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Research Article
Distribution, diversity patterns and faunogenesis of the millipedes (Diplopoda) of the Himalayas
expand article infoSergei I. Golovatch, Jochen Martens§
‡ Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Moscow, Russia
§ Johannes Gutenberg University of Mainz, Mainz, Germany
Open Access

Abstract

The Himalayas support a highly rich, diverse, multi-layered, mostly endemic diplopod fauna which presently contains >270 species, 53 genera, 23 families and 13 orders. This is the result of mixing the ancient, apparently Tertiary and younger, Plio-Pleistocene elements of various origins, as well as the most recent anthropochore (= man-mediated) introductions. At the species and, partly, generic levels, the fauna is largely autochthonous and sylvicolous, formed through abounding in situ radiation and vicariance events. In general, the species from large genera and families tend to occupy a wide range of altitudes, but nearly each of the constituent species shows a distribution highly localized both horizontally and altitudinally, yet quite often with sympatry or even syntopy involved. The bulk of the fauna is Indo-Malayan in origin, with individual genera or families shared with those of SE Asia (mostly) and/or S India (few). Sino-Himalayan and, especially, Palaearctic components are subordinate, but also clearly distinguishable.

Keywords

Diplopoda, faunistics, Plio-Pleistocene, Asia, Palaearctic

Introduction

The Himalaya Range, or Himalayas for short, meaning “the abode of snow” in Sanskrit, is the mountain range in Asia that separates the Indian subcontinent from the Tibetan Plateau. Sometimes by extension, it is also the name of a massive mountain system that includes the Karakoram, the Hindu Kush, and other, lesser, ranges that reach out from the Pamir Knot (http://maps.thefullwiki.org/Himalayas). However, below the Himalayas is treated in the strict sense. The main Himalayan Range runs, northwest to southeast, from the Indus River valley to the Brahmaputra River valley, forming an arc which varies in width from 400 km in the western Kashmir-Xinjiang region to 150 km in the southeastern Tibet-Arunachal Pradesh region. The range consists of three extensive sub-ranges, with the northernmost, and highest, known as the Great Himalayas.

The Himalayan mountain system is the Earth’s highest and home to the world’s highest peaks, the Eight-thousanders, which include Mount Everest and K2. The system, which includes various outlying sub-ranges, stretches across five countries: India, Nepal, Bhutan, China and Pakistan. The Himalayan Range is bordered on the northwest by the Karakoram and Hindu Kush ranges, on the north by the Tibetan Plateau, and on the south by the Indo-Gangetic Plain. The region is roughly delimited by 74°E in the west and 95°E in the east. Some of the world’s major rivers, the Indus, the Ganges, and the Tsangpo-Brahmaputra, rise in the Himalayas, and their combined drainage basin is home to some 600 million people. The Himalayas have profoundly shaped the cultures of South Asia, having united and separated them as well; many Himalayan peaks are sacred in Hinduism and Buddhism (https://en.wikipedia.org/wiki/Himalayas). An orographic map of the Himalayas is presented in Fig. 1.

Figure 1. 

Orographic map of the Himalayan region.

From a biologist’s viewpoint, the Himalayas represent a highly important biogeographical barrier between the cold and arid uplands of Central Asia and the largely tropical South and Southeast Asia. During the southwestern monsoon period, precipitation mainly occurs on the southern slopes, being greatly reduced on the northern ones. This picture is especially typical of the Central Himalayas, more or less within Kumaon, Nepal, Sikkim and Bhutan, as more to the west the aridity of Central Asia extends across the southern slopes while in the eastern parts of the system heavy monsoon rains, though declining in amount and frequency, reach beyond the main ridge as far as southeastern Tibet (Troll 1967, Miehe 2015).

The drastic climatic gradient within the Central Himalayas is of utmost importance in affecting the distribution of various organisms. Although phyto- and zoogeographical regions delimited differ to some degree, they both emphasize the role of the Himalayas as a contact zone between two major biogeographical realms, the Palaearctic and Oriental, which meet and intermesh here in various combinations. All areas lying north of the Central Himalayas obviously belong to the Palaearctic, as do the highest parts of the inhabited southern macroslope. The lower and lowest elevations of the southern macroslope are largely attributable to the Oriental, or Indo-Malayan realm. In addition, a third realm, the Sino-Himalayan biogeographical region, can be distinguished, bringing old faunal elements into the Himalayan chain. However, the border between both regions is generally neither striking nor abrupt, forming more (especially in the eastern Himalayas) or less (in their central parts) vast transition areas, numerous inversions or anomalies. In other words, the otherwise manifest rule “(sub)tropical organisms for (sub)tropical environments only” is very often violated in the Himalayas, particularly in the central parts of the system and as regards animals in general (Martens 1984, 1993, 2015). Even the pattern of vertical zonation of the tree plant cover in the region is rather conventional (Dobremez 1972) (Fig. 2).

Figure 2. 

The vegetation belts and most important plant communities in the Nepal Himalayas. The Roman numerals at the bottom indicate the floral regions of Nepal (modified, after Dobremez 1972).

The first, provisional review of the millipede fauna of the Himalayas (Golovatch and Martens 1996) was based on a fauna of approximately 200 species or subspecies. Now, the list has reached more than 270 species or subspecies belonging to 53 genera distributed in 23 families and 13 orders (Table 1). As before, several species and even genera remain unidentified. The main increase is due to the omnipresent family Paradoxosomatidae, especially as regards the fauna of Nepal. The objective of this paper is to critically list the actually known Himalayan diplopod fauna and to discuss the different faunal and evolutionary influences that made this fauna so rich and complex.

Material and methods

These results mostly rely on published records, which have grown considerably over the past two decades. The resultant checklist (Table 1) is not just a literature compilation, as it is largely based on the vast experience of the first author who has described numerous diplopod species from the Himalayas. Most of the recent advance has appeared, based on material collected by the second author and/or his collaborators during numerous, often long-term expeditions to Nepal, India and elsewhere. The trips to Nepal covered all seasons, focused on the exploration of local forest soil/litter fauna and concerned all forest biomes ranging from the terai lowlands to and beyond the timber line.

Diplopoda of the Himalayas. In addition to the taxonomic composition of the fauna, data on the vertical and geographical distribution of species in the region are also presented, largely with key references.

Fauna Altitude (m a.s.l.) Country/state and main reference(s)
Order Polyxenida 1585–2400
Family Polyxenidae 1585–2400
Genus Polyxenus Latreille, 1802–03
1. Polyxenus sp. 1585 India, Jammu & Kashmir (Silvestri 1936)
Genus Monographis Attems, 1907
2. Monographis mira (Turk, 1947) 1600–2400 Nepal1 & India, Almora (Turk 1947, Condé 1962, Golovatch and Wesener 2016)
Genus Unixenus Jones, 1944
3. Unixenus sp. 2400–4550 Nepal2 (Condé and Jacquemin-Nguyen Duy 1968, Golovatch and Martens 1996)
Order Sphaerotheriida 140–2700
Family Zephroniidae 140–2700
Genus Indosphaera Attems, 1935
4. Indosphaera feae Attems, 1935 ? India, Assam (Attems 1936, Golovatch and Wesener 2016)
Genus Kophosphaera Attems, 1935 1650–2100
5. Kophosphaera brevilamina Attems, 1936 1700 India, West Bengal & Darjeeling Distr. (Golovatch and Martens 1996, Golovatch and Wesener 2016)
6. K. devolvens Attems, 1936 1700–2050 India, Sikkim & Darjeeling Distr. (Golovatch and Martens 1996, Golovatch and Wesener 2016)
7. K. excavata (Butler, 1874) ? Nepal, Chitlang; India, Sikkim, Darjeeling Distr. & Assam (Golovatch and Martens 1996, Golovatch and Wesener 2016)
8. K. mammifera Attems, 1936 ? India, Darjeeling Distr. & Assam (Golovatch and Martens 1996, Golovatch and Wesener 2016)
9. K. martensi Wesener, 2015 2100 Nepal (Wesener 2015)
10. K. politissima Attems, 1935 1650–1870 India, Darjeeling Distr. & Nepal (Wesener 2015)
11. K. shivapuri Wesener, 2015 1700–2100 Nepal (Wesener 2015)
Genus Zephronia Gray, 1832 140–2700
12. Zephronia alticola alticola Attems, 1936 400–1700 India, Darjeeling Distr. & Assam (Attems 1936, Golovatch and Martens 1996)
13. Z. alticola bengalica Attems, 1936 ? India, West Bengal (Attems 1936, Golovatch and Martens 1996)
14. Z. debilis Attems, 1936 1700 India, Darjeeling Distr. (Attems 1936, Golovatch and Martens 1996)
15. Z. densipora Attems, 1936 ? India, Assam (Attems 1936, Golovatch and Martens 1996)
16. Z. disparipora Attems, 1936 140 India, Assam (Attems 1936, Golovatch and Martens 1996)
17. Z. hirta Attems, 1936 1700 India, Darjeeling Distr. (Attems 1936, Golovatch and Martens 1996)
18. Z. hysophila Attems, 1936 ? India, Assam (Attems 1936, Golovatch and Martens 1996)
19. Z. juvenis Attems, 1936 ? India, Assam (Attems 1936, Golovatch and Martens 1996)
20. Z. lignivora Attems, 1936 180–330 India, Assam (Attems 1936, Golovatch and Martens 1996)
21. Z. manca Attems, 1936 1000–1700 Vietnam & India, Darjeeling Distr. (Attems 1936, Golovatch and Martens 1996)
22. Z. montana (Karsch, 1881) ? “Himalaya” (Wesener 2015)
23. Z. nepalensis Wesener, 2015 1700–2600 Nepal (Wesener 2015)
24. Z. nigrinota Butler, 1872 2300–2700 India, Darjeeling Distr. (Golovatch and Martens 1996, Golovatch and Wesener 2016)
25. Z. specularis Attems, 1936 ? India, Assam (Attems 1936, Golovatch and Martens 1996)
26. Z. tigrina Butler, 1872 ? India, Darjeeling Distr. (Golovatch and Wesener 2016)
27. Z. tigrinoides Attems, 1936 170 India, Darjeeling Distr. (Attems 1936, Golovatch and Martens 1996)
28. Z. tumida Butler, 1882 ? India, Assam & Darjeeling Distr.; Myanmar (Wesener 2015)
Order Glomerida 150–3300
Family Glomeridae 150–3300
Genus Hyleoglomeris Verhoeff, 1910 150–3300
29. Hyleoglomeris crassipes Golovatch, 1987 2450–2720 Nepal (Golovatch 1987b, Golovatch and Martens 1996)
30. H. electa (Silvestri, 1917) 500–1700 India, Darjeeling Distr. (Attems 1936, Golovatch and Martens 1996)
31. H. gorkhalis Golovatch, 1987 1200 Nepal (Golovatch 1987b, Golovatch and Martens 1996)
32. H. khumbua Golovatch, 1987 3250–3300 Nepal (Golovatch 1987b, Golovatch and Martens 1996)
33. H. modesta Silvestri, 1917 150 India, Assam (Golovatch and Martens 1996)
34. H. nagarjunga Golovatch, 1987 1600–2100 Nepal (Golovatch 1987b, Golovatch and Martens 1996, Golovatch et al. 2006)
35. H. tinjurana Golovatch, 1987 2450 Nepal (Golovatch 1987b, Golovatch and Martens 1996)
36. H. venustula Silvestri, 1917 ? India, Assam (Golovatch and Martens 1996)
Order Siphonophorida
Family Siphonorhinidae 500–1700
Genus Siphonorhinus Pocock, 1894 500–1700
37. Siphonorhinus cingulatus (Attems, 1936) 500–1700 Vietnam and India, Darjeeling Distr. (Attems 1936, Golovatch and Wesener 2016)
38. S. coniceps (Attems, 1936) 1700 India, Darjeeling Distr. (Attems 1936, Golovatch and Wesener 2016)
39. S. larwoodi (Turk, 1947) 1600 India, Almora (Golovatch and Martens 1996, Golovatch and Wesener 2016)
Order Siphonocryptida
Family Siphonocryptidae 2450
Genus Hirudicryprus Enghoff & Golovatch, 1995
40. Hirudicryprus quintumelementum Korsós, Geoffroy & Mauriès, 2009 2450 Nepal (Korsós et al. 2009)
Order Platydesmida <2000
Family Andrognathidae <2000
Genus Pseudodesmus Pocock, 1887
41. ?Pseudodesmus sp. <2000 Nepal (Golovatch and Martens 1996)
Order Polyzoniida 4700–4800
Family Hirudisomatidae 4700–4800
Genus Nepalozonium Shelley, 1996
42. Nepalozonium trimaculatum Shelley, 1996 4700–4800 Nepal (Shelley 1996)
Order Chordeumatida 900–4100
Family Kashmireumatidae 2600–4100
Genus Kashmireuma Mauriès, 1982 2600–4100
43. Kashmireuma nepalensis Mauriès, 1988 3600–4100 Nepal (Mauriès 1988, Golovatch and Martens 1996)
44. K. nielseni Mauriès, 1982 2600–3500 India, Kashmir (Mauriès 1982, Golovatch and Martens 1996)
45. K. schawalleri Shear, 1987 3450–3600 Nepal (Shear 1987, Golovatch and Martens 1996)
Family Cleidogonidae 900–3900
Genus Tianella Attems, 1904 900–3900
46. Tianella ausobskyi Shear, 1987 2500–3050 Nepal (Shear 1987, Golovatch and Martens 1996)
47. T. bobanga Shear, 1979 2460–2500 Nepal (Shear 1979, 1987, Golovatch and Martens 1996)
48. T. daamsae Shear, 1987 3600–3900 Nepal (Shear 1987, Golovatch and Martens 1996)
49. T. gitanga Shear, 1987 2550 Nepal (Shear 1987, Golovatch and Martens 1996)
50. T. jaljalensis Mauriès, 1988 2350 Nepal (Mauriès 1988, Golovatch and Martens 1996)
51. T. kathmandua Mauriès, 1988 1700 Nepal (Mauriès 1988, Golovatch and Martens 1996)
52. T. lughla Shear, 1979 2950–3300 Nepal (Shear 1979, 1987, Golovatch and Martens 1996)
53. T. managa Shear, 1987 2550 Nepal (Shear 1987, Golovatch and Martens 1996)
54. T. mangsingma Mauriès, 1988 2250 Nepal (Mauriès 1988, Golovatch and Martens 1996)
55. T. martensi Shear, 1979 1150–2900 Nepal (Shear, 1979, 1987, Golovatch and Martens 1996)
56. T. smetanai Mauriès, 1988 3250 Nepal (Mauriès 1988, Golovatch and Martens 1996)
57. Tianella sp. 900–1400 India, Darjeeling Distr. (Golovatch and Martens 1996)
Family Megalotylidae 1900–4100
Genus Nepalella Shear, 1979 1900–4100
58. Nepalella deharvengi Mauriès, 1988 2900–3500 Nepal (Mauriès 1988, Golovatch and Martens 1996)
59. N. gairiensis Mauriès, 1988 3000 Nepal (Mauriès 1988, Golovatch and Martens 1996)
60. N. gunsa Shear, 1987 3600–3800 Nepal (Shear 1987, Golovatch and Martens 1996)
61. N. jaljalae Mauriès, 1988 2200 Nepal (Mauriès 1988, Golovatch and Martens 1996)
62. N. khumbua Shear, 1979 3350–3300 Nepal (Shear 1979, 1987, Golovatch and Martens 1996)
63. N. phulcokia Mauriès, 1988 2250 Nepal (Mauriès 1988, Golovatch and Martens 1996)
64. N. ringmoensis Mauriès, 1988 2750–3000 Nepal (Mauriès 1988, Golovatch and Martens 1996)
65. N. taplejunga Shear, 1987 3000–3300 Nepal (Shear 1987, Golovatch and Martens 1996)
66. N. thodunga Shear, 1979 3200 Nepal (Shear 1979, 1987, Golovatch and Martens 1996)
67. N. tragsindola Mauriès, 1988 2450–3300 Nepal (Mauriès 1988, Golovatch and Martens 1996)
68. Nepalella sp. 1900–4100 Nepal (Golovatch and Martens 1996)
Order Callipodida 650
Family Caspiopetalidae 650
Genus Bollmania Silvestri, 1896
69. Bollmania kohalana (Attems, 1936) 650 Pakistan, Punjab (Attems 1936, Golovatch and Wesener 2016)
Order Julida 1680–4800
Family Julidae 1680–4800
Genus Anaulaciulus Pocock, 1895 1900–4500
70. Anaulaciulus acaudatus Korsós, 1996 3990 India, Sikkim (Korsós 1996, Golovatch and Martens 1996)
71. A. bilineatus Korsós, 1996 3300–4300 Nepal (Korsós 1996, Golovatch and Martens 1996)
72. A. kashmirensis Korsós, 1996 3100–3200 India, Kashmir (Korsós 1996, Golovatch and Martens 1996)
73. A. nepalensis Korsós, 1996 2600–3400 Nepal (Korsós 1996, Golovatch and Martens 1996)
74. A. niger Korsós, 1996 2600–4500 Nepal (Korsós 1996, Golovatch and Martens 1996)
75. A. tibetanus Korsós, 1996 3700 China, Tibet; India, Assam (Korsós 1996, Golovatch and Martens 1996)
76. A. topali Korsós, 1996 2300 India, Kashmir (Korsós 1996, Golovatch and Martens 1996)
Genus Nepalmatoiulus Mauriès, 1983 1680–4800
77. Nepalmatoiulus appendiculatus Enghoff, 1987 1900–2100 India, Uttar Pradesh (Enghoff 1987, Golovatch and Martens 1996)
78. N. deharvengi (Mauriès, 1983) 2550–3350 Nepal (Mauriès 1983, Golovatch and Martens 1996)
79. N. dhaulagiri Enghoff, 1987 3000–3350 Nepal (Enghoff 1987, Golovatch and Martens 1996)
80. N. generalis Enghoff, 1987 3400 Nepal (Enghoff 1987, Golovatch and Martens 1996)
81. N. hyalilobus Enghoff, 1987 3600–3800 Nepal (Enghoff 1987, Golovatch and Martens 1996)
82. N. ivanloebli Enghoff, 1987 2200–4800 Nepal (Mauriès 1983, Enghoff 1987, Golovatch and Martens 1996)
83. N. juctapositus Enghoff, 1987 2800–3050 Nepal (Mauriès 1983, Golovatch and Martens 1996)
84. N. martensi Enghoff, 1987 3250–3300 Nepal (Enghoff 1987, Golovatch and Martens 1996)
85. N. mauriesi Enghoff, 1987 3600 Nepal (Enghoff 1983, Golovatch and Martens 1996)
86. N. nigrescens Enghoff, 1987 2300 Bhutan (Enghoff 1983, Golovatch and Martens 1996)
87. N. pineti Enghoff, 1987 2900 Nepal (Enghoff 1987, Golovatch and Martens 1996)
88. N. rugiflagrum Enghoff, 1987 3300 Bhutan (Enghoff 1987, Golovatch and Martens 1996)
89. N. smetanai (Mauriès, 1983) 1900–2700 Nepal (Enghoff 1983, Golovatch and Martens 1996)
90. N. sympatricus Enghoff, 1987 3000 Nepal (Enghoff 1987, Golovatch and Martens 1996)
88. N. uncus Enghoff, 1987 2550 Nepal (Enghoff 1987, Golovatch and Martens 1996)
91. N. wuermlii Enghoff, 1987 1680–2600 Bhutan (Enghoff 1987, Golovatch and Martens 1996)
92. N. zachonoides Enghoff, 1987 2450–2600 Nepal (Enghoff 1987, Golovatch and Martens 1996)
Order Spirostreptida 200–2500
Family Cambalopsidae <1000–1200
Genus Podoglyphiulus Attems, 1909 <1000–1200
93. Podoglyphiulus elegans nepalensis Mauriès, 1983 <1000 Nepal3 (Mauriès 1983, Golovatch and Martens 1996)
Genus Trachyjulus Peters, 1864
94. Trachyjulus mimus Silvestri, 1924 1200 India, Assam (Silvestri 1924, Golovatch and Martens 1996, Golovatch and Wesener 2016)
95. T. wilsonae Mauriès, 1983 <1000 Nepal (Mauriès 1983, Golovatch and Martens 1996)
Family Harpagophoridae 200–2500
Genus Dametus Attems, 1942
96. Dametus falcatus (Attems, 1936) 400–500 India, Assam (Attems 1936, Golovatch and Wesener 2016)
Genus Gonoplectus Chamberlin, 1921 200–2500
97. Gonoplectus alius Demange, 1961 ? India, Assam (Demange 1961, Golovatch and Martens 1996)
98. G. bhutanensis Demange, 1988 350–450 Bhutan (Demange 1988, Golovatch and Martens 1996)
99. G. broelemanni Demange, 1961 1800–2300 Nepal (Demange 1961, Golovatch and Martens 1996)
100. G. corniger (Attems, 1936) ? India, Assam (Attems 1936, Golovatch and Martens 1996)
101. G. gracilis (Attems, 1936) 1200 India, Darjeeling Distr. (Attems 1936, Golovatch and Martens 1996)
102. G. hyatti Demange, 1961 1200 Nepal (Demange 1961, Golovatch and Martens 1996)
103. G. malayus malayus (Carl, 1909) 200–2500 Indonesia, Java; Nepal, Bhutan & India, Madhya Pradesh, Uttar Pradesh, Himachal Pradesh, West Bengal (Golovatch and Martens 1996, Golovatch and Wesener 2016)
104. G. malayus lindbergi (Carl, 1909) 350 Bhutan & India, Darjeeling Distr. (Golovatch and Martens 1996, Golovatch and Wesener 2016)
105. G. probus (Attems, 1936) 1000 India, Darjeeling Distr. (Attems 1936, Golovatch and Martens 1996, Golovatch and Wesener 2016)
106. G. remyi Demange, 1961 ? India, Assam (Demange 1961, Golovatch and Martens 1996, Golovatch and Wesener 2016)
107. G. repertus (Attems, 1936) 900 India, Darjeeling Distr. (Attems 1936, Golovatch and Wesener 2016)
108. G. sulcatus (Attems, 1936) 2400 India, Darjeeling Distr. (Attems 1936, Golovatch and Wesener 2016)
Order Spirobolida <1000–1800
Family Pachybolidae
Genus Trigoniulus Pocock, 1894 <1000
109. Trigoniulus corallinus (Gervais, 1847) <1000 Pantropical, in India nearly throughout, including Assam (Golovatch and Wesener 2016)
Family Pseudospirobolellidae
Genus Physobolus Attems, 1936
110. Physobolus olivaceus Attems, 1936 1800 India, Darjeeling Distr. (Attems 1936, Golovatch and Wesener 2016)
Order Polydesmida 150–4500
Family Cryptodesmidae
Genus Trichopeltis Pocock, 1894 350–1000
111. Trichopeltis watsoni Pocock, 1895 350–1000 Bangladesh, Myanmar, Bhutan and India, Darjeeling Distr., West Bengal, Assam & near Kolkata (Attems 1936, Golovatch and Martens 1996, Golovatch and Wesener 2016)
Family Haplodesmidae 150–1750
Genus Koponenius Golovatch & VandenSpiegel, 2014 150–1750
112. Koponenius biramus Golovatch & VandenSpiegel, 2014 1750 Nepal (Golovatch and VandenSpiegel 2014)
113. K. schawalleri Golovatch & VandenSpiegel, 2016 150 Nepal (Golovatch and VandenSpiegel 2016)
114. K. unicornis Golovatch & VandenSpiegel, 2014 880 India, Darjeeling Distr. (Golovatch and VandenSpiegel 2014, Golovatch and Wesener 2016)
Family Opisotretidae 1100–2440
Genus Martensodesmus Golovatch, 1987 1100–2440
115. Martensodesmus bicuspidatus Golovatch, 1988 1650–2000 Bhutan (Golovatch 1988a, Golovatch and Martens 1996, Golovatch et al. 2013)
116. M. excornis Golovatch, 1988 2440 Bhutan (Golovatch 1988a, Golovatch and Martens 1996, Golovatch et al. 2013)
117. M. himalayensis Golovatch, 1987 1100–1300 Nepal (Golovatch 1987a, Golovatch and Martens 1996, Golovatch et al. 2013)
118. M. nagarjungicus Golovatch, 1987 1900–2100 Nepal (Golovatch 1987a, Golovatch and Martens 1996, Golovatch et al. 2013)
119. M. sherpa Golovatch, 1987 1200 Nepal (Golovatch 1987a, Golovatch and Martens 1996, Golovatch et al. 2013)
120. Martensodesmus sp. 1300–2150 Nepal, Bhutan (Golovatch and Martens 1996)
Family Paradoxosomatidae 150–4500
Genus Anoplodesmus Carl, 1932 1000–3600
121. Anoplodesmus affinis (Golovatch, 1990) 2475–2700 Nepal (Golovatch 1990a, Golovatch and Martens 1996)
122. A. cylindricus (Carl, 1935) 1650–2850 Nepal & India, Darjeeling Distr. (Carl 1935, Golovatch 1984, Golovatch and Martens 1996, Golovatch and Wesener 2016)
123. A. elongissimus (Golovatch, 1984) 1000 India, Darjeeling Distr. (Golovatch 1984, Golovatch and Martens 1996, Golovatch and Wesener 2016)
124. A. magnus Golovatch, 2015 2700 Nepal (Golovatch 2015a)
125. A. martensi (Golovatch, 1990) 2250–3600 Nepal (Golovatch 1990a, 2014b, 2016a, Golovatch and Martens 1996)
126. A. schawalleri (Golovatch, 1990) 2050–2150 Nepal (Golovatch 1990a, Golovatch and Martens 1996)
127. A. similis (Golovatch, 1990) 2300–3000 Nepal (Golovatch 1990a, Golovatch and Martens 1996)
128. A. spinosus Golovatch, 2016 2500 Nepal (Golovatch 2016a)
129. A. subcylindricus (Carl, 1932) ? S India & Nepal (Nguyen 20104, Golovatch and Wesener 2016)
Genus Beronodesmoides Golovatch, 2015 1650–4250
130. Beronodesmoides anteriporus Golovatch, 2015 1650–3350 Nepal (Golovatch 2015a, 2016c)
131. B. bifidus Golovatch, 2015 3100 Nepal (Golovatch 2015a)
132. B. lobatus Golovatch, 2015 4000–4250 Nepal (Golovatch 2015a, Golovatch et al. 2016)
133. B. longifemoratus Golovatch, 2015 2700–2800 Nepal (Golovatch 2016a)
134. B. martensi Golovatch, 2016 2700 Nepal (Golovatch 2016c)
135. B. montigena Golovatch, 2016 3550 Nepal (Golovatch 2016c)
136. B. typicus Golovatch, 2016 3400 Nepal (Golovatch 2016c)
Genus Beronodesmus Golovatch, 2014 1650–4500
137. Beronodesmus curtispinus Golovatch, 2015 4500 Nepal (Golovatch 2015a)
138. B. distospinosus Golovatch, 2015 1650–3080 Nepal (Golovatch 2015a, 2016c)
139. B. gorkhalis Golovatch, 2015 3050–3600 Nepal (Golovatch 2015a, Golovatch et al. 2016)
140. B. latispinosus Golovatch, 2015 1900–3500 Nepal (Golovatch 2015a, 2016c, Golovatch et al. 2016)
141. B. longispinus Golovatch, 2015 2550–4270 Nepal (Golovatch 2015a, 2016c, Golovatch et al. 2016)
142. B. martensi Golovatch et al., 2016 2650 Nepal (Golovatch et al. 2016)
143. B. minutus Golovatch, 2015 3300–3500 Nepal (Golovatch 2015a)
144. B. pallidus Golovatch, 2014 3800–4100 Nepal (Golovatch 2014b)
145. B. serratus Golovatch et al., 2016 3300–3500 Nepal (Golovatch et al. 2016)
146. B. simplex Golovatch, 2016 2100 Nepal (Golovatch 2016c, Golovatch et al. 2016)
147. B. sinuatospinus Golovatch, 2015 2150–2250 Nepal (Golovatch 2015a, 2016c)
Genus Delarthrum Attems, 1936 600–4100
148. Delarthrum aberrans (Golovatch, 1996) 1000–2600 Nepal (Golovatch 1996, 2014b, 2016a, Golovatch and Martens 1996)
149. D. affine (Golovatch, 1994) 1400 Nepal (Golovatch 1994a, 2014b, Golovatch and Martens 1996)
150. D. alatum (Golovatch, 1996) 1900–2100 Nepal (Golovatch 1996, 2014b, Golovatch and Martens 1996)
151. D. andreevi Golovatch, 2014 1800 Nepal (Golovatch 2014b)
152. D. arunense (Golovatch, 1994) 1850–2150 Nepal (Golovatch 1994a, 2014b, 2016a, Golovatch and Martens 1996)
153. D. beroni Golovatch, 2014 600–1000 Nepal (Golovatch 2014b)
154. D. bifidum (Golovatch, 1996) 2550–2650 Nepal (Golovatch 1996, 2014b, Golovatch and Martens 1996)
155. D. chulingense (Golovatch, 1994) 3000–3700 Nepal (Golovatch 1994a, 2014b, Golovatch and Martens 1996)
156. D. communicans (Golovatch, 1992) 2650 Nepal (Golovatch 1992, 2014b, Golovatch and Martens 1996)
157. D. curtisoma Golovatch, 2015 2050–2150 Nepal (Golovatch 2015a)
158. D. curtum Golovatch, 2014 600–1000 Nepal (Golovatch 2014b)
159. D. densesetosum Golovatch, 2015 2400 Nepal (Golovatch 2015a)
160. D. elegans (Golovatch, 1992) 1350 Nepal (Golovatch 1992, 2014b, Golovatch and Martens 1996)
161. D. extremum (Golovatch, 1996) 2450 Nepal (Golovatch 1996, 2014b, Golovatch and Martens 1996)
162. D. facile (Golovatch, 1996) 2200–2400 Nepal (Golovatch 1996, 2014b, Golovatch and Martens 1996)
163. D. fechteri (Golovatch, 1990) 2330–3150 Nepal (Golovatch 1990a, 2014b, Golovatch and Martens 1996)
164. D. foveatum (Golovatch, 1996) 1800–2000 Nepal (Golovatch 1992, 2014b, Golovatch and Martens 1996)
165. D. furcatum (Golovatch, 1996) 600–2000 Nepal (Golovatch 1996, 2014b, 2016c, Golovatch and Martens 1996)
166. D. gracile Golovatch, 2015 1750 Nepal (Golovatch 2015a)
167. D. granulosum (Golovatch, 1994) 2000 Nepal (Golovatch 1994a, 2014b, 2016c, Golovatch and Martens 1996)
168. D. heterotergale Golovatch, 2014 600–1000 Nepal (Golovatch 2014)
169. D. hingstoni (Carl, 1935) 3400 China, Tibet (Carl 1935, Golovatch and Martens 1996)
170. D. hirsutum (Golovatch, 1994) 2400–4100 Nepal (Golovatch 1994a, 2014b, 2015a, Golovatch and Martens 1996)
171. D. intermedium (Golovatch, 1994) 1000–1100 Nepal (Golovatch 1994a, 2014b, Golovatch and Martens 1996)
172. D. invocatum (Golovatch, 1996) 2600–2800 Nepal (Golovatch 1996, 2014b, Golovatch and Martens 1996)
173. D. kuznetsovi (Golovatch, 1994) 3000 Nepal (Golovatch 1994a, 2014b, Golovatch and Martens 1996)
174. D. longisetum (Golovatch, 1994) 1400–1600 Nepal (Golovatch 1994a, 2014b, 2016c, Golovatch and Martens 1996)
175. D. longispinum (Golovatch, 1996) 2150–2250 Nepal (Golovatch 1996, 2016c, Golovatch and Martens 1996)
176. D. modestum (Golovatch, 1996) 3450–3600 Nepal (Golovatch 1996, 2014b, Golovatch and Martens 1996)
177. D. nyakense (Golovatch, 1992) 2270–2400 Nepal (Golovatch 1992, 2014b, Golovatch and Martens 1996)
178. D. obscurum Attems, 1936 ca 2800 N Pakistan, Punjab (Attems 1936, Golovatch and Wesener 2016)
179. D. philosophicum (Golovatch, 1994) 1650–2450 Nepal (Golovatch 1994a, 2014b, Golovatch and Martens 1996)
180. D. planifemur Golovatch, 2015 2200 Nepal (Golovatch 2015a)
181. D. prolixum (Golovatch, 1996) 2550–2650 Nepal (Golovatch 1996, 2014b, Golovatch and Martens 1996)
182. D. pumilum (Attems, 1944) ? India, Uttar Pradesh (Attems 1944, Golovatch and Wesener 2016)
183. D. quadridentatum Golovatch, 2016 2600–2800 Nepal (Golovatch 2016b)
184. D. schawalleri (Golovatch, 1992) 1000–2150 Nepal (Golovatch 1992, 1994a, 2014, Golovatch and Martens 1996)
185. D. setosum Golovatch, 2014 ? Nepal (Golovatch 2014b)
186. D. silvestre (Golovatch, 1994) 2000–3400 Nepal (Golovatch 1994a, 2014b, 2016a, Golovatch and Martens 1996)
187. D. simile (Golovatch, 1992) 2300–2700 Nepal (Golovatch 1992, 2014b, Golovatch and Martens 1996
188. D. simplex (Golovatch, 1996) 1650 Nepal (Golovatch 1996, 2014b, Golovatch and Martens 1996)
189. D. simulans (Carl, 1935) 3700 Nepal & China, Tibet (Carl 1935, Golovatch and Martens 1996)
190. D. spectabile (Golovatch, 1994) 2650 Nepal (Golovatch 1994a, 2014b, 2016c, Golovatch and Martens 1996)
191. D. spiniger (Attems, 1936) 1000–2200 India, West Bengal & Darjeeling Distr. (Attems 1936, Golovatch 1984, Golovatch and Wesener 2016)
192. D. spinigerum (Golovatch, 1992) 600–1400 Nepal (Golovatch 1992, 2014b)
193. D. splendens (Golovatch, 1992) 1650–2150 Nepal (Golovatch 1992, 1994a, 2014, Golovatch and Martens 1996)
194. D. subalatum (Golovatch, 1996) 2600–2800 Nepal (Golovatch 1996, 2014b, Golovatch and Martens 1996)
195. D. subsimulans (Golovatch, 1996) 3100–3300 Nepal (Golovatch 1996, 2014b, Golovatch and Martens 1996)
196. D. tenuitergale Golovatch, 2014 3250 Nepal (Golovatch 2014)
197. D. tergale (Golovatch, 1994) 2650 Nepal (Golovatch 1994a, 2014b, Golovatch and Martens 1996)
198. D. tuberculatum (Golovatch, 1994) 3000–3300 Nepal (Golovatch 1992, 1994a, 2014b, Golovatch and Martens 1996)
199. D. typicum Golovatch, 2014 3100 Nepal (Golovatch 2014)
200. D. uncum (Golovatch, 1996) 2100–3420 Nepal (Golovatch 1996, 2014, 2015a, Golovatch and Martens 1996)
201. D. unicolor (Attems, 1936) 1200–1700 India, Assam & Darjeeling Distr. (Attems 1936, Golovatch and Wesener 2016)
Genus Hirtodrepanum Golovatch, 1994
202. Hirtodrepanum latigonopum Golovatch, 1994 2100–2600 Nepal (Golovatch 1994b, 2015a, Golovatch et al. 2016)
Genus Kaschmiriosoma Schubart, 1935 1000–3300
203. Kaschmiriosoma contortipes Schubart, 1935 2000–3300 N Pakistan & India, Jammu & Kashmir (Schubart 1935, Silvestri 1936, Golovatch 1983, Golovatch and Martens 1996, Jeekel 2003, Shelley 2014)
204. K. nulla (Attems, 1936) 1000 India, Himachal Pradesh (Attems 1936, Golovatch and Martens 1996, Jeekel 2003, Golovatch and Wesener 2016)
205. K. pleuropterum (Attems, 1936) 2800 N Pakistan, Punjab (Attems 1936, Golovatch and Martens 1996, Jeekel 2003, Golovatch and Wesener 2016)
Genus Kronopolites Attems, 1914
206. Kronopolites coriaceus Golovatch, 2015 2000 Nepal (Golovatch 2015a)
207. K. occidentalis Golovatch, 1983 1500 India, Jammu & Kashmir (Golovatch 1983, Golovatch and Martens 1996, Golovatch and Wesener 2016)
Genus Orthomorpha Bollman, 1893
208. Orthomorpha coarctata (de Saussure, 1860) 600–650 Nepal & India, pantropical anthropochore (Golovatch and Martens 1996, Golovatch and Wesener 2016)
209. “O.almorensis Turk, 1947 1600 India, Almora (Turk 1947, Golovatch and Martens 1996, Golovatch and Wesener 2016)
Genus Oxidus Cook, 1911
210. Oxidus gracilis (C.L. Koch, 1847) 570–1200 Nepal & India, subcosmopolitan anthropochore (Golovatch and Martens 1996, Golovatch and Wesener 2016)
Genus Pocockina Jeekel, 1965
211. Pocockina schawalleri Golovatch, 2016 150 Nepal (Golovatch 2016a)
Genus Streptogonopus Attems, 1914
212. Streptogonopus phipsoni (Pocock, 1892) ≤2700 Pakistan, Bangladesh, Nepal & India, West Bengal (Golovatch 2015a, Golovatch and Wesener 2016)
Genus Substrongylosoma Golovatch, 1984 1000–2200
213. Substrongylosoma bifurcatum Golovatch, 2016 2000 Nepal (Golovatch 2016a)
214. S. distinctum Golovatch, 1984 1200–1500 India, Darjeeling Distr. (Golovatch 1984, Golovatch and Martens 1996, Golovatch and Wesener 2016)
215. S. falcatum Golovatch, 1984 1000–1400 India, Darjeeling Distr. (Golovatch 1984, Golovatch and Martens 1996, Golovatch and Wesener 2016)
216. S. exiguum Golovatch, 2016 1900 Nepal (Golovatch 2016a)
217. S. montigena (Carl, 1935) 1200–2200 India, Darjeeling Distr. (Carl 1935, Golovatch 1984, Golovatch and Martens 1996, Golovatch and Wesener 2016)
218. S. schawalleri Golovatch, 1993 1620–2000 Nepal (Golovatch 1993, Golovatch and Martens 1996)
Genus Sundanina Attems, 1914
219. “Sundaninaseptentrionalis Turk, 1947 ca 1700 India, Almora (Turk 1947, Golovatch and Wesener 2016)
Genus Topalosoma Golovatch, 1984 900
220. Topalosoma setiferum Golovatch, 1984 900 India, Darjeeling Distr. (Golovatch 1984, Golovatch and Martens 1996, Golovatch and Wesener 2016)
Genus Trogodesmus Pocock, 1895
221. Trogodesmus uncinatus (Attems, 1936) ? India, Assam (Attems 1936, Golovatch and Wesener 20165)
Genus Touranella Attems, 1937 2300–2800
222. Touranella himalayaensis Golovatch, 1994 2300–2700 Nepal (Golovatch 1994b, Golovatch and Martens 1996)
223. T. pilosa Golovatch, 2016 2600–2800 Nepal (Golovatch 2016b)
Family Polydesmidae 350–4250
Genus Bhutanodesmus Golovatch, 1988
224. Bhutanodesmus velatus Golovatch, 1988 350–450 Bhutan (Golovatch 1988, Golovatch and Martens 1996)
Genus Epanerchodus Attems, 1901 2300–4250
225. Epanerchodus buddis (Golovatch, 1986) 3300–3400 Nepal (Golovatch 1986, Golovatch and Martens 1996, Golovatch et al. 2011)
226. E. occultus (Golovatch, 1986) 2300–2800 Nepal (Golovatch 1986, Golovatch and Martens 1996, Golovatch et al. 2011)
227. E. sacer (Golovatch, 1987) 3300–3400 Nepal (Golovatch 1987a, Golovatch and Martens 1996, Golovatch et al. 2011)
228. E. theocraticus (Golovatch, 1990) 2600–2800 Nepal (Golovatch 1990b, Golovatch and Martens 1996, Golovatch et al. 2011)
229. E. theosophicus (Golovatch, 1986) 3200 Nepal (Golovatch 1986, Golovatch et al. 2011, Golovatch and Martens 1996)
230. Epanerchodus sp. 3450–4250 Nepal & Bhutan (Golovatch and Martens 1996)
Genus Glenniea Turk, 1945 350–2800
231. Glenniea bhotiaensis Golovatch, 1988 350–450 Bhutan (Golovatch 1988a, Golovatch and Martens 1996)
232. G. indica Turk, 1945 2800 India, Himachal Pradesh (Turk 1945a, 1945b, Golovatch 1988a, Golovatch and Martens 1996, Golovatch and Wesener 2016)
233. G. martensi (Golovatch, 1987) 1200 Nepal (Golovatch 1987b, 1988a, Golovatch and Martens 1996)
234. G. minuscula Golovatch, 1988 1900–2300 Bhutan (Golovatch 1988a, Golovatch and Martens 1996)
235. G. perarmata Golovatch, 1988 1680 Bhutan (Golovatch 1988a, Golovatch and Martens 1996)
Genus Himalodesmus Golovatch, 1986 1000–3400
236. Himalodesmus audax Golovatch, 1986 2650 Nepal (Golovatch 1986, Golovatch and Martens 1996)
237. H. benefactor Golovatch, 1987 2600–3400 Nepal (Golovatch 1987a, Golovatch and Martens 1996)
238. H. faustus Golovatch, 1987 1000–1750 Nepal (Golovatch 1987a, Golovatch and Martens 1996)
239. H. parvus Golovatch, 1987 2200 Nepal (Golovatch 1987a, Golovatch and Martens 1996)
240. H. prosperus Golovatch, 1990 2600–2800 Nepal (Golovatch 1990b, Golovatch and Martens 1996)
241. H. pulcher Golovatch, 1987 2450 Nepal (Golovatch 1987a, Golovatch and Martens 1996)
242. H. pygmaeus Golovatch, 1986 3300–3400 Nepal (Golovatch 1986, Golovatch and Martens 1996)
243. H. vigens Golovatch, 1987 2150–2250 Nepal (Golovatch 1987a, Golovatch and Martens 1996)
Genus Typhlopygmaeosoma Turk, 1972
244. Typhlopygmaeosoma hazeltonae Turk, 1972 1850 India, Himachal Pradesh (Turk 1972, Shear 1986, Golovatch 1988b, Golovatch et al. 2014)
Family Trichopolydesmidae 450–4500
Genus Assamodesmus Manfredi, 1955
245. Assamodesmus lindbergi Manfredi, 1954 ? India, Assam (Manfredi 1954, Golovatch 1988b, Golovatch and Martens 1996, Golovatch et al. 2014)
Genus Hingstonia Carl, 1935 2000–4500
246. Hingstonia beatae Golovatch, 1990 2400–3500 Nepal (Golovatch 1990b, Golovatch and Martens 1996, Golovatch et al. 2014)
247. H. dorjulana Golovatch, 1988 2450–3100 Bhutan (Golovatch 1988a, Golovatch and Martens 1996, Golovatch et al. 2014)
248. H. eremita Carl, 1935 2000 Nepal (Carl 1935, Golovatch 1986, Golovatch and Martens 1996, Golovatch et al. 2014)
249. H. falcata Golovatch, 1986 2650 Nepal (Golovatch 1986, Golovatch and Martens 1996, Golovatch et al. 2014)
250. H. fittkaui Golovatch, 1990 3350–3450 Nepal (Golovatch 1990b, Golovatch and Martens 1996, Golovatch et al. 2014)
251. H. gogonana Golovatch, 1988 3650–4000 Bhutan (Golovatch 1988a, Golovatch and Martens 1996, Golovatch et al. 2014)
252. H. pahakholana Golovatch, 1990 2600–2800 Nepal (Golovatch 1990b, Golovatch and Martens 1996, Golovatch et al. 2014)
253. H. pelelana Golovatch, 1988 3300–3400 Bhutan (Golovatch 1988a, Golovatch and Martens 1996, Golovatch et al. 2014)
254. H. perarmata Golovatch, 1986 3150 Nepal (Golovatch 1986, Golovatch and Martens 1996, Golovatch et al. 2014)
255. H. serrata Golovatch, 1987 3400–3600 Nepal (Golovatch 1987a, Golovatch and Martens 1996, Golovatch et al. 2014)
256. H. sympatrica Golovatch, 1990 3550–3650 Nepal (Golovatch 1990b, Golovatch and Martens 1996, Golovatch et al. 2014)
257. H. variata Golovatch, 1987 2600–4500 Nepal (Golovatch 1987a, 1990b, Golovatch and Martens 1996, Golovatch et al. 2014)
258. H. yeti Golovatch, 1988 1600–2600 Bhutan (Golovatch 1988a, Golovatch et al. 2014)
259. Hingstonia sp. 2200–3900 Nepal (Golovatch and Martens 1996)
Genus Magidesmus Golovatch, 1988 3100–3400
260. Magidesmus affinis Golovatch, 1988 3300–3400 Bhutan (Golovatch 1988a, Golovatch et al. 2014)
261. M. bhutanensis Golovatch, 1988 3100 Bhutan (Golovatch 1988a, Golovatch et al. 2014)
Genus Pseudosphaeroparia Carl, 1932
262. Pseudosphaeroparia cavernicola Turk, 1945 2800 India, Uttar Pradesh (Turk 1945a, 1945b, Golovatch and Martens 1996, Golovatch et al. 2014)
Genus Sholaphilus Carl, 1932 1100–2200
263. Sholaphilus asceticus Golovatch, 1986 1300–1650 Nepal (Golovatch 1986, Golovatch and Martens 1996, Golovatch et al. 2014)
264. S. dalai Golovatch, 1986 2400 Nepal (Golovatch 1986, Golovatch and Martens 1996, Golovatch et al. 2014)
265. S. gompa Golovatch, 1990 2000–2100 Nepal (Golovatch 1990b, Golovatch and Martens 1996, Golovatch et al. 2014)
266. S. lama Golovatch, 1986 1800–2000 Nepal (Golovatch 1986, Golovatch and Martens 1996, Golovatch et al. 2014)
267. S. martensi Golovatch, 1986 1100–1850 Nepal (Golovatch 1986, Golovatch and Martens 1996, Golovatch et al. 2014)
268. S. monachus Golovatch, 1990 2050–2150 Nepal (Golovatch 1990b, Golovatch and Martens 1996, Golovatch et al. 2014)
Genus Topalodesmus Golovatch, 1988
269. Topalodesmus communis Golovatch, 1988 2000–2200 India, Darjeeling Distr. (Golovatch 1988b, Golovatch and Martens 1996, Golovatch et al. 2014)
Family Pyrgodesmidae 450–1200
270–275? Several genera and species (including at least 2 species of Cryptocorypha Attems, 1907) 450–1200 Nepal (Golovatch and Martens 1996)

Results

Species of Diplopoda

Species concepts are only little addressed in diplopod taxonomy. To think about species limits and species definitions is not at all trivial; in nearly every case, a morphological species concept is used with the background idea that these entities, defined by external characters, fit well to the Biological Species Concept. In practical alpha-taxonomy it circumscribes reproductively isolated groups of specimens. Diplopod taxonomists largely base their identifications on adult male samples. Differences in male genitalic structure usually provide the basic characters that allow us to safely determine millipede species. In most cases this raises no problems. In the Himalayas, however, we have to tackle with numerous populations in a wide array of forest habitats found at various altitudes and in remote and secluded valleys. Hardly surprisingly, the Himalayas do support quite a number of examples of species swarms among Diplopoda as well. Species delimitation may then cause problems like those described by Martens (1978) for the polymorphous biantid harvestman Biantes pernepalicus Martens, 1978. Such situations also resemble the few known cases of insular species swarms in millipedes of Macaronesia (e.g. Enghoff 1992).

The following examples can be given and easily added to the roster of similar observations that Martens (2015) made or compiled for Himalayan mammals, birds, arachnids, insects and several other animal groups. Such a distribution pattern can be termed fanned (see below) and is also found in the endemic Himalayan diplopod genera Beronodesmus and Beronodesmoides containing 11 and seven species, respectively (Golovatch 2016c, Golovatch et al. 2016). The main species-specific characters are in minor details of gonopodal structure, in particular, the shapes of the various outgrowths (Fig. 3). Vicariance speciation must have taken place in situ, with several of the congeners forming pairs or trios that can occur sympatrically or even syntopically and thus implying a series of secondary dispersal events.

Figure 3. 

Gonopodal structural variations between several species of Beronodesmus: B. martensi Golovatch et al., 2016 (1–3), B. serratus Golovatch et al., 2016 (4, 5), B. simplex Golovatch, 2016 (6, 7), B. distospinosus Golovatch, 2015 (8, 9), B. latispinosus Golovatch, 2015 (10, 11), B. sinuatospinus Golovatch, 2015 (12, 13) and B. gorkhalis Golovatch, 2015 (14). Scale bars: 1.0 mm (4–5, 14), 0.5 mm (1–3, 8–13) or 0.4 mm (6, 7). After Golovatch (2015a, 2016c) and Golovatch et al. (2016).

Zoogeographical patterns and origins

Ecology and dispersal abilities

The vast majority of Himalayan diplopod species are highly localized in distribution, both geographical and altitudinal. There are only few relatively widespread species like Zephronia manca, Siphonorhinus cingulatus (both recorded from Vietnam & Darjeeling District, India) or Trichopeltis watsoni (Bangladesh, Myanmar, Bhutan and India, Darjeeling Distr., West Bengal, Assam & near Kolkata). Despite extended collection acitivities over most parts of Nepal during several decades, most millipedes in the Himalayas remain known from only a single or very few localities. This particularly concerns the best-explored fauna of Nepal, Central Himalayas (ca 160 spp.), including species of the dominant family Paradoxosomatidae (82 spp., or >50%).

In contrast, most genera occur through a range of altitudes (Fig. 4), but are more or less restricted to forest habitats. The alpine (= nival) zone of the Central Himalayas which lies above closed forests appears to only marginally be populated by Diplopoda (Table 1, Figs 2, 4), the bulk of the fauna being confined to the tropical and subtropical forest belts. The highest records belong to Nepalozonium trimaculatum and Nepalmatoiulus ivanloebli, both found at 4800 m a.s.l. This pattern conforms to general knowledge that millipedes are basically a class of forest-dwelling terrestrial arthropods both trophecologically and historically largely associated with woodlands and ranging from nemoral (= broadleaved forest) and coniferous forest in temperate regions in the north to rainforest tropical areas in the south (Golovatch 1997a, 1997b). Such a background is accepted and it serves as the basis for faunogenetic reconstructions using phyto- and palaeogeographical evidence.

Figure 4. 

Vertical distribution of several genera of Diplopoda in the Himalayas (modified, after Golovatch and Martens 1996).

According to Martens (1993, 2015), broad vertical distribution belts appear to be exceptional in animals at least in the Central Himalayas, in contrast to fanned ones which are especially characteristic of species-rich groups, both vertebrates and invertebrates. Yet the vertical width of a distributional belt varies very considerably between taxa. Diplopods in their natural habitats seem to be particularly restricted to their forest habitats, with little capacity for enlarging their areas, both vertically and horizontally. There is hardly any other larger land arthropod group that shows a similarly strong specialization and relation to a habitat. In many cases, congeners occupy only limited vertical belts and such patterns are the result of multi-layered speciation processes that occurred in secluded valleys and mountain massifs. There are obvious altitudinal adaptations resulting from the interaction between Pliocene/Pleistocene climate oscillations and speciation processes. Whilst species of different clades occupy narrow vertical belts, close congeners, i.e. sister species, are mostly horizontally allopatric because of their allopatric vicariance speciation. Sympatry or syntopy are quite common among congeners (see above for Beronodesmus, Fig. 3), this alone implying a series of secondary dispersal events. On the other hand, all 13 species of Nepalmatoiulus known from Nepal and Bhutan are nearly exclusively allopatric in distribution (Enghoff 1987).

Narrow belts seem to be rare, when each individual species has been recorded from a single or very few localities, but even the whole species set combined remains restricted to a narrow altitudinal range. Much more common, rather usual are the situations when the vertical and horizontal distribution of a species is highly restricted, but that of the corresponding species-group or genus is very considerable (Table 1, Fig. 4).

Among the Diplopoda of the Himalayas, most if not all of the rather to highly species-rich genera show fanned vertical distribution patterns. Such are all genera at least in the orders Sphaerotheriida, Glomerida, Chordeumatida, Julida and Spirostreptida, as well as most in the order Polydesmida. Their origins seem to be very different, but profound in situ speciation is their general characteristic. No evident narrow belts seem to be distinguishable in the combined vertical distribution of millipede congeners in the Himalayas (Table 1, Figs 2, 4). Even within Nepal, a gradual east-west decline in diplopod diversity is clearly observed towards the country’s central regions, apparently following the climate aridity gradient and increased absence of humid forest.

As noted above, zoogeographically the Himalayas are traditionally viewed as a vast, yet clear-cut transitional zone between the Palaearctic and Oriental (= Indo-Malayan) realms. Martens (2015, p. 212) writes that “The renowned species diversity of the Himalayan fauna results from the area’s position between tropical SE Asia to the south and southeast, temperate High Asia to the north and dry Central Asia to the northwest, in addition to its proximity to endemic-rich SW China, which harbours many Tertiary relicts. Elements from all these areas contributed to and influenced the present faunal composition, creating one of the world’s 25 biodiversity hotspots, Indo-Burma and the adjacent South Central China.” He discriminated a predominantly immigrant fauna with five main sources: SW Chinese Himalayan from the northeastern Himalayan Arc; Indochinese Himalayan from the southeast; tropical Indian from the south; central Asian from the north via Tibet; and W Asian Himalayan from mountain ranges westwards to the Mediterranean. This fauna is mainly composed of species with good dispersal abilities such as bats, birds, butterflies, dragonflies and many other groups. Since millipedes are poor dispersers and only relatively few species are anthropochores, we refrain from enlisting any as belonging to this category, even though a few widespread Oriental species do reach the Himalayas from the east and/or southeast (e.g. the above Zephronia manca, Siphonorhinus cingulatus, Streptogonopus phipsoni, Gonoplectus malayus malayus or Trichopeltis watsoni). Human introductions cannot be excluded in such cases, these being especially apparent as regards the few unquestioned anthropochores like Trigoniulus corallinus, Orthomorpha coarctata or Oxidus gracilis.

In addition, a Sino-Himalayan fauna (or even a Sino-Himalayan region) is distinguished, which is characterized by remarkable diversity, partially of Indo-Malayan origins and partially Palaearctic in nature, but with marked peculiarities. Holt et al. (2012), based on the modern distributions and phylogenies of amphibians, birds and non-marine mammals, defined 20 distinct zoogeographic regions grouped into 11 realms, including a Sino-Japanese realm which appears to show closer phylogenetic affinities to the Palaearctic than to the Oriental realm. It includes Japan, Tibet and nearly all of China. Eventually, that paper represents one of the most consistent, but no less unsuccessful attempts at uncritically combining the landscape-typological (= zonal) and faunogenetic approaches to biogeography which, however, must be clearly separated at least as regards the biotas of older biomes (e.g. Chernov 1975, Golovatch 2015b). To reiterate Chernov (1975) briefly, from the “viewpoint” of a biome or more local landscape it only matters whether the constituent species properly function as its biotic elements, regardless of their origins. In contrast, from a faunogenetic point of view, the more ancient the biome or landscape, the more ancient its biota and the more complex its history. Thus both approaches must be clearly distinguished, especially as regards the relatively younger zonal biomes like tundra or taiga (= boreal forest) from the particularly ancient, rather regional than zonal, subtropical and tropical ones.

In terms of its faunal composition, the Sino-Himalayan region represents a mixed zone of elements derived from both the Palaearctic and Oriental realms, but it includes moreover a wealth of endemics with surprisingly small and often relict distributions (Martens 2015). It is within this category that many of the Himalayan Diplopoda seem best to place. Perhaps the most conspicuous example of such a pattern is represented by the definitely relict order Siphonocryptida which globally contains only two genera and seven species. Thus, the genus Siphonocryptus Pocock, 1894, comprises three species: one in Sumatra, Indonesia, the other two in continental Western Malaysia. In contrast, the distribution pattern of Hirudicryptus Enghoff & Golovatch, 1995 is trans-Palaearctic (Fig. 5). The type species occurs only on Madeira and the Canaries, where it is largely confined to the relict, subtropical, laurisilva biome. One species each is endemic to Taiwan, to Nepal and to the NW Caucasus (Golovatch et al. 2015, Zuev 2017). It may well be that the distribution pattern under consideration dates back at least to the Oligocene times of the so-called “Warm Earth” to have highly probable explanations rooted in palaeobotanical evidence. These imply a gradual shrinkage and disruption ever since of the previously dominating and continuous subtropical biome (Golovatch 1997a, 1997b, Zherikhin 2003). Being so vastly disjunct, the present-day distribution of Siphonocryptida is best accounted for by extinction events (Shelley and Golovatch 2011).

Figure 5. 

Distribution of the genus Hirudicyptus (Siphonocryptidae, Siphonocryptida). After Golovatch et al. (2015).

Genus- and species-level relations

Further possible examples of the Sino-Himalayan pattern seem to be represented by Glenniea (Polydesmidae, a largely Holarctic family), with five species from the Himalayas of Nepal and Bhutan, and three in S China (Golovatch 2015b), Nepalella (Megalotylidae, the genus being the westernmost in that temperate to tropical Asian family), with 23 species or subspecies from Nepal, S China and SE Asia (Minelli 2015, Table 1), Hirtodrepanum (Paradoxosomatidae, a subcosmopolitan family absent only from N America) with one species each in Nepal and S China (Golovatch 2014a), and Martensodesmus (Opisotretidae, the genus being the westernmost in that Indo-Australian family), with at least five species from the Himalayas of Nepal and Bhutan, two in S China, and one in S Vietnam (Golovatch et al. 2013). The latter example, however, may likewise illustrate the predominating zoogeographical connections of the Himalayan fauna with the Indo-Malayan one. Such are also the genera Cryptocorypha (Pyrgodesmidae), with 14 described species from S India, Myanmar, S China and Australasia, eastwards to Vanuatu, Melanesia (Golovatch and VandenSpiegel 2015, Golovatch and Wesener 2016), Zephronia (Sphaerotheriida), with 38 species or subspecies from Nepal, NE India, Myanmar, Thailand, Cambodia, Vietnam, Western (= mainland) Malaysia and Hong Kong (Wesener 2016), Siphonorhinus (Siphonophorida), with eight species from Indonesia, Vietnam, Cambodia, Laos, India and Madagascar (Minelli 2015) and Pocockina (Paradoxosomatidae), with three species from Nepal and Myanmar (Golovatch 2016b). The small genus Trichopeltis (Cryptodesmidae) includes nine described species, largely from Indochina, Sumatra, Indonesia, Myanmar and S China; only the much more widespread T. watsoni and Gonoplectus malayus malayus reach as far west as the Himalayas of India (Table 1). Similarly, amongst the Paradoxosomatidae the genus Trogodesmus contains four species in Myanmar and one more in Assam, India (Nguyen and Sierwald 2013); Touranella harbours two species from Nepal and further four from Vietnam (Golovatch 2016b); Kronopolites has one species in Jammu & Kashmir, India, one in Thailand, one in Laos, and another nine in China, including one in Taiwan (Likhitrakarn et al. 2015, Golovatch 2015a); Anoplodesmus is a large genus which, regardless of a few pantropical anthropochores, comprises approximately 35 species in S (India and Sri Lanka), E (Taiwan) and SE Asia (eastwards up to Sumatra, Indonesia), including nine congeners confined to the Himalayas (Golovatch 2016a, Table 1); Streptogonopus contains not only S. phipsoni, a species quite widespread in the Himalayas and certain adjacent countries (Table 1), but also two more in India, and one each in Eritrea, Thailand and Vietnam (Nguyen et al. 2016); Delarthrum is one of the most species-rich genera of Diplopoda (56 spp.), with most (55 spp.) of its diversity confined to the Himalayas of Pakistan, Nepal and India (Table 1), and only one outlier congener in S India (Golovatch and Wesener 2016, Golovatch 2016b). Much like Delarthrum, the genus Sholaphilus demonstrates faunal connections between the Himalayas (6 spp., Table 1) and S India (1 sp., Golovatch and Wesener 2016).

Suprageneric relations

At these taxonomic levels, the fully to largely tropical orders Sphaerotheriida, Siphonophorida and Spirostreptida, as well as most of Spirobolida, Platydesmida and Polydesmida (at least some Haplodesmidae, Cryptodesmidae, Paradoxosomatidae, Pyrgodesmidae and Trichopolydesmidae) seem to represent Indo-Malayan elements in the Himalayan millipede fauna. A siphonophoridan and a cryptodesmid species, both have been recorded as far north as N Pakistan (Golovatch 1991). However, at the species level the bulk of diversity is endemic and highly localized, both horizontally and vertically, clearly due to allopatric vicariance (cf. Golovatch and Martens 1996). There are several Himalayan endemic to subendemic diplopod genera, but not a single suprageneric taxon: Kophosphaera (Sphaerotheriida, seven species, Wesener 2016 & Table 1), Koponenius (Polydesmida, four species from Nepal, NW India and Myanmar (Golovatch and VandenSpiegel 2014, 2016), Kashmireuma (Chordeumatida), with three species from Nepal and N India (Table 1), Himalodesmus (Polydesmidae, Table 1), Beronodesmoides, Beronodesmus, Substrongylosoma (all Paradoxosomatidae, Table 1), Magidesmus (Trichopolydesmidae, Table 1), as well as the monotypic Assamodesmus, Bhutanodesmus, Topalodesmus, Topalosoma and Typhlopygmaeosoma (Table 1).

Faunal connections to the northwest and/or north are demonstrated by a few examples only. Even at the generic level, not all of them could unequivocally be treated as likely Palaearctic components in the Himalayan millipede fauna. Thus, the genus Kaschmiriosoma (Paradoxosomatidae) is composed of three species endemic to N Pakistan, and one to both N Pakistan and N India (Jeekel 2003). Such a pattern might seem to have been accounted for by an originally northwestwards dispersal. Even in the absence of a phylogenetic reconstruction, the gonopodal structure in Kaschmiriosoma, especially the particularly complex, strongly coiled and thus apomorphous solenophores as observed in the southernmost, Himalayan K. contortipes and K. nulla, may rather be evidence of a Palaearctic origin of the genus. Its deemed closest relatives within the tribe Sulciferini, also often showing particularly complex and strongly twisted gonopodal solenophores, are observed in the genera Gonobelus Attems, 1936, Inversispina Zhang, in Zhang et al. 1997 and a few others, all confined to southern China, occasionally including Taiwan (Jeekel 1980, Golovatch 2012, 2016b).

Ties to the north are much better pronounced, e.g., in the genera Tianella (Cleidogonidae), Epanerchodus (Polydesmidae), Bollmania (Caspiopetalidae) and Anaulaciulus (Julidae). Tianella has two described and a number on still undescribed species in Kyrgyzstan and Kazakhstan, Central Asia, as well as 11 named species in Nepal and a few undescribed ones from both Nepal and N India (Mauriès 1988, Read and Golovatch 1994, Table 1). Epanerchodus is a very large genus comprising 70+ species from Central and E Asia: Russian Far East, Korea, China, Taiwan and Japan, as well as several species from Nepal (Minelli 2015, Table 1). Bollmania is composed of eight described and a few undescribed species ranging from Turkmenistan, Iran, Uzbekistan, Tajikistan and Afghanistan in Central Asia to S China; one species is known from the Himalayas of Pakistan (Stoev et al. 2008, Table 1). Anaulaciulus is also a highly speciose Asian genus which contains nearly 50 species ranging from across the Himalayas, through China, to the Russian Far East, Korea, Japan and Taiwan (Korsós 2001, Table 1). It is partly sympatric with still another, similarly large, Asian genus Nepalmatoiulus (Julidae), which harbours 55 species also distributed across the Himalayas, but then extending more to the east and southeast (S China, Taiwan, S Ryukyus, Myanmar, Indochina and peninsular Malaysia) (Enghoff 1987, Korsós and Lazányi 2013, Table 1). In the Himalayas, many species from these genera are high-montane (Table 1), thus reinforcing their presumed Palaearctic origins. The occurrence of the sole known species of Nepalozonium (Polyzoniida) at 4700–4800 a.s.l., i.e. among the highest records in the entire class, coupled with the family Hirudisomatidae where it belongs being strictly Holarctic (Minelli 2015), is clearly evidence of its Palaearctic roots.

The pattern demonstrated by the very large genus Hyleoglomeris (Glomeridae) strongly resembles that of the family Siphonocryptidae (see above and Fig. 5), but in no way is it relictual. Indeed, its 100+ species range from the Balkans and Greek islands in the west, through Anatolia, the Caucasus, Central Asia and the Himalayas, to China, Korea, Japan, and Taiwan in the east, and through Indochina to the Philippines and Sulawesi, Indonesia in the southeast (Golovatch et al. 2006, Table 1). This picture actually reflects one of the fundamental patterns of historical biogeography as evidenced in the entire class Diplopoda (Shelley and Golovatch 2011). Generally, west-east trans-Himalayan faunal connections are traced in numerous millipede higher taxa: Glomerida, Julida, Chordeumatida, Callipodida, Siphonocryptida, Spirostreptida, Polydesmida etc. Southeast Asia is the only corner in the world where all 16 extant orders of Diplopoda are still to be found. In many cases, the Himalayas might have served as a paramount stepping stone and refugium in linking, much more in the past than at present, the faunas of SE Asia to those of Europe and W Asia. In the past, the Himalayas started rising and absorbing surrounding faunal elements often already present in the area. After having gained a certain height, the Himalayas functioned more as a trap, a “prison”, mountain ridges and deep valley systems hindering further faunal exchange, with the tremendous speciation process that came into action. Old migration routes both ways are thereby evident, although the influence of the Oriental fauna is by far greater. Since the uplift of the Himalayas started in the early Oligocene (about 27 Mya), the diplopod fauna of the region has gradually acquired its own, highly characteristic, very rich and diverse composition, multi-layered and very complex, with profound in situ radiations and vicariance events (cf. Golovatch and Martens 1996, Martens 2015). At least some of the oldest components are presently highly disjunct and clearly relict, as is the order Siphonocryptida (see above & Fig. 5). The most recent faunal layer is certainly represented by the few pantropical or subcosmopolitan introductions like Trigoniulius corallinus, Orthomorpha coarctata or Oxidus gracilis.

Conclusions

Since the previous review of millipede chorology and faunogenesis in the Himalayas (Golovatch and Martens 1996), our knowledge of the Himalayan fauna has become considerably enriched (ca 200 vs >270 spp.) and often refined taxonomically. This is particularly true of the composition of the dominant family Paradoxosomatidae. However, the main results and trends remain unchanged.

The Himalayas support a highly rich, diverse, multi-layered, mostly endemic diplopod fauna. This is the result of mixing the ancient, apparently Tertiary and younger, Plio-Pleistocene elements of various origins, as well as the most recent anthropochore introductions. At the species and, partly, generic levels, the fauna is largely autochthonous and sylvicolous, formed through abounding in situ radiation and vicariance events, when overall the species from large genera and families tend to occupy a wide range of altitudes, but nearly each of the constituent species shows a distribution highly localized both horizontally and altitudinally, yet quite often with sympatry or even syntopy involved. The bulk of the fauna is Indo-Malayan in origin, with individual genera or families shared with those of SE Asia (mostly) and/or S India (few) (Fig. 6). Their constituent species tend to be lowland to mid-montane, but the general rule “(sub)tropical organisms for (sub)tropical environments only” fails very often.

Figure 6. 

General schematic picture of the faunogenesis of Himalayan Diplopoda. Arrows reflect the main pathways of faunal migration or exchange, their thickness roughly corresponding to the degree of influence. The thickest arrow 1 clearly emphasizes the dominant roles the Indo-Malayan core fauna may have played in the present-day composition of the Himalayan fauna, its most ancient layers extending westwards to reach central and western Asia, as well as Europe (by default also northwards up to eastern Asia and even North America). The considerably less thick arrows 2 and 3 are to reflect the more subordinate roles the Sino-Himalayan and Palaearctic elements, respectively, could have played in the modern Himalayan fauna. Arrows 4 and, especially, 5 are even less thick and demonstrate the relatively minor faunal exchanges to be presumed between the Indian and Himalayan faunas.

The Palaearctic influence is modest (Fig. 6), but still can be traced in several genera and families. Collectively, their species tend to be high montane, but with numerous exceptions as well. The trans-Himalayan faunal connections at higher taxonomic levels, generic to ordinal, that link SE Asia to Europe are manifest. They show routes and directions of ancient dispersal both ways, but the one from SE Asia seems to have prevailed over the opposite one. Moreover, certain fragments or offshoots of such ancient, obviously Tertiary, opposite migrations more or less along the southern coasts of the receding Tethys Sea may have been left en route in S and SW China, as well as in N Pakistan and Central Asia.

One must also take into account that a number of presumably Himalayan species groups might have originated from the times when Tibet was still forest-covered and the Himalayan chain still in its infancies. According to Schmidt (2006), stem species invaded the raising Himalayas from the north where they developed to presently known species swarms. After the Himalayas and Tibetan Plateau had raised sufficiently high, Tibet became drier and the forests vanished including their fauna, the Himalayas becoming their exile. Schmidt coined the term “Tibeto-Tertiary element” of paleo-Tibetan origin with present Himalayan distributions (Schmidt 2006, Schmidt et al. 2012).

The particularly rich Himalayan diplopod fauna with its numerous small-ranging species confined to permanent forest sheds new light on a much disputed controversy among geographers, zoologists, taxonomists, climatologists and glacialogists (Kuhle 1982, 2015 and figure 4 therein). Was the central Himalayan chain, at least at certain sections of the Pleistocene, covered by a complete shield of ice? Taxonomists dealing with low-dispersal soil/litter-dwelling arthropods have a clear response. Any ice cover would have been detrimental to the local soil arthropod fauna and would have driven its larger part or entirely to extinction. Only a long and steady evolution under more or less constant, albeit slightly varying, conditions would have allowed the biota to develop gradually over long geological periods (Martens 2015). This scenario certainly applies to all Himalayan Diplopoda.

The above picture of faunal connections (Fig. 6) is consistent both with general wisdom (e.g. Martens 2015) and our previous analysis (Golovatch and Martens 1996), the salient aspects of Diplopoda, contrary to many other animal groups in the Himalayas, being their pronounced sylvicoly, extremely diverse and small-ranging species endemism, and mostly Oriental and/or Indian origins, while some of the rather ostensible influence of the Palaearctic may have come not only from the north and/or northwest, but also from the currently subtropical regions of S China. Reciprocal migrations from the Himalayan faunal knot as a major refugium and secondary diversification centre also seem quite plausible, but documenting such requires detailed phylogenies which unfortunately are still almost missing.

Acknowledgements

J.M. thanks all of the many co-workers who accompanied him and made the various expeditions as successful as they finally turned out. His wife Beate especially is to be mentioned; she participated in the 1983 extended expedition and, most importantly, encouraged him to leave for Asia nearly annually for many weeks. The Feldbausch-Stiftung and the Wagner-Stiftung at the Fachbereich Biologie of Mainz University granted financial aid to J.M. We both heartily thank all colleagues, friends and institutions.

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