Research Article |
Corresponding author: Somsak Panha ( somsak.pan@chula.ac.th ) Academic editor: Robert Mesibov
© 2018 Ruttapon Srisonchai, Henrik Enghoff, Natdanai Likhitrakarn, Somsak Panha.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Srisonchai R, Enghoff H, Likhitrakarn N, Panha S (2018) A revision of dragon millipedes IV: the new genus Spinaxytes, with the description of nine new species (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys 797: 19-69. https://doi.org/10.3897/zookeys.797.29510
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Nine new species constituting the ‘spiny’ group of dragon millipedes are assigned to the new genus Spinaxytes Srisonchai, Enghoff & Panha, gen. n. Seven new species are described from Thailand: S. biloba Srisonchai, Enghoff & Panha, sp. n. and S. palmata Srisonchai, Enghoff & Panha, sp. n. from Surat Thani Province, S. hasta Srisonchai, Enghoff & Panha, sp. n. from Chumphon Province, S. krabiensis Srisonchai, Enghoff & Panha, sp. n. (type species) and S. sutchariti Srisonchai, Enghoff & Panha, sp. n. from Krabi Province, S. uncus Srisonchai, Enghoff & Panha, sp. n., and S. macaca Srisonchai, Enghoff & Panha, sp. n. from Phang Nga Province; as well as one from Malaysia, S. tortioverpa Srisonchai, Enghoff & Panha, sp. n., and one from Myanmar, S. efefi Srisonchai, Enghoff & Panha, sp. n. The new genus is endemic to South Myanmar, South Thailand, and Malaysia, and all new species are restricted to limestone habitats. All were exclusively found living on humid rock walls and/or inside small caves. Complete illustrations of external morphological characters, an identification key, and a distribution map are provided.
dragon millipede, endemic, new species, taxonomy, Thailand
This is the fourth paper in a series of articles about revision of the dragon millipedes.
The new genus is narrowly distributed in the Malay Peninsula (Malaysia, Myanmar, and Thailand).
Specimens were hand-collected from many localities throughout South Thailand, in some parts of Malaysia and in southern Myanmar. We also observed the habitats of all specimens. Specimens were mostly stored in 70% ethanol for morphological study and partly in 95% ethanol for molecular analysis. Latitude, longitude, and elevation were recorded by using a Garmin GPSMAP 60 CSx, and all coordinates and elevations were checked with Google Earth.
The main collectors in this work were staff and students of the Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University which we here refer to as ‘ASRU members’.
All living specimen photos were taken with a Nikon D700 equipped with a AF-S VR Micro-Nikkor 105 mm lens during fieldwork. Newly collected specimens preserved in ethanol were imaged with an Olympus DP72 camera on an Olympus SZX16 stereomicroscope, using image stacking Cell-D auto-montage software. Scanning electron micrographs were generated with a JEOL – JSM–5410 LV. All samples studied with SEM were carefully dissected under a microscope, mounted on aluminium stubs, and coated with gold. After imaging with SEM, all objects were removed and kept in dry condition. Drawings were outlined under a stereo microscope (Leica Wild M10) with a drawing tube and finished using dot-line technique (stipple). Plates were composed in Adobe Photoshop CS6.
We use morphological terminology according to previous taxonomic publications (
Spinaxytes krabiensis Srisonchai, Enghoff & Panha, gen. et sp. n.
The genus Spinaxytes gen. n. is characterized by:
1. Paraterga spiniform.
2. Metaterga with two rows of tubercles/cones/spines.
3. Postfemoral part of gonopod conspicuous, demarcated from femur by deep mesal and lateral sulci.
4. Lamina lateralis distinctly demarcated from lamina medialis.
5. Lamina medialis very long, curved, larger and longer than lamina lateralis.
The name is a noun in apposition; from the Latin ‘spina’, referring to the spine-like paraterga of all constituent species; ‘–xytes’ ensures harmony with Desmoxytes (and its synonym ‘Pteroxytes’).
1. Spinaxytes biloba Srisonchai, Enghoff & Panha, sp. n.
2. Spinaxytes efefi Srisonchai, Enghoff & Panha, sp. n.
3. Spinaxytes hasta Srisonchai, Enghoff & Panha, sp. n.
4. Spinaxytes krabiensis Srisonchai, Enghoff & Panha, sp. n.
5. Spinaxytes macaca Srisonchai, Enghoff & Panha, sp. n.
6. Spinaxytes palmata Srisonchai, Enghoff & Panha, sp. n.
7. Spinaxytes sutchariti Srisonchai, Enghoff & Panha, sp. n.
8. Spinaxytes tortioverpa Srisonchai, Enghoff & Panha, sp. n.
9. Spinaxytes uncus Srisonchai, Enghoff & Panha, sp. n.
The new genus is easily distinguished from other genera of dragon millipedes by having spiniform paraterga, lamina lateralis smaller and shorter than lamina medialis, lamina medialis long and curved. Some species of the genus Hylomus Cook & Loomis, 1924, share spine-like paraterga; however, the gonopod details are totally different.
General description of the genus Spinaxytes. The description applies to adult males and females, except for the gonopods and when “male” is specified (Figs
Size. Body length ca. 18–33 mm (male) ca. 16–33 mm (female), width 1.0–2.2 mm (male) 1.3–2.9 mm (female), size varies between species, usually female a bit longer than male.
Colour. Most species in life with dark brown colour. Colour in alcohol: all specimens partly faded to pale brown after 5 years’ preservation in alcohol; specimens kept in darkness faded more slowly.
Antennae (Figure
Head. Delicately setose; vertex, labrum and genae sparsely setose; epicranial suture conspicuous as a deep, brown or black stripe.
Collum (Figure
Tegument. Quite dull, sometimes shining; collum, metaterga and surface below paraterga smooth/microgranulate; prozona finely shagreened; paraterga, epiproct and sterna smooth. Stricture between prozona and metazona shallow, wide.
Metaterga (Figure
Pleurosternal carinae (Figure
Paraterga (Figs
Telson (Figure
General body characters of Spinaxytes gen. n. (S. palmata sp. n., ♂ paratype, CUMZ-pxDGT00216) – SEM images A, B anterior body part (arrowheads point to pleurosternal carinae) C collum D, E body rings 9–11 F telson G posteriormost rings and telson H body ring 10 I mouth parts, ventral view J gonopods K sternal lobe between coxae 4 L, N tip of epiproct M tip of tarsus and claw of leg 13.
Sterna (Figs
Legs (Figs
Gonopods (Figure
All species of the new genus are allopatric. However, some of the new species can be found in syntopy with some species of Desmoxytes. The nine species described here are confined to limestone habitats and have narrow distributions; we therefore regard them as endemic. All species blend perfectly with habitat environment, mostly living on humid rock walls and/or inside small caves.
According to the current knowledge, Spinaxytes gen. n. is distributed only in the Malay Peninsula (south Myanmar, south Thailand, and north Malaysia): Myanmar: Thanintharyi Region; Thailand: Chumphon Province (Thung Tako, Mueng Chumphon, Lang Suan and Sawi Districts), Surat Thani Province (Phanom District), Phang Nga Province (Mueng Phang Nga and Takua Thung Districts), Krabi Province (Ao Luek and Muaeng Krabi Districts); Malaysia: Perak State (Figure
1 | Male femora 6–9 without modification (e.g., Figs |
2 |
– | Male femora 6 and 7, or 7, or 8 and 9 humped/with apophyses (e.g., Figs |
6 |
2 |
Collum with 5+5 tubercles in anterior row, 2+2 tubercles in intermediate row, 3+3 tubercles in posterior row (Figure |
S. efefi sp. n. |
– | Collum with 4+4 tubercles/cones in anterior row, 1+1 in intermediate row, 2+2 in posterior row (e.g., Figs |
3 |
3 | Sternal lobe between male coxae 4 bilobed/bifurcate/subtrapeziform (e.g., Figs |
4 |
– | Sternal lobe between male coxae 4 spear-like (not bilobed, not bifurcate, not subtrapeziform) (Figs |
S. hasta sp. n. |
4 | Postfemoral part very large, angled 90 degrees with femoral part (Figs |
S. tortioverpa sp. n. |
– | Postfemoral part small, not angled 90 degrees with femoral part (e.g., Figs |
5 |
5 | Lamina lateralis small (Figs |
S. palmata sp. n. |
– | Lamina lateralis large (Figs |
S. sutchariti sp. n. |
6 | Only male femora 7 modified (Figs |
S. uncus sp. n. |
– | Male femora 6 and 7, or 8 and 9 modified (e.g. Figs |
7 |
7 | Lamina medialis with process-like lobe at base (Fig. |
S. biloba sp. n. |
– | Lamina medialis without process-like lobe at base (Figs |
8 |
8 |
Paraterga extremely long (Fig. |
S. krabiensis sp. n. |
– | Paraterga moderately long (Fig. |
S. macaca sp. n. |
Holotype. ♂, THAILAND, Surat Thani Province, Phanom District, near Khlong Phanom National Park, Pha Daeng, 8°53'41"N, 98°33'12"E, ca. 67 m a.s.l., 7 Aug. 2016, ASRU members leg. (CUMZ-pxDGT00205). Paratypes. 17 ♂♂, 24 ♀♀, same data as for holotype (CUMZ- pxDGT00206); 1 ♂, 1 ♀, same data as for holotype (ZMUC00040249); 1 ♂, 1 ♀, same data as for holotype (NHMW9423). Further specimens, not paratypes. 5 ♂♂, 3 ♀♀, THAILAND, Surat Thani Province, Phanom District, near Khlong Phanom National Park, Pha Daeng, 8°53'41"N, 98°33'12"E, ca. 67 m a.s.l., 6 Aug. 2015, ASRU members leg. (CUMZ).
The species name is an adjective, refers to the two additional process-like lobes on the solenophore (one on lamina lateralis and one on lamina medialis).
Differs from other species by having: metaterga 5–19 with 2+2 cones in anterior row and 3+3 cones in posterior row; sternal lobe between male coxae 4 subtrapeziform; male femora 8 and 9 with apophyses distally; lamina lateralis with an additional process-like protruding lobe; lamina medialis basally with an additional protruding process-like lobe.
SIZE. Length 15–17 mm (male), 16–18 mm (female); width of midbody metazona 1.0–1.2 mm (male), 1.3–1.5 mm (female). Width of rings 2 = 3 < 4 < collum < 5 < head = 6–17, thereafter body gradually tapering towards telson.
Colour (Figure
Antennae. Reaching to body ring 7 or 8 (male) and 6 (female) when stretched dorsally.
Collum (Figure
Tegument. Quite shining; collum coarsely microgranulate; metaterga and surface below paraterga finely microgranulate.
Metaterga (Figure
Paraterga (Figure
Telson (Figure
Sterna (Figs
Legs (Figs
Gonopods (Figs
(Figure
Among all Spinaxytes species, S. biloba sp. n. is obviously the smallest (length 15–18 mm, width of midbody metazona 1.0–1.5 mm), and the live pale brown colouration is lighter than that of other species.
Male femora and sternal lobe between male coxae 4 of Spinaxytes gen. n. A–C S. biloba sp. n. (A Sternal lobe B Femur 8 C Femur 9) D, E S. efefi sp. n. (D Sternal lobe E Femur 5 or 6) F, G S. hasta sp. n. (F Sternal lobe G Femur 5 or 6) H–J S. krabiensis sp. n. (H Sternal lobe I Femur 6 J Femur 7) K–M S. macaca sp. n. (K Sternal lobe L Femur 6 M Femur 7) N, O S. palmata sp. n. (N Sternal lobe O Femur 5 or 6) P, Q S. sutchariti sp. n. (P Sternal lobe Q Femur 5 or 6) R, S S. tortioverpa sp. n. (R Sternal lobe S Femur 5 or 6) T–V S. uncus sp. n. (T Sternal lobe U Femur 6 V Femur 7).
Holotype. ♂, MYANMAR, Tanintharyi Region, Myeik, 20 km northeast of Monoron, Lenya National Park, limestone mountain near Ngawun Chaung River, 11°40'20"N, 99°13'30"E, ca. 64 m a.s.l., 9 Jun. 2015, FFI staff and ASRU members leg. (CUMZ-pxDGT00207). Paratypes. 20 ♂♂, 25 ♀♀, same data as for holotype (CUMZ-pxDGT00208); 1 ♂, 1 ♀, same data as for holotype (ZMUC00040250); 1 ♂, 1 ♀, same data as for holotype (ZMUM); 1 ♂, 1 ♀, same data as for holotype (NHMW9422); 1 ♂, 1 ♀, same data as for holotype (NHMUK).
The name is an artificially constructed homophone (efefi = FFI) honouring FFI (Fauna and Flora International, Myanmar), an organization for biodiversity conservation; in recognition of their hard work to protect wildlife including invertebrates.
Sternal lobe between male coxae 4 not bilobed and male femora without modification. Similar in this respect to S. hasta sp. n., but differs by having: collum with 5(4)+5(4) tubercles in anterior row, 2+2 tubercles in intermediate row and 3+3 tubercles in posterior row; metaterga 2–8 with 3+3 cones in anterior row and 3+3 cones in posterior row; metaterga 9–18 with 3+3 cones in anterior row and 4+4 cones in posterior row; metatergum 19 with 3+3 tubercles/cones in anterior row and 4+4 tubercles/cones in posterior row; postfemoral part of gonopod with a triangular process and a triangular ridge.
SIZE. Length 26–30 mm (male), 30–32 mm (female); width of midbody metazona 2.1–2.2 mm (male), 2.7–2.9 mm (female). Width of collum = ring 2 = 3 = 4 < head = 5–17, thereafter body gradually tapering towards telson.
Colour (Figure
Antennae. Reaching to body ring 9 or 10 (male) and 7 or 8 (female) when stretched dorsally.
Collum (Figure
Tegument. Quite dull; collum, metaterga (posterior part) and surface below paraterga coarsely microgranulate; metaterga (anterior part) smooth.
Metaterga (Figure
Paraterga (Figure
Telson (Figure
Sterna (Figs
Legs (Figs
Gonopods (Figs
(Figure
No variation was found. Body ring 19 of S. efefi sp. n. seems to be shorter than in other species, and the tip of paraterga on collum is obtuse whereas in other species (except S. biloba sp. n.) it is sharp.
Holotype. ♂, THAILAND, Chumphon Province, Thung Tako District, Khao Ma Ngaen, 10°05'27"N, 99°04'25"E, ca. 28 m a.s.l., 23 Oct. 2016, ASRU members leg. (CUMZ-pxDGT00209). Paratypes. 5 ♂♂, 6 ♀♀, same data as for holotype (CUMZ-pxDGT00210); 1 ♂, 1 ♀, same data as for holotype (ZMUC00040251). Further specimens, not paratypes, all from THAILAND, Chumphon Province. Mueang Chumphon District: 8♂♂, 1 ♀, Wat Tham Sanook, 10°28'52"N, 99°04'29"E, ca. 54 m a.s.l., 3 Jul. 2017, ASRU members leg. (CUMZ). Lang Suan District: 2 ♀♀, Wat Ratcha Burana School, 9°56'21"N, 99°02'26"E, ca. 34 m a.s.l., 10 Sep. 2016, ASRU members leg. (CUMZ); 1 ♂, 5 ♀♀, Wat Tham Khao Kriap (Khao Kriap Cave), 9°49'08"N, 99°02'22"E, ca. 102 m a.s.l., 5 Jun. 2009, ASRU members leg. (CUMZ). Sawi District: 8 ♂♂, 3 ♀♀, Wat Nam Cha, 10°17'54"N, 99°01'57"E, ca. 95 m a.s.l., 5 Jun. 2009, ASRU members leg. (CUMZ).
The name is a Latin noun in apposition meaning spear, referring to the shape of the sternal lobe between male coxae 4 which is somewhat similar to a spear.
Sternal lobe between male coxae 4 not bilobed, not bifurcate; male femora without modification. Similar in this respect to S. efefi sp. n., but differs by having: collum with 4+4 tubercles in anterior row, 1+1 tubercles in intermediate row and 2+2 tubercles in posterior row; metaterga 2–8 with 2+2 cones in anterior row and 2+2 cones in posterior row; metaterga 9–18 with 2+2 cones in anterior row and 2+2 cones in posterior row; metatergum 19 with 2+2 tubercles/cones in anterior row and 2+2 tubercles/cones in posterior row; lamina medialis (lm) with a large lobe in middle part.
SIZE. Length 23–33 mm (male), 26–33 mm (female); width of midbody metazona 1.7–2.2 mm (male), 2.1–2.8 mm (female). Width of collum = ring 2 = 3 = 4 < head = 5–16, thereafter body gradually tapering towards telson.
Colour (Figure
Antennae (Figure
Collum (Figure
Tegument. Quite dull; collum, metaterga and surface below paraterga finely microgranulate.
Metaterga (Figure
Paraterga (Figure
Telson (Figure
Sterna (Figs
Legs (Figs
Gonopods (Figs
(Figure
There are some variations: the lateral setiferous tubercles of the epiproct are conspicuous in some specimens, inconspicuous in others; the hypoproct is subtrapeziform in some individuals, subsemicircular in others; the caudal margin of the hypoproct is rounded in some specimens, angular in others.
Holotype. ♂, THAILAND, Krabi Province, Mueang Krabi District, Wat Tham Sue (Tiger Cave), valley behind Tiger Cave, 8°07'38"N, 98°55'26"E, ca. 87 m a.s.l., 9 Jul. 2017, ASRU members leg. (CUMZ-pxDGT00211). Paratypes. 5 ♂♂, 9 ♀♀, 1 juvenile, same data as for holotype (CUMZ-pxDGT00212); 1 ♂, 1 ♀, same data as for holotype (ZMUC00040252). Further specimens, not paratypes, all from THAILAND, Krabi Province. Ao Luek District: 2 ♂♂, 1 ♀, Than Bok Khorani, 8°23'28"N, 98°44'07"E, ca. 46 m a.s.l., 14 Jan. 2013, ASRU members leg. (CUMZ); 5 ♂♂, 1 ♀, Than Bok Khorani, 8°23'28"N, 98°44'07"E, ca. 46 m a.s.l., 23 Aug. 2014, ASRU members leg. (CUMZ); 12 ♂♂, 7 ♀♀, 1 juvenile, Than Bok Khorani, 8°23'28"N, 98°44'07"E, ca. 46 m a.s.l., 30 Aug. 2015, ASRU members leg. (CUMZ); 3 ♂♂, Than Bok Khorani, 8°23'28"N, 98°44'07"E, ca. 46 m a.s.l., Jan. 2016, ASRU members leg. (CUMZ); 1 ♂, 5 ♀♀, P.N. Mountain Resort, 8°24'09"N, 98°44'18"E, ca. 46 m a.s.l., 30 Aug. 2015, ASRU members leg. (CUMZ); 1 ♂, 1 broken ♂, Tham Sa Yuan Thong (Sa Yuan Thong Cave), 8°23'29"N, 98°46'17"E, ca. 7 m a.s.l., 9 Oct. 2006, ASRU members leg. (CUMZ). Muaeng Krabi District: 1 broken ♂, Wat Tham Sue (Tiger Cave), valley behind Tiger Cave, 8°07'38"N, 98°55'26"E, ca. 87 m a.s.l., 25 Oct. 2007, ASRU members leg. (CUMZ); 1 broken ♂, 1 ♀, Wat Tham Sue (Tiger Cave), valley behind Tiger Cave, 8°07'38"N, 98°55'26"E, ca. 87 m a.s.l., 7 Oct. 2009, ASRU members leg. (CUMZ); 1 ♂, Wat Tham Sue (Tiger Cave), valley behind Tiger Cave, 8°07'38"N, 98°55'26"E, ca. 87 m a.s.l., 24 Aug. 2014, ASRU members leg. (CUMZ); 1 ♀, Wat Tham Sue (Tiger Cave), valley behind Tiger Cave, 8°07'38"N, 98°55'26"E, ca. 87 m a.s.l., 30 Aug. 2015, P. Pimvichai, P. Prasankok and N. Natarat leg. (CUMZ); 2 ♂♂, 1 ♀, 1 broken ♀, Wat Tham Sue (Tiger Cave), valley behind Tiger Cave, 8°07'38"N, 98°55'26"E, ca. 87 m a.s.l., 25 Jul. 2017, ASRU members leg. (CUMZ).
The new species is named after the province where the type locality lies.
Male femora 6 and 7 humped distally. Similar in this respect to S. macaca sp. n., but differs by having: paraterga orange, longer; male femora 6 smaller; tip of lamina lateralis round, not protuding as digitiform; tip of lamina medialis terminating in two lobes.
SIZE. Length 28–31 mm (male), 30–33 mm (female); width of midbody metazona 1.8–2.0 mm (male), 2.2–2.5 mm (female). Width of rings 2 = 3 = 4 < collum < head = 5–16, thereafter body gradually tapering towards telson.
Colour (Figure
Antennae (Figure
Collum (Figure
Tegument. Quite shining; collum coarsely microgranulate; metaterga and surface below paraterga smooth.
Metaterga (Figure
Paraterga (Figure
Telson (Figure
Sterna (Figs
Legs (Figs
Gonopods (Figs
(Figure
We found variations in the lateral setiferous tubercles of the epiproct: conspicuous in some specimens, inconspicuous in others.
Holotype. ♂, THAILAND, Phang Nga Province, Takua Thung District, Wat Suwan Khuha (Monkey Cave), 8°25'42"N, 98°28'22"E, ca. 27 m a.s.l., 8 Aug. 2016, ASRU members leg. (CUMZ-pxDGT00213). Paratypes. 7 ♂♂, 2 ♀♀, same data as for holotype (CUMZ-pxDGT00214); 1 ♂, 1 ♀, same data as for holotype (ZMUC00040253); 1 ♂ same data as for holotype (ZMUM).
The species is named after the monkey, long-tailed macaque (Macaca fascicularis) living at the type locality (Monkey Cave).
Male femora 6 and 7 humped distally. Similar in this respect to S. krabiensis sp. n., but differs from it by having: paraterga brownish white, shorter; male femora 6 larger; tip of lamina lateralis (ll) protruding as a small lobe, digitiform; tip of lamina medialis (lm) bent, terminating in one lobe.
SIZE. Length 27–29 mm (male), 29–32 mm (female); width of midbody metazona 1.8–1.9 mm (male), 2.0–2.3 mm (female). Width of collum = 2 = 3 < 4 < head = 5–16, thereafter body gradually tapering towards telson.
Colour (Figure
Antennae (Figure
Collum (Figure
Tegument. Quite shining; collum, metaterga and surface below paraterga smooth.
Metaterga (Figure
Paraterga (Figure
Telson (Figure
Sterna (Figs
Legs (Figs
Gonopods (Figs
(Figure
S. macaca sp. n. is morphologically similar to S. krabiensis sp. n. and S. uncus sp. n. with which it shares a fork-like sternal lobe between male coxae 4 and a small lamina lateralis.
Holotype. ♂, THAILAND, Surat Thani Province, Phanom District, Wat Tham Wararam, 8°53'07"N, 98°40'01"E, ca. 51 m a.s.l., 6 Aug. 2016, ASRU members leg. (CUMZ-pxDGT00215). Paratypes. 23 ♂♂, 22 ♀♀, same data as for holotype (CUMZ-pxDGT00216); 1 ♂, 1 ♀, same data as for holotype (ZMUC00040254); 1 ♂, 1 ♀, same data as for holotype (ZMUM); 1 ♂, 1 ♀, same data as for holotype (NHMW9425); 1 ♂, 1 ♀, same data as for holotype (NHMUK). Further specimens, not paratypes. THAILAND: 1 ♂, 2 ♀♀, Surat Thani Province, Phanom District, Wat Tham Wararam, 8°53'07"N, 98°40'01"E, ca. 51 m a.s.l., 5 Aug. 2014, ASRU members leg. (CUMZ).
The species name is a Latin adjective, referring to the tip of lamina medialis which is somewhat hand-shaped.
Male femora without modification. Similar in this respect to S. efefi sp. n., S. hasta sp. n., S. sutchariti sp. n. and S. tortioverpa sp. n., but differs from them by having: anterior part of sternal lobe between male coxae 4 bifurcate, fork-like; tip of lamina medialis expanded, hand-shaped.
SIZE. Length 26–30 mm (male), 27–32 mm (female); width of midbody metazona 1.9–2.2 mm (male), 2.0–2.4 mm (female). Width of collum = 2 = 3 = 4 < head < 5–16, thereafter body gradually tapering towards telson.
Colour (Figure
Antennae (Figure
Collum (Figure
Tegument. Very shining; collum coarsely microgranulate; metaterga and surface below paraterga smooth.
Metaterga (Figure
Paraterga (Figure
Telson (Figure
Sterna (Figs
Legs (Figs
Gonopods (Figs
(Figure
There are variations in the hypoproct: subtrapeziform in some specimens, subtriangular in the others; caudal margin in some individuals round, angular in the others.
Holotype. ♂, THAILAND, Krabi Province, Muang Krabi District, Tham Na Mee (Na Mee Cave), 8°08'12"N, 98°48'23"E, ca. 70 m a.s.l., 31 Aug. 2015, C. Sutcharit leg. (CUMZ-pxDGT00217). Paratypes. 7 ♂♂, 4 ♀♀, same data as for holotype (CUMZ-pxDGT00218).
The name honours associate professor Dr. Chirasak Sutcharit, malacologist of ASRU (CUMZ), collector of this new species and numerous other dragon millipedes.
Male femora without modification, sternal lobe between male coxae 4 incompletely bilobed. Similar in these respects to S. palmata sp. n., but differs by having: a large and round lamina lateralis; tip of lamina medialis terminating in two spines; distal part of solenomere circular in tranverse section.
SIZE. Length 20–25 mm (male), 23–27 mm (female); width of midbody metazona 1.5–1.8 mm (male), 1.9–2.2 mm (female). Width of collum = 2 = 3 = 4 < head = 5–16, thereafter body gradually tapering towards telson.
Colour (Figure
Antennae. Reaching to body ring 8 or 9 (male) and 6 (female) when stretched dorsally.
Collum. With three transverse rows of setiferous tubercles, 4+4 tubercles in anterior row, 1+1 tubercles in intermediate row and 2+2 tubercles in posterior row; with one inconspicuous setiferous notch at lateral margin; paraterga spiniform, quite short, tip sharp, elevated at ca. 15°–20° in both male and female, directed caudolaterad.
Tegument. Quite dull; collum and metaterga (posterior part) coarsely microgranulate; metaterga (anterior part) and surface below paraterga smooth.
Metaterga. With two transverse rows of setiferous tubercles and setiferous spines; metaterga 2–19 with 2+2 tubercles in anterior row and 2+2 spines in posterior row; lateral spines of posterior row bigger and longer than mesal ones, subequal in size and length on all body rings.
Paraterga. Long; directed dorsolaterad on body rings 2–16, elevated at ca. 45°–50° (male) 40°–50° (female), directed dorsocaudad on ring 17, directed increasingly caudad on body rings 18 and 19. Ozopore visible in lateral view.
Telson. Epiproct quite long; tip subtruncate; lateral setiferous tubercles conspicuous; apical tubercles inconspicuous. Hypoproct subtrapeziform (in some specimens subtriangular); caudal margin round (in some specimens angular), with inconspicuous setiferous tubercles.
Sterna (Figure
Legs (Figure
Gonopods (Figs
(Figure
We found variation in the hypoproct: in some specimens subtrapeziform, in others subtriangular; caudal margin in some individuals round, in others angular. Parasitic mite larvae, probably of the genus Leptus Latreille, 1796, were found attached to the anterior body part of some female specimens. Larvae of ?Leptus have previously been found on species of Desmoxytes (D. cervina) and Nagaxytes (N. acantherpestes (Golovatch & Enghoff, 1994)) (
Holotype. ♂, MALAYSIA, Perak State, Ipoh City, Gua Tempurung, limestone mountain, 4°24'58"N, 101°11'16"E, ca. 92 m a.s.l., 27 Sep. 2007, B. W. Ng and ASRU members leg. (CUMZ-pxDGT00219). Paratypes. 1 ♂, 1 ♂ gonopods missing, 1 ♀, same data as for holotype (CUMZ-pxDGT00220).
The name is a noun in apposition, from the Latin tortio meaning torsion and verpa for penis, refers to the distal part of gonopod (postfemoral part, solenophore and solenomere) which makes a 90 degrees torsion relative to the femoral part.
Male femora without modification; collum with 4+4 tubercles in anterior row, 1+1 tubercles in intermediate row and 2+2 tubercles in posterior row. Similar in these respects to S. hasta sp. n. and S. palmata sp. n., but differs from them by having: a completely bilobed sternal lobe between male coxae 4; postfemoral part, solenophore and solenomere angled 90 degrees with femoral part; lamina lateralis with two lobes – the lateral one spine-like, the mesal one shorter and ridge-like; solenomere very long, longer than lamina medialis.
SIZE. Length 28–30 mm (male), 30–33 mm (female); width of midbody metazona 2.0 mm (male), 2.8 mm (female). Width of rings 2 = 3 = 4 < head = 5–16, thereafter body gradually tapering towards telson.
Colour. Specimens in life with body black/brownish black. Colour in alcohol: after 10 years changed to brown; paraterga brownish white; antennae brown (except whitish distal part of antennomeres 7 and 8); head, collum, metaterga and prozona blackish brown; surface below paraterga brown/blackish brown; sterna, epiproct and legs pale brown; a few basal podomeres whitish brown.
Antennae. Reaching to body ring 8 or 9 (male) and 6 or 7 (female) when stretched dorsally.
Collum. With three transverse rows of setiferous tubercles, 4+4 tubercles in anterior row, 1+1 tubercles in intermediate row and 2+2 tubercles in posterior row; anterior margin truncate; with one inconspicuous setiferous notch at lateral margin; paraterga wing-like, long and broad, tip sharp, elevated at ca. 15°–20° in both male and female, directed caudolaterad.
Tegument. Quite dull; collum, metaterga and surface below paraterga finely microgranulate.
Metaterga. With two transverse rows of setiferous tubercles/cones and spines; metaterga 2–8 with 2+2 cones in anterior row and 2+2 spines in posterior row; metaterga 9–19 with 2+2 tubercles/cones in anterior row and 2+2 spines in posterior row; mesal spines of posterior row bigger and longer than lateral ones, gradually reduced in length and size on posterior rings.
Paraterga. Very long; directed dorsolaterad on body rings 2–17, elevated at ca. 65°–70° (male) 60°–70° (female), directed increasingly caudad on body rings 18 and 19. Ozopore visible in dorsolateral view.
Telson. Epiproct quite short; tip subtruncate; lateral setiferous tubercles inconspicuous; apical tubercles inconspicuous. Hypoproct subsemicircular; caudal margin round, with inconspicuous setiferous tubercles.
Sterna (Figure
Legs (Figure
Gonopods (Figs
Known only from the type locality which is currently a tourist attraction (cave). We regard this species as endemic to Malaysia.
A photograph of a live specimen was not taken during the field survey, but our collector noticed its black or brownish black colour. All specimens were seen crawling on rock walls where they seem to blend perfectly with the substrate. No variation in morphological characters was found.
Holotype. ♂, THAILAND, Phang Nga Province, Mueang Phang Nga District, Phung Chang Cave, 8°26'34"N, 98°30'59"E, ca. 24 m a.s.l., 8 Aug. 2016, ASRU members leg. (CUMZ-pxDGT00221). Paratypes. 15 ♂♂, 16 ♀♀, same data as for holotype (CUMZ-pxDGT00222); 1 ♂, 1 ♀, same data as for holotype (ZMUC00040255); 1 ♂, 1 ♀, same data as for holotype (ZMUM); 1 ♂, 1 ♀, same data as for holotype (NHMW9424). Further specimens, not paratypes, all from THAILAND, Phang Nga Province. Muaeng Phang Nga District: 4 ♂♂, 2 ♀♀, Phung Chang Cave, 8°26'34"N, 98°30'59"E, ca. 24 m a.s.l., 6 Aug. 2014, ASRU members leg. (CUMZ); 4 ♂♂, Phung Chang Cave, 8°26'34"N, 98°30'59"E, ca. 24 m a.s.l., 5 Aug. 2015, ASRU members leg. (CUMZ); 2 ♂♂, 4 broken ♀♀, Tham Nam Pud, 8°27'50"N, 98°32'36"E, ca. 58 m a.s.l., 8 Oct. 2006, ASRU members leg. (CUMZ); 3 ♂♂, 2 ♀♀, Tham Nam Pud, 8°27'50"N, 98°32'36"E, ca. 58 m a.s.l., 5 Aug. 2015, ASRU members leg. (CUMZ); 2 ♂♂, 1 ♀, Wat Tham Bang Toei, 8°27'52"N, 98°34'10"E, ca. 24 m a.s.l., 10 Jul. 2017, ASRU members leg. (CUMZ); 8 ♂♂, 2 ♀♀, Tham Pha Phueng Bureau of Monks, 8°28'24"N, 98°32'15"E, ca. 78 m a.s.l., 10 Jul. 2017, ASRU members leg. (CUMZ). Thap Put District: 11 ♂♂, 3 ♀♀, Wat Kerewong (Tham Koab), 8°31'52"N, 98°34'39"E, ca. 76 m a.s.l., 9 Jul. 2017, ASRU members leg. (CUMZ).
The name is a Latin noun in apposition (uncus), meaning hook, and refers to the hook-like lamina medialis of gonopod.
Differs from other species by having only male femora 7 strongly humped distally, in combination with the distal part of lamina medialis hook-like, tip long and sharp.
SIZE. Length 20–27 mm (male), 25–29 mm (female); width of midbody metazona 1.4–1.6 mm (male), 2.1–2.3 mm (female). Width of collum < 2 = 3 < 4 < head = 5–16, thereafter body gradually tapering towards telson.
Colour (Figure
Antennae (Figure
Collum (Figure
Tegument. Quite shining; collum, metaterga (posterior part) and surface below paraterga finely microgranulate; metaterga (anterior part) coarsely microgranulate.
Metaterga (Figure
Paraterga (Figure
Telson (Figure
Sterna (Figs
Legs (Figs
Gonopods (Figs
(Figure
Some variation in the hypoproct was observed in this species: in some specimens subtrapeziform, in others subsemicircular; caudal margin in some individuals conspicuous, in others inconspicuous. In addition, specimens from Wat Kerewong (Tham Koab) have smaller and shorter paraterga than other specimens.
The new genus Spinaxytes, defined by
Based on a comparatively large number of specimens and species of the new genus, our study confirmed that gonopod characters can be used confidently to discriminate the species, just as we found in Desmoxytes and Nagaxytes (
Almost all specimens collected by us were found on humid rock walls in small caves. Therefore, we strongly suspect that all species in this genus prefer to live on rock walls. The black or dark brown body colour makes them difficult to see against dark-coloured rocks. Quite often some specimens of Desmoxytes and Gigaxytes species are encountered in the same habitat as species of the new genus, but it seems likely that those species live on leaf litter, on the ground or on tree branches instead of rock walls. Considering currently known distributions of species of Spinaxytes gen. n., and their restriction to small limestone areas (Figure
The discovery rate of new dragon millipede species has been increasing in recent years (
The main funding for this work is from the Thailand Research Fund, the TRF Senior Research Scholar (2015–2018), RTA 5880002 and BDC-PG2-161002 to SP. RS’s visit for one year to the Natural History Museum of Denmark, University of Copenhagen, was granted by the Human Resource Development in Science Project (Science Achievement Scholarship of Thailand (SAST)) and an internal grant from the Natural History Museum of Denmark. We gratefully acknowledge the Plant Genetic Conservation Project under the Initiative of Her Royal Highness Maha Chakri Sirindhorn and Center of Excellence on Biodiversity for permission to field work in several restricted/remote areas. This work would not have been possible at all without the impressive collections made by ASRU members, especially Dr. Chirasak Sutcharit for his kind help in collecting numerous specimens. Thanks are due to B. W. Ng. (Malaysia) and Fauna & Flora International (Myanmar) for helping us to collect specimens in Malaysia and Myanmar. We are the most grateful to the subject editor R Mesibov, and journal referees, SI Golovatch and N Akkari, for all valuable comments and advices that greatly improved the manuscript. The first author also thanks to Ms Thita Krutchen for her kind teaching the drawing skills.