Abstract
This paper summarizes the taxonomic and biogeographical knowledge of Trechus species known so far from the Transhimalaya of Central Tibet and from the southern adjacent Tibetan Himalaya of Tibet and Nepal. Nine species groups are proposed, 25 new species as well as three additional new subspecies are described: The species group of Trechus antonini Deuve, 1997, with ten species newly described: T. astrophilus sp. n., T. budhaensis sp. n., T. lama sp. n., T. rarus sp. n., T. religiosus sp. n., T. singularis sp. n., T. tsampa sp. n., T. tseringi sp. n., T. yak sp. n., with an additional subspecies T. yak shogulaensis ssp. n., and T. yeti sp. n., all from South Central Tibet; the monotypic species group of the newly described Trechus chaklaensis sp. n. from South Central Tibet; the species group of Trechus dacatraianus Deuve, 1996, with two species newly described: T. bastropi sp. n., and T. mieheorum sp. n., both from South Central Tibet; the species group of Trechus franzianus Mateu & Deuve, 1979, with four species newly described: T. aedeagalis sp. n. from Far West Nepal, T. eremita sp. n. from West Nepal, T. muguensis sp. n. from West Nepal, and T. sculptipennis sp. n. from Far West Nepal; the monotypic species group of the newly described Trechus rolwalingensis sp. n. from the upper Rolwaling Valley of Central Nepal, with an additional subspecies T. rolwalingensis daldunglana ssp. n. from the lower Rolwaling Valley; the monotypic species group of the newly described Trechus solhoeyi sp. n. from South Central Tibet; the monotypic species group of the newly described Trechus stratiotes sp. n. from north eastern Saipal Himal of Far West Nepal, with an additional subspecies T. stratiotes malikasthana ssp. n. from south eastern Saipal Himal; the species group of Trechus thibetanus Jeannel, 1928, with three species newly described: T. dongulaensis sp. n., T. glabratus sp. n., and T. namtsoensis sp. n., all from South Central Tibet; the species group of Trechus wrzecionkoi Deuve, 1996, with two species newly described: T. korae sp. n., and T. martinae sp. n., both from South Central Tibet. The following two synonymies are proposed: Trechus franzianus Mateu & Deuve, 1979 = Trechus surdipennis Mateu & Deuve, 1979, syn. n.; Trechus thibetanus Jeannel, 1928 = Trechus pseudocameroni Deuve, 1996, syn. n. A key to all species known of South Central Tibet and the Tibetan Himalaya is presented for the first time, and the distributional data of all these species are mapped. The distributional maps highlight the extremely limited distribution of all wingless Trechus species. In situ speciation following the geographical separation of the range of the ancestral species and lack of subsequent range expansion of strictly edaphic species is postulated. Trechus species do not only exhibit a stronger local endemism, but the individual species groups are also endemic to several parts of the Himalayan-Tibetan Orogen. This indicates that the evolution of these Trechus species groups is directly linked to separate geological formations. Based on geological knowledge, the evolution of the species groups endemic to the Tibetan Himalaya and the Transhimalaya started already in the Miocene after these mountains were lifted up to high montane elevations. The recent distributional area of the species can therefore not be the result of range expansion during the Holocene from Pleistocene refugia outside the Tibetan Himalaya or the Transhimalaya. Instead the existence of glacial refugia can be postulated to be in the lower parts of the same mountain slope on which the species occur today. These results clearly challenge the theory of a Tibetan inland ice sheet stretching through the Himalayan transverse valleys during the Last Glacial Maximum.References
Andrewes, H.E. (1930) The Carabidae of the third Mount Everest Expedition, 1924. Transactions of the Entomological Society of London, 78, 1–44 + 1 map.
Barr, T.C. (1979) Revision of Appalachian Trechus (Coleoptera: Carabidae). Brimleyana, 2, 29–75.
Baumbach, H. (2003) Vegetation und Landnutzung in der Karnali-Region (West-Nepal) unter besonderer Berücksichtigung der Distrikte Jumla, Mugu und Humla. In: Hartmann, M. & Baumbach, H. (Eds.), Biodiversität und Naturausstattung im Himalaya. Verein der Freunde und Förderer des Naturkundemuseums Erfurt, Erfurt, pp. 23–35.
Belousov, I.A. (1990) New carabids of the tribe Trechini (Coleoptera, Carabidae) from the Caucasus. III. 18 new species and a new subspecies of the genus Trechus Clairv. Revue d’Entomologie de l’URSS, 69, 609–632 [in Russian].
Belousov, I.A. & Kabak, I.I. (1996) To the knowledge of the Asiatic species of the genus Trechus Clairville (Insecta: Coleoptera: Carabidae). Annalen des Naturhistorischen Museums in Wien, 98B, 361–398.
Belousov, I.A. & Kabak, I.I. (2000) Contribution to the knowledge of species of the genus Trechus Clairv. (Coleoptera, Carabidae) from China. Entomological Review, 80, 12–37.
Belousov, I.A. & Kabak, I.I. (2001) New Duvalius and Trechus species from China (Coleoptera: Carabidae: Trechini). Folia Heyrovskyana, 9, 1–33.
Casale, A. (1979) Trechinae nouveaux du Népal (Col. Carabidae). Entomologica Basiliensia, 4, 187–192.
Derbyshire, E., Shi, Y., Li, J., Zheng, B., Li, S. & Wang, J. (1991) Quaternary glaciation of Tibet: The geological evidence. Quaternary Science Reviews, 10, 485–510.
Deuve, T. (1992) Contribution à la connaissance des Trechidae asiatiques (Coleoptera). Bulletin de la Société entomologique de France, 97 (2), 171–184.
Deuve, T. (1996) Nouveaux Trechus du Tibet et des régions limitrophes (Coleoptera, Trechidae). Bulletin de la Société entomologique de France, 101 (1), 65–70.
Deuve, T. (1997) Nouveaux Trechini du Tibet et des régions limitrophes (Coleoptera, Trechidae). Revue Francaise d’Entomologie (Nouvelle série), 19 (3–4), 139–154.
Deuve, T. & Schmidt, J. (2003) Sur la variation géographique de Carabus (Meganebrius) epsteini Heinertz, 1978, dans les massifs du Dhaulagiri et des Annapurna (Insecta: Coleoptera: Carabidae). In: Hartmann, M. & Baumbach, H. (Eds.), Biodiversität und Naturausstattung im Himalaya. Verein der Freunde und Förderer des Naturkundemuseums Erfurt, Erfurt, pp. 63–66.
Deuve, T. & Schmidt, J. (2005) Le genre Cychropsis Boileau, 1901, dans l’ouest du Népal Central (Coleoptera, Carabidae). L’Entomologiste, Paris, 61 (1), 3–15.
Fort, M. (1995) The himalayan glaciation: Myth and reality. Journal of Nepal Geological Society, 11, 257–272.
Fort, M. (1996) Late cenozooic environmental changes and uplift of the northern side of the central Himalaya: a reappraisal from field data. Palaeogeography, Palaeoclimatology, Palaeoecology, 120, 123–145.
Hartmann, M., Weipert, J. & Weigel, A. (1998): Die zoologischen Nepal-Expeditionen des Naturkundemuseum Erfurt 1992 bis 1997. Veröffentlichungen Naturkundemuseum Erfurt, 17, 15–30.
Hieke, F. (2003) Die Amara-Arten des Subgenus Bradytulus Tschitscherin, 1894 (Insecta: Coleoptera: Carabidae). In: Hartmann, M. & Baumbach, H. (Eds.), Biodiversität und Naturausstattung im Himalaya. Verein der Freunde und Förderer des Naturkundemuseums Erfurt, Erfurt, pp. 107–194.
Jeannel, R. (1926) Monographie des Trechinae. Morphologie comparée et distribution géographique d’un groupe de Coléoptères (Première livrasion). Abeille, Paris, 32, 221–550.
Jeannel, R. (1927) Monographie des Trechinae. Morphologie comparée et distribution géographique d’un groupe de Coléoptères (Deuxième livrasion). Abeille, Paris, 33, 1–592.
Jeannel, R. (1928) Les Trechus (Coleoptera, Carabidae) de l’Everest expedition, 1924, et remarques sur quelques espèces de l’Himalaya. The Annals and Magazine of Natural History, tenth series, 1, 283–291.
Jeannel, R. (1936) Mission scientifique de l'Omo, Coleoptera, I. Carabidae: Trechinae et Perigoninae. Mission scientifique de l'Omo, Paris, 2 [1935], 23–75.
Jeannel, R. (1954) Les Trechus du Mont Elgon (Coleoptera, Trechidae). Mémoires du Muséum National d’Histoire Naturelle, Paris, 7, 147–177.
Jeannel, R. (1960) Révision des “Trechini” du Caucase (Coleoptera, Trechidae). Mémoires du Muséum National d’Histoire Naturelle, Paris, 17, 155–216.
Klinge, M. & Lehmkuhl, F. (2004) Pleistocene glaciations in southern and eastern Tibet. In: Ehlers, J. & Gibbart, P.L. (Eds.), Quaternary glaciations – extent and chronology, Part III: South America, Asia, Africa, Australia, Antarctica. Elsevier, Amsterdam, pp. 243–254.
König, O. (2004) The glaciation of the Rolwaling Himal and the Kangchenjunga Himal during the Last Glacial Maximum (Nepal, E-Himalaya). In: Ehlers, J. & Gibbart, P.L. (Eds.), Quaternary glaciations – extent and chronology, Part III: South America, Asia, Africa, Australia, Antarctica. Elsevier, Amsterdam, pp. 279–284.
Kuhle, M. (1982) Der Dhaulagiri- und Annapurna-Himalaya. Ein Beitrag zur Geomorphologie extremer Hochgebirge. Textteil. Zeitschrift für Geomorphologie, Supplementband, 41, 1–229.
Kuhle, M. (1983) Der Dhaulagiri- und Annapurna-Himalaya. Ein Beitrag zur Geomorphologie extremer Hochgebirge. Empirische Grundlage. Gebrüder Borntraeger, Berlin, Stuttgart, 383 pp.
Kuhle, M. (2001) The Glaciation of High Asia and its Causal Relation to the Onset of Ice Age. Die Erde, 132, 339–359.
Kuhle, M. (2004) The High Glacial (Last Ice Age and LGM) ice cover in High and Central Asia. In: Ehlers, J. & Gibbart, P.L. (Eds.), Quaternary glaciations – extent and chronology, Part III: South America, Asia, Africa, Australia, Antarctica. Elsevier, Amsterdam, pp. 175–199.
Lehmkuhl, F., Klinge, M. & Lang, A. (2002) Late Quaternary glacier advances, lake level fluctuations and aeolian sedimentation in Southern Tibet. Zeitschrift für Geomorphologie, Supplementband, 126, 183–218.
Lorenz, W. (2005) A systematic list of extant ground beetles of the world (Coleoptera “Geadephaga”: Trachypachidae and Carabidae incl. Paussinae, Cicindelinae, Rhysodinae), Second edition. Published by the author, Tutzing, 530 pp.
Martens, J. (1987) Remarks on my Himalayan expeditions. Courier Forschungsinstitut Senckenberg, 93, 7–31.
Mateu, J. & Deuve, T. (1979) Nouveaux Trechus Clairville des montagnes d’Asie. Nouvelle Revue d’Entomologie, 9 (2), 97–105.
Miehe, G. (1990) Langtang Himal. Flora und Vegetation als Klimazeiger und -zeugen im Himalaya. A Prodromus of the Vegetation Ecology of the Himalayas. Dissertationes Botanicae, 158, 1–529 + 3 maps.
Miehe, G. (1991) Der Himalaya, eine multizonale Gebirgsregion. In: Walter, H. & Breckle, S.-W. (Eds.), Ökologie der Erde, Band 4. Spezielle Ökologie der Gemäßigten und Arktischen Zonen außerhalb Euro-Nordasiens. Gustav Fischer Verlag, Stuttgart, pp. 181–230.
Miehe, G. & Miehe, S. (2000) Environtal changes in the pastures of Xizang. Marburger Geographische Schriften, 135, 282–311.
Miehe, G., Winiger, M., Böhner, J. & Yili, Z. (2001) Climatic Diagram Map of High Asia 1 : 4 000 000. Erdkunde, 55, 94–97 + 1 map.
Owen, L.A. & Benn, D.I. (2005) Equilibrium-line altitudes of the Last Glacial Maximum for the Himalaya and Tibet: an assessment and evaluation of results. Quaternary International, 138–139, 55–78.
Pawlowski, J. (1973) Espéces bulgares du genre Trechus Clair. (Coleoptera, Carabidae). Acta Zoologica Cracoviensia, 18, 217–270.
Pawlowski, J. (1979) Révision du genre Trechus Clairv. (Coleoptera, Carabidae) du Proche Orient. Acta Zoologica Cracoviensia, 23, 247–476.
Penev, L., Casale, A. & Turin, H. (2003) Biogeography. In: Turin, H., Penev, L., & Casale, A. (Eds.), The genus Carabus in Europe. A Synthesis. Pensoft Publishers, Sofia, Moscow, pp. 327–425.
Royden, L.H., Burchfiel, B.C. & van der Hilst, R.D. (2008) The Geological Evolution of the Tibetan Plateau. Science, 321, 1054–1058.
Schmidt, J. (1994) Beschreibungen neuer Arten der Carabidae aus Nepal (1): Gattungen Trechus, Deltomerodes, Pterostichus (Insecta: Coleoptera). Reichenbachia, 30 (21), 129–135.
Schmidt, J. (1999) Revision of the genus Calathus from Nepal (Descriptions of new species of Carabidae from Nepal Himalayas, part 6) (Coleoptera Carabidae Sphodrini). In: Zamotajlov, A. & Sciaky, R. (Eds.), Advances in Carabidology. Papers dedicated to the memory of Prof. Dr. Oleg L. Kryzhanovskij. MUISO Publishers, Krasnodar, pp. 299–318.
Schmidt, J. (2006) Die Pterostichus-Arten des Subgenus Pseudethira Sciaky, 1996, in Zentral- und West-Nepal (Coleoptera, Carabidae): Taxonomie, Phylogenie, Biogeographie. In: Hartmann, M. & Weipert, J. (Eds.), Biodiversität und Naturausstattung im Himalaya. Verein der Freunde und Förderer des Naturkundemuseums Erfurt, Erfurt, pp. 179–243.
Schmidt, J. (2007) Neue Pseudethira-Funde im Annapurna-Massiv im westlichen Zentral-Nepal-Himalaya mit Hinweisen zur Lage von Glazialrefugien im oberen Kali Gandaki- und Marsyangdi Khola-Tal (Coleoptera, Carabidae, Pterostichus). Veröffentlichungen Naturkundemuseum Erfurt, 26, 13–25.
Schmidt, J. & Hartmann, M. (2001) Revision der Notiophilus-Arten Nepals (Coleoptera, Carabidae). Veröffentlichungen Naturkundemuseum Erfurt, 20, 151–164.
Schmidt, J. & Hartmann, M. (2009) Pristosia Motschulsky, 1865 from the Nepal Himalaya: Taxonomy and Biogeography (Coleoptera: Carabidae: Sphodrini). Zootaxa, 2009, 1–26.
Shi, Y., Zheng, B. & Li, S. (1992) Last glaciation and maximum glaciation in the Qinghai-Xizang (Tibet) Plateau: A controversy to M. Kuhle’s ice sheet hypothesis. Zeitschrift für Geomorphologie, N.F., Supplement, 84, 19–35.
Shilenkov, V.G. (1992) The high altitude fauna of South Siberian Mountains and its Origin (Coleoptera: Carabidae). In: Noonan, G.R., Ball, G.E. & Stork, N.E. (Eds.), The Biogeography of Ground Beetles of Mountains and Islands. Intercept, Andover-Hampshire, pp. 53–65.
Spicer, R.A., Harris, N.B.W., Widdowson, M., Herman, A.B., Guo, S., Valdes, P.J., Wolfe, J.A., Kelley, S.P. (2003) Constant elevation of southern Tibet over the past 15 million years. Nature, 421, 622–623.
Uéno, S.-I. (1967) The Trechus (Coleoptera, Trechinae) from the Rolwaling Himal. Bulletin of the National Science Museum Tokyo, 10, 241–246.
Yin, A. & Harrison, T.M. (2000) Geologic evolution of the Himalayan-Tibetan Orogen. Annual Review of Earth Planetary Sciences, 28, 211–280.
Zhou’Li, J., Zhang, S.Q., Zhao, J.D., Cui, J.X. (2004) Quaternary glaciations in China. In: Ehlers, J. & Gibbart, P.L. (Eds.), Quaternary glaciations – extent and chronology, Part III: South America, Asia, Africa, Australia, Antarctica. Elsevier, Amsterdam, pp. 105–113.