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Research Article
Diversity, distribution patterns, and fauno-genesis of the millipedes (Diplopoda) of mainland China
expand article infoSergei I. Golovatch, Weixin Liu§
‡ Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Moscow, Russia
§ South China Agricultural University, Guangzhou, China
Open Access

Abstract

Based on all available information, 339 species from 71 genera, 26 families, and eleven orders of Diplopoda have hitherto been recorded from mainland China, the fauna thus being very rich, albeit far from completely known, comprising various zoogeographic elements and populating very different environments. Diplopods mainly occur in various woodlands, in caves, and high in the mountains. Most species (> 90 %, usually highly localised, including 160 cavernicoles), 18 genera, and one family are strictly endemic to continental China. Mapping not only the horizontal, but also the vertical distributions of Diplopoda in China shows the bulk of the fauna to be expectedly restricted to forested lowland and mountain biomes or their remnants. Yet some Chordeumatida, Callipodida, Polydesmida, Julida, and even Spirobolida seem to occur only in the subalpine to alpine environments and thus may provisionally be considered as truly high-montane. The long-acknowledged notions of China being a great biogeographic zone transitional between the Palaearctic and Oriental regions generally find good support in millipede distributions, in particular at the higher taxonomic levels (generic, familial, and ordinal). While the Palaearctic/Holarctic components expectedly dominate the fauna of the northern parts of the country, the Oriental ones prevail in its south and along the Pacific coast. Both realms are increasingly mixed and intermingled towards China’s centre. However, in addition to the above traditional views, based on distribution patterns alone, southern China seems to harbour a rather small, but highly peculiar faunal nucleus or origin centre of its own, whence Himalaya, Myanmar, Thailand, Indochina and/or Taiwan could have become populated by younger lineages. The millipede fauna of continental China is thus a tangled mixture of zoogeographic elements of various origins and ages, both relict and more advanced. The few anthropochores must have been the latest faunal “layer” to populate China.

Keywords

continental China, Diplopod fauna, zoogeography

Introduction

Millipedes (Diplopoda) form a highly diverse, yet strongly understudied arthropod class with > 11,000 described species (Minelli 2015). Apparently, only ca. 20 % of the global species diversity of millipedes are currently known, with the actual number of species being estimated between 50,000 and 80,000 species (Minelli and Golovatch 2013). Being mainly represented by mesophilous forest-dwelling detritivores, millipedes have long been recognised as playing important ecological roles, mostly in temperate and tropical land ecosystems where their diversity is especially pronounced (Golovatch and Kime 2009).

The class encompasses 16 extant orders, 140+ families, and ca. 2,000 genera (Minelli and Golovatch 2013), while the distributions of higher taxa fully agree with the major biogeographic divisions of Earth into the Holarctic (Palaearctic + Nearctic), Afrotropical, Oriental, Neotropical and Australian regions which are accepted since Alfred Russel Wallace and Joseph Dalton Hooker. Antarctica is completely devoid of diplopods, whereas the Oriental Region appears to be the sole one to harbour all 16 orders. Being very ancient (Silurian, early Palaeozoic) and diverse taxonomically, widespread (present on all continents except Antarctica), virtually fully terrestrial (even fossils show spiracles), poorly vagile (with highly limited dispersal capacities) and highly limited in compensatory ecological faculties (strongly restricted by a single limiting ecological factor even if the others are favourable), Diplopoda have long been considered as an exemplary group for biogeographic studies and reconstructions (e.g. Shelley and Golovatch 2011).

China has long been considered as a huge territory lying between and linking the Palaearctic and Oriental realms, with very considerable areas of southern China representing not only a marked transitional zone (e.g. Wulf 1944; Zherikhin 2003; Holt et al. 2012), but also the largest karst belt of the world particularly rich in cavernicoles, including millipedes (Golovatch 2015a). Continental China as conventionally understood here includes Hainan and Hong Kong but excludes Taiwan. The territory in question covers ca. 9,326 million sq. km, spanning ca. 5,500 km from north to south and ca. 5,200 km from west to east. China’s topography is very complex. The outline descends step by step from west to east: mountains, high plateaus and hilly land prevail and take up nearly 70 % of the total area, with deserts also located in the west, but mostly plains, deltas and hills in the east. The climates are likewise varied, ranging from sharply continental in the north, through temperate in the middle, to monsoon subtropical and tropical in the south, with a warm humid influence along the eastern sea coasts (https://en.wikipedia.org/wiki/Geographic_information_systems_in_China).

China with its highly varied climates and relief (ca. 70 % national land area being mountains or plateaus) is exceptionally rich in ecological conditions and it supports as many as 18 natural latitudinal belts or biomes (Ni et al. 2000). They range from Polar desert and Alpine tundra in Tibet, through grasslands (savanna, steppe) or desert in the northern parts, to various woodlands (scrub, boreal forest, temperate forest, tropical forest etc.) (Fig. 1). Nature zonation is generally well-expressed, forested biomes prevailing in total area and forming a succession of boreal forest in the north, through temperate (conifer, deciduous and evergreen), to tropical rainforest in the far south. Altitudinal zonation follows the same general pattern which varies depending on location and grows increasingly complex from seven vegetation or eco-geographic belts in the Tianshan Mountains in the northwest or Tibetan Plateau in the southwest to 14 in Yunnan in the south (review by Zhang et al. 2004).

Figure 1. 

Nature zonation and the main biomes of China (after Ni et al. 2000).

Even though the millipede fauna of China enjoys a very long history of taxonomic study, dating back to 1833 (Wang and Mauriès 1996), it still remains far from well-known. Based on all available information, 339 species from 71 genera, 26 families, and eleven orders of Diplopoda have hitherto been recorded from mainland China (Table 1), but there can be no doubt that our review will soon be out of date.

The present paper is an attempt not only to summarise the Chinese species list (as of the end of 2019), but also to provide an analysis of the distribution patterns revealed, both altitudinal and horizontal, and to hypothesise the main sources, routes and stages of fauno-genesis. A very similar approach has recently been applied to treating the millipedes of the Himalaya (Golovatch and Martens 2018).

Materials and methods

Only described species and published records are considered in our paper, while dubious taxa and those not identified to the species level have been omitted from both checklist and bibliography.

Several broken transects have been chosen to grossly reflect the macro relief of mainland China that accompanies the usual mapped distributions (Figs 215). The maps and their corresponding transects at the bottom show both horizontal and vertical distributions of all or most species in a number of largely speciose genera from different families and of various origins across China. The species on the maps and along transects are arranged from west to east and/or north to south. The generic level has been chosen as the most suitable to be accepted in historical biogeography (Kryzhanovsky 2002). The above novel approach to a graphic presentation of faunistic data allows us to combine the horizontal and vertical distributions of millipedes in the easiest and most vivid way on the same map. Mapping largely concerns endemic species and only the territory of mainland China.

The colour maps were generated using Google Earth Pro version 7.3.2.5495 and Adobe Photoshop CS6. The final images were processed with Adobe Photoshop CS6.

Results

The diplopod fauna of continental China at any higher level is basically a mixture of various zoogeographic elements. At the species level, most diplopods encountered in China are not only endemic to the country, but they are also more or less narrowly localised. This holds especially true for cave-dwellers which are usually presumed troglobionts restricted to a single or few adjacent caves. Generally, as the real diversity of millipedes in China has been estimated to amount to no less than 1,000 species (Golovatch 2015a), the list in Table 1, however impressive, seems to represent only ca. 1/3 of the fauna. It is thereby noteworthy that epigean Diplopoda remain especially badly understudied, since much of the collecting and taxonomic exploration efforts still focus on cavernicoles (Golovatch 2015a).

Table 1.

The millipede fauna of continental China, with data on distributions and basic literature sources.

Taxa Altitude (m a.s.l.) Distribution, province/region (main reference/s)
Order Polyxenida Verhoeff, 1934 Global
Family Polyxenidae Lucas, 1840 Global
Genus Eudigraphis Silvestri, 1948 East Asia
1. Eudigraphis sinensis Ishii & Liang, 1990 ca. 100 Zhejiang, Hangzhou, Lake Xihu (Ishii and Liang 1990)
Genus Polyxenus Latreille, 1802–03 Global
2. Polyxenus hangzoensis Ishii & Liang, 1990 ca. 100 Zhejiang, Hangzhou, Lake Xihu (Ishii and Liang 1990)
Order Glomerida Brandt, 1833 Holarctic and SE Asia
Family Glomeridae Leach, 1816 Holarctic and SE Asia
Genus Hyleoglomeris Verhoeff, 1910 145–2810 Balkans, Anatolia, Caucasus, Central, E and SE Asia
3. Hyleoglomeris albicorporis Zhang & Zhang, 1995 ca. 1660 Yunnan, Baoshan City, Cave Shihua Dong (Zhang and Zhang 1995)
4. H. aschnae Makhan, 2010 ca. 730 Chongqing, Beibei, Mt. Jinyunshan (Makhan 2010b)
5. H. baxian Liu & Tian, 2015 ca. 145 Guangxi, Du’an County, Chengjiang Town, Cave Baxian Park Dong (Liu and Tian 2015a)
6. H. bicolor (Wood, 1865) 210 Hong Kong, Mt. Taimoshan (Golovatch et al. 2006b)
7. H. curtisulcata Golovatch, Liu & Geoffroy, 2012 420 Guangxi, Huanjiang County, Mulun, Cave Gang Lai Dong (Golovatch et al. 2012b)
8. H. emarginata Golovatch, 1981 310 Jiangsu, Nanjing City, Mt. Zijinshan (Golovatch 1981, Golovatch et al. 2006b)
9. H. eusulcata Golovatch, Geoffroy & Mauriès, 2006 ca. 410 Guizhou, Libo County, caves Latai Dong and Shuijiang Dong (Golovatch et al. 2006b)
10. H. generalis Liu & Tian, 2015 550 Guizhou, Cengong County, Shuiwei Town, Cave Jiangjun Dong (Liu and Tian 2015a)
11. H. getuhensis Liu & Tian, 2015 ca. 910 Guizhou, Ziyun County, Getuhe National Geopark, Cave Miaoting Dong (Liu and Tian 2015a)
12. H. grandis Liu & Tian, 2015 ca. 280 Guangxi, Dahua County, Qibainong Geopark, Cave Qiaoxu Dong (Liu and Tian 2015a)
13. H. gudu Golovatch, Liu & Geoffroy, 2012 1365 Guizhou, Anlong County, Cave Hei Dong (Golovatch et al. 2012b)
14. H. heshang Golovatch, Liu & Geoffroy, 2012 ca. 700 Guangxi, Xilin County, Cave Zhoubang Dong (Golovatch et al. 2012b)
15. H. kunnan Golovatch, Liu & Geoffroy, 2012 420 Guangxi, Huanjiang County, Mulun, Cave Ganxiao Dong (Golovatch et al. 2012b)
16. H. lii Golovatch, Liu & Geoffroy, 2012 190 Guangxi, Fuchuan County, Cave Baifu Dong (Golovatch et al. 2012b)
17. H. maculata Golovatch, Geoffroy & Mauriès, 2006 ca. 1315 Yunnan, Mengzi County, Cave Laoshao Dong (Golovatch et al. 2006b)
18. H. mashanorum Golovatch, Liu & Geoffroy, 2012 ca. 210 Guangxi, Huanjiang County, Mulun, Cave Mashan Dong (Golovatch et al. 2012b)
19. H. multistriata Liu & Tian, 2015 ca. 400 Guizhou, Jiangkou County, Nuxi Town, Cave I Dong (Liu and Tian 2015a)
20. H. mulunensis Golovatch, Liu & Geoffroy, 2012 ca. 210 Guangxi, Huanjiang County, Mulun, Cave Xia Dong (Golovatch et al. 2012b)
21. H. nigu Golovatch, Liu & Geoffroy, 2012 ca. 1120 Guizhou, Qianxi County, Cave Luo Sai Dong (Golovatch et al. 2012b)
22. H. qiyi Golovatch, Liu & Geoffroy, 2012 ca. 210 Guangxi, Huanjiang County, Mulun, Cave MinLi Dong (Golovatch et al. 2012b)
23. H. reducta Golovatch, Geoffroy & Mauriès, 2006 ca. 1315 Yunnan, Jianshui County, Cave Yan Dong (Golovatch et al. 2006b)
24. H. rhinoceros Liu & Tian, 2015 ca. 1025 Guizhou, Anlong County, Dushan Town, Cave Xiniu Dong (Liu and Tian 2015a)
25. H. rukouqu Liu & Wynne, 2019 190 Guangxi, Yangshuo County, Cave Shangshuiyan Dong (Liu and Wynne 2019)
26. H. sinensis (Brölemann, 1896) 1540–2810 Sichuan, Kangding County, and Tibet (Golovatch et al. 2006b, Liu and Tian 2015a); New record: Sichuan, W of Ningnan County, 3.3 km WSW of Xiaotiancun village
27. H. tiani Golovatch, Liu & Geoffroy, 2012 ca. 300 Hunan, Linwu County, Huatang Town, Cave Long Dong (Golovatch et al. 2012b)
28. H. variabilis Liu & Tian, 2015 830 Guizhou, Cengong County, Pingle Town, Cave Wanfuchangcheng Dong (Liu and Tian 2015a)
29. H. wuse Golovatch, Liu & Geoffroy, 2012 ca. 425 Guizhou, Maolan County, Cave Dongge Dong (Golovatch et al. 2012b)
30. H. xia Golovatch, Liu & Geoffroy, 2012 ca. 300 Hunan, Linwu County, Sanhe Town, Tianhe Village, Cave 1 Dong (Golovatch et al. 2012b)
31. H. xueju Golovatch, Liu & Geoffroy, 2012 ca. 140 Guangxi, Du’an County, Cave Yaonan Dong (Golovatch et al. 2012b)
32. H. xuxiakei Liu & Wynne, 2019 190 Guangxi, Yangshuo County, Cave Guanshan No. 4 Dong (Liu and Wynne 2019)
33. H. yinshi Golovatch, Liu & Geoffroy, 2012 1205 Guizhou, Kaiyang County, Cave Xianyan Dong (Golovatch et al. 2012b)
34. H. youhao Golovatch, Liu & Geoffroy, 2012 ca. 300 Hunan, Linwu County, Sanhe Town, near Changshali Village, Cave 2 Dong (Golovatch et al. 2012b)
Order Sphaerotheriida Brandt, 1833 S and E Africa, Madagascar, Seychelles, Sri Lanka, S India, Himalayas, E China, SE Asia, Australia, New Zealand
Family Zephroniidae Gray, in Jones, 1843 Seychelles, Himalayas, E China, SE Asia, Sumatra, Java, Borneo, Sulawesi, Philippines
Genus Prionobelum Verhoeff, 1924 10–1500 Vietnam, E China
35. Prionobelum hainani (Gressitt, 1941) 375 Hainan, Tai-Pin-ts’uen (Dwa Bi), foot of Mt. Loi Mother (Mauriès 2001)
36. P. joliveti Mauriès, 2001 145 Hainan, W of Danzhou (Mauriès 2001)
37. P. maculosum (Attems, 1935) 10 Fujian, Fuzhou City (Attems 1935, Mauriès 2001, Wesener 2016)
38. P. majorinum (Zhang & Li, 1982) 1200 Hainan, Mt. Diaoluoshan (Zhang and Li 1982c, Mauriès 2001)
39. P. multidentata (Wang & Zhang, 1993) 1500 Fujian, Jiangle County, Mt. Longqi (Wang and Zhang 1993b, Wesener 2016)
Genus Zephronia Gray, 1832 Himalayas, Myanmar, E China, Indochina, Thailand, Malaysia
40. Zephronia profuga Attems, 1936 ? Hong Kong (Attems 1936, Wesener 2016)
Order Platydesmida Cook, 1895 Mediterranean, Near East, E and SE Asia, Nearctic, Central America
Family Andrognathidae Cope, 1869 E and SE Asia, USA, Mexico
Genus Brachycybe Wood, 1964 E and SE Asia, USA
41. Brachycybe cooki (Loomis, 1942) ca. 1090 Jiangxi, S of Jiujiang, Lushan City, Guling Town (Loomis 1942, Shelley et al. 2005)
Order Polyzoniida Cook, 1895 Global
Family Polyzoniidae Gervais, 1844 Holarctic
Genus Angarozonium Shelley, 1997 N Asia
42. Angarozonium amurense (Gerstfeldt, 1859) 100–1800 Heilongjiang, mouth of Songari River; also Siberia and Mongolia (Mikhaljova 2017)
Order Julida Brandt, 1833 Holarctic, E and SE Asia
Family Julidae Leach, 1814 Holarctic, E and SE Asia
Genus Anaulaciulus Pocock, 1895 10–3350 Himalaya and E Asia
43. Anaulaciulus enghoffi Korsós, 2001 2700 Gansu, Karyn Valley, S wall of Latshi-san Pass (Korsós 2001)
44. A. otigonopus Zhang, 1993 ca. 200 Hunan, Changsha City, Mt. Yuelushan (Zhang 1993a)
45. A. paludicola (Pocock,1895) 10 Zhejiang, 25 mi of Ningo (Ningbo), Lake Wo-Lee (Causey 1966)
46. A. tibetanus Korsós, 2001 2700–3350 Tibet, Dü Chu Valley; Assam, India, 11,000 feet (Korsós 2001)
47. A. tonginus (Karsch, 1881) ? Hong Kong; Taiwan; ? Hunan (Korsós 1994)
48. A. vallicola (Pocock, 1895) ? Zhejiang, 60 mi inland from Sam-moon Bay, Da-zeh Valley (Causey 1966)
Genus Nepalmatoiulus Mauriès, 1983 275–3650 Himalaya, E and SE Asia
49. Nepalmatoiulus brachymeritus Enghoff, 1987 2810 Sichuan, Kangding (Enghoff 1987b)
50. N. eulobos Enghoff, 1987 320 Guangdong, Meizhou City, Mt. Qingliangshan (Enghoff 1987b)
51. N. fraterdraconis Enghoff, 1987 ca. 1045 Jiangxi, Jiujiang City, Mt. Lushan, road to Guling (Enghoff 1987b)
52. N. polyakis Enghoff, 1987 ca. 275 Sichuan, Suining City (Enghoff 1987b)
53. N. rhaphimeritus Enghoff, 1987 2810 Sichuan, Kangding City (Enghoff 1987b)
54. N. tibetanus Enghoff, 1987 2750–3650 SE Tibet, Do-Chu Valley, Pasho Distr., near Rombe Gompa (Enghoff 1987b)
55. N. yunnanensis Enghoff, 1987 ? Yunnan (Enghoff 1987b)
Genus Pacifiiulus Mikhaljova, 1982 Siberia
56. Pacifiiulus amurensis (Gerstfeldt, 1859) 100–2500 Heilongjiang, between mouths of Ussuri and Garyn rivers; also Siberia and the Russian Far East (Mikhaljova 2017)
Family Mongoliulidae Pocock, 1903 E Asia
Genus Skleroprotopus Attems, 1901 125–1190 E Asia
57. Skleroprotopus confucius Attems, 1901 ca. 490 Hebei, Zhangjiakou City (Attems 1901)
58. S. laticoxalis Takakuwa, 1942 395 Liaoning, Shenyang City (Takakuwa 1942)
59. S. membramipedalis Zhang, 1985 ca. 125–150 Beijing, Fangshan, caves Shihua and Yunshui (Zhang 1985a, Vagalinski et al. 2018)
60. S. serratus Takakuwa & Takashima, 1949 ca. 1190 Shanxi, Yantou village (Takakuwa and Takashima 1949)
Family Nemasomatidae Bollman, 1893 Holarctic
Genus Orinisobates Lohmander, 1933 Holarctic E of Ural Mountains
61. Orinisobates gracilis (Verhoeff, 1934) ? Xinjiang, Urumqi, Mt. Tianshan (Verhoeff 1934, Enghoff 1985)
Genus Sinostemmiulus Chamberlin & Wang, 1953 China
62. Sinostemmiulus simplicior Chamberlin & Wang, 1953 ? Zhejiang, Chenghsien (Cheng County?) (Chamberlin and Wang 1953, Hoffman 1966)
Family Parajulidae Bollman, 1893 Nearctic and E Asia
Genus Karteroiulus Attems, 1909 Nearctic and E Asia
63. Karteroiulus niger Attems, 1909 ? Jiangxi, Tai-an-Long (Enghoff 1987a)
Order Spirobolida Cook, 1895 Pantropical
Family Spirobolidae Bollman, 1893 E Asia
Genus Spirobolus Brandt, 1833 305–3350 E Asia
64. Spirobolus bungii Brandt, 1833 ? North of Beijing (Keeton 1960)
65. S. cincinnalis Wang & Zhang, 1993 1500 Fujian, Jiangle County, Mt. Longqi (Wang and Zhang 1993b)
66. S. grahami Keeton, 1960 ca. 305–3350 Sichuan, Suifu; S of Suifu on the Yunnan border; Mupin; near Yueh-Shi, Granham; Mt. Omeishan; Kweichow: Shih Men Kan (Keeton, 1960); Hubei, Jianshi County (Wang and Zhang 1993b)
67. S. umbobrochus Keeton, 1960 ca. 915 Sichuan, Yongshien; Kueichow, Shih Men Kan (Keeton 1960)
68. S. walkeri Pocock, 1895 ca. 150–760 Zhejiang, Chusan Island, “Da-laen-Saen”, 30 mi SW of Ningpo (Keeton 1960)
Order Spirostreptida Brandt, 1833 Pantropical
Family Cambalopsidae Cook, 1895 Himalaya, E and SE Asia, Java, Borneo
Genus Glyphiulus Gervais, 1847 105–4150 E and SE Asia, Java, Borneo
69. Glyphiulus acutus Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2011 ca. 210 Guangxi, Huanjiang County, Mulun, caves Ganglai Dong and Huobayun Dong (Golovatch et al. 2011a)
70. G. adeloglyphus Zhang & Li, 1982 ca. 120 Guangxi, Yangshuo County, Xingping Town (Zhang and Li 1982b)
71. G. anophthalmus (Loksa, 1960) ca. 105 Guangxi (Loksa 1960)
72. G. balazsi (Loksa, 1960) ca. 990 or 835 Guizhou, Luodian County or Longping Town (Loksa 1960)
73. G. basalis Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 ca. 4150 Sichuan, Xinlong County, Cave Ganchuan Dong (Golovatch et al. 2007a)
74. G. beroni Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 ca. 1315 Yunnan, Jianshui County, Cave Baguo Dong; and Tonghai County, Cave Xianren Dong (Golovatch et al. 2007a)
75. G. calceus Jiang, Guo, Chen & Xie, 2018 900 Guangxi, Tian’e County, Bala Town, Madong village, Hanyaotun, Cave Xianren Dong (Jiang et al. 2018)
76. G. deharvengi Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 730 Hunan, Longshan County, Huoyan Village, Cave Feihu Dong, Cave Baiyan Dong, Cave Remi Dong (Golovatch et al. 2007a)
77. G. difficilis Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2011 ca. 925 Guangxi, Leye County, Yachang Town, Huaping, Cave She Dong and Cave Xiayan Dong (Golovatch et al. 2011b)
78. G. echinoides Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2011 ca. 270 Guangxi, Fushui County, Bapen village, Cave II Dong (Golovatch et al. 2011b)
79. G. foetidus Jiang, Guo, Chen & Xie, 2018 690–820 Guangxi, Xilin County, Zhoubang village, Cave Zhoubang Dong; Yunnan, Guangnan County, Bamei Town, Ake village, Cave Miaopu Dong (Jiang et al. 2018)
80. G. formosus (Pocock, 1895) ca. 135 Hong Kong (Pocock 1895)
81. G. granulatus Gervais, 1847 135–440 Pantropical; Guangxi, Longzhou; Hong Kong; Taiwan (Golovatch et al. 2007a)
82. G. guangnanensis Jiang, Guo, Chen & Xie, 2018 690 Yunnan, Guangnan County, Bamei Town, Ake village, Cave Miaopu Dong (Jiang et al. 2018)
83. G. impletus Jiang, Guo, Chen & Xie, 2018 320–830 Guangxi, Lingyun County, сaves (Jiang et al. 2018)
84. G. intermedius Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 ca. 485 Sichuan, Chengdu, Cave Huanlong Dong (Golovatch et al. 2007b)
85. G. latellai Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 ca. 1495 Guizhou, Qianxi County, Honglin village, Cave Hangtu Dong, Cave Xiao Dong, Cave Xixiang Dong, Cave Dayan Dong, Cave Tiaoshui Dong, Cave Ludiaoai Dong, Cave Shuhuayan Dong, Cave Shuiluo Dong (Golovatch et al. 2007a)
86. G. latus Jiang, Lv, Guo & Chen, 2017 ca. 410 Sichuan, Leshan City, Muchuan County, Cave Longgong Dong (Jiang et al. 2017)
87. G. liangshanensis Jiang, Lv, Guo & Chen, 2017 ca. 470–1155 Sichuan, Liangshan Yi Autonomous Prefecture, Xichang City, Xixi, Xianren Cave; Miyi County, Baima Town, Cave Zhuanxulong Dong (Jiang et al. 2017)
88. G. lipsorum Mauriès & Nguyen Duy-Jacquemin, 1997 ca. 430 Hubei, cave (Mauriès and Nguyen Duy-Jacquemin 1997)
89. G. maocun Liu & Wynne, 2019 180 Guangxi, Lingchuan County, Maocun Village, Cave Liangfeng Dong (Liu and Wynne 2019)
90. G. melanoporus Mauriès & Nguyen Duy-Jacquemin, 1997 ca. 180 Guangxi, near Guilin, cave (Mauriès and Nguyen Duy-Jacquemin 1997); Xiufeng District, Cave Maomaotou (Liu and Wynne 2019)
91. G. mulunensis Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2011 ca. 270 Guangxi, Huanjiang County, Mulun, caves Mashan Dong and Ganglai II Dong (Golovatch et al. 2011a)
92. G. obliteratoides Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 1400 Guizhou, Anshun City, Liangshuijing, Cave Tianxian Dong (Golovatch et al. 2007b)
93. G. obliteratus Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 ca. 1315 Yunnan, Mile County, caves Bailong Dong and Houshan Dong (Golovatch et al. 2007b)
94. G. paracostulifer Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 ca. 1495 Guizhou, Qianxi County, Honglin Town, Cave Laohu Dong (Golovatch et al. 2007b)
95. G. paragranulatus Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 ca. 1315 Yunnan, Jianshui County, Cave Yan Dong (Golovatch et al. 2007a)
96. G. paramulunensis Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2011 211 Guangxi, Huanjiang County, Mulun, caves Shui Dong and Xialan Dong (Golovatch et al. 2011a)
97. G. parobliteratus Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 ca. 725–860 Guizhou, Suiyang County, Wenquan Town, Shuanghe, Cave Dafeng Dong (Golovatch et al. 2007b)
98. G. pergranulatus Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 ca. 1065 Guizhou, Guanling County, Huajiang, Cave Da Dong and Cave Anjiada Dong (Golovatch et al. 2007a)
99. G. proximus Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2011 ca. 210 Guangxi, Huanjiang County, Mulun, caves Ganxiao Dong and Dongtu Dong (Golovatch et al. 2011a)
100. G. pulcher Loksa, 1960 ca. 105 Guangxi, Daxin County, Fulong Town, a cave (Loksa 1960, Jiang et al. 2018)
101. G. quadrohamatus Chen & Meng, 1991 ca. 1110 Guizhou, Zhenning County, several caves (Chen and Meng 1991)
102. G. rayrouchi Mauriès & Nguyen Duy-Jacquemin, 1997 ca. 390 Guizhou, Maguan, Cave Heiyan Dong (Mauriès and Nguyen Duy-Jacquemin 1997)
103. G. recticullus Zhang & Li, 1982 ca. 325 Zhejiang, Qingyuan County (Zhang and Li 1982b)
104. G. semigranulatus Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 ca. 1315 Yunnan, Mile County, Cave Bailong Dong; Jianshui County, Cave Yanzi Dong (Golovatch et al. 2007a)
105. G. septentrionalis Murakami, 1975 ca. 170 Guangxi, Guilin; Japan, Ryukyus, Okinawa Island (Golovatch et al. 2007a)
106. G. sinensis (Meng & Zhang, 1993) ca. 1065 Guizhou, Guanling County, cave (Meng and Zhang 1993)
107. G. speobius Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2011 ca. 310 Guangxi, Huanjiang County, caves Xialan Dong and Shenlong Dong (Golovatch et al. 2011a)
108. G. subgranulatus Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 1313 Yunnan, Mengzi County, cave near footpath to plateau, Pothole No. 2 (Golovatch et al. 2007a)
109. G. subobliteratus Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2007 ca. 1685 Yunnan, Shilin County, Cave Zhiyun Dong (Golovatch et al. 2007b)
110. G. tiani Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2011 ca. 210 Guangxi, Huanjiang County, Mulun, Cave Dongzai Dong (Golovatch et al. 2011a)
111. G. zorzini Mauriès & Nguyen Duy-Jacquemin, 1997 ca. 1105 Guizhou, Shuicheng County, Cave Anjia Yan (Mauriès and Nguyen Duy-Jacquemin 1997)
Genus Hypocambala Silvestri, 1897 SE Asia
112. Hypocambala polytricha Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2011 ca. 110 Guangxi, Longzhou County, Nonggang, Cave Biji Dong (Golovatch et al. 2011c)
Family Pericambalidae Silvestri, 1909 China, Indochina
Genus Bilingulus Zhang & Li, 1981 China, Vietnam
113. Bilingulus sinicus Zhang & Li, 1981 165 Guangxi, Guilin City, a cave (Zhang and Li 1981a); Yangshuo County, Cave Shangshuiyan; Xiufeng District, Cave Maomaotou; Lingchuan County, Cave Liangfeng Dong (Liu and Wynne 2019)
Genus Parabilingulus Zhang & Li, 1981 105–120 China
114. Parabilingulus aramulus Zhang & Li, 1981 ca. 120 Guangxi, Yangshuo County, Xingping Town (Zhang and Li 1981a)
115. P. simplicius Mauriès & Jacquemin-Nguyen Duy, 1997 ca. 105 Guangxi, Gongcheng County, Cave Heiyan Dong (Mauriès and Nguyen Duy-Jacquemin 1997)
Family Harpagophoridae Attems, 1909 Afrotropical, Himalaya, Sri Lanka, S India, E and SE Asia, Sunda Archipelago
Genus Agaricogonopus Zhang & Zhang, 1997 China
116. Agaricogonopus acrotrifoliolatus Zhang & Zhang, 1997 ca. 870 Yunnan, Xishuangbanna, Mengla County, tropical rainforest (Zhang and Zhang 1997; Pimvichai et al. 2010)
Genus Junceustreptus Demange, 1961 650–1895 China
117. Junceustreptus brevispinus Zhang, 1985 ca. 650 Yunnan, Xishuangbanna, Mengman (Zhang 1985b; Pimvichai et al. 2010)
118. J. browningi Demange, 1962 ca. 1895 Yunnan (Demange 1962; Pimvichai et al. 2010)
119. J. retrorsus Hoffman, 1980 ca. 1890 Sichuan, Ning Gyuen Nfu (Hoffman 1980; Pimvichai et al. 2010)
Genus Prominulostreptus Pimvichai, Enghoff & Panha, 2010 ? China
120. Prominulostreptus prominulus (Demange, 1962) ? Yunnan, Lou-Fou-Tsouen (Ing-Ka-Tsoue) (Demange 1962; Pimvichai et al. 2010)
Genus Uriunceustreptus Zhang & Chang, 1990 650–1750 China, Vietnam
121. Uriunceustreptus afemorispinus Zhang & Chang, 1990 ca. 1750 Yunnan, Gejiu City (Zhang and Chang 1990; Pimvichai et al. 2010)
122. U. bilamellatus Zhang, 1997 ca. 650 Sichuan (now Chongqing), Youyang County (Zhang et al. 1997)
Order Chordeumatida Pocock, 1894 Mostly Holarctic, but also Central and SW South America, Madagascar, Sri Lanka, S India, E and SE Asia, Sunda Archipelago, Philippines, New Guinea, Australia, New Zealand
Family Guizhousomatidae Mauriès, 2005 China
Genus Guizhousoma Mauriès, 2005 ca. 1495 China
123. Guizhousoma latellai Mauriès, 2005 ca. 1495 Guizhou, Qianxi County, Honglin, caves Changtu Dong, Shujiayan, Luosai Dong, Shuiluo Dong, Tiaoshui Dong; and Dafang County, Cave Hei Dong (Mauriès 2005)
Family Kashmireumatidae Mauriès, 1982 Himalaya, E and SE Asia
Genus Lipseuma Golovatch, Geoffroy & Mauriès, 2006 435–1405 China
124. Lipseuma bernardi Golovatch, Geoffroy & Mauriès, 2006 ca. 435 Sichuan, Xinlong County, Three Eyes Cave (Golovatch et al. 2006a, 2007)
125. L. josianae Golovatch Geoffroy & Mauriès, 2006 1405 Hubei, Banqiao Town, Cave ChuanDongZi (Golovatch et al. 2006a)
Genus Vieteuma Golovatch, 1984 2100–2300 China, Vietnam
126. Vieteuma hubeiense Mauriès & Nguyen Duy-Jacquemin, 1997 ca. 2130 Hubei, Shennongjia, Yanziya, Cave Yanzi Dong (Mauriès and Nguyen Duy-Jacquemin 1997)
127. V. longi Shear, 2002 2100–2300 Yunnan, Baoshan City, Mt. Gaoligongshan, Nankang, 36 air km SE of Tengchong; and LuoshuiDong, 28 air km SE of Teng Chong (Shear 2002)
Family Megalotylidae Golovatch, 1978 Himalaya, Myanmar, E and SE Asia
Genus Nepalella Shear, 1979 750–4530 Himalaya, Myanmar, E and SE Asia
128. Nepalella caeca Shear, 1999 1795 Guizhou, Shuicheng County, Cave Anjia Yan (Shear 1999, Liu, Wesener et al. 2017c)
129. N. grandis Golovatch, Geoffroy & Mauriès, 2006 ca. 1670 Yunnan, Zhenxiong County, Cave Baiyin Dong (Golovatch et al. 2006a)
130. N. grandoides Golovatch, Geoffroy & Mauriès, 2006 ca. 750 Sichuan, Beichuan County, caves Yuan Dong and Black Wind Dong (Golovatch et al. 2006a, Liu, Wesener et al. 2017d)
131. N. griswoldi Shear, 2002 2100–2300 Yunnan, Baoshan City, Mt. Gaoligongshan, Luoshuidong, 28 air km of Tengcheng (Shear 2002)
132. N. jinfoshan Liu, in Liu, Wesener et al., 2017 2017 1500–2100 Chongqing, Jinfoshan, Cave Houshan Dong; Cave Lingguan Dong (Liu, Wesener et al. 2017d)
133. N. kavanaughi Shear, 2002 2500 Yunnan, Nujiang, Pianma, native forest on Mt. Gaoligongshan (Shear 2002)
134. N. lobata Liu, in Liu, Wesener et al., 2017 1000 Sichuan, Mianyang City, Beichuan County, Cave Liangshui Dong (Liu, Wesener et al. 2017d)
135. N. magna Shear, 2002 2300 Yunnan, Baoshan City, Mt. Gaoligongshan, Luoshuidong, 28 air-km of Tengchong (Shear 2002)
136. N. marmorata Golovatch, Geoffroy & Mauriès, 2006 ca. 4350 Sichuan, Xinlong County, caves Snake Mouth Dong and Three Eyes Dong (Golovatch et al. 2006a, 2007)
137. N. pianma Shear, 2002 2500 Yunnan, Nujiang, Pianma, Mt. Gaoligongshan, native forest (Shear 2002)
138. N. troglodytes Liu, in Liu, Wesener et al., 2017 1200–1300 Guizhou, Guiyang City, Xifeng County, Hejiadong village, Cave Hejia Dong; same County, Mushan village, Cave Zhangkou Dong; Guizhou, Qiannan, Longli County, Cave Feilong Dong; Guizhou, Qiannan, Fuquan County, Cave Sanlou Dong (Liu, Wesener et al. 2017d)
139. N. wangi Liu, in Liu, Wesener et al., 2017 1300 Chongqing, Wulong County, Huangying Town, Qimenxia, Cave I Dong (Liu, Wesener et al. 2017d)
Order Callipodida Pocock, 1894 Holarctic, E and SE Asia
Family Caspiopetalidae Lohmander, 1931 Central Asia and China
Genus Bollmania Silvestri, 1896 Central Asia and China
140. Bollmania beroni Stoev & Enghoff, 2005 ca. 1315 Yunnan, Jianshui County, Cave Yan Dong (Stoev and Enghoff 2005)
Family Paracortinidae Wang & Zhang, 1993 China and Vietnam
Genus Angulifemur Zhang, 1997 1315 China
141. Angulifemur tridigitis Zhang, 1997 ca. 1315 Yunnan, Mengzi City, Cave Niupeng-yanzi Dong (Zhang 1997)
142. A. unidigitis Zhang, 1997 ca. 1315 Yunnan, Mengzi City, caves Longbaopo Dong and Laoxiao Dong (Zhang 1997)
Genus Paracortina Wang & Zhang, 1993 865–3300 China and N Vietnam
143. Paracortina carinata Wang & Zhang, 1993 3300 Yunnan, Shangrila (= Zhongdian) County (Wang and Zhang 1993a)
144. P. chinensis Stoev & Geoffroy, 2004 ca. 1670 Yunnan, Zhenxiong County, caves Ke Ma Dong, Da Hei Dong and Liao Jun Dong (Stoev and Geoffroy 2004)
145. P. leptoclada Wang & Zhang, 1993 3300 Yunnan, Shangrila (= Zhongdian) County (Wang and Zhang 1993a)
146. P. serrata Wang & Zhang, 1993 ca. 1845 Yunnan, Deqin County (Wang and Zhang 1993a)
147. P. stimula Wang & Zhang, 1993 3300 Yunnan, Shangrila (= Zhongdian) County (Wang and Zhang 1993a)
148. P. thallina Wang & Zhang, 1993 3300 Yunnan, Shangrila (= Zhongdian) County; Sichuan, Batang County (Wang and Zhang 1993a)
149. P. viriosa Wang & Zhang, 1993 3300 Yunnan, Shangrila (= Zhongdian) County (Wang and Zhang 1993a); Tibet, Mangkang County (Stoev et al. 2008)
150. P. voluta Wang & Zhang, 1993 ca. 2690 Sichuan, Yajiang County (Wang and Zhang 1993a)
151. P. yinae Liu & Tian, 2015 ca. 865 Guangxi, Baise City, Longlin County, Tianshengqiao Town, Yanchang village, Cave I (Liu and Tian 2015c)
152. P. zhangi Liu & Tian, 2015 ca. 965 Guizhou, Qianxinan Autonomous Prefecture, Ceheng County, Rongdu village, Cave Qiaoxia Dong (Liu and Tian 2015c)
Family Sinocallipodidae Zhang, 1993 China and Indochina
Genus Sinocallipus Zhang, 1993 1860 China, Laos and Vietnam
153. Sinocallipus simplopodicus Zhang, 1993 1860 Yunnan, Hehou City, Cave Xiao Dong (Zhang 1993b)
Order Polydesmida Pocock, 1887 Global
Family Cryptodesmidae Karsch, 1880 Pantropical
Genus Trichopeltis Pocock, 1894 165–1890 Himalaya, E and SE Asia, Malaysia, Sunda Archipelago
154. Trichopeltis bellus Liu, Golovatch & Tian, 2017 1530 Yunnan, Qujing City, Luoping County, Machang village, Cave Shuiyuan Dong (Liu et al. 2017a)
155. T. intricatus Liu, Golovatch & Tian, 2017 1890 Yunnan, Kunming City, Shilin County, Guishan Town, Cave Haiyi I Dong (Liu et al. 2017a)
156. T. latellai Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2010 ca. 1495 Guizhou, Qianxi County, Honglin Town, caves Tiaoshui Dong and Changtu Dong (Golovatch et al. 2010c)
157. T. liangfengdong Liu & Wynne, 2019 180 Guangxi, Lingchuan County, Cave Liangfeng Dong (Liu and Wynne 2019)
158. T. reflexus Liu, Golovatch & Tian, 2017 ca. 165 Hunan, Chenzhou City, Linwu County, Xianghualing Town, II Dong Cave (Liu et al. 2017a)
Family Haplodesmidae Cook, 1895 Himalaya, Myanmar, E and SE Asia, Malaysia, Sunda Archipelago, New Guinea, Melanesia, Australia
Genus Doratodesmus Cook, 1895 Sunda Archipelago, China
159. Doratodesmus grandifoliatus Zhang, in Zhang & Wang, 1993 ca. 1315 Yunnan, Mengzi County, Cave Longbaopo Dong (Zhang and Wang 1993)
Genus Eutrichodesmus Silvestri, 1910 65–1495 E and SE Asia, Sunda Archipelago, Melanesia
160. Eutrichodesmus anisodentus (Zhang, 1995) ca. 385 Fujian, Mt. Wuyishan (Zhang 1995b)
161. E. apicalis Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2015 ca. 75 Hubei, Yichang, Yichang County, Grotte des Araignées (Golovatch et al. 2015)
162. E. arcicollaris Zhang, in Zhang & Wang, 1993 ca. 170 Yunnan, Hekou County, Cave Huayu Dong (Zhang and Wang 1993, Golovatch et al. 2009a, 2009b)
163. E. digitatus Liu & Tian, 2013 ca. 65 Guangdong, Qingyuan City, Jintan Town, Cave Mi Dong (Liu and Tian 2013)
164. E. distinctus Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2009 ca. 105 Guangxi, Fusui County, Bapen, Cave 4 Dong (Golovatch et al. 2009b)
165. E. dorsiangulatus (Zhang, in Zhang & Wang, 1993) ca. 635 Yunnan, Mengla County, Cave Baoniujiao Dong (Zhang and Wang 1993, Golovatch et al. 2009a, 2009b)
166. E. incisus Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2009 ca. 1495 Guizhou, Qianxi County, Honglin, caves Tiaoshui Dong, Cave Liaojingling Dong, Jiayan Dong, Dakong Dong and Luosai Dong (Golovatch et al. 2009a)
167. E. jianjia Liu & Wynne, 2019 190 Guangxi, Yangshuo County, Cave Guanshan No. 4 (Liu and Wynne 2019)
168. E. latellai Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2015 1060 Guizhou, Zhenfeng County, Beipanjiang Town, Cave Shui Chi Dong (Water Pool Cave) (Golovatch et al. 2015)
169. E. latus Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2009 ca. 560 Guangxi, Leye County, Yachang Nature Reserve, caves Yanwu Dong, Xiayan Dong, Xiaoshui Dong and She Dong (Golovatch et al. 2009a)
170. E. lipsae Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2015 ca. 160 Guangxi, Guilin, Grotte des Squelettes (Golovatch et al. 2015)
171. E. monodentus (Zhang, in Zhang & Wang, 1993) ca. 650 Yunnan, Mengla County, Cave Caiyun Dong (Zhang and Wang 1993, Golovatch et al. 2009a, 2009b)
172. E. obliteratus Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2015 ca. 1065 Guizhou, Guanling County, Huajiang Town, Cave Huashiban Dong (Slippery Cave) (Golovatch et al. 2015)
173. E. pectinatidentis (Zhang, 1995) ca. 1010 Zhejiang, Lin’an County, Mt. Tianmu (Zhang 1995a)
174. E. planatus Liu & Tian, 2013 ca. 550 Guangxi, Hechi City, Liujia Town, Cave Zhenzhuyan Dong (Liu and Tian 2013)
175. E. sketi Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2015 730 Hunan, Longshan County, Huoyan, Cave Feihu Dong (Golovatch et al. 2015)
176. E. similis Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2009 ca. 310–420 Guangxi, Huanjiang County, Mulun Nature Reserve, caves Gui II Dong and Shenlong Dong (Golovatch et al. 2009a)
177. E. simplex Liu & Tian, 2013 130 Jiangxi, Fenyi County, Cave Taoyuan Dong (Liu and Tian 2013)
178. E. soesilae Makhan, 2010 ca. 735 Chongqing, Beibei, Mt. Jinyunshan (Makhan 2010a)
179. E. spinatus Liu & Tian, 2013 ca. 875 Hunan, Guidong County, Sidu Town, Sidu Caves (Liu and Tian 2013)
180. E. tenuis Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2015 ca. 1065 Guizhou, Guanling County, Yongning Town, Cave Yun Dong (Cloud Cave) (Golovatch et al. 2015)
181. E. triangularis Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2015 ca. 750 Sichuan, Beichuan County, Cave Yan Dong (Golovatch et al. 2015)
182. E. troglobius Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2015 1205 Guizhou, Kaiyang County, Cave Xianyan Dong (Golovatch et al. 2015)
183. E. trontelji Golovatch, Geoffroy, Mauriès & VandenSpiegel, 2015 ca. 410 Guizhou, Libo County, caves Shui Jiang Dong, Shuipu Da Dong, Shuipa, Latai Dong, Jia Ban and Feng Dong (Golovatch et al. 2015)
Family Paradoxosomatidae Daday, 1889 Global except for N America
Genus Anoplodesmus Pocock, 1895 S, E and SE Asia
184. Anoplodesmus chinenis Golovatch, 2013 1700–2400 Shaanxi, Mt. Taibaishan, southern slopes, above Houshenzi, primary broadleaved forest (Golovatch 2013a)
Genus Antheromorpha Jeekel, 1968 E and SE Asia
185. Antheromorpha rosea Golovatch, 2013 1200–1700 Yunnan, S of Pianma; Baoshan District, near Hemu, Mt. Gaoligongshan, near Cave Bianfu II Dong (Golovatch 2013a, 2013b); also N Thailand and Laos (Likhitrakarn et al. 2019)
Genus Belousoviella Golovatch, 2012 China
186. Belousoviella kabaki Golovatch, 2012 3360 Sichuan, SW of Mianning, right tributary of Yalongjiang River canyon (Golovatch 2012)
Genus Cawjeekelia Golovatch, 1980 100–2110 E and SE Asia
187. Cawjeekelia nova Golovatch, 2011 2110 Chongqing, Dabashan Mt. Range, NE of Heyu, Betula forest (Golovatch 2011)
188. C. pallida Golovatch, 1996 100–200 Hong Kong, Tai Po Kau Nature Reserve (Golovatch 1996)
189. C. propria (Mikhaljova & Korsós, 2003) 500 Jilin, Mt. Changbaishan National Park; also N Korea (Mikhaljova and Korsós 2003, Golovatch 2013a)
Genus Desmoxytes Chamberlin, 1923 E and SE Asia
190. Desmoxytes planata (Pocock, 1895) 560 Nearly pantropical; Yunnan, Xishuangbanna, Menglun, Tropical Botanical Garden (Srisonchai et al. 2018; Golovatch 2018)
Genus Enghoffosoma Golovatch, 1993 E and SE Asia
191. Enghoffosoma longipes Golovatch, 2011 3150 Yunnan, NW slope of Mt. Yulongxueshan (Golovatch 2011)
Genus Gonobelus Attems, 1936 995–2615 China
192. Gonobelus belousovi Golovatch, 2014 995 Sichuan, NE of Shimian, Xiangshuigou River, Tianpingzi (Golovatch 2014a)
193. G. martensi Golovatch, 2013 1700–2600 Shaanxi, Mt. Taibaishan (Golovatch 2013a)
194. G. pentaspinus Golovatch, 2013 2475 Sichuan, NW of Mianning (Golovatch 2013b)
195. G. sinensis Attems, 1936 2615 Yunnan, Mt. Laojunshan, 3.7 km ENE of Segengsheng (Golovatch 2017)
Genus Hedinomorpha Verhoeff, 1934 1300–4490 Central Asia and China
196. Hedinomorpha affinis Golovatch, 2014 2870 Gansu, Mt. Lianhuashan (Golovatch 2014a)
197. H. altiterga Golovatch, 2019 1445 Gansu, WWS of Longnan (Wudu), 2.4 km NW of Zhongzhaixiang (Golovatch 2019a)
198. H. bifida Golovatch, 2019 3665 Sichuan, 7.3 km S of Ganzi (Golovatch 2019a)
199. H. biramipedicula Zhang & Tang, 1985 ca. 1360 Shaanxi, Qinling, Mt. Taibaishan (Zhang and Tang 1985)
200. H. circofera Golovatch, 2013 ca. 2735 Qinghai, Beishan National Park, 120 km N of Xining (Golovatch 2013a)
201. H. circularis (Takakuwa & Takashima, 1949) ? Shanxi, Chinkaiji (Takakuwa and Takashima 1949, Golovatch 2019a)
202. H. crassiterga Golovatch, 2019 4490 Sichuan, 16.8 km SSW Ganzi (Golovatch 2019a)
203. H. flavobulbus Golovatch, 2019 3650 Gansu, WWS of Longnan (Wudu), Yin Duoguosa & Aounang divide (Golovatch 2019a, 2019b)
204. H. hummelii Verhoeff, 1934 ? Gansu, Tan-Chang (Verhoeff 1934)
205. H. jeekeli (Golovatch, 2009) 1300–2600 Shaanxi, Foping Nature Reserve, Panda area (Golovatch 2009); Shaanxi, Mt. Taibaishan, S slopes, above Houshenzi, primary and secondary broadleaved forests (Golovatch 2013a)
206. H. martensi Golovatch, 2014 3510 Sichuan, Langmusi, remnants of a moist Abies forest above town (Golovatch 2014a)
207. H. montana Golovatch, 2016 3080–3695 Yunnan, NNE of Weixi City, 8.15 km ESE of Shajiama; N of Weixi City, 2.95 km NW of Xugongqingshang Village; NW of Jianchuan, 4.7 km WNW of Damaidi; Mt. Laojunshan, NE of Liming, 4.2 km S of Muzhengdu (Golovatch 2016b, 2017)
208. H. nigra Golovatch, 2013 3530–4000 Sichuan, Jiuzhaigou County, N of Dajisi (Golovatch 2013b)
209. H. proxima Golovatch, 2016 3570 Yunnan, Mt. Tianbaoshan between Shangrila and Mt. Habaxueshan, E slope, NW of Bengla (Golovatch 2016b)
210. H. reducta Golovatch, 2012 2900 Sichuan, SW of Mianning, Right tributary of Yalongjiang River canyon, ca. 9 km SW of Mofanggou (Golovatch 2012)
211. H. subnigra Golovatch, 2013 3910 Yunnan, W of Lake Lugu (Golovatch 2013b)
212. H. yunnanensis Golovatch, 2016 3480 Yunnan, NNE of Weixi City, right tributary of Lapugon River, 5.2 km ENE of Jizong (Golovatch 2016b)
Genus Helicorthomorpha Attems, 1914 E and SE Asia
213. Helicorthomorpha holstii (Pocock, 1895) 340 Widespread in SE Asia; Yunnan; Guangdong, Dinghushan Mt., 86 km W of Guangzhou (Attems 1936, Golovatch 1981)
Genus Hirtodrepanum Golovatch, 1994 Himalaya and China
214. Hirtodrepanum chinense Golovatch, 2014 1990–2015 Yunnan, Deqin, Dewei Line, E of Aqiku; Mekong Valley, 2 km E of Yezhixiang (Golovatch 2014a, 2019a)
Genus Hylomus Cook & Loomis, 1924 ca. 140–910 E and SE Asia
215. Hylomus cornutus (Zhang & Li, 1982) ca. 140 Guangxi, Guilin, Yangshuo (Zhang and Li 1982a)
216. H. draco Cook & Loomis, 1924 ca. 400 Jiangxi, Jiujiang City, Mt. Lushan (Cook and Loomis 1924, Srisonchai et al. 2018)
217. H. eupterygotus (Golovatch, Li, Liu & Geoffroy, 2012) ca. 260 Hunan, Linwu County, Tianhe, Cave I Dong and Changshali Cave I Dong (Golovatch et al. 2012a)
218. H. getuhensis (Liu, Golovatch & Tian, 2014) ca. 910 Guizhou, Ziyun County, Getuhe National Geopark, caves Suidao Dong and Taiyang Dong (Liu et al. 2014)
219. H. laticollis (Liu, Golovatch & Tian, 2016) 450 Guangdong, Yingde City, Huanghua Town, Yanbei village, Cave Yangyan Dong (Liu et al. 2016)
220. H. lingulatus (Liu, Golovatch & Tian, 2014) ca. 140 Guangxi, Guilin, Pingle County, Cave Chaotianyan (Liu et al. 2014)
221. H. longispinus (Loksa, 1960) ? Guangxi, a cave (no exact locality known) (Loksa 1960)
222. H. lui (Golovatch, Li, Liu & Geoffroy, 2012) ca. 155 Guangxi, Yongfu County, Shangxiao, Cave Dachong Dong (Golovatch et al. 2012a)
223. H. minutuberculus (Zhang, 1986) ca. 295 Guangxi, Tianlin County (Zhang 1986)
224. H. nodulosus (Liu, Golovatch & Tian, 2014) ca. 350 Guangxi, Du’an County, Xia’ao Town, near Xia’ao Middle School, Cave II Dong; same county, Yong’an Town, Yong’an village, Cave I Dong; same town, Anju Village, Cave Suidao Dong; same county, Longwan Town, Qunle village, entrance to Cave I Dong (Liu et al. 2014)
225. H. parvulus (Liu, Golovatch & Tian, 2014) ca. 350 Guangxi, Du’an, Xia’ao (Liu et al. 2014)
226. H. phasmoides (Liu, Golovatch & Tian, 2016) ca. 445 Guangxi, Lingyun County, Jiayou Town, Yangli village, Cave Fengliu Dong (Liu et al. 2016)
227. H. scolopendroides (Golovatch, Geoffroy & Mauriès, 2010) ca. 210–350 Guangxi, Huanjiang County, Dacai Town, Cave Shenlong Dong; Du’an County, Gaoling Town, Jinzhu village, Cave I Dong, Cave II Dong; Xia’ao Town, Cave I Dong (Golovatch et al. 2010b, Liu et al. 2014)
228. H. scutigeroides (Golovatch, Geoffroy & Mauriès, 2010) ca. 310 Guangxi, Huanjiang County, Cave Ganglai Dong, Cave Mashan II Dong, Cave Gonglu Dong, Cave Shui Dong, and Du’an County, Disu Town, Dading village, Cave II Dong, same county, Longwan Town, Nongqu village, Cave I Dong, (Golovatch et al. 2010b, Liu et al. 2014)
229. H. similis (Liu, Golovatch & Tian, 2016) 230 Guangdong, Yingde City, Qingkeng Town, Bangjiao village, Cave Bangjiao Dong (Liu et al. 2016)
230. H. simplipodus (Liu, Golovatch & Tian, 2016) 140 Guangdong, Qingyuan City, Yangshan County, Chengjia Town, Dabei Village, Cave Kuangzhanyan (Liu et al. 2016)
231. H. spinissimus (Golovatch, Li, Liu & Geoffroy, 2012) 190 Guangxi, Fuchuan County, Guanyuan, Cave Guanyuan Dong (Golovatch et al. 2012a)
232. H. spinitergus (Liu, Golovatch & Tian, 2016) ca. 210 Guangxi, Huanjiang County, near Cave Gui DongII, secondary forest (Liu et al. 2016)
233. H. variabilis (Liu, Golovatch & Tian, 2016) 500 Guangxi, Fengshan County, numerous caves (Liu et al. 2016)
234. H. yuani Liu & Wynne, 2019 180 Guangxi, Lingchuan County, Cave Liangfeng (Liu and Wynne 2019)
Genus Inversispina Zhang, 1997 510–4150 China and Taiwan
235. Inversispina erectispina Golovatch, 2012 2400–4150 Sichuan, SW of right tributary of Yalongjiang River, canyon; Sichuan, NW of Mianning, broadleaved forest; Sichuan, Jiulong County, SW of Wulaxixiang, broadleaved forest; Yunnan, between Tianbaoshan and Luzilashan, between Shuimofang and Xipazi; Yunnan, N of Lijiang, NW of Baoshanxiang, W of Bengluo village (Golovatch 2012, 2013b, 2016a, 2016b)
236. I. multispina Golovatch, 2016 2360 Sichuan, SSE of Shimian, S of Zhuma (Golovatch 2016a)
237. I. tortiapicalis Zhang, 1997 510 Hubei, Hefeng Tu jiazu County, Yien (Zhang et al. 1997)
238. I. trispina Golovatch, 2013 1050 Sichuan, Mt. Emeishan, Wannian Monastery (Golovatch 2013a)
Genus Kronopolites Attems, 1914 35–3600 Himalaya, E and SE Asia
239. Kronopolites biagrilectus Hoffman, 1963 35–3600 Jiangxi, 10 mi S of Jiujiang (oHoffman 1963); Sichuan, SSE of Shimian, S of Zhuma; Yunnan, Mt. Laojunshan,NE Liming, 2.5 km SE of Yankuluo; N of Lanping, 10.3 km SW of Hexi; N of Lanping, 11.3 km SW of Hexi; Yunnan, SE of Deqen City, 3.3 km S of Gejiancun; Yunnan, Mt. Laojunshan, NE Liming, 2.5 km SE of Yankuluo; N of Lanping, 10.3 km SW of Hexi; N of Lanping, 11.3 km SW of Hexi (Golovatch 2016a, 2016b, 2017)
240. K. davidiani Golovatch, 2014 3365 Sichuan, Wenchuan City, 214 National Road, WSW of Edi (Golovatch 2014a)
241. K. swinhoei (Pocock, 1895) 1300–1700 Shaanxi, Mt. Taibaishan (oHoffman 1963); Shaanxi, Panda area, Foping Nature Reserve; Gansu, WWS of Longnan (Wudu), 2.4 km NW of Zhongzhaixiang (Golovatch 2017, 2019a)
Genus Mandarinopus Verhoeff, 1934 700–2955 China
242. Mandarinopus corticinus (Attems, 1936) ? Yunnan (Attems 1936, Golovatch 2019a)
243. M. gracilipes Verhoeff, 1934 700–2195 Gansu, Baishui Jiang River; WWS of Longnan (Wudu), 3 km W of Jiejiaonuocun, Yin Duoguosa (Verhoeff 1934, Golovatch 2019a)
244. M. hirsutus Golovatch, 2019 2315 Yunnan, NW of Lijiang, W of Chang Jiang (= Yangtze) River, NW of Jinzhuang, 2.5 km N of Tuozhi village (Golovatch 2019a)
245. M. rugosus (Golovatch, 2013) 2400 Yunnan, N of Lijiang (Golovatch 2013a, 2019a)
246. M. semirugosus (Golovatch, 2013) 2955 Sichuan, NW of Mianning (Golovatch 2013b, 2019a)
Genus Nedyopus Attems, 1914 170–450 E and SE Asia
247. Nedyopus beroni (Golovatch, 1995) 350–450 Jiangsu, Nanjing City, Mt. Zijin (Golovatch 1995)
248. N. picturatus (Golovatch, 1995) ca. 170 Guangxi, Guilin (Golovatch 1995)
Genus Orthomorpha Bollman, 1893 E and SE Asia, Sunda Archipelago
249. Orthomorpha coarctata (de Saussure, 1860) ca. 20 Pantropical; Hainan, Sanya (Golovatch 1994)
250. “Orthomorpha” endeusa Attems, 1898 ? China (Attems 1898)
Genus Orthomorphella Hoffman, 1963 China
251. Orthomorphella pekuensis (Karsch, 1881) ca. 40–165 Hebei, Shanlin, 70 km of Peking (Golovatch 1981); Hunan, Yuanling County, Mumaling (Zhang et al. 1997); New record: Jilin, Changchun City.
Genus Oxidus Cook, 1911 E and SE Asia
252. Oxidus gracilis C. L. Koch, 1847 200–1300 Subcosmopolitan, anthropochore; near Beijing; Shaanxi, Xi’an City; Guangxi, near Guilin; Sichuan, Maoxian County, NE of Shimian (Golovatch 2013a, 2014a)
Genus Piccola Attems, 1953 China and Vietnam
253. Piccola golovatchi Liu & Tian, 2015 ca. 840 Guangxi, Baise City, Tianlin County, Langping Town, Cave Shizikou Dadong (Liu and Tian 2015b)
Genus Polylobosoma Jeekel, 1980 10–1600 China and Vietnam
254. Polylobosoma panda (Golovatch, 2009) 1600 Shaanxi, Foping Nature Reserve, Panda area (Golovatch 2009, 2014a)
255. P. roseipes (Pocock, 1895) 10 Zhejiang, Ningpo (Jeekel 1980)
Genus Sellanucheza Enghoff, Golovatch & Nguyen, 2004 995–3155 E and SE Asia
256. Sellanucheza jaegeri Golovatch, 2013 1300–1700 Shaanxi, Mt. Taibaishan (Golovatch 2013a)
257. S. tenebra (Hoffman, 1961) ? Sichuan, Wushan (Hoffman 1961)
258. S. typica Golovatch, 2013 995–3155 Sichuan, Maoxian County, SE of Nanxizhen (Golovatch 2013b); Sichuan, NE of Shimian, Xiangshuigou River, Tianpingzi (Golovatch 2014a)
Genus Sigipinius Hoffman, 1961 2810–4195 China
259. Sigipinius campanuliformis Golovatch, 2013 3910 Yunnan, W of Lake Lugu, N of Dajisi (Golovatch 2013b)
260. S. complex Golovatch, 2013 3780–4120 Sichuan, S of Muli (Golovatch 2013b)
261. S. dentiger Golovatch, 2016 3570 Yunnan, Mt. Tianbaoshan between Shangrila and Habaxue Shan, E slope, NW of Bengla (Golovatch 2016b)
262. S. grahami Hoffman, 1961 2810–4170 Sichuan, Lixi County, SW of Tonghua; Jiuzhaigou County, N of Dajisi; Maoxian County, SE of Nanxizhen; Lixian, NNW of Xuecheng, Ertaizi; N of Lixian, Mengdonggou & Lianghekou divide, W of Xing Fanweizi; Gansu, WWS of Longnan (Wudu), Yin Duoguosa & Aounang divide; WWS of Longnan (Wudu), Yin Duoguosa & Yaxielu, W of Zhagazu, WWS of Longnan (Wudu), Wushenggou & Line Chaping divide; NNE Zhugqu, Minjiang Bas, 3 km ENE Xiaohuangya, Qinyugou (Golovatch 2013b, 2019a)
263. S. kabaki Golovatch, 2013 3330–3550 Xinjiang, Koeksu Basin (Golovatch 2013b)
264. S. montanus (Golovatch, 2011) 3710–4090 Yunnan, S of Nixi, near upper timber-line of a humid montane Abies forest; WNW of Zhongdian, humid mid-montane Abies forest with admixture of broad-leaved hardwood species (Golovatch 2011, 2013b)
265. S. pinnifer Golovatch, 2016 3625 Sichuan, SSE of Shimian, S of Zhuma (Golovatch 2016a)
266. S. simplex Golovatch, 2013 3915–4195 Sichuan, Jiulong County, SW of Wulaxixiang; Muli County, SW of Wulaxixiang (Golovatch 2013b); Sichuan, Kangding NNE of Yalaxiang, Shuangyanwo (Golovatch 2014a)
267. S. spiniger Golovatch, 2014 3690–3960 Yunnan, from Lijiang to Shangrila, 214 National Road, WSW of Edi (Golovatch 2014a)
Genus Sinomorpha Golovatch, 2013 China
268. Sinomorpha setosa Golovatch, 2013 1050 Sichuan, Mt. Emeishan, Wannian Monastery (Golovatch 2013a)
Genus Tetracentrosternus Pocock, 1895 Myanmar, Thailand and Indochina
269. Tetracentrosternus hoffmani Golovatch, 2013 1610 Yunnan, Mt. Gaolinggongshan, S of Pianma (Golovatch 2013a)
Genus Tonkinosoma Jeekel, 1953 500–1250 China and Vietnam
270. Tonkinosoma flexipes Jeekel, 1953 500 Guangxi, Hechi City, Fengshan County, Jinya Town, Hangdong village (Liu and Golovatch 2018a); also N Vietnam (Jeekel 1953)
271. T. tiani Liu & Golovatch, 2018 1250 Guizhou, Qianxinan, Anlong County, Sayu Town, Ganhan Dong Cave (Liu and Golovatch 2018a)
Genus Tylopus Jeekel, 1968 350–4025 Myanmar, China, Thailand and Indochina
272. Tylopus deharvengi Liu & Luo, 2013 350 Guangxi, Du’an County, Xia’ao Town, Cave I Dong (Liu and Luo 2013)
273. T. kabaki Golovatch, 2014 3575–4025 Yunnan, Deqen, Tuoxia Highway, Mt. Xiaruolisuzuxiang & Yezhizhen; same province, NW of Lijiang, W of Chang Jiang (Yangtze River), NW of Jinzhuang, 6 km of Tuozhi village; N of Lijiang, W of Maguwa, 4.2 km SE of Shanggaohan village; N of Lijiang, W of Maguwa, 4.4 km ENE of Shanggaohan village; Mekong Valley, ENE of Yezhixiang, 3 km NE of Houqing (Golovatch 2014a, 2019b)
274. T. nigromarginatus Golovatch, 2018 835 Chongqing, Mt. Jinyunshan, secondary forest, stump, trees, small cave (Golovatch 2018)
275. T. reductus Golovatch, 2013 1600–1800 Yunnan, Mt. Gaoligongshan, S of Pianma (Golovatch 2013a)
276. T. schawalleri Golovatch, 2013 2500–2700 Yunnan, Mt. Dincangshang, above Dali (Golovatch 2013a)
277. T. similis Golovatch, 2014 1670 Yunnan, from Lijiang to Shangrila, E of Guojie Luocun (Golovatch 2014a)
278. T. sinensis Golovatch, 1995 1315 Yunnan, Mengzi County, Cave Hafatiao Dong (Golovatch 1995)
Genus Wulingina Zhang, 1997 China
279. Wulingina macroloba Zhang, 1997 510 Hubei, Hefeng Tu jiazu County (Zhang 1997)
280. W. miniloba Zhang, 1997 510 Hubei, Hefeng Tu jiazu County (Zhang 1997)
Genus Yuennanina Attems, 1936 1915–1920 China
281. Yuennanina aceratogaster Zhang & Li, 1977 1920 Yunnan, Kunming City (Zhang and Li 1977)
282. Y. ceratogaster Attems, 1936 1920 Yunnan, Kunming City (Attems 1936)
283. Y. petalolobodes Chang & Zhang, 1989 1915 Yunnan, Kunming, Chenggong County (Chang and Zhang 1989)
Family Polydesmidae Leach, 1815 Palaearctic and SE Asia
Genus Epanerchodus Attems, 1901 35–3090 Central and E Asia, marginally N Vietnam
284. Epanerchodus belousovi Golovatch, 2014 2810 Sichuan, Kangding City (Golovatch 2014c)
285. E. chutou Liu & Golovatch, 2018 680 Guizhou, Shiqian County, Cave Feng Dong (Liu and Golovatch 2018b)
286. E. coniger Liu & Golovatch, 2018 ca. 1620 Guizhou, Bijie City, Zhijin County, Chengguan Town, Dongshan village, Cave Houshan Dong (Liu and Golovatch 2018b)
287. E. draco Geoffroy & Golovatch, 2004 ca. 1670 Yunnan, Zhenxiong County, a cave; Guizhou, Liupanshui City, Shuicheng County, Cave Shendongmigong Dong (Geoffroy and Golovatch 2004, Liu and Golovatch 2018b)
288. E. eurycornutus Zhang & Wang, 1992 885 Zhejiang, Mt. Tianmu (Zhang and Wang 1992)
289. E. frater Geoffroy & Golovatch, 2004 ca. 1670 Yunnan, Zhenxiong County, Cave Dahei Dong (Geoffroy and Golovatch 2004)
290. E. fuscus Golovatch, 2015 ca. 2450 Yunnan, Lanping County (Golovatch 2015b)
291. E. gladiatus Liu & Golovatch, 2018 920 Guizhou, Wuchuan County, Huangdu Town, Gaodong village, Cave Yinshi Dong (Liu and Golovatch 2018b)
292. E. jaegeri Golovatch, 2014 ca. 2345 Shaanxi, Mt. Taibaishan (Golovatch 2014b)
293. E. jiangxiensis Liu & Golovatch, 2018 475 Jiangxi, Lianhua County, Gaotan village, Cave Shuilian Dong (Liu and Golovatch 2018b)
294. E. koreanus Verhoeff, 1937 2230 Jilin, Mt. Changbaishan (Golovatch 2014b)
295. E. latus Liu & Golovatch, 2018 ca. 1330 Chongqing, Wushan County, Luoping Town, Qinglong village, Cave Qinglong Dong (Liu and Golovatch 2018b)
296. E. lipsae Golovatch & Geoffroy, 2014 ca. 750 Sichuan, Beichuan and Jiangyou counties, numerous caves (Golovatch and Geoffroy 2014, Liu and Golovatch 2018b)
297. E. martensi Golovatch, 2014 ca. 2345 Shaanxi, Mt. Taibaishan (Golovatch 2014b)
298. E. orientalis Attems, 1901 ca. 205 Guangxi, Fuchuan County, Cave Banbianshan Dong (Golovatch et al. 2012c), also Japan and Taiwan
299. E. parvus Liu & Golovatch, 2018 830 Guizhou, Cengong County, Pingzhuang Town, Cave Wanfuchangcheng Dong (Liu and Golovatch 2018b)
300. E. potanini Golovatch, 1991 Sichuan, Gansu and Yunnan provinces (Golovatch 1991a, 2014b)
301. E. schawalleri Golovatch, 2014 ca. 1550 Sichuan, Mt. Emeishan (Golovatch 2014b)
302. E. soror Geoffroy & Golovatch, 2004 ca. 1670 Yunnan, Zhenxiong County, caves Hama Dong, Dahei Dong and Xianren Dong (Geoffroy and Golovatch 2004, Liu and Golovatch 2018b)
303. E. sphaerisetosus Zhang & Chen, 1983 ca. 35 Zhejiang, 10 mi S of Jinhua City, Gaocun village (Zhang and Chen 1983)
304. E. stylotarseus Chen & Zhang, 1990 ca. 1220 Guizhou, Guanling County, several caves (Chen and Zhang 1990, Golovatch et al. 2007, 2012)
305. E. tujiaphilus Liu & Golovatch, 2018 730 Hunan, Longshan County, Huoyan village, Cave Tujiamei Dong (Liu and Golovatch 2018b)
306. E. typicus Golovatch, 2014 ca. 3030 Yunnan, Deqin County (Golovatch 2014c)
307. E. varius (Geoffroy & Golovatch, 2004) ca. 755–3090 Numerous caves in Hubei, Banqiao Town; and Sichuan, Xinlong and Beichuan counties (Geoffroy and Golovatch 2004, Golovatch et al. 2007, Golovatch and Geoffroy 2014)
308. E. yunnanensis Golovatch, 2014 1995 Yunnan, Dali City (Golovatch 2014b)
Genus Glenniea Turk, 1945 170–1510 Himalaya and China
309. Glenniea blanca Golovatch & Geoffroy, 2014 600 Sichuan, Tongjiang County, Cave Lou Fang Dong (= Grotte de la Maison) (Golovatch and Geoffroy 2014)
310. G. lagredae Golovatch & Geoffroy, 2014 1360–1510 Sichuan, Beichuan County, Cave Yuan Dong (= La grotte du Rocher); Sichuan, Huajiaoling County, Cave Zhangjiayankoukeng Dong (Golovatch and Geoffroy 2014)
311. G. prima Golovatch, Li, Liu & Geoffroy, 2012 ca. 170 Guangxi, Longzhou County, Shanglong Town, Lenglei Nonggang Forest (Golovatch et al. 2012c, Golovatch and Geoffroy 2014)
Genus Pacidesmus Golovatch, 1991 ca. 180–1865 China and N Thailand
312. Pacidesmus armatus Golovatch, Geoffroy & Mauriès, 2010 ca. 310 Guangxi, Huanjiang County, Cave Xialan Dong, caves Shui Dong and Shenglong Dong (Golovatch et al. 2010a)
313. P. bedosae Golovatch, Geoffroy & Mauriès, 2010 ca. 310 Guangxi, Huanjiang County, caves Dongtu Dong, Huoka Dong and Ganxiao Dong (Golovatch et al. 2010a)
314. P. bifidus Golovatch & Geoffroy, 2014 ca. 495 Guangxi, near Fengshan County, Cave Henglixin Dong (Golovatch and Geoffroy 2014, Liu and Golovatch 2019)
315. P. martensi Golovatch & Geoffroy, 2006 ca. 1495 Guizhou, Dafang County, Cave Hei Dong; Qianxi County, Honglin Town, caves Luoshui Dong and Luosai Dong (Golovatch and Geoffroy 2006, Golovatch et al. 2007, Liu and Golovatch 2019)
316. P. sinesis (Golovatch & Hoffman, 1989) ca. 1285 Guizhou, Zhenning County, Cave Kaikou Dong (Loksa 1960, Golovatch and Hoffman 1989, Chen and Meng 1990, Liu and Golovatch 2019)
317. P. superdraco Golovatch, Geoffroy & Mauriès, 2007 ca. 410 Guizhou, Libo County, Cave Laitai Dong (Golovatch et al. 2007)
318. P. tiani Golovatch, Geoffroy & Mauriès, 2010 ca. 310 Guangxi, Huanjiang County, caves Ganglai Dong I and II (Golovatch et al. 2010a)
319. P. trifidus Golovatch & Geoffroy, 2014 ca. 180 Guangxi, Guilin City, Cave Kulou Dong (Golovatch and Geoffroy 2014); Yangshuo County, Cave Guanshan No. 4; Xiufeng District, Cave Maomaotou; Yangshuo County, Cave Shangshuiyan (Liu and Wynne 2019)
320. P. trilobatus Liu & Golovatch, 2020 ca. 1315 Yunnan, Wenshan County, Liujing Town, Laozhai village, Cave I Dong (Liu and Golovatch 2020)
321. P. uncatus Liu & Golovatch, 2020 ca. 1865 Yunnan, Qujing City, Zhanyi County, Cave Tianshengqiao Dong (Liu and Golovatch 2020)
322. P. whitteni Liu & Golovatch, 2020 ca. 755 Guangxi, Fengshan County, Jinya Town, Hangdong village, Cave I Dong (Liu and Golovatch 2020)
Genus Polydesmus Latreille, 1802–03 Amphi-Palaearctic
323. Polydesmus liber Golovatch, 1991 ca. 140 Hong Kong (Golovatch 1991a)
Family Pyrgodesmidae Silvestri, 1896 Pantropical
Genus Cryptocorypha Attems, 1907 Old World, up to Melanesia in the east
324. Cryptocorypha spinicoronata (Zhang & Li, 1981) ca. 1110 Guangxi, Tianlin County, Langping Town (Zhang and Li 1981b)
Family Xystodesmidae Cook, 1895 Holarctic, E and SE Asia up to N Vietnam in the south
Genus Kiulinga Hoffman, 1956 10–1080 China
325. Kiulinga jeekeli Hoffman, 1956 1080 Jiangxi, Jiujiang City, Jiguling (Hoffman 1956, Zhang and Mao 1984)
326. K. lacustris (Pocock, 1895) 10 Zhejiang, 25 mi S of Ninghsien, Lake Wo-Lee (Hoffman 1956)
327. K. lobosa Zhang & Mao, 1984 ca. 30 Zhejiang, Zhoushan City, Daishan Island (Zhang and Mao 1984)
Genus Riukiaria Attems, 1938 170–4440 E Asia up to N Vietnam in the south
328. Riukiaria belousovi Golovatch, 2014 4100 Sichuan, Muli County, SW of Wulaxixiang (Golovatch 2014d)
329. R. capaca Wang & Zhang, 1993 170 Fujian, Jiangle County (Wang and Zhang 1993b)
330. R. chinensis Tanabe, Ishii & Yin, 1996 885 Zhejiang, Mt. Tianmu (Tanabe et al. 1996)
331. R. davidiani Golovatch, 2014 2810 Sichuan, Lixian County, SW of Tonghua (Golovatch 2014d)
332. R. kabaki Golovatch, 2014 4440 Sichuan, Kangding City, NNE of Walaxiang, NE of Yusicun (Golovatch 2014d)
333. R. korolevi Golovatch, 2014 2900 Sichuan, W of Jiuzhaigou (Golovatch 2014d)
334. R. martensi Golovatch, 2014 1700 Shaanxi, Mt. Taibaishan, southern slopes, above Houzhenzi, primary broadleaved forest (Golovatch 2014d)
335. R. spatuliformis Golovatch, 2015 2525 Sichuan, N of Luding City, N of Lanan (Golovatch 2015b)
336. R. tianmu (Tanabe, Ishii & Yin, 1996) 885 Zhejiang, Mt. Tianmu (Tanabe et al. 1996, Golovatch 2014d)
Family Opisotretidae Hoffman, 1980 Himalaya, Myanmar, Indochina, Indonesia, New Guinea, Ryukyu Islands, Japan and Christmas Island, Australia, Indian Ocean (Golovatch et al. 2013)
Genus Carlotretus Hoffman, 1980 S China and Sumatra, Indonesia (Golovatch et al. 2013)
337. Carlotretus triramus Golovatch, Geoffroy, Stoev & VandenSpiegel, 2013 ca. 200 Guangxi, Chongzuo City, Longzhou County, Shanglong Town, Nonggang Forest (Golovatch et al. 2013)
Genus Martensodesmus Golovatch, 1987 150–200 Himalaya, Indochina and S China (Golovatch et al. 2013)
338. Martensodesmus bedosae Golovatch, Geoffroy, Stoev & VandenSpiegel, 2013 ca. 150 Guangxi, Hechi City, Du’an County, Baling karst hill (Golovatch et al. 2013)
339. M. spiniger Golovatch, Geoffroy, Stoev & VandenSpiegel, 2013 ca. 200 Guangxi, Chongzuo City, Longzhou County, Shanglong Town, Nonggang Forest (Golovatch et al. 2013)

As noted above, according to the ordinal and supra-ordinal distributions in the Diplopoda and a purely biogeographic reconstruction of their origins and early evolution by Shelley and Golovatch (2011), the Oriental Region is the only biogeographic realm of the globe that supports all 16 extant orders of the class. Amongst them, eleven orders are known to occur in mainland China, with the distribution patterns of their constituent families and genera available in Table 1. The remaining five orders, albeit formally excluded from consideration, are added to the roster (Table 2), because representatives of the orders Glomeridesmida, Siphonophorida, Siphonocryptida, Siphoniulida, and Stemmiulida occur or occurred in the adjacent parts of East, Southeast and/or Central Asia. Thus, one extant species of Glomeridesmida and Siphonocryptida each is known from northern Thailand and Taiwan, respectively (Shelley and Golovatch 2011, Golovatch 2015a), several Siphonophorida have been recorded from Vietnam, Laos and northern Pakistan (Jeekel 2001), while fossil Siphoniulida have recently been described from northern Myanmar (Liu et al. 2017c). Two very small orders, Siphoniulida and Siphonocryptida, are considered relict, in a stage of evolutionary decline, whereas most if not all of the remaining orders of Diplopoda are far more diverse and currently in an expansive stage of their evolution (Shelley and Golovatch 2011, Shelley 2011, Golovatch 2015a).

Table 2.

Distribution patterns of all 16 extant millipede orders, those presently known to occur in mainland China being marked with an asterisk.

Orders Distribution pattern Orders Distribution pattern
Polyxenida* Cosmopolitan Siphonophorida Pantropical
Glomeridesmida Pantropical Chordeumatida* Holarctic + Neotropical + Oriental
Glomerida* Holarctic + Oriental Callipodida* Holarctic + Oriental
Sphaerotheriida* Old World Julida* Holarctic + Oriental
Platydesmida* Subcosmopolitan Stemmiulida Pantropical
Polyzoniida* Subcosmopolitan Spirostreptida* Pantropical
Siphoniulida Neotropical + Oriental Spirobolida* Pantropical
Siphonocryptida Palaearctic + Oriental Polydesmida* Cosmopolitan

The greatest and about equal shares in the diplopod fauna of mainland China expectedly belong to Holarctic/Palaearctic or Oriental elements, with the former naturally dominating the northern, the latter the southern, parts of the country, and both thoroughly mixed and intermingled mainly in the more central parts. The orders Polyxenida, Polyzoniida, Platydesmida, Glomerida, Callipodida, Chordeumatida, and Julida, the families Polydesmidae and Xystodesmidae, as well as certain genera of Paradoxosomatidae seem best to be attributed to Holarctic/Palaearctic components in the fauna of China. In contrast, most of the remaining higher taxa such as the largely tropical orders Sphaerotheriida, Spirobolida, and Spirostreptida, the families Cryptodesmidae, Haplodesmidae, Opisotretidae, and Pyrgodesmidae, as well as several genera of Paradoxosomatidae seem to represent the Oriental stem. Only two families (of 25, or 8%) are endemic or subendemic to China: the monobasic Guizhousomatidae (Chordeumatida), an apparently relict troglobiont from Guizhou Province, and the Paracortinidae (Callipodida) with two genera (maybe just one, see Stoev and Geoffroy 2004) and a handful of species (including two from northern Vietnam). The number of endemic genera is quite high, 16 (of 65, or ca. 25 %): Sinostemmiulus (Julida), Parabilingulus, Agaricogonopus, Junceustreptus, Prominulostreptus (all Spirostreptida), Lipseuma (Chordeumatida), Angulifemur (Callipodida), Belousoviella, Gonobelus, Mandarinopus, Orthomorphella, Sigipinius, Sinomorpha, Wulingina, Yuennanina (all Polydesmida: Paradoxosomatidae), and Kiulinga (Polydesmida: Xystodesmidae). One might think the higher the altitude, the more likely the taxon’s Holarctic or Palaearctic origin and, vice versa, the lower the elevation, the more probable a tropical descent. However, the vertical distributions usually fail to provide a clear-cut support to attributing a higher taxon to this or that stem. The following examples can serve to show this.

The huge, Eurasian, warm-temperate to tropical genus Hyleoglomeris (Glomeridae, Glomerida) currently contains 100+ species, including numerous cavernicoles. Unlike the glomerid fauna of the adjacent Indochina which harbours a considerable proportion of endemic genera (60 % in Vietnam), continental China currently supports only 32 species of Hyleoglomeris, most of which occur in caves alone (Golovatch 2015a). The genus ranges from the Balkans in the west, though Anatolia, the Caucasus, Central Asia, the Himalaya, Myanmar and Indochina, to Taiwan, the Philippines and Sulawesi, Indonesia in the east. Importantly, a fossil congener is known from Baltic amber (Eocene, 44 Mya) (Wesener et al. 2019). Hyleoglomeris spp. are widespread across China and occur at various elevations, from nearly sea-level to high mountains (Fig. 2), the highest record belonging to H. sinensis (2810 m a.s.l.) (Table 1). In the Himalaya of Nepal, one species occurs even higher in the mountains, being high-montane: H. khumbua Golovatch, 1987 (3250–3300 m a.s.l.) (Golovatch and Martens 2018).

Figure 2. 

Distribution of the family Glomeridae, genus Hyleoglomeris in mainland China. Red lines show the transect Baoshan – Kangding – Honghe – Chongqing – Libo – Du’an – Tongren – Guilin – Linwu – Hong Kong – Nanjing, along which the elevations are crudely indicated below. 1 H. albicorporis 2 H. sinensis 3 H. reducta 4 H. maculata 5 H. heshang 6 H. gudu 7 H. rhinceros 8 H. nigu 9 H. getuhensis 10 H. aschnae 11 H. yinshi 12 H. grandis 13 H. eusulcata 14 H. xueju 15 H. wuse 16 H. qiyi 17 H. curtisulcata 18 H. mulunensis 19 H. kunnan 20 H. mashanorum 21 H. baxian 22 H. variabilis 23 H. multistriata 24 H. generalis 25 H. rukouqu 26 H. xuxiakei 27 H. lii 28 H. xia 29 H. youhao 30 H. tiani 31 H. bicolor, 32 H. emarginata.

A very similar pattern is demonstrated by the subendemic genus Paracortina (Paracortinidae, Callipodida), with 12 species, of which ten (Fig. 3) are confined to the mountains of southwestern China (Liu and Tian 2015c), mostly high-montane (3300 m a.s.l., Table 1). Only a few are cavernicoles.

Nepalmatoiulus (Julidae, Julida) is another very large genus which presently comprises 55 species that span from the central Himalaya in the west, through Bhutan, Myanmar, Indochina, Thailand and West Malaysia, to the Ryukyus, Japan and Taiwan in the east (Enghoff 1987b). Seven species range across the southern parts of China (Fig. 4), including two high-montane ones (2750–3650 m a.s.l., Table 1). Although Beron (2008) reported closer unidentified Diplopoda from up to 5300 m a.s.l. from Nepal, the world’s highest record for a known species belongs to N. ivanloebli Enghoff, 1987, also from Nepal: 4800 m a.s.l. (Enghoff 1987b, Shelley and Golovatch 2011). The same general pattern is observed in the similarly speciose (ca. 50 spp.), but more boreal genus Anaulaciulus (Julidae), the distribution of which covers northern Pakistan and India, the Himalaya, northern Myanmar, the Far East of Russia, all Japan and Korea, Taiwan, as well as central and eastern China. The highest record belongs to A. bilineatus Korsós, 2001 from Nepal: 3600–4300 m a.s.l. (Korsós 2001). Unlike Nepalmatoiulus, no Anaulaciulus spp. are known to occur in southern China, both these genera being allo- to parapatric. Among the Julidae in China, only very few are cavernicoles.

Figure 3. 

Distribution of the family Paracortinidae, genus Paracortina in mainland China. Red lines show the transect Batang – Shangrila – Yajiang – Zhenxiong – Baise, along which the elevations are crudely indicated below. 1 P. viriosa 2 P. serrata 3 P. thallina 4 P. carrinata 5 P. leptoclada 6 P. stimula 7 P. voluta 8 P. chinensis 9 P. yinae 10 P. zhangi.

Figure 4. 

Distribution of the family Julidae, genus Nepalmatoiulus in mainland China. Red lines show the transect Linzhi – Kangding – Suining – Jiujiang – Meizhou, along which the elevations are crudely indicated below. 1 N. tibetanus 2 N. brachymeritus 3 N. rhaphimeritus 4 N. polyakis 5 N. fraterdraconis 6 N. eulobos. Nepalmatoiulus yunnanensis is not mapped because no exact locality in Yunnan is known.

Particularly clear Palaearctic origins are observed in the large genus Skleroprotopus (Mongoliulidae, Julida), most species of which inhabit the Russian Far East, Korea, Japan and China (Table 1), the small Siberian genus Angarozonium (Polyzoniidae, Polyzoniida) only marginally encountered in northern China (Table 1), the rather small Siberio-Nearctic genus Orinisobates (Nemasomatidae, Julida) represented in China by a single species endemic to the southern Tianshan Mountains (Table 1) (Mikhaljova 2017). The same concerns Polydesmus (Polydesmidae, Polydesmida), a very large genus with ca. 80 species, most of which occur in Europe, the Mediterranean area, Anatolia and the western Caucasus, but a few are known from Japan, and one each in northern Vietnam and Hong Kong (Table 1) (Golovatch 1991a, Nguyen 2009).

The large genus Nepalella (Megalotylidae, Chordeumatida), with its 27 species spanning from Nepal (10 species) in the west, through Myanmar (two species) and Thailand (two species), to Vietnam (one species) in the south, and southwestern China (12 species, including several presumed troglobionts) in the north (Liu, Wesener et al. 2017d), shows the same general pattern (Fig. 5). Most congeners are mid-montane, but one, N. marmorata, has been recorded from ca. 4350 m a.s.l. (Table 1).

Figure 5. 

Distribution of the family Megalotylidae, genus Nepalella in mainland China. Red lines show the transect Baoshan – Xinlong – Beichuan – Shuicheng – Guiyang – Jinfoshan, along which the elevations are crudely indicated below. 1 N. kavanaughi 2 N. pianma 3 N. magna 4 N. griswoldi 5 N. marmorata 6 N. lobata 7 N. grandoides 8 N. grandis 9 N. caeca 10 N. troglodytes 11 N. jinfoshan 12 N. wangi.

Basically the same picture is revealed in the distribution of the huge Central to East Asian genus Epanerchodus (Polydesmidae, Polydesmida) which presently encompasses 118 species or subspecies, both epi- and endogean, including 25 across almost entire continental China (Liu and Golovatch 2018b) (Table 1, Fig. 6). Their vertical distributions range from nearly sea-level to high-montane (3090 m a.s.l.), but a few congeners from the Himalaya occur even up to 4250 m a.s.l. (Golovatch and Martens 2018).

The genus Pacidesmus (Polydesmidae, Polydesmida) shows a highly peculiar distribution (Fig. 7), with all of its eleven Chinese species being low- to mid-montane and restricted to karst caves in the south (Liu and Golovatch in press), whereas the type species, P. shelleyi Golovatch, 1991, comes from the summit (2200–2500 m a.s.l.) of Mount Doi Inthanon, northern Thailand (Golovatch 1991a). Similarly, the small genus Glenniea (Polydesmidae) contains five lowland to mid-montane epigean species from the Himalaya of India and Bhutan (Golovatch and Martens 2018), as well as another three species (including two cavernicoles) from southern China (Golovatch and Geoffroy 2014) (Table 1, Fig. 8).

Figure 6. 

Distribution of the family Polydesmidae, genus Epanerchodus in mainland China. Red lines show the transect Shangrila – Dali – Xingyi – Beichuan – Zhengxiong – Tongren – Xi’an – Wushan – Fuchuan – Lianhua – Hangzhou – Changbaishan, along which the elevations are crudely indicated below. 1 E. potanini 2 E. typicus 3 E. fuscus 4 E. yunnanensis 5 E. belousovi 6 E. schawalleri 7 E. stylotarseus 8 E. lipsae 9 E. varius 10 E. frater 11 E. soror 12 E. draco 13 E. coniger 14 E. gladiatus 15 E. chutou 16 E. parvus 17 E. jaegeri 18 E. martensi 19 E. tujiaphiulus 20 E. latus 21 E. orientalis 22 E. jiangxiensis 23 E. enrycornutus 24 E. sphaerisetosus 25 E. koreanus.

Figure 7. 

Distribution of the family Polydesmidae, genus Pacidesmus in mainland China. Red lines show the transect Qujing – Wenshan – Dafang – Fengshan – Huanjiang – Guilin, along which the elevations are crudely indicated below. 1 P. uncatus 2 P. trilobatus 3 P. martensi 4 P. sinensis 5 P. whitteni 6 P. bifidus 7 P. superdraco 8 P. tiani 9 P. bedosae 10 P. armatus 11 P. trifidus.

Figure 8. 

Distribution of the family Polydesmidae, genus Glenniea in mainland China. Red lines show the transect Beichuan – Longzhuan – Tongjiang, along which the elevations are crudely indicated below. 1 G. lagredae 2 G. prima 3 G. blanca.

The great Holarctic family Xystodesmidae (Polydesmida) presently encompasses 66 genera and ca. 410 species, most of which occur in the Nearctic. Only few genera and species are known from Central and northern South America (to Ecuador in the south), the Antilles, the Mediterranean region and East Asia (Shelley and Smith 2018). The largest East Asian genus Riukiaria currently contains 35 species or subspecies from southern Japan, southern Korea, Taiwan and China (Korsós et al. 2011, Golovatch 2014d, 2015b, Nguyen 2016). We disagree with Nguyen (2016), who split Riukiaria into two genera and created a new genus, Parariukiaria Nguyen, 2016, to accommodate a new species from northern Vietnam and three previously described ones from China. To our mind, Riukiaria and Parariukiaria show all transitional stages in the reduction of a gonoprefemoral process and, albeit without formal synonymy advanced here, both may well be regarded as representing a single large genus, in which several peripheral, southernmost congeners demonstrate a more or less strongly suppressed process on the gonopodal prefemur, from relatively small to totally missing. All nine Riukiaria species in China are epigean and span across the central and southern parts of the country, occurring in lowland to high-montane habitats (170–4440 m a.s.l., Table 1, Fig. 9).

Figure 9. 

Distribution of the family Xystodesmidae, genus Riukiaria in mainland China. Red lines show the transect Muli – Kangding – Jiuzhaigou – Xi’an – Jiangle – Hangzhou, along which the elevations are crudely indicated below. 1 R. belousovi 2 R. kabaki 3 R. spatuliformis 4 R. davidiani 5 R. korolevi 6 R. martensi 7 R. capaca 8 R. chinensis 9 R. tianmu.

As noted above, in China the great family Paradoxosomatidae, which is amongst the largest in the class (200+ genera, 1,000+ species), dominates most of the tropical faunas across the world, but is absent from the Nearctic, contains remarkably few troglobionts (Golovatch 2015a) and comprises genera of various origins. Some seem to be rooted in the Palaearctic (including several endemic or subendemic ones), the others are likely to be Oriental. Among the former elements, the following two rather species-rich genera can be taken as examples.

The genus Hedinomorpha is subendemic to China, with most of its 17 species known from the country being high-montane (up to 4490 m a.s.l., Table 1, Fig. 10), and only one more restricted to Tajikistan, Central Asia (Golovatch 2019b). The genus Sigipinius is strictly endemic to mainland China and contains nine high-montane species (2810–4195 m a.s.l., Table 1, Fig. 11). Such paradoxosomatid genera as Cawjeekelia, Kronopolites, Mandarinopus and Orthomorphella likewise seem best to be attributed to Palaearctic elements in the fauna of China.

Figure 10. 

Distribution of the family Paradoxosomatidae, genus Hedinomorpha in mainland China. Red lines show the transect Shangrila – Ganzi – Liangshan – Xining – Lanzhou – Jiuzhaigou – Xi’an, along which the elevations are crudely indicated below. 1 H. montana 2 H. yunnanensis 3 H. proxima 4 H. crassiterga 5 H. bifida 6 H. subnigra 7 H. reducta 8 H. martensi 9 H. circofera 10 H. affinis 11 H. nigra 12 H. altiterga 13 H. flavobulbus 14 H. biramipedicula 15 H. jeekeli; neither H. circularis nor H. hummelii is mapped because their exact type localities remain unknown.

Figure 11. 

Distribution of the family Paradoxosomatidae, genus Sigipinius in mainland China. Red lines show the transect Aksu – Shangrila – Lijiang – Kangding – Jiuzhaigou, along which the elevations are crudely indicated below. 1 S. kabaki 2 S. montana 3 S. spiniger 4 S. dentiger 5 S. campanuliformis 6 S. complex 7 S. simplex 8 S. pinnifer 9 S. grahami.

In contrast, Paradoxosomatidae also contain a good number of presumed Oriental components, mostly tropical to subtropical. Thus, the genus Hylomus presently comprises 36 species from Myanmar, Thailand, Laos, Vietnam and China (Srisonchai et al. 2018, Liu and Wynne 2019, Golovatch 2019b). Many of them are presumed troglobionts. The distributions of all 20 Hylomus spp. recorded from China cover much of the southern and eastern parts of the country and are only confined to lowland to mid-montane habitats (ca. 140–910 m a.s.l., Table 1, Fig. 12). At the moment, with its 73 species (Golovatch 2019b) that range from southern China in the north, through most of Indochina, to Myanmar in the south, Tylopus remains the largest genus of Paradoxosomatidae globally. However, the altitudinal distributions vary from lowland to high-montane (350–4025 m a.s.l., Table 1), cavernicoles are few, while the Chinese congeners mark the northern range limit of the genus and are confined to the southwestern parts of the country (Fig. 13). Because Tylopus and Hedinomorpha seem to be particularly similar morphologically and co-occur, albeit probably never strictly sympatric, in southwestern China (at least Yunnan, Figs 10, 13), these areas seem to mark the southern range limit of Hedinomorpha.

Figure 12. 

Distribution of the family Paradoxosomatidae, genus Hylomus in mainland China. Red lines show the transect Tianlin – Ziyun – Du’an – Huanjiang – Guilin – Linwu – Qingyuan – Jiujiang, along which the elevations are crudely indicated below. 1 H. minutuberculus 2 H. getuhensis 3 H. phasmoides 4 H. variabilis 5 H. parvulus 6 H. scolopendroides 7 H. nodulosus 8 H. scutigeroides 9 H. spinitergus 10 H. lui 11 H. yuani 12 H. cornutus 13 H. lingulatus 14 H. spinissimus 15 H. eupterygotus 16 H. laticollis 17 H. simplipodus 18 H. similis 19 H. draco; H. longispinus is not mapped because its exact type locality remains unknown.

Figure 13. 

Distribution of the family Paradoxosomatidae, genus Tylopus in mainland China. Red lines show the transect Shangrila – Dali – Mengzi – Jinyunshan – Du’an, along which the elevations are crudely indicated below. 1 T. reductus 2 T. kabaki 3 T. similis 4 T. schawalleri 5 T. sinensis 6 T. nigromarginatus 7 T. deharvengi.

The relatively large genera Anoplodesmus, Antheromorpha, Enghoffosoma, Nedyopus and Sellanucheza also seem best to refer to as Oriental components in the fauna of China, because it is southern China that marks their northern range limits. The same concerns the small genera Hirtodrepanum, Inversispina, Piccola, Polylobosoma and Tetracentrosternus, all of which show one or a few congeners either in the Himalaya and/or Myanmar, or northern Vietnam, or Taiwan. The mono- or oligotypic Belousoviella, Gonobelus, Sinomorpha, Wulingina, and Yuennanina are all strictly endemic to China, mostly to its southwestern parts, but their Oriental stem is clear-cut due to their closest affinities.

The immediately above paradoxosomatid genera endemic or subendemic to southern China which all seem to be of Oriental stock, together with some other polydesmidans like Carlotretus and Martensodesmus (both Opisotretidae), Glenniea and Pacidesmus (both Polydesmidae, Figs 7, 8), as well as several others (e.g. Cryptodesmidae, Haplodesmidae, Pyrgodesmidae), regardless of whether they are Oriental or Palaearctic in origin, seem to be sufficiently numerous and manifest to warrant the recognition of a separate, albeit secondary, subordinate, southern Chinese diversity and faunogenetic centre which must have seriously contributed to at least the faunas of the adjacent parts of the Himalaya, Myanmar, Thailand, Indochina and Taiwan. The influence of that southern Chinese centre in the Himalaya has recently been emphasized (Golovatch and Martens 2018).

The Oriental realm as one of the main sources for the formation of the millipede fauna of China can also be exemplified by the basically tropical to subtropical orders Sphaerotheriida, Spirobolida and Spirostreptida, as well as the families Cryptodesmidae, Haplodesmidae, Opisotretidae, Pyrgodesmidae (all Polydesmida) and Sinocallipodidae (Callipodida), some of which often vary a lot in altitudinal distributions just like numerous Holarctic/Palaearctic groups. The often presumed rule “tropical elements for low elevations only” does not always work.

The genus Glyphiulus, the largest in the family Cambalopsidae (Spirostreptida), presently comprises 60+ species in East and Southeast Asia (to Borneo in the east), 42 of which are encountered at 105–4150 m a.s.l. across China (Fig. 14). Most of them are cavernicoles (Liu and Wynne 2019). A similarly large and even more widespread genus, Eutrichodesmus (Haplodesmidae), presently encompasses 50 species (Liu et al. 2017b, Liu and Wynne 2019) which range from southern Japan and Taiwan in the north, through entire Southeast Asia, to Vanuatu, Melanesia in the south. The distributions of all 24 species that populate continental China seem to be more typical, much better agreeing with the above rule: 65–1495 m a.s.l. (Table 1, Fig. 15). At least half of them are also cavernicoles.

Figure 14. 

Distribution of the family Cambalopsidae, genus Glyphiulus in mainland China. Red lines show the transect Xinlong – Liangshan – Honghe – Chengdu – Guiyang – Hechi – Longshan – Guilin – Hong Kong – Qingyuan, along which the elevations are crudely indicated below. 1 G. basalis 2 G. liangshanensis 3 G. beroni 4 G. paragranulatus 5 G. semigranulatus 6 G. subobliteratus 7 G. subgranulatus 8 G. obliteratus 9 G. latus 10 G. intermedius 11 G. zorzini 12 G. guangnanensis 13 G. foetidus 14 G. sinensis 15 G. pergranulatus 16 G. quadrohamatus 17 G. paracostulifer 18 G. latellai 19 G. obliteratoides 20 G. rayrouchi 21 G. difficilis 22 G. impletus 23 G. granulatus 24 G. basazsi 25 G. calceus 26 G. parobliteratus 27 G. pulcher 28 G. echinoides 29 G. acutus 30 G. mulunensis 31 G. proximus 32 G. tiani 33 G. paramulunensis 34 G. speobius 35 G. deharvengi 36 G. melanoporus 37 G. septentrionalis 38 G. adeloglyphus 39 G. maocun 40 G. formosus 41 G. recticulus; G. anophthalmus and G. lipsorum are not mapped because their exact type localities remain unknown, whereas G. granulatus is mapped, but it is pantropical.

Figure 15. 

Distribution of the family Haplodesmidae, genus Eutrichodesmus in mainland China. Red lines show the transect Mengla – Hekou – Beichuan – Guiyang – Huanjiang – Guilin – Yichang – Qingyuan – Guidong – Fenyi – Wuyishan – Hangzhou, along which the elevations are crudely indicated below. 1 E. dorsiangulatus 2 E. monodentus 3 E. arcicollaris 4 E. triangularis 5 E. tenuis 6 E. latellai 7 E. obliteratus 8 E. incisus 9 E. latus 10 E. soesilae 11 E. triglobius 12 E. distinctus 13 E. trontelji 14 E. planatus 15 E. similis 16 E. sketi 17 E. lipsae 18 E. jianjia 19 E. apicalis 20 E. digitatus 21 E. spinatus 22 E. simplex 23 E. anisodentus 24 E. pectinatidentis.

Discussion

The diversity estimates presented in Table 1, i.e. 339 species, 71 genera, 26 families, and eleven orders, are much or significantly higher than those reported from the main adjacent areas. The similarly huge territories of Siberia and the Russian Far East that lie north of China support only ca. 130 species, 46 genera, 18 families and five orders of Diplopoda, while the fauna is reasonably well known (Mikhaljova 2017). This is hardly surprising because the prevailing permafrost and sharply continental climates of Asian Russia are largely too harsh to sustain a rich millipede fauna. The even harsher, mostly arid Mongolia is extremely poor in millipedes, with some nine species, five genera and families, and three orders involved (Mikhaljova 2012, Nefediev et al. 2015).

In contrast, the great Himalayan Range spanning for >2,300 km from northwest to southeast and mostly lying south of China supports >275 species, 53 genera, 23 families and 13 orders of diplopods (Golovatch and Martens 2018). Similarly, the fauna of India presently amounts to > 270 species, at least 90 genera, 25 families, and eleven orders (Golovatch and Wesener 2016), vs. 92 species from 34 genera, 13 families, and eight orders recorded from Myanmar (Likhitrakarn et al. 2017) or ca. 230 species in Thailand (Likhitrakarn et al. 2019). A direct correlation between area and latitude is clear: the larger the area and the closer it lies to the equator, the richer the biota, including the diplopod faunas. However, the more southerly, the greater the diversity, and the more incomplete and fragmentary is our knowledge.

Certainly the Chinese millipede fauna still remains strongly understudied, given the country’s great size and habitat diversity, including the globe’s greatest karst areas. It may well amount to 1,000 species (Golovatch 2015a), chiefly due to the still particularly poorly studied micropolydesmidans, as well as cavernicoles. Southern China’s karsts are unique in often harbouring up to 5–6 diplopod species per cave (Golovatch 2015a). At least some of the remaining orders such as Glomeridesmida, Siphonocryptida, Siphonophorida, Siphoniulida, and Stemmiulida that occur in the Oriental Region (Table 2), including areas immediately adjacent to mainland China, may also be expected to populate the country. For example, Jiang et al. (2019) have recently described a fossil Siphonophorida from Cretaceous amber (ca. 99 Mya) in northern Myanmar, and an extant species is long known to occur in northern Pakistan (Golovatch 1991b). In addition, the same Burmese amber contains still undescribed Stemmiulida (Stoev et al. 2019) and two described species of Siphoniulida (Liu et al. 2017c). Likewise, as noted above, an extant species of Siphonocryptida and Glomeridesmida each is known from Taiwan and northern Thailand, respectively (Korsós et al. 2008, Shelley 2011).

While the Palaearctic/Holarctic components expectedly dominate the fauna of the northern parts of the country, the Oriental ones prevail in its south and along the Pacific coast. Both realms are increasingly mixed and intermingled towards China’s centre. However, in addition to the above traditional views, based on millipede distribution patterns alone, southern China seems to harbour a subordinate, but highly peculiar faunal nucleus, or origin centre of its own, whence the adjacent Himalaya, Indochina and/or Taiwan could have become populated by younger lineages. The presence of a family (the monobasic Guizhousomatidae) and numerous genera endemic or subendemic to southern China, both apparently relict and relatively advanced, seems to be evidence of this. Within the order Callipodida alone, the family Sinocallipodidae seems to be the basalmost and representing a suborder of its own, the Paracortinidae is a more advanced subendemic, same as the mostly Central Asian Caspiopetalidae (Stoev and Geoffroy 2004, Stoev and Enghoff 2011). More importantly, a fossil family representing a separate suborder has recently been discovered in the Cretaceous Burmese amber, ca. 99 Mya (Stoev et al. 2019).

The millipede fauna of mainland China is thus a tangled mixture of zoogeographic elements of various origins and ages, apparently both relict and more advanced. The few anthropochores/introductions must have been the latest faunal “layer” to populate China.

Acknowledgements

We are grateful to the caving team of the South China Agricultural University (SCAU), Guangzhou, China, for their assistance. The second author was sponsored by the National Natural Science Foundation of China (Grant no. 31801956 and Grant no. 41871039). Special thanks go to Robert Mesibov (Tasmania, Australia) and Pavel Stoev (Sofia, Bulgaria), who provided very helpful reviews of the manuscript and thus considerably improved it. The first author was supported by the Presidium of the Russian Academy of Sciences, Program No. 41 “Biodiversity of natural systems and biological resources of Russia”.

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