Research Article |
Corresponding author: Weixin Liu ( da2000wei@163.com ) Academic editor: Didier Vanden Spiegel
© 2014 Weixin Liu, Sergei I. Golovatch, Mingyi Tian.
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:
Liu W, Golovatch SI, Tian M (2014) A review of the dragon millipede genus Desmoxytes Chamberlin, 1923 in China, with descriptions of four new species (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys 448: 9-26. https://doi.org/10.3897/zookeys.448.8081
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Four new species of Desmoxytes are described from southern China: D. lingulata sp. n., D. parvula sp. n., and D. nodulosa sp. n., from Guangxi Zhuang Autonomous Region, and D. getuhensis sp. n. from Guizhou Province. In addition, new records of D. scutigeroides Golovatch, Geoffroy & Mauriès, 2010 and D. scolopendroides Golovatch, Geoffroy & Mauriès, 2010 are provided, with a modified key to Desmoxytes species currently known to occur in China. Two of the new species, D. nodulosa sp. n. and D. getuhensis sp. n., seem to be troglobites.
Diplopod, Desmoxytes , new species, cave, troglobite, key, China
Desmoxytes Chamberlin, 1923 is a large, common, rather well defined, southeast Asian genus of the basically oriental millipede tribe Orthomorphini, subfamily Paradoxosomatinae, family Paradoxosomatidae (
At present, China supports 10 species of Desmoxytes, including 7 presumed troglobites. Unlike the epigean congeners usually demonstrating bright live colorations, the cavernicolous Desmoxytes are typically poorly pigmented and appear to be confined to caves in southern China while the genus is the sole among oriental Paradoxosomatidae to contain troglobites (
D. cornuta Zhang & Li, 1982, from Guangxi, Guilin, Yangshuo (
D. draco Cook & Loomis, 1924, from Jiangxi, Jiujiang, Lushan Mountains (
D. eupterygota Golovatch, Li, Liu & Geoffroy, 2012, from two caves in Hunan, Chenzhou, Linwu (
D. longispina Loksa, 1960, from a cave in Guangxi (no exact locality is known) (
D. lui Golovatch, Li, Liu & Geoffroy, 2012, from a cave in Guangxi, Yongfu (
D. minutubercula Zhang, 1986, from Guangxi, Tianlin (
D. planata Pocock, 1895, from a cave in Yunnan, Luxi, but basically nearly pantropical (
D. scolopendroides Golovatch, Geoffroy & Mauriès, 2010, from a cave in Guangxi, Huanjiang (
D. scutigeroides Golovatch, Geoffroy & Mauriès, 2010, from several caves in Guangxi, Huanjiang (
D. spinissima Golovatch, Li, Liu & Geoffroy, 2012, from a cave in Guangxi, Fuchuan. (
The present paper describes a further four new species of Desmoxytes, two of which seem to be troglobites, as well as provides new records of two known presumed troglobitic congeners.
The holotypes and a number of paratypes are deposited in the zoological collection of the South China Agricultural University, Guangzhou, China (SCAU), with some material also to be housed in the Institute of Zoology, Chinese Academy of Sciences, Beijing, China (IZAS), and Zoological Museum, State University of Moscow, Russia (ZMUM). The methods and terminology used are after
♂ (SCAU), China, Guangxi, Guilin City, Pingle County, Ertang Town, Chaotianyan, 24°37.075'N, 110°45.501'E, 257 m, 29.IV.2013, leg. Tian Mingyi, Liu Weixin, Sun Feifei & Yin Haomin.
3 ♂ (SCAU), same locality and collecting data as of the holotype.
To emphasize a peculiar, paramedian, linguiform, sternal process between ♂ coxae 5.
Differs from congeners in the paraterga being antler-shaped, the humped ♂ femur 6, combined with small, setose tubercles between ♂ coxae 3 and a peculiar sternal process between ♂ coxae 5, as well as the shout and curved gonopod femorite and a condensed solenophore.
Length ca 18.0–18.5 mm (♂), width of pro- and metaterga together with paraterga 0.8–1.0 and 1.8–2.0 mm (♂), respectively. Holotype 18.0 mm long, 0.8 and 2.0 mm wide on midbody pro- and metazonae, respectively. Head broadest, 1.2–1.4 mm (♂) (Fig.
Desmoxytes lingulata sp. n., ♂ paratype from Ertang Twon, Chaotianyan. A, D anterior part of body, ventral and dorsal views, respectively B, E midbody segments, ventral and dorsal views, respectively C, F posterior part of body, ventral and dorsal views, respectively G sternal process in the middle of ♂ sternum 5 in situ, ventral view H gonopods in situ, ventral view I femur 6, lateral view. Scale bars: A–F, I = 0.5 mm; G, H = 0.2 mm.
Prozonae very delicately microalveolate; surface below paraterga 2–4 rather shagreened and microspinulate (Fig.
Desmoxytes lingulata sp. n., ♂ paratype from Ertang Twon, Chaotianyan. A Collum B metatergum 2, dorsal view C segment 10, dorsal view D sternal cones between coxae 3, ventral view E sternal processes between coxae 5, ventral view F femur 6, front view. Scale bar: A–B, D–E = 0.5 mm; C, F = 1.0 mm.
Sterna sparsely setose, cross-impressions faint (Fig.
Gonopods (Figs
This species seems to be especially similar to D. cornuta Zhang & Li, 1982, from Guangxi, Guilin, Yangshuo. Obvious differences lie in a peculiar linguiform sternal process between ♂ coxae 5, combined with the stout, curved gonopod femorite and a condensed solenophore in D. lingulata sp. n., as opposed to an elongated and suberect one in D. cornuta (cf.
♂ (SCAU), China, Guangxi, Hechi City, Du’an County, Xia’ao Town, cave I, 24°15.144'N, 107°56.272'E, 347 m, 2.V.2013, leg. Tian Mingyi, Liu Weixin, Sun Feifei & Yin Haomin.
1 ♀ (SCAU), same locality and collecting data as of the holotype.
To emphasize the small size of this species.
Differs from congeners in the combination of spiniform paraterga, a paramedian pair of subtrapzoidal processes between ♂ coxae 4, the humped ♂ femur 6, and certain details of gonopod structure.
Length ca 18 (♂) or 19 mm (♀), width of pro- and metaterga together with paraterga 0.8 and 1.2 (♂), or 1.0 and 1.4 mm (♀), respectively. Head broadest, 1.3 mm (♂) or 1.5 mm (♀) wide. Coloration of material rather uniformly brownish, antennae and lateral body parts dark brown, venter and a few basal podomeres yellowish, basal parts paraterga pink (Fig.
Desmoxytes parvula sp. n., ♂ holotype from Xia’ao Town, cave I. A, D anterior part of body, lateral and dorsal views, respectively B, E midbody segments, dorsal and ventral views, respectively C, F posterior part of body, dorsal and ventral views, respectively G cross-section of a midbody segment, frontal view H sternal process between coxae 4 in situ, ventral view I femur 6, lateral view. Scale bars: A–G, I = 0.5 mm; H = 0.2 mm.
Pro- and metazonae very delicately microalveolate, metaterga finely shagreened and transversely rugulose, surface below paraterga finely shagreened (Fig.
Sterna moderately setose, cross-impressions very weak (Fig.
Gonopods (Fig.
Even though this species has been taken from a cave, it hardly represents a true cavernicole as it is rather strongly pigmented and shows short antennae and legs.
♂ (SCAU), China, Guangxi, Hechi City, Du’an County, Xia’ao Town, near Xia’ao Middle School, cave II, 24°17.987'N, 107°57.146'E, 317 m, 3.V.2013, leg. Tian Mingyi, Liu Weixin, Sun Feifei & Yin Haomin.
3 ♂, 4 ♀ (SCAU), 1 ♂, 1 ♀ (IZAS), 1 ♂, 1 ♀ (ZMUM), same locality, and collecting data as of the holotype. 1 ♀ (SCAU), same county, Yong’an Town, Yong’an Village, cave I, 24°14.659'N, 108°03.032'E, 287 m; 1 ♀ (SCAU), same town, Anju Village, cave Suidao Dong, 24°13.340'N, 108°05.694'E, 311 m, 3.V.2013, leg. Tian Mingyi, Liu Weixin, Sun Feifei & Yin Haomin; 1 ♂, 2 ♀ (SCAU), same county, Longwan Town, Qunle Village, cave I, 23°56.021'N, 108°10.962'E, 459 m, 27.VI.2013, leg. Tian Mingyi, Lin Wei, Liu Weixin, Yin Haomin & Huang Sunbin.
To emphasize the humped ♂ femora 5–7.
Differs from congeners in most of the paraterga being wing-shaped, combined with the humped ♂ femora 5–7, the sternal process present between ♂ coxae 4, occasionally also between ♂ coxae 3, as well as a short gonopod femorite and a strongly condensed solenophore.
Length ca 19–22 (♂) or 20–23 mm (♀), width of midbody pro- and metaterga together with paraterga 1.0–1.5 and 2.2–2.8 (♂), or 1.8–2.0 and 2.8–3.0 mm (♀), respectively. Holotype 21.0 mm long, 1.5 and 2.5 mm wide on midbody pro- and metaterga, respectively. Coloration of material varying from pallid to rather uniformly dark brownish (Fig.
Desmoxytes nodulosa sp. n., ♂ paratype from Xia’ao Town, near Xia’ao Middle School, cave II. A, D anterior part of body, subdorsal and ventral views, respectively B, E midbody segments, dorsal and ventral views, respectively C posterior part of body, dorsal view F gonopods in situ, ventral view G c ross-section of a midbody segment, caudal view H sternal processes between coxae 3 and 4, ventral view I femora 5–7, ventral view. Scale bars: 0.5 mm.
Prozonae very delicately microalveolate, but shining; collum, metaterga, paraterga and surface below paraterga finely shagreened and microgranulate, moderately setose in posterior parts of metaterga (Fig.
Desmoxytes nodulosa sp. n., ♂ paratype from Xia’ao Town, near Xia’ao Middle School, cave II. A Collum B matatergum 10, dorsal view C–E femora 5–7, lateral view F sternal processes between coxae 3, ventral view G sternal processes between coxae 4, front view. Scale bar: A–E = 1.0 mm; F–G = 0.5 mm.
Sterna sparsely setose, cross-impressions visible (Fig.
Gonopods (Figs
Although the coloration of this species is quite variable, based on several troglomorphic traits such as some individuals being completely unpigmented, and the antennae and legs clearly elongated, this species may well be a troglobite.
♂ (SCAU), China, Guizhou Prov., Anshun City, Ziyun County, Getuhe National Geopark, cave Suidao Dong, 25°41.32'N, 106°18.26'E, 950 m, 28.XII.2012, leg. Tian Mingyi, Liu Weixin, Sun Feifei & Yin Haomin.
2 ♂, 5 ♀, 1 ♂ juv., 1 ♀ juv. (SCAU), 1 ♂, 1 ♀ (IZAS), 1 ♂, 1 ♀ (ZMUM), same locality and collectiong data as of the holotype. 1 ♂, 6 ♀ (SCAU), same locality, cave Taiyang Dong, 25°41.55'N, 106°14.27'E, 1056 m, 28.XII.2012, leg. Tian Mingyi, Liu Weixin, Sun Feifei & Yin haomin.
To emphasize the location of the new species within the Getuhe National Geopark.
Differs from congeners in the paraterga being long and spiniform throughout, and the antennae and legs very long, combined with setose tubercles between ♂ coxae 4, the humped ♂ femur 6, and the gonopods strongly condensed.
Length ca 23–27 (♂) or 25–28 mm (♀); width of pro- and metaterga together with paraterga 1.2–1.4 and 2.5–3.0 (♂) or 1.5–2.0 and 2.8–3.0 mm (♀), respectively. Holotype 26.0 mm long, 1.2 and 2.5 mm wide on midbody pro- and metaterga, respectively. Coloration of material rather uniformly light brownish to nearly pallid, anterior body part a little darker, some specimens pinkish (Fig.
Desmoxytes getuhensis sp. n., ♂ paratype from Getuhe National Geopark, cave Taiyang Dong. A, D anterior part of body, sublateral and dorsal views, respectively B, E midbody segments, lateral and dorsal views, respectively C, F posterior part of body, lateral and dorsal views, respectively G sternal process between coxae 4 in situ, ventral view H gonopods in situ, ventral view I femur 6, lateral view. Scale bars: A–F, I = 1.0 mm, G, H = 0.5 mm.
Tegument rather shining and smooth, prozonae delicately microalveolate, metaterga and surface below paraterga finely shagreened to microgranulate (Fig.
Sterna quite spasely setose, cross-impressions weak (Fig.
Gonopods (Figs
Based on several troglomorphic traits such as some individuals being nearly unpigmented, and the antennae and legs very strongly enlongated, this species seems to be a troglobite.
Desmoxytes scutigeroides Golovatch, Geoffroy & Mauriès, 2010: 58.
Desmoxytes scutigeroides –
1 ♂, 1 ♀ (SCAU), China, Guangxi, Du’an County, Disu Town, Dading Village, cave II, 23°56.34'N, 108°0.32'E, 26.VI.2013, leg. Tian Mingyi, Lin Wei, Yin Haomin & Huang Sunbin; 1 ♂, 1 ♀, 1 ♀ fragment (SCAU), same county, Longwan Town, Nongqu Village, cave I, 23°56.021 N, 108°10.962 E, 459 m, 27.VI.2013, leg. Tian Mingyi, Liu Weixin, Lin Wei, Yin Haomin & Huang Sunbin.
This species has been described from a few caves in Huanjiang County, Guangxi, China while the new samples derive from two caves in the neighbouring Du’an County, Guangxi. The above material is in good agreement with the original description by
Desmoxytes scolopendroides Golovatch, Geoffroy & Mauriès, 2010: 60.
Desmoxytes scolopendroides –
1 ♀ (SCAU), China, Guangxi, Du’an County, Gaoling Town, Jinzhu Village, cave I, 24°06.547'N, 108°04.785'E, 190 m, 3.V.2013, leg. Tian Mingyi; 1 ♀ (SCAU), same locality, cave II, 24°06.514'N, 108°04.695'E, 218 m, 3.V.2013, leg. Liu Weixin; 5 ♂, 5 ♀ (SCAU), same county, Xia’ao Town, cave I, 24°15.144'N, 107°56.272'E, 347 m, 2.V.2013, leg. Tian Mingyi, Liu Weixin, Sun Feifei & Yin Haomin; 1 ♀, 4 ♂ juv., 8 ♀ juv. (SCAU), same cave, 28.VI.2013, leg. Tian Mingyi, Liu Weixin, Lin Wei, Yin Haomin & Huang Sunbin; 3 ♂, 3 ♀ (SCAU), same cave, 28.XII.2013, leg. Tian Mingyi, Liu Weixin, Yin Haomin & Luo Xiaozhu.
This species has been described from a cave in Huanjiang County, Guangxi, China while the new samples come from a few more caves in the neighbouring Du’an County, Guangxi. The above material is in good agreement with the original description by
1 | Paraterga spiniform, mostly very long and directed evidently more dorsad than laterad (Figs |
2 |
– | Paraterga wing- (Fig. |
8 |
2 | Adult body relatively small, length <20 mm | D. parvula sp. n. |
– | Adult body much larger, length >20 mm | 3 |
3 | Paraterga long and spiniform only on collum and following four segments, evidently shorter on segment 5, small and coni- to tuberculiform thereafter | D. lui |
– | Paraterga subequally long and spiniform at least in segments 2–18 (Fig. |
4 |
4 | Only ♂ femur 7 very evidently humped distoventrally | D. longispina |
– | Either ♂ femur 6 or both femora 6 and 7 very evidently humped distoventrally | 5 |
5 | Both ♂ femora 6 and 7 very evidently humped ventrally in distal quarter | D. spinissima |
– | Only ♂ femur 6 very evidently humped distoventrally | 6 |
6 | Metaterga not only with normally arranged setigerous tubercles, but also with a row of similar tubercles along posterior rim | D. minutubercula |
– | Metaterga 2–4 with several transverse rows of setigerous spines, following metaterga smooth, sculpture gradually disappearing | 7 |
7 | Gonopods telopodite subfalcate, femorite stouter relative to a condensed solenophore; Guangxi | D. scutigeroides |
– | Gonopods (Figs |
D. getuhensis sp. n. |
8 | Paraterga wing-shaped | 9 |
– | Paraterga antler-shaped | 12 |
9 | ♂ femora unmodified | D. eupterygota |
– | At least a pair of ♂ femora (5–7) humped | 10 |
10 | Metaterga 2–19 with only two transverse rows of 2+2(3) setigerous spines | 11 |
– | Metaterga 2–19 with more than two transverse rows of setigerous spines | D. scolopendroides |
11 | ♂ femora 5–7 very evidently humped distoventrally (Figs |
D. nodulosa sp. n. |
– | ♂ femora 5 and 6 slightly humped distoventrally; two processes between ♂ coxae 4 | D. planata |
12 | Paraterga 2–18 antler-shaped, evidently branched; Jiangxi | D. draco |
– | Anterior paraterga evidently antler-shaped, posterior paraterga rather long and spiniform, evidently 2- or 3-dentate laterally; Guangxi | 13 |
13 | A pair of setose tubercles between ♂ coxae 3, and a peculiar linguiform sternal process between ♂ coxae 5 (Figs |
D. lingulata sp. n. |
– | A pair of bristle-like tubercles between ♂ coxae 3, and a very deeply divergent sternal process between ♂ coxae 5 | D. cornuta |
We thank Ms. Bo Jiang, Biodiversity Conservation official, Department of Environmental Protection, Guangxi Regional Government, Nanning, for her support and encouragement. Our thanks are also due to Mr. Weihu Yang, Getuhe National Geopark, and members of our team in SCAU, for their assistance. Two reviewers kindly provided their suggestions and critical remarks which were helpful to improve the manuscript. This study was sponsored by Nanjing Institute of Environmental Sciences, Ministry of Environmental Protection, through a biodiversity conservation project in 2013 which was focused on cave biodiversity, and the Specialized Research Fund for the Doctoral Program of Higher Education of China (Grant no. 20134404110026).