Desmoxytes cervina (Pocock, 1895)

Srisonchai, Ruttapon, Enghoff, Henrik, Likhitrakarn, Natdanai & Panha, Somsak, 2018, A revision of dragon millipedes I: genus Desmoxytes Chamberlin, 1923, with the description of eight new species (Diplopoda, Polydesmida, Paradoxosomatidae), ZooKeys 761, pp. 1-177 : 1

Desmoxytes cervina (Pocock, 1895) Figs 16, 17, 18, 19, 20, 21, 22

Prionopeltis cervinus Pocock, 1895: 831. Attems 1914: 203; 1936: 215.

Pratinus cervinus - Attems 1937: 120. Jeekel 1964: 63. Jeekel 1968: 61.

Desmoxytes cervina - Jeekel 1980a: 654. Golovatch and Enghoff 1994: 61. Nguyen and Sierwald 2013: 1241. Likhitrakarn et al. 2017: 19.

Desmoxytes pterygota Golovatch & Enghoff, 1994: 55, syn. n. Enghoff 2005: 96. Decker 2010: 30. Nguyen and Sierwald 2013: 1242.

Desmoxytes sp. - Golovatch and Enghoff 1994: 60.

Material examined.

Lectotype. Male (NHMUK, Bm 1892.5.4.76), MYANMAR, south Tenasserim, leg. E. W. Oates, most legs missing. Lectotype here designated.

Holotype ( D. pterygota ): Male (ZMUC), THAILAND, Ranong Province, Kapoe District, in forest at big waterfalls south of Kapoe (Khao Phra Narai Waterfall?), 15 November 1990, leg. M. Andersen and A. R. Rasmussen.

Paratypes ( D. pterygota ): 2 males (ZMUC), 1 male (ZMUM), THAILAND, Ranong Province, Kapoe District, in forest at big waterfalls south of Kapoe, 15 November 1990, leg. M. Andersen and A. R. Rasmussen.

Other material examined.

MYANMAR: 10 males, 7 females (CUMZ), Tanintharyi Region, Lenya National Park, Phayarhtan Cave (Buddha Cave), approximately 10 km from Ban Nam Yen Village, inside the deep rainforest near limestone mountain, on decaying wood and under bark, 11°13'50"N, 99°10'35"E, ca. 85 m a.s.l., 6 June 2015, leg. C. Sutcharit, R. Chanabun and R. Srisonchai.

THAILAND: Chumphon Province: 1 male, 2 females (CUMZ), Mueang Chumphon District, Tham Chang Phuek Bureau of Monks, 10°26'47"N, 99°02'06"E, ca. 93 m a.s.l., 13 March 2017, leg. C. Sutcharit, R. Srisonchai and ASRU members. 1 male (CUMZ), Sawi District, Wat Nam Cha, 10°17'54"N, 99°01'57"E, ca. 105 m a.s.l., 3 July 2017, leg. C. Sutcharit, R. Srisonchai and ASRU members.

Krabi Province: 1 male (CUMZ), Ao-Luek District, P.N. Moutain Resort, 8°24'09"N, 98°44'18"E, ca. 60 m a.s.l., 30 August 2015, leg. C. Sutcharit and ASRU members. 1 male remaining rings 7-20 (CUMZ), Khlong Thom District, Emerald Blue Pool, 7°55'30"N, 99°16'05"E, ca. 67 m a.s.l., 15 January 2009, leg. S. Panha and ASRU members. 1 female (CUMZ), Muang Krabi District, Wat Tham Sue (Tiger Cave), valley behind Tiger Cave, 8°07'38"N, 98°55'27"E, ca. 86 m a.s.l., 18 May 2010, leg. S. Panha and ASRU members. 2 males, 3 females, 1 broken male and missing gonopods (CUMZ), Muang Krabi District, Wat Tham Sue (Tiger Cave), valley behind Tiger Cave, 8°07'38"N, 98°55'27"E, ca. 86 m a.s.l., 7 October 2006, leg. S. Panha and ASRU members. 1 male, 4 females (CUMZ), Muang Krabi District, Wat Tham Sue (Tiger Cave), valley behind Tiger Cave, 8°07'38"N, 98°55'27"E, ca. 86 m a.s.l., 24 August 2014, leg. S. Panha and ASRU members. 2 males, 1 female, 1 juvenile (CUMZ), Muang Krabi District, Wat Tham Sue (Tiger Cave), valley behind Tiger Cave, 8°07'38"N, 98°55'27"E, ca. 86 m a.s.l., 30 August 2015, leg. C. Sutcharit, R. Srisonchai and ASRU members. 15 males, 3 females, 1 juvenile (CUMZ), Muang Krabi District, Wat Tham Sue (Tiger Cave), valley behind Tiger Cave, 8°07'38"N, 98°55'27"E, ca. 86 m a.s.l., 9 July 2017, leg. C. Sutcharit, R. Srisonchai and ASRU members. 11 males, 6 females, 1 juvenile (CUMZ), Muang Krabi District, Wat Tham Sue (Tiger Cave), valley behind Tiger Cave, 8°07'38"N, 98°55'27"E, ca. 86 m a.s.l., 25 July 2017, leg. C. Sutcharit, R. Srisonchai and ASRU members.

Nakhon Si Thammarat Province: 2 males, 2 females, 4 juveniles (ZMUC), Sichon District, Khao Lark Waterfall, 25 August 2007, leg. ASRU members. 1 female (CUMZ), Thung Song District, Yong Waterfall, 8°10'21"N, 99°44'34"E, ca. 138 m a.s.l., 20 July 2008, leg. S. Panha and ASRU members. 1 male missing gonopods (CUMZ), Nopphitam District, Krung Ching Waterfall, 8°43'27"N, 99°40'04"E, ca. 173 m a.s.l., 17 January 2013, leg. C. Sutcharit, R. Srisonchai and ASRU members. 1 female (CUMZ), Khanom District, Nai Plao Beach, 9°07'26"N, 99°52'60"E, ca. 20 m a.s.l., 4 December 2015, leg. S. Panha and ASRU members. Many specimens (CUMZ), Tham Phannara District, Wat Tham Kanlaya Namit, 8°30'48"N, 99°22'52"E, ca. 51 m a.s.l., 4 July 2017, leg. C. Sutcharit, R. Srisonchai and ASRU members. Many specimens (CUMZ), Tham Phannara District, Wat Tham Thong Phannara, 8°25'21"N, 99°22'47"E, ca. 32 m a.s.l., 4 July 2017, leg. C. Sutcharit, R. Srisonchai and ASRU members.

Phang Nga Province: 1 male missing gonopods, 3 males, 1 female (CUMZ), Thai Mueang District, Khaolak-Lumru National Park, 8°37'35"N, 98°14'25"E, ca. 72 m a.s.l., 7 October 2006, leg. S. Panha and ASRU members. 1 male, 1 female (ZMUC), Thap Put District, Highway No. 4 Phet Kasem Road ca. 0.5 km north of the Headquarters of the Khao Lak–Lamru National Park, on the street next to secondary rainforest. 8°37'N, 98°14'E, ca. 30-40 m a.s.l., 29 August– 12 September 2008, leg. N. Laufer. 1 male missing gonopods, 1 male (CUMZ), Khura Buri District, Mu Koh Surin National Park, Koh Surin Nuea, 9°26'27"N, 97°52'11"E, ca. 39 m a.s.l., 8 April 2012, leg. S. Panha and ASRU members. 1 broken male and missing gonopods (CUMZ), Mueang Phang Nga District, Tham Nam Pud, 8°27'50"N, 98°32'36"E, ca. 58 m a.s.l., 7 October 2006, leg. S. Panha and ASRU members. Many specimens (CUMZ), Mueang Phang Nga District, Tham Nam Pud, 8°27'50"N, 98°32'36"E, ca. 58 m a.s.l., 5 August 2015, leg. C. Sutcharit, R. Srisonchai and ASRU members. 1 male (CUMZ), Mueang Phang Nga District, Tham Pha Sue Bureau of Monks, 8°28'24"N, 98°32'15"E, ca. 78 m a.s.l., 10 July 2017, leg. C. Sutcharit, R. Srisonchai and ASRU members. 2 females (CUMZ), Mueang Phang Nga District, Tham Nam Pud Bureau of Monks, 8 October 2006, leg. S. Panha and ASRU members. 1 male, 1 female (CUMZ), Mueang Phang Nga District, Tao Thong Waterfall, 8°29'08"N, 98°35'09"E, ca. 25 m a.s.l., 7 October 2006, leg. S. Panha and ASRU members. 1 female (CUMZ), Mueang Phang Nga District, Phung Chang Cave, 8°26'34"N, 98°30'59"E, ca. 24 m a.s.l., 26 September 2009, leg. S. Panha and ASRU members. 3 males, 2 females, 14 juveniles (CUMZ), Mueang Phang Nga District, Phung Chang Cave, 8°26'34"N, 98°30'59"E, ca. 24 m a.s.l., 6 August 2014, leg. C. Sutcharit, R. Srisonchai and ASRU members. Many specimens (CUMZ), Mueang Phang Nga District, Phung Chang Cave, 8°26'34"N, 98°30'59"E, ca. 24 m a.s.l., 5 August 2015, leg. C. Sutcharit, R. Srisonchai and ASRU members. 5 males, 4 females (CUMZ), Mueang Phang Nga District, Phung Chang Cave, 8°26'34"N, 98°30'59"E, ca. 24 m a.s.l., 8 August 2016, leg. C. Sutcharit, R. Srisonchai, and ASRU members. 3 males, 2 juveniles (CUMZ), Takua Thung District, Wat Suwan Khuha (Monkey Cave), 8°25'42"N, 98°28'22"E, ca. 25 m a.s.l., 8 August 2016, leg. C. Sutcharit, R. Srisonchai, and ASRU members. Many specimens (CUMZ), Thap Put District, Wat Khiri Wong (Tham Kob), 8°31'57"N, 98°34'40"E, ca. 97 m a.s.l., 9 July 2017, leg. C. Sutcharit, R. Srisonchai, and ASRU members.

Phuket Province: 1 male of Desmoxytes sp. (ZMUC), Thalang District, Thepkrasattree Subdistrict, Tonsai Waterfall, 8°01'44"N, 98°21'45"E, ca. 67 m a.s.l., 12 October 1991, leg. M. Anderson, O. Martin, and N. Scharff. 1 female (CUMZ), Mueang Phuket District, Panwa Cave, 7°48'04"N, 98°24'34"E, ca. 25 m a.s.l., June 2007, leg. S. Panha and ASRU members. 1 female (CUMZ), Mueang Phuket District, Panwa Cave, 7°48'04"N, 98°24'34"E, ca. 25 m a.s.l., 5 November 2007, leg. S. Panha and ASRU members.

Ranong Province: 1 male (CUMZ), Kra Buri District, Tham Phra Khayang, 10°19'35"N, 98°45'54"E, ca. 51 m a.s.l., 21 November 2015, leg. S. Panha, P. Tongkerd, and A. Pholyotha. 1 female (CUMZ), Kra Buri District, Bok Krai Waterfall, 10°22'35"N, 98°51'22"E, ca. 106 m a.s.l., 3 January 2013, leg. S. Panha, P. Tongkerd, and A. Pholyotha.

Surat Thani Province: 2 broken and mixed females, 2 females remaining rings 13-20, 1 broken female (CUMZ), Phanom District, Khlong Phanom National Park, 8°52'44"N, 98°40'26"E, ca. 68 m a.s.l., 28 August 2007, leg. S. Panha and ASRU members. 1 male, 6 juveniles (CUMZ), Phanom District, Khlong Phanom National Park, Pha Daeng, 8°53'41"N, 98°33'12"E, ca. 67 m a.s.l., 7 August 2016, leg. C. Sutcharit, R. Srisonchai, and ASRU members. 8 males, 3 females (CUMZ), Phanom District, Khlong Phanom National Park, Pha Daeng, 8°53'41"N, 98°33'12"E, ca. 67 m a.s.l., 1 August 2017, leg. C. Sutcharit, R. Srisonchai, and ASRU members. 4 males (CUMZ), Phanom District, Ban Song Phi Nong, 8°50'51"N, 98°44'16"E, ca. 74 m a.s.l., 7 August 2016, leg. C. Sutcharit, R. Srisonchai, and ASRU members. 10 males, 3 females, 1 broken male missing gonopods (CUMZ), Ban Ta Khun District, Ratchaprapa Dam, 8°57'22"N, 98°48'22"E, ca. 53 m a.s.l., 8 October 2008, leg. S. Panha and ASRU members. Many specimens (CUMZ), Ban Ta Khun District, Ratchaprapa Dam, 8°57'22"N, 98°48'22"E, ca. 53 m a.s.l., 4 August 2014, leg. C. Sutcharit, R. Srisonchai, and ASRU members. Many specimens (CUMZ), Ban Ta Khun District, Ratchaprapa Dam, 8°57'22"N, 98°48'22"E, ca. 53 m a.s.l., 3 August 2015, leg. C. Sutcharit, R. Srisonchai, and ASRU members. 1 male (CUMZ), Ban Ta Khun District, Ratchaprapa Dam, 8°57'22"N, 98°48'22"E, ca. 53 m a.s.l., 5 August 2016, leg. C. Sutcharit, R. Srisonchai, and ASRU members. 1 male (CUMZ), Ban Ta Khun District, Khlong Hoi, 8 October 2008, leg. S. Panha and ASRU members. 2 males, 1 male missing gonopods, 1 juvenile, 1 broken juvenile (CUMZ), Ban Ta Khun District, Khao Wong Water Supply Station, 8°55'47"N, 98°56'25"E, ca. 97 m a.s.l., 9 October 2008, leg. S. Panha and ASRU members. 2 males (CUMZ), Ban Ta Khun District, Wat Khao Pang (Suspension Bridge), 8°56'54"N, 98°49'21"E, ca. 24 m a.s.l., 5 May 2017, leg. S. Panha and ASRU members. 1 male, 1 male missing gonopods, 2 females, 2 broken females, 1 juvenile (CUMZ), Khirirat Nikhom District, Wat Satit Khirirom, 9°01'48"N, 98°59'12"E, ca. 47 m a.s.l., 8 October 2008, leg. S. Panha and ASRU members. 1 male (CUMZ), Khirirat Nikhom District, Wat Satit Khirirom, 9°01'48"N, 98°59'12"E, ca. 47 m a.s.l., 5 September 2009, leg. S. Panha and ASRU members. 27 males, 10 females (CUMZ), 2 males, 1 female (ZMUC), 1 male, 1 female (ZMUM), 1 male, 1 female (NHMW), 1 male, 1 female (NHMUK), Khirirat Nikhom District, Wat Satit Khirirom, 9°01'48"N, 98°59'12"E, ca. 47 m a.s.l., 10 July 2017, leg. C. Sutcharit, R. Srisonchai, and ASRU members. 1 male missing right gonopod (CUMZ), Khirirat Nikhom District, km3 near Khirirat Nikhom City, 9 October 2008, leg. S. Panha and ASRU members. 1 male, 1 female (CUMZ), Khirirat Nikhom District, Tham Wang Badan Bureau of Monks, 8°56'09"N, 98°57'28"E, ca. 69 m a.s.l., 3 August 2017, leg. C. Sutcharit, R. Srisonchai, and ASRU members. 3 broken males (CUMZ), Ko Samui District, Na Muang Waterfall, 9°27'58"N, 99°59'02"E, ca. 53 m a.s.l., 27 January 2006, leg. S. Panha and ASRU members. 1 female (CUMZ), Ko Samui District, Na Muang Waterfall, 9°27'58"N, 99°59'02"E, ca. 53 m a.s.l., 4 December 2015, leg. S. Panha, P. Tongkerd and A. Pholyotha. 1 male remaining rings 1-11, 1 female (ZMUM), Ko Samui, Thailand, June 2013, leg. Korabushkin Daniil. 10 males, 5 females, 4 juveniles (CUMZ), Wiang Sa District, Khiri Rat Pattana Bureau of Monks (Wat Khao Poon), 8°31'37"N, 99°22'59"E, ca. 68 m a.s.l., 4 July 2017, leg. C. Sutcharit, R. Srisonchai, and ASRU members. 1 male, 1 female (CUMZ), Ban Na San District, Khao Kok Maharat Bureau of Monks, 8°41'33"N, 99°22'45"E, ca. 71 m a.s.l., 4 July 2017, leg. C. Sutcharit, R. Srisonchai, and ASRU members.

Diagnosis.

Differs from other Desmoxytes species by the combination of the following characters: body colour brownish red/brown/pale brown; paraterga brownish red/red/yellowish brown; metaterga 2-4 with 2+2 (anterior row) and 2+2 (posterior row) tubercles, metaterga 5-19 with 2(1)+2(1) (anterior row) and 2+2 (posterior row) tubercles; lamina lateralis (ll) stout; anterolaterally with a long, distinct, deep and wide furrow; ventral ridge (vrl) of lamina lateralis very long, wide, conspicuous; process (plm) of lamina medialis long, distinctly demarcated from distal lobe, irregularly shaped, directed mesodorsad; distal lobe (dlm) of lamina medialis distally with one distinct lamella; broad lobe (blm) slightly thick at the edge, distinctly demarcated from distal lobe (dlm) by a wide and shallow indentation.

Type locality.

MYANMAR, southern Myanmar, Tenasserim [Tanintharyi Region].

Redescription.

SIZE: Length 25-31 mm (male), 33-39 mm (female); width of midbody metazona ca. 2.1 mm (male), 3.6 mm (female). Width of head <collum <body ring 2 ≥ 3 = 4 <5-16, thereafter body gradually tapering towards telson.

COLOUR (Figs 16, 17 A–E): In life with two colour morphs. Brownish red morph - body brownish red (testaceous); head and antenna brownish black (except distal part of antennomere 7 and antennomere 8 whitish); collum, epiproct and legs brown; rings 2-3 brownish red or brown; metaterga, surface below paraterga and sterna brownish red; paraterga brownish red or red; a few basal podomeres reddish brown. Brown morph - body brown (female pale brown); head and antenna (except distal part of antennomere 7 and antennomere 8 whitish) brownish black or black; collum and rings 2-3 brownish black; metaterga, surface below paraterga, epiproct and legs brown; paraterga yellowish brown; sterna pale brown; a few basal podomeres brownish white. Colour in alcohol: after 100 years changed to greenish dark or greenish brown, after 5-10 years changed to pale brown.

ANTENNAE (Fig. 18D): Very long and slender, reaching to body ring 6 or 7 (male) and 5 or 6 (female) when stretched dorsally.

COLLUM (Fig. 18A): With 1 transverse row of setae, 3+3 anterior setae; paraterga of collum low, elevated at ca. 15°-20°, directed caudolaterad, with two inconspicuous notches on lateral margin.

TEGUMENT: Moderately shining; collum coarsely microgranulate; prozona finely shagreened; metaterga and surface below paraterga finely microgranulate; paraterga, sterna and epiproct smooth.

METATERGA (Fig. 18 A–C): With 2 transverse rows of tubercles; metaterga 2-4 with 2+2 anterior tubercles and 2+2 posterior tubercles; metaterga 5-19 with 2(1)+2(1) anterior tubercles and 2+2 posterior tubercles (lateral setae of anterior row in some specimens very distinct, in some specimens poorly developed).

PARATERGA (Fig. 18E, F): Long, strongly developed; directed caudolaterad on body rings 2-17, elevated at ca. 45° (male) 40° (female); directed increasingly caudad on body rings 18 and 19; anterior margin with 2 distinct notches, on lateral margin of body rings 9, 10, 12, 13, 15-18 with tiny denticle near the tip.

TELSON (Fig. 19 C–G): Epiproct: tip usually truncate (in some specimens slightly emarginate); lateral setiferous tubercles and apical setiferous tubercles conspicuous, coniform. Hypoproct subtrapeziform; caudal margin slightly round (in some specimens subtruncate), with inconspicuous setiferous tubercles.

STERNA (Fig. 20): Cross-impressions shallow (in some specimens slightly deep). Sternal lobe between male coxae 4 swollen, usually subtrapeziform (in some specimens subsemicircular), usually thick when seen in lateral view (in some specimens thin), slightly attenuated near tip, tip round/subtruncate/slightly emarginate.

LEGS (Fig. 19 H–J): Very long and slender. Male femora 5 and 6 strongly humped ventrally in middle portion.

GONOPODS (Figs 21, 22): Coxa (cx) longer than prefemur. Cannula (ca) long and slender. Prefemur (pfe) ca. 2/3 as long as femur. Femur (fe) long and slender. Mesal sulcus (ms) and lateral sulcus (ls) conspicuous, very deep and narrow. Postfemur (pof) conspicuous, ventrally narrow. Solenophore (sph) well-developed: lamina lateralis (ll) stout; anterolaterally with a long, distinct, deep and wide furrow; with a long, wide and conspicuous ventral ridge (vrl): lamina medialis (lm) well-developed; process (plm) long, of irregular shape (varies within population), distinctly demarcated from distal lobe, tip blunt (in some specimens terminating in spines), directed mesodorsad; distal lobe (dlm) distally with one distinct lamella; broad lobe (blm) slightly thick at the edge, distinctly demarcated from distal lobe (dlm) by a wide and shallow indentation. Solenomere (sl) long.

Distribution and habitat.

Desmoxytes cervina is known from Myanmar (Lenya National Park) and Thailand (Chumphon, Krabi, Nakhon Si Thammarat, Phang Nga, Phuket, Ranong, and Surat Thani Provinces). The type locality is in south of Tenasserim, but we do not know the exact location, probably somewhere near Taninthayi township. The locality of the paralectotype (see below) "Malewoon, Tenasserim", is currently known as Maliwan Village.

Almost all specimens were collected from limestone habitats, a few specimens were collected from granitic areas. Interestingly, D. cervina was also found in some islands in both the Andaman Sea and the Gulf of Thailand (Mu Koh Surin National Park, Phuket, and Ko Samui).

Decker (2010) classified " D. pterygota " as endemic for Thailand due to its narrow range near the type locality. After intensive collecting and synonymising of D. cervina and " D. pterygota ", the known distribution range has expanded to southern Myanmar and southern Thailand.

This species was found living together with D. delfae and D. corythosaurus sp. n. According to our observations, it is probable that they may even share microhabitat: humid rocks, branches of trees and rock walls.

Note on material.

This species was described based on two males, one of which was collected by L. Fea (Malewoon, Tenasserim - MSNG) and the other collected by E. W. Oates (south Tenasserim - NHMUK).

Jeekel (1964) revised this species by examining the male in MSNG; he stated that "in anticipation of the designation as holotype of the specimen collected by Oates in the British Museum, I have labelled this specimen as paratype". However, the holotype has never been designated in the original description. R.L. Hoffman visited NHMUK and labelled the specimen collected by Oates as paratype, but he did not publish this designation, which is in conflict with Jeekel’s (1964) paratype designation. Therefore, we here designate the male collected by Oates and belonging to NHMUK as lectotype to stabilise the name. The male in MSNG (which we have not examined) is considered to be a paralectotype.

Some specimens kept in MHNG and identified as " D. pterygota " by Decker (2010), are probably D. cervina because the localities fall within the distribution range of this species.

Remarks.

This is the first report of the colour of living specimens for this species; the brownish red colour is apparently aposematic. Pocock (1895) and Golovatch and Enghoff (1994) did not mention the colour of living specimens. We found two colour morphs of D. cervina : brownish red and brown. The majority of specimens are brownish red, and the minority are brown; the latter colour can be found in a few specimens within a population.

Interestingly, brownish red and brown morphs occur in the same habitat in the valley behind Tiger Cave and Ban Song Phi Nong. The brown morph was found at Wat Tham Kanlaya Namit, Wat Tham Thong Phannara, Tham Nam Pud, Phung Chang Cave and Wat Suwan Khuha (Monkey Cave). Specimens from the remaining localities are of the brownish red morph. We examined the morphological characters of all specimens of both colour morphs; all specimens exhibit the same morphology, especially in the gonopods which are identical. Perhaps the difference in colour is caused by environmental factors and/or genetic variation.

We assume that the type material of both D. cervina , collected by Oates and Fea a hundred years ago, and " D. pterygota " collected by M. Andersen and A.R. Rasmussen 17 years ago, did probably exhibit brownish red colour because all specimens near the type localities are brownish red. We found additionally that the colour of some females is pale brownish red or pale brown, and the colour of juveniles is pale brown.

Jeekel (1964) wrote in his redescription that D. cervina shows collum without setae, metaterga 2-19 with 1+1 anterior tubercles and 2+2 posterior tubercles. Golovatch and Enghoff (1994) distinguished " D. pterygota " from D. cervina by having smooth metaterga. They also described " D. pterygota " as having no pleurosternal carinae, antenna reaching to ring 4 in male, collum with 2 rows of 3+3 anterior setae and 1+1 intermediate setae, metaterga 2-19 with 2+2 anterior cones and 2+2 posterior cones. After examination of all type material and newly collected specimens of D. cervina and " D. pterygota ", we found that:

- all specimens of D. cervina and " D. pterygota " display fine microgranulation on the metaterga.

- all specimens have pleurosternal carinae, in body ring 2 very distinct and crest-like, in ring 3 very small, thereafter absent.

- antenna reaches to ring 6-7 in male and to ring 5-6 in female of both D. cervina and " D. pterygota ".

- collum has 3+3 anterior setae in all specimens - because Jeekel studied old preserved specimens, the setae may have been lost over time.

- metaterga 5-19 varies within populations; metaterga 2-4 with 2+2 anterior tubercles and 2+2 posterior tubercles, metaterga 5-19 with 2+2/1+1 anterior and 2+2 posterior tubercles.

Several other characters show variability, as follows:

I. variation within populations

- size of tubercles on metaterga: tubercles conspicuous in some specimens, inconspicuous in the others (bigger and more obvious in the holotype of " D. pterygota " than in the lectotype of D. cervina ).

- tip of process (plm) of lamina medialis: in some specimens terminating in one blunt process, in others terminating in a sharp spine.

- shape of sternal lobe between male coxae 4: in some individuals subtrapeziform, in others subsemicircular.

- tip of sternal lobe between male coxae 4: in some specimens round, in some subtruncate, in others emarginate.

- cross-impressions on sternum: in some individuals shallow and faint, in others slightly deep.

- tip of epiproct: in some individuals truncate, in others slightly emarginate.

- caudal margin of hypoproct: in some specimens slightly round, in others subtruncate.

- size of sternal lobe between male coxae 4 when seen in lateral view in specimens from Wat Satit Khirirom: in some specimens thick, in others thin.

II. variation between populations

- colour: all individuals in the same population usually have the same colour: brown or brownish red. However, in some populations (valley behind Tiger Cave and Ban Song Phi Nong) brownish red and brown individuals coexist.

Although the male paralectotype of D. cervina (in MSNG) has not been examined by us, the morphological characters for this specimen as redescribed by Jeekel (1964) perfectly match the morphology of the numerous other specimens we have seen. Based on our analysis of morphology and variation of these specimens we have synonymised " D. pterygota " under D. cervina .

Distribution data support the synonymisation: the type localities of " D. pterygota " (Ranong Province) and D. cervina (Tenasserim = Taninthayi) are very close to each other. Our intensive surveys prove that this species is distributed quite widely, but nevertheless is found in south Myanmar and south Thailand only.

During the field survey, we noticed several adult males of D. cervina which were infested with red mites. The mites are probably larvae of the genus Leptus Latreille, 1796 ( Prostigmata , family Erythraeidae ). Associations between mites and millipedes may be of a phoretic or a parasitic nature ( Gerdeman et al. 2000, Swafford and Bond 2010, Farfan and Klompen 2012, Mwabvu 2014). In Fig. 17C and D, several engorged mites are seen along with a few small, non-engorged ones, and we therefore assume that the mite species found in D. cervina is parasitic, like other Leptus larvae. Southcott (1992) described Leptus millipedius from julid millipedes, but this is the first record of a parasitic prostigmatan mite from a paradoxosomatid millipede. The only record of a mite from Paradoxosomatidae concerns the widespread Oxidus gracilis (C. L. Koch, 1847) which was reported as associated with Cosmolaelaps hortensis Ishikawa, 1986 ( Mesostigmata , family Laelapidae ) ( Ishikawa 1986).

Golovatch and Enghoff (1994) reported one broken male of " Desmoxytes sp." from Phuket Province, Thailand. According to the remarks of Golovatch and Enghoff (1994), the gonopod characters of this specimen were similar to D. delfae , but the paraterga showed a higher degree of elevation. We examined this specimen again and found it to share gonopod and other characters with D. cervina . We therefore treat " Desmoxytes sp." as D. cervina .

Coexisting species.

D. delfae at Wat Tham Sue (Tiger Cave), Krung Ching Waterfall, Khiri Rat Pattana Bureau of Monks (Wat Khao Poon); D. corythosaurus sp. n. at Ban Song Phi Nong.