. Figure 50. Scolopendra dawydoffi (Syntypes NHMW 8234): A Cephalic plate and trunk segments 1-3 B Tergites 9-11 C Tooth-plates D Forcipular segment E-G Spiracles 3, 5 and 8, respectively. Figure 51. Scolopendra dawydoffi (Syntypes NHMW 8234): A Sternites 9-11 B Sternite of ultimate leg-bearing segment and coxopleura C Tergite of ultimate leg-bearing segment D Lateral view of coxopleuron E-F Dorsal and ventral view of ultimate leg prefemora. Additional material. Thailand - ZMUC 1/7.59, one adult female and 13 juvenile spms., labeled as " Scolopendra subspinipes ", Phu Kradueng, Loei, 1300 m, evergreen forest, 24/11/1958, leg. B. Degerbol. CUMZ 00294 two adult spms., Wat Thang Biang, Pak Chong, Nakhon Ratchsrima ( 14°32'22.0"N , 101°21'54.6"E ). CUMZ 00290, one spm., Sakaerat Biosphere Reserve Center, Nakon Ratchasima ( 14°30'36.5"N , 101°55'51.5"E ). CUMZ 00272.1-2, two spms., Saphan Hin Waterfall, Khlong Yai, Trad ( 12°06'06.0"N , 102°42'39.2"E ) Malaysia - NHMUK.1950.4.19.12, one spm., Kelantan, Malay Peninsula (labeled as " Scolopendra subspinipes "). Diagnosis. 17-18 antennal articles, 6 basal articles glabrous dorsally. Each tooth-plate with 5-10 teeth. Tergites 2(3)-20 with paramedian sutures. Complete tergite margination from TT11-21. Tergite of ultimate leg-bearing segment without depression or suture. Paramedian sutures on anterior 15-60% of sternites. Coxopleural process with 2-3 apical+subapical spines. Ultimate leg prefemora with 1-2 VL, 0-2 M, 0-2 DM, prefemoral process with 1-5 spines. One tarsal spur on legs 1-19. Composite description. Body length up to 16.2 cm (14.7 and 15.1 cm in syntypes). Reddish colouration on entire body. Cephalic plate and tergites dichromatic. Cephalic plate and tergites reddish orange; posterior border of tergites with dark band. Cephalic plate with small punctae; median sulcus present. Posterior part of cephalic plate without paramedian sulci. Antenna usually with 18 articles (sometimes 17 on one side in some specimens), basal 6 subcylindrical and glabrous dorsally (Fig. 52A), 5-5.5 articles glabrous ventrally. Antennae reach segment 4. Forcipular trochanteroprefemoral process with denticles in two groups, one apical and 2-3 inner. Anterior part of coxosternite with tooth-plates, wider than long or nearly equivalent, 5-7 robust teeth (Figs 48A, 52 D-F ); atypically with 10 teeth (CUMZ 00272). Tooth-plate with straight, transverse suture. Coxosternal surface smooth, without median suture (Figs 48B, 52B). Article 2 of second maxillary telopodite with spur. Figure 52. Scolopendra dawydoffi (CUMZ 00272): A Cephalic plate and basal antennal articles B Forcipular segment C Tergites 9-11 D-F Variation in teeth on tooth-plates and trochanteroprefemoral process. Anterior margin of T1 underlying cephalic plate (Fig. 48C). Complete paramedian sutures on TT2-3; margination typically from TT11-14. Tergite surfaces (Figs 49A, 52C) smooth, without median sulci. Tergite of ultimate leg-bearing segment (Figs 49B, 53E) curved posteriorly, without median furrow or depression; ratio of width: length of tergite of ultimate leg-bearing segment 0.7:1. Anterior part of sternites (Figs 48D, 53A) with short paramedian sutures reaching approximately 15-30% (atypically, to 60% in one specimen: CUMZ 00272). Surface of sternites smooth, mostly with depression (small circular pit present on posterior median part of sternite in one specimen: CUMZ 00294). Sternite of ultimate leg-bearing segment (Fig. 49D) with sides converging posteriorly. Pore-field on coxopleuron terminating beneath margin of tergite of ultimate leg-bearing segment, dorsal margin of pore area sinuous, most elevated anteriorly. Figure 53. Scolopendra dawydoffi (CUMZ 00272): A Sternites 9-11 B-C Variation in numbers of spines on prefemural process of ultimate leg D Sternite of ultimate leg-bearing segment, coxopleura and ultimate legs, showing lack of ventral and median spines on prefemora E Tergite of ultimate leg-bearing segment F Lateral view of coxopleuron. Coxopleural process moderately long or short with two apical spines and one subapical spine (atypically only two apical spines; Fig. 53F); pore-free area extending 65-90% length from distal part of coxopleural process to margin of sternite of ultimate leg-bearing segment (Figs 49D, 53D). All legs without setae and tibial spur. One tarsal spur on legs 1-19. Ultimate legs: thick and moderately long (Figs 49C, 53D), with ratios of lengths of prefemur and femur 1.3:1, femur and tibia 1.3:1, tibia and tarsus 2 1.8:1, tarsus 1 and tarsus 2 1.7:1. Prefemora flattened dorsally (atypically rounded; Figs 49 F-G , 53B), with robust blackish spines. Prefemoral spines (Figs 49E, 51 E-F ): 1-2 VL, 0-2 M, 0-2 DM, prefemoral process with 1-3 spines, atypically with 5 spines (Figs 49G, 53B). Posterior margin of prefemur with shallow 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 (Fig. 28 C-D ). Sternite of genital segment 2 developed. Gonopod absent in male. Lamina subanalis between genitalia and anal valve; lamina analis between anal valve and tergite of genital segment. Tergite and sternite of genital segments with small setae. Penis with apical bristle. Discussion. This species is distinguished from Scolopendra subspinipes by its short, robust ultimate legs and three apical/subapical spines on the coxopleural process. The characteristic of incomplete paramedian sutures on the sternites further distinguishes it from Scolopendra subspinipes and Scolopendra japonica (which have complete paramedian sutures on the sternites). However, Scolopendra dawydoffi is similar to Scolopendra multidens in the absence of gonopods in the male. The distribution of Scolopendra dawydoffi is restricted to mainland Southeast Asia whereas Scolopendra multidens occurs in temperate regions of Asia, including both inland and insular parts. A specimen identified as Scolopendra multidens from Vietnam is genetically differentiated from Thai populations (see discussion of Scolopendra multidens above for molecular arguments in favour of the two taxa being separate species). Moreover, to test the hypothesis that characteristics of Scolopendra dawydoffi might indicate affinities to the cingulata group (with reference to the Mediterranean species Scolopendra cingulata ; Attems 1930a ), as implied by the original " cingulatoides " name for Scolopendra dawydoffi , our phylogenetic analysis included Scolopendra cingulata sequences from Spain. The result (Fig. 1) demonstrated that Scolopendra dawydoffi was not grouped together with Scolopendra cingulata but should be recognized as a distinct species based on its genetic distance and geographical distribution. A morphological comparison between these two species is presented in Table 10. Table 10. Morphological comparison of Scolopendra dawydoffi and Scolopendra multidens ; data from present study and previous taxonomic studies, i.e., Attems (1938) , Chao (2008) , Kronmueller (2012) .? Character not present, L left side.

Characters	Scolopendra dawydoffi		Scolopendra multidens
Characters	Laos (Syntypes)	Thailand	China (Holotype)	Hong Kong and Taiwan
ULBS
APSAP	APSAP	APSAP	APSAP
VLMDMSP	VLMDMSP	VLMDMSP	VLMDMSP

Distribution. This species was formerly reported from only two occurrences in Southeast Asia, one in each of Laos and Vietnam ( Attems 1938 ). We add more material from Thailand and some adjacent areas based on museum collections. The updated distribution range (Fig. 29) is as follows: Southeast Asia: Thailand (Nakhon Ratchasima, Loei and Trad), Laos (Thakhek), Malaysia (Kelantan state) and Vietnam (fide Tran et al. 2013 : Ha Giang, Hanoi, Thai Nguyen and Ha Tinh).