Research Article |
Corresponding author: Sergei I. Golovatch ( sgolovatch@yandex.ru ) Academic editor: Pavel Stoev
© 2018 Sergei I. Golovatch, Jochen Martens.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Golovatch SI, Martens J (2017) Distribution, diversity patterns and faunogenesis of the millipedes (Diplopoda) of the Himalayas. In: Stoev P, Edgecombe GD (Eds) Proceedings of the 17th International Congress of Myriapodology, Krabi, Thailand. ZooKeys 741: 3-34. https://doi.org/10.3897/zookeys.741.20041
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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.
Diplopoda , faunistics, Plio-Pleistocene, Asia, Palaearctic
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.
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 (
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 (
The first, provisional review of the millipede fauna of the Himalayas (
These results mostly rely on published records, which have grown considerably over the past two decades. The resultant checklist (Table
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 ( |
Genus Monographis Attems, 1907 | ||
2. Monographis mira (Turk, 1947) | 1600–2400 | Nepal1 & India, Almora ( |
Genus Unixenus Jones, 1944 | ||
3. Unixenus sp. | 2400–4550 | Nepal2 ( |
Order Sphaerotheriida | 140–2700 | |
Family Zephroniidae | 140–2700 | |
Genus Indosphaera Attems, 1935 | ||
4. Indosphaera feae Attems, 1935 | ? | India, Assam ( |
Genus Kophosphaera Attems, 1935 | 1650–2100 | |
5. Kophosphaera brevilamina Attems, 1936 | 1700 | India, West Bengal & Darjeeling Distr. ( |
6. K. devolvens Attems, 1936 | 1700–2050 | India, Sikkim & Darjeeling Distr. ( |
7. K. excavata (Butler, 1874) | ? | Nepal, Chitlang; India, Sikkim, Darjeeling Distr. & Assam ( |
8. K. mammifera Attems, 1936 | ? | India, Darjeeling Distr. & Assam ( |
9. K. martensi Wesener, 2015 | 2100 | Nepal ( |
10. K. politissima Attems, 1935 | 1650–1870 | India, Darjeeling Distr. & Nepal ( |
11. K. shivapuri Wesener, 2015 | 1700–2100 | Nepal ( |
Genus Zephronia Gray, 1832 | 140–2700 | |
12. Zephronia alticola alticola Attems, 1936 | 400–1700 | India, Darjeeling Distr. & Assam ( |
13. Z. alticola bengalica Attems, 1936 | ? | India, West Bengal ( |
14. Z. debilis Attems, 1936 | 1700 | India, Darjeeling Distr. ( |
15. Z. densipora Attems, 1936 | ? | India, Assam ( |
16. Z. disparipora Attems, 1936 | 140 | India, Assam ( |
17. Z. hirta Attems, 1936 | 1700 | India, Darjeeling Distr. ( |
18. Z. hysophila Attems, 1936 | ? | India, Assam ( |
19. Z. juvenis Attems, 1936 | ? | India, Assam ( |
20. Z. lignivora Attems, 1936 | 180–330 | India, Assam ( |
21. Z. manca Attems, 1936 | 1000–1700 | Vietnam & India, Darjeeling Distr. ( |
22. Z. montana (Karsch, 1881) | ? | “Himalaya” ( |
23. Z. nepalensis Wesener, 2015 | 1700–2600 | Nepal ( |
24. Z. nigrinota Butler, 1872 | 2300–2700 | India, Darjeeling Distr. ( |
25. Z. specularis Attems, 1936 | ? | India, Assam ( |
26. Z. tigrina Butler, 1872 | ? | India, Darjeeling Distr. ( |
27. Z. tigrinoides Attems, 1936 | 170 | India, Darjeeling Distr. ( |
28. Z. tumida Butler, 1882 | ? | India, Assam & Darjeeling Distr.; Myanmar ( |
Order Glomerida | 150–3300 | |
Family Glomeridae | 150–3300 | |
Genus Hyleoglomeris Verhoeff, 1910 | 150–3300 | |
29. Hyleoglomeris crassipes Golovatch, 1987 | 2450–2720 | Nepal ( |
30. H. electa (Silvestri, 1917) | 500–1700 | India, Darjeeling Distr. ( |
31. H. gorkhalis Golovatch, 1987 | 1200 | Nepal ( |
32. H. khumbua Golovatch, 1987 | 3250–3300 | Nepal ( |
33. H. modesta Silvestri, 1917 | 150 | India, Assam ( |
34. H. nagarjunga Golovatch, 1987 | 1600–2100 | Nepal ( |
35. H. tinjurana Golovatch, 1987 | 2450 | Nepal ( |
36. H. venustula Silvestri, 1917 | ? | India, Assam ( |
Order Siphonophorida | ||
Family Siphonorhinidae | 500–1700 | |
Genus Siphonorhinus Pocock, 1894 | 500–1700 | |
37. Siphonorhinus cingulatus (Attems, 1936) | 500–1700 | Vietnam and India, Darjeeling Distr. ( |
38. S. coniceps (Attems, 1936) | 1700 | India, Darjeeling Distr. ( |
39. S. larwoodi (Turk, 1947) | 1600 | India, Almora ( |
Order Siphonocryptida | ||
Family Siphonocryptidae | 2450 | |
Genus Hirudicryprus Enghoff & Golovatch, 1995 | ||
40. Hirudicryprus quintumelementum Korsós, Geoffroy & Mauriès, 2009 | 2450 | Nepal ( |
Order Platydesmida | <2000 | |
Family Andrognathidae | <2000 | |
Genus Pseudodesmus Pocock, 1887 | ||
41. ?Pseudodesmus sp. | <2000 | Nepal ( |
Order Polyzoniida | 4700–4800 | |
Family Hirudisomatidae | 4700–4800 | |
Genus Nepalozonium Shelley, 1996 | ||
42. Nepalozonium trimaculatum Shelley, 1996 | 4700–4800 | Nepal ( |
Order Chordeumatida | 900–4100 | |
Family Kashmireumatidae | 2600–4100 | |
Genus Kashmireuma Mauriès, 1982 | 2600–4100 | |
43. Kashmireuma nepalensis Mauriès, 1988 | 3600–4100 | Nepal ( |
44. K. nielseni Mauriès, 1982 | 2600–3500 | India, Kashmir ( |
45. K. schawalleri Shear, 1987 | 3450–3600 | Nepal ( |
Family Cleidogonidae | 900–3900 | |
Genus Tianella Attems, 1904 | 900–3900 | |
46. Tianella ausobskyi Shear, 1987 | 2500–3050 | Nepal ( |
47. T. bobanga Shear, 1979 | 2460–2500 | Nepal ( |
48. T. daamsae Shear, 1987 | 3600–3900 | Nepal ( |
49. T. gitanga Shear, 1987 | 2550 | Nepal ( |
50. T. jaljalensis Mauriès, 1988 | 2350 | Nepal ( |
51. T. kathmandua Mauriès, 1988 | 1700 | Nepal ( |
52. T. lughla Shear, 1979 | 2950–3300 | Nepal ( |
53. T. managa Shear, 1987 | 2550 | Nepal ( |
54. T. mangsingma Mauriès, 1988 | 2250 | Nepal ( |
55. T. martensi Shear, 1979 | 1150–2900 | Nepal (Shear, 1979, 1987, |
56. T. smetanai Mauriès, 1988 | 3250 | Nepal ( |
57. Tianella sp. | 900–1400 | India, Darjeeling Distr. ( |
Family Megalotylidae | 1900–4100 | |
Genus Nepalella Shear, 1979 | 1900–4100 | |
58. Nepalella deharvengi Mauriès, 1988 | 2900–3500 | Nepal ( |
59. N. gairiensis Mauriès, 1988 | 3000 | Nepal ( |
60. N. gunsa Shear, 1987 | 3600–3800 | Nepal ( |
61. N. jaljalae Mauriès, 1988 | 2200 | Nepal ( |
62. N. khumbua Shear, 1979 | 3350–3300 | Nepal ( |
63. N. phulcokia Mauriès, 1988 | 2250 | Nepal ( |
64. N. ringmoensis Mauriès, 1988 | 2750–3000 | Nepal ( |
65. N. taplejunga Shear, 1987 | 3000–3300 | Nepal ( |
66. N. thodunga Shear, 1979 | 3200 | Nepal ( |
67. N. tragsindola Mauriès, 1988 | 2450–3300 | Nepal ( |
68. Nepalella sp. | 1900–4100 | Nepal ( |
Order Callipodida | 650 | |
Family Caspiopetalidae | 650 | |
Genus Bollmania Silvestri, 1896 | ||
69. Bollmania kohalana (Attems, 1936) | 650 | Pakistan, Punjab ( |
Order Julida | 1680–4800 | |
Family Julidae | 1680–4800 | |
Genus Anaulaciulus Pocock, 1895 | 1900–4500 | |
70. Anaulaciulus acaudatus Korsós, 1996 | 3990 | India, Sikkim ( |
71. A. bilineatus Korsós, 1996 | 3300–4300 | Nepal ( |
72. A. kashmirensis Korsós, 1996 | 3100–3200 | India, Kashmir ( |
73. A. nepalensis Korsós, 1996 | 2600–3400 | Nepal ( |
74. A. niger Korsós, 1996 | 2600–4500 | Nepal ( |
75. A. tibetanus Korsós, 1996 | 3700 | China, Tibet; India, Assam ( |
76. A. topali Korsós, 1996 | 2300 | India, Kashmir ( |
Genus Nepalmatoiulus Mauriès, 1983 | 1680–4800 | |
77. Nepalmatoiulus appendiculatus Enghoff, 1987 | 1900–2100 | India, Uttar Pradesh ( |
78. N. deharvengi (Mauriès, 1983) | 2550–3350 | Nepal ( |
79. N. dhaulagiri Enghoff, 1987 | 3000–3350 | Nepal ( |
80. N. generalis Enghoff, 1987 | 3400 | Nepal ( |
81. N. hyalilobus Enghoff, 1987 | 3600–3800 | Nepal ( |
82. N. ivanloebli Enghoff, 1987 | 2200–4800 | Nepal ( |
83. N. juctapositus Enghoff, 1987 | 2800–3050 | Nepal ( |
84. N. martensi Enghoff, 1987 | 3250–3300 | Nepal ( |
85. N. mauriesi Enghoff, 1987 | 3600 | Nepal (Enghoff 1983, |
86. N. nigrescens Enghoff, 1987 | 2300 | Bhutan (Enghoff 1983, |
87. N. pineti Enghoff, 1987 | 2900 | Nepal ( |
88. N. rugiflagrum Enghoff, 1987 | 3300 | Bhutan ( |
89. N. smetanai (Mauriès, 1983) | 1900–2700 | Nepal (Enghoff 1983, |
90. N. sympatricus Enghoff, 1987 | 3000 | Nepal ( |
88. N. uncus Enghoff, 1987 | 2550 | Nepal ( |
91. N. wuermlii Enghoff, 1987 | 1680–2600 | Bhutan ( |
92. N. zachonoides Enghoff, 1987 | 2450–2600 | Nepal ( |
Order Spirostreptida | 200–2500 | |
Family Cambalopsidae | <1000–1200 | |
Genus Podoglyphiulus Attems, 1909 | <1000–1200 | |
93. Podoglyphiulus elegans nepalensis Mauriès, 1983 | <1000 | Nepal3 ( |
Genus Trachyjulus Peters, 1864 | ||
94. Trachyjulus mimus Silvestri, 1924 | 1200 | India, Assam ( |
95. T. wilsonae Mauriès, 1983 | <1000 | Nepal ( |
Family Harpagophoridae | 200–2500 | |
Genus Dametus Attems, 1942 | ||
96. Dametus falcatus (Attems, 1936) | 400–500 | India, Assam ( |
Genus Gonoplectus Chamberlin, 1921 | 200–2500 | |
97. Gonoplectus alius Demange, 1961 | ? | India, Assam ( |
98. G. bhutanensis Demange, 1988 | 350–450 | Bhutan ( |
99. G. broelemanni Demange, 1961 | 1800–2300 | Nepal ( |
100. G. corniger (Attems, 1936) | ? | India, Assam ( |
101. G. gracilis (Attems, 1936) | 1200 | India, Darjeeling Distr. ( |
102. G. hyatti Demange, 1961 | 1200 | Nepal ( |
103. G. malayus malayus (Carl, 1909) | 200–2500 | Indonesia, Java; Nepal, Bhutan & India, Madhya Pradesh, Uttar Pradesh, Himachal Pradesh, West Bengal ( |
104. G. malayus lindbergi (Carl, 1909) | 350 | Bhutan & India, Darjeeling Distr. ( |
105. G. probus (Attems, 1936) | 1000 | India, Darjeeling Distr. ( |
106. G. remyi Demange, 1961 | ? | India, Assam ( |
107. G. repertus (Attems, 1936) | 900 | India, Darjeeling Distr. ( |
108. G. sulcatus (Attems, 1936) | 2400 | India, Darjeeling Distr. ( |
Order Spirobolida | <1000–1800 | |
Family Pachybolidae | ||
Genus Trigoniulus Pocock, 1894 | <1000 | |
109. Trigoniulus corallinus (Gervais, 1847) | <1000 | Pantropical, in India nearly throughout, including Assam ( |
Family Pseudospirobolellidae | ||
Genus Physobolus Attems, 1936 | ||
110. Physobolus olivaceus Attems, 1936 | 1800 | India, Darjeeling Distr. ( |
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 ( |
Family Haplodesmidae | 150–1750 | |
Genus Koponenius Golovatch & VandenSpiegel, 2014 | 150–1750 | |
112. Koponenius biramus Golovatch & VandenSpiegel, 2014 | 1750 | Nepal ( |
113. K. schawalleri Golovatch & VandenSpiegel, 2016 | 150 | Nepal ( |
114. K. unicornis Golovatch & VandenSpiegel, 2014 | 880 | India, Darjeeling Distr. ( |
Family Opisotretidae | 1100–2440 | |
Genus Martensodesmus Golovatch, 1987 | 1100–2440 | |
115. Martensodesmus bicuspidatus Golovatch, 1988 | 1650–2000 | Bhutan ( |
116. M. excornis Golovatch, 1988 | 2440 | Bhutan ( |
117. M. himalayensis Golovatch, 1987 | 1100–1300 | Nepal ( |
118. M. nagarjungicus Golovatch, 1987 | 1900–2100 | Nepal ( |
119. M. sherpa Golovatch, 1987 | 1200 | Nepal ( |
120. Martensodesmus sp. | 1300–2150 | Nepal, Bhutan ( |
Family Paradoxosomatidae | 150–4500 | |
Genus Anoplodesmus Carl, 1932 | 1000–3600 | |
121. Anoplodesmus affinis (Golovatch, 1990) | 2475–2700 | Nepal ( |
122. A. cylindricus (Carl, 1935) | 1650–2850 | Nepal & India, Darjeeling Distr. ( |
123. A. elongissimus (Golovatch, 1984) | 1000 | India, Darjeeling Distr. ( |
124. A. magnus Golovatch, 2015 | 2700 | Nepal ( |
125. A. martensi (Golovatch, 1990) | 2250–3600 | Nepal ( |
126. A. schawalleri (Golovatch, 1990) | 2050–2150 | Nepal ( |
127. A. similis (Golovatch, 1990) | 2300–3000 | Nepal ( |
128. A. spinosus Golovatch, 2016 | 2500 | Nepal ( |
129. A. subcylindricus (Carl, 1932) | ? | S India & Nepal ( |
Genus Beronodesmoides Golovatch, 2015 | 1650–4250 | |
130. Beronodesmoides anteriporus Golovatch, 2015 | 1650–3350 | Nepal ( |
131. B. bifidus Golovatch, 2015 | 3100 | Nepal ( |
132. B. lobatus Golovatch, 2015 | 4000–4250 | Nepal ( |
133. B. longifemoratus Golovatch, 2015 | 2700–2800 | Nepal ( |
134. B. martensi Golovatch, 2016 | 2700 | Nepal ( |
135. B. montigena Golovatch, 2016 | 3550 | Nepal ( |
136. B. typicus Golovatch, 2016 | 3400 | Nepal ( |
Genus Beronodesmus Golovatch, 2014 | 1650–4500 | |
137. Beronodesmus curtispinus Golovatch, 2015 | 4500 | Nepal ( |
138. B. distospinosus Golovatch, 2015 | 1650–3080 | Nepal ( |
139. B. gorkhalis Golovatch, 2015 | 3050–3600 | Nepal ( |
140. B. latispinosus Golovatch, 2015 | 1900–3500 | Nepal ( |
141. B. longispinus Golovatch, 2015 | 2550–4270 | Nepal ( |
142. B. martensi Golovatch et al., 2016 | 2650 | Nepal ( |
143. B. minutus Golovatch, 2015 | 3300–3500 | Nepal ( |
144. B. pallidus Golovatch, 2014 | 3800–4100 | Nepal ( |
145. B. serratus Golovatch et al., 2016 | 3300–3500 | Nepal ( |
146. B. simplex Golovatch, 2016 | 2100 | Nepal ( |
147. B. sinuatospinus Golovatch, 2015 | 2150–2250 | Nepal ( |
Genus Delarthrum Attems, 1936 | 600–4100 | |
148. Delarthrum aberrans (Golovatch, 1996) | 1000–2600 | Nepal ( |
149. D. affine (Golovatch, 1994) | 1400 | Nepal ( |
150. D. alatum (Golovatch, 1996) | 1900–2100 | Nepal ( |
151. D. andreevi Golovatch, 2014 | 1800 | Nepal ( |
152. D. arunense (Golovatch, 1994) | 1850–2150 | Nepal ( |
153. D. beroni Golovatch, 2014 | 600–1000 | Nepal ( |
154. D. bifidum (Golovatch, 1996) | 2550–2650 | Nepal ( |
155. D. chulingense (Golovatch, 1994) | 3000–3700 | Nepal ( |
156. D. communicans (Golovatch, 1992) | 2650 | Nepal ( |
157. D. curtisoma Golovatch, 2015 | 2050–2150 | Nepal ( |
158. D. curtum Golovatch, 2014 | 600–1000 | Nepal ( |
159. D. densesetosum Golovatch, 2015 | 2400 | Nepal ( |
160. D. elegans (Golovatch, 1992) | 1350 | Nepal ( |
161. D. extremum (Golovatch, 1996) | 2450 | Nepal ( |
162. D. facile (Golovatch, 1996) | 2200–2400 | Nepal ( |
163. D. fechteri (Golovatch, 1990) | 2330–3150 | Nepal ( |
164. D. foveatum (Golovatch, 1996) | 1800–2000 | Nepal ( |
165. D. furcatum (Golovatch, 1996) | 600–2000 | Nepal ( |
166. D. gracile Golovatch, 2015 | 1750 | Nepal ( |
167. D. granulosum (Golovatch, 1994) | 2000 | Nepal ( |
168. D. heterotergale Golovatch, 2014 | 600–1000 | Nepal (Golovatch 2014) |
169. D. hingstoni (Carl, 1935) | 3400 | China, Tibet ( |
170. D. hirsutum (Golovatch, 1994) | 2400–4100 | Nepal ( |
171. D. intermedium (Golovatch, 1994) | 1000–1100 | Nepal ( |
172. D. invocatum (Golovatch, 1996) | 2600–2800 | Nepal ( |
173. D. kuznetsovi (Golovatch, 1994) | 3000 | Nepal ( |
174. D. longisetum (Golovatch, 1994) | 1400–1600 | Nepal ( |
175. D. longispinum (Golovatch, 1996) | 2150–2250 | Nepal ( |
176. D. modestum (Golovatch, 1996) | 3450–3600 | Nepal ( |
177. D. nyakense (Golovatch, 1992) | 2270–2400 | Nepal ( |
178. D. obscurum Attems, 1936 | ca 2800 | N Pakistan, Punjab ( |
179. D. philosophicum (Golovatch, 1994) | 1650–2450 | Nepal ( |
180. D. planifemur Golovatch, 2015 | 2200 | Nepal ( |
181. D. prolixum (Golovatch, 1996) | 2550–2650 | Nepal ( |
182. D. pumilum (Attems, 1944) | ? | India, Uttar Pradesh ( |
183. D. quadridentatum Golovatch, 2016 | 2600–2800 | Nepal ( |
184. D. schawalleri (Golovatch, 1992) | 1000–2150 | Nepal ( |
185. D. setosum Golovatch, 2014 | ? | Nepal ( |
186. D. silvestre (Golovatch, 1994) | 2000–3400 | Nepal ( |
187. D. simile (Golovatch, 1992) | 2300–2700 | Nepal ( |
188. D. simplex (Golovatch, 1996) | 1650 | Nepal ( |
189. D. simulans (Carl, 1935) | 3700 | Nepal & China, Tibet ( |
190. D. spectabile (Golovatch, 1994) | 2650 | Nepal ( |
191. D. spiniger (Attems, 1936) | 1000–2200 | India, West Bengal & Darjeeling Distr. ( |
192. D. spinigerum (Golovatch, 1992) | 600–1400 | Nepal ( |
193. D. splendens (Golovatch, 1992) | 1650–2150 | Nepal ( |
194. D. subalatum (Golovatch, 1996) | 2600–2800 | Nepal ( |
195. D. subsimulans (Golovatch, 1996) | 3100–3300 | Nepal ( |
196. D. tenuitergale Golovatch, 2014 | 3250 | Nepal (Golovatch 2014) |
197. D. tergale (Golovatch, 1994) | 2650 | Nepal ( |
198. D. tuberculatum (Golovatch, 1994) | 3000–3300 | Nepal ( |
199. D. typicum Golovatch, 2014 | 3100 | Nepal (Golovatch 2014) |
200. D. uncum (Golovatch, 1996) | 2100–3420 | Nepal ( |
201. D. unicolor (Attems, 1936) | 1200–1700 | India, Assam & Darjeeling Distr. ( |
Genus Hirtodrepanum Golovatch, 1994 | ||
202. Hirtodrepanum latigonopum Golovatch, 1994 | 2100–2600 | Nepal ( |
Genus Kaschmiriosoma Schubart, 1935 | 1000–3300 | |
203. Kaschmiriosoma contortipes Schubart, 1935 | 2000–3300 | N Pakistan & India, Jammu & Kashmir ( |
204. K. nulla (Attems, 1936) | 1000 | India, Himachal Pradesh ( |
205. K. pleuropterum (Attems, 1936) | 2800 | N Pakistan, Punjab ( |
Genus Kronopolites Attems, 1914 | ||
206. Kronopolites coriaceus Golovatch, 2015 | 2000 | Nepal ( |
207. K. occidentalis Golovatch, 1983 | 1500 | India, Jammu & Kashmir ( |
Genus Orthomorpha Bollman, 1893 | ||
208. Orthomorpha coarctata (de Saussure, 1860) | 600–650 | Nepal & India, pantropical anthropochore ( |
209. “O.” almorensis Turk, 1947 | 1600 | India, Almora ( |
Genus Oxidus Cook, 1911 | ||
210. Oxidus gracilis (C.L. Koch, 1847) | 570–1200 | Nepal & India, subcosmopolitan anthropochore ( |
Genus Pocockina Jeekel, 1965 | ||
211. Pocockina schawalleri Golovatch, 2016 | 150 | Nepal ( |
Genus Streptogonopus Attems, 1914 | ||
212. Streptogonopus phipsoni (Pocock, 1892) | ≤2700 | Pakistan, Bangladesh, Nepal & India, West Bengal ( |
Genus Substrongylosoma Golovatch, 1984 | 1000–2200 | |
213. Substrongylosoma bifurcatum Golovatch, 2016 | 2000 | Nepal ( |
214. S. distinctum Golovatch, 1984 | 1200–1500 | India, Darjeeling Distr. ( |
215. S. falcatum Golovatch, 1984 | 1000–1400 | India, Darjeeling Distr. ( |
216. S. exiguum Golovatch, 2016 | 1900 | Nepal ( |
217. S. montigena (Carl, 1935) | 1200–2200 | India, Darjeeling Distr. ( |
218. S. schawalleri Golovatch, 1993 | 1620–2000 | Nepal ( |
Genus Sundanina Attems, 1914 | ||
219. “Sundanina” septentrionalis Turk, 1947 | ca 1700 | India, Almora ( |
Genus Topalosoma Golovatch, 1984 | 900 | |
220. Topalosoma setiferum Golovatch, 1984 | 900 | India, Darjeeling Distr. ( |
Genus Trogodesmus Pocock, 1895 | ||
221. Trogodesmus uncinatus (Attems, 1936) | ? | India, Assam ( |
Genus Touranella Attems, 1937 | 2300–2800 | |
222. Touranella himalayaensis Golovatch, 1994 | 2300–2700 | Nepal ( |
223. T. pilosa Golovatch, 2016 | 2600–2800 | Nepal ( |
Family Polydesmidae | 350–4250 | |
Genus Bhutanodesmus Golovatch, 1988 | ||
224. Bhutanodesmus velatus Golovatch, 1988 | 350–450 | Bhutan (Golovatch 1988, |
Genus Epanerchodus Attems, 1901 | 2300–4250 | |
225. Epanerchodus buddis (Golovatch, 1986) | 3300–3400 | Nepal ( |
226. E. occultus (Golovatch, 1986) | 2300–2800 | Nepal ( |
227. E. sacer (Golovatch, 1987) | 3300–3400 | Nepal ( |
228. E. theocraticus (Golovatch, 1990) | 2600–2800 | Nepal ( |
229. E. theosophicus (Golovatch, 1986) | 3200 | Nepal ( |
230. Epanerchodus sp. | 3450–4250 | Nepal & Bhutan ( |
Genus Glenniea Turk, 1945 | 350–2800 | |
231. Glenniea bhotiaensis Golovatch, 1988 | 350–450 | Bhutan ( |
232. G. indica Turk, 1945 | 2800 | India, Himachal Pradesh ( |
233. G. martensi (Golovatch, 1987) | 1200 | Nepal ( |
234. G. minuscula Golovatch, 1988 | 1900–2300 | Bhutan ( |
235. G. perarmata Golovatch, 1988 | 1680 | Bhutan ( |
Genus Himalodesmus Golovatch, 1986 | 1000–3400 | |
236. Himalodesmus audax Golovatch, 1986 | 2650 | Nepal ( |
237. H. benefactor Golovatch, 1987 | 2600–3400 | Nepal ( |
238. H. faustus Golovatch, 1987 | 1000–1750 | Nepal ( |
239. H. parvus Golovatch, 1987 | 2200 | Nepal ( |
240. H. prosperus Golovatch, 1990 | 2600–2800 | Nepal ( |
241. H. pulcher Golovatch, 1987 | 2450 | Nepal ( |
242. H. pygmaeus Golovatch, 1986 | 3300–3400 | Nepal ( |
243. H. vigens Golovatch, 1987 | 2150–2250 | Nepal ( |
Genus Typhlopygmaeosoma Turk, 1972 | ||
244. Typhlopygmaeosoma hazeltonae Turk, 1972 | 1850 | India, Himachal Pradesh ( |
Family Trichopolydesmidae | 450–4500 | |
Genus Assamodesmus Manfredi, 1955 | ||
245. Assamodesmus lindbergi Manfredi, 1954 | ? | India, Assam ( |
Genus Hingstonia Carl, 1935 | 2000–4500 | |
246. Hingstonia beatae Golovatch, 1990 | 2400–3500 | Nepal ( |
247. H. dorjulana Golovatch, 1988 | 2450–3100 | Bhutan ( |
248. H. eremita Carl, 1935 | 2000 | Nepal ( |
249. H. falcata Golovatch, 1986 | 2650 | Nepal ( |
250. H. fittkaui Golovatch, 1990 | 3350–3450 | Nepal ( |
251. H. gogonana Golovatch, 1988 | 3650–4000 | Bhutan ( |
252. H. pahakholana Golovatch, 1990 | 2600–2800 | Nepal ( |
253. H. pelelana Golovatch, 1988 | 3300–3400 | Bhutan ( |
254. H. perarmata Golovatch, 1986 | 3150 | Nepal ( |
255. H. serrata Golovatch, 1987 | 3400–3600 | Nepal ( |
256. H. sympatrica Golovatch, 1990 | 3550–3650 | Nepal ( |
257. H. variata Golovatch, 1987 | 2600–4500 | Nepal ( |
258. H. yeti Golovatch, 1988 | 1600–2600 | Bhutan ( |
259. Hingstonia sp. | 2200–3900 | Nepal ( |
Genus Magidesmus Golovatch, 1988 | 3100–3400 | |
260. Magidesmus affinis Golovatch, 1988 | 3300–3400 | Bhutan ( |
261. M. bhutanensis Golovatch, 1988 | 3100 | Bhutan ( |
Genus Pseudosphaeroparia Carl, 1932 | ||
262. Pseudosphaeroparia cavernicola Turk, 1945 | 2800 | India, Uttar Pradesh ( |
Genus Sholaphilus Carl, 1932 | 1100–2200 | |
263. Sholaphilus asceticus Golovatch, 1986 | 1300–1650 | Nepal ( |
264. S. dalai Golovatch, 1986 | 2400 | Nepal ( |
265. S. gompa Golovatch, 1990 | 2000–2100 | Nepal ( |
266. S. lama Golovatch, 1986 | 1800–2000 | Nepal ( |
267. S. martensi Golovatch, 1986 | 1100–1850 | Nepal ( |
268. S. monachus Golovatch, 1990 | 2050–2150 | Nepal ( |
Genus Topalodesmus Golovatch, 1988 | ||
269. Topalodesmus communis Golovatch, 1988 | 2000–2200 | India, Darjeeling Distr. ( |
Family Pyrgodesmidae | 450–1200 | |
270–275? Several genera and species (including at least 2 species of Cryptocorypha Attems, 1907) | 450–1200 | Nepal ( |
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
The following examples can be given and easily added to the roster of similar observations that
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
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.
According to
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
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
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.
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.
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 (
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 (
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 (
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 (
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 (
The pattern demonstrated by the very large genus Hyleoglomeris (Glomeridae) strongly resembles that of the family Siphonocryptidae (see above and Fig.
Since the previous review of millipede chorology and faunogenesis in the Himalayas (
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.
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.
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
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 (
The above picture of faunal connections (Fig.
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.