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Bonn zoological Bulletin 61 (2): 142-209 December 2012 A revision of the Lathrobium species of the Himalaya (Coleoptera: Staphylinidae: Paederinae) VoikerAssing Gabelsbergerstr. 2. D-30163 Hannover, Germany; E-mail: [email protected]. Abstract. In all, 48 namedLathrobium species, all ofthemmicropterous, microphthalmous, and locally endemic, arere- cognised in the Himalaya, among them 32 species new to science: L. aciforme sp. n. (C-Nepal), L. amiapwnense sp. n. (C-Nepal), L. apalatum sp. n. (C-Nepal), L. attritum sp. n. (C-Nepal), L. barbatum sp. n. (C-Nepal), L. barbulatum sp. n. (C-Nepal),L. bibarbatum sp. n. (NE-Nepal),L. calcaratum sp. n. (C-Nepal),L. cavicrus sp. n. (C-Nepal),L. compres- sicrus sp. n. (C-Nepal),L. compressum sp. n. (C-Neapl), L. cumim sp. n. (C-Nepal),L. diremptum sp. n. (NE-Nepal),L. discissiim sp. n. (N-lndia; E-Nepal), L. excisum sp. n. (E-Nepal), L. exsertum sp. n. (C-Nepal), L. fodens sp. n. (W-Ne- pal),L. ignoratiim sp. n. (C-Nepal), L. inexcisum sp. n. (NE-Nepal),L. infractiim sp. n. (E-Nepal),L. kleebergi sp. n. (E- Nepal), L. lamjunense sp. n. (C-Nepal), L. milkeense sp. n. (E-Nepal), L. muguicum sp. n. (NW-Nepal), L. palatum sp. n. (C-Nepal), L. planissimum sp. n. (W-Nepal), L. privum sp. n. (W-Nepal), L. rupinaicum sp. n. (C-Nepal), L. separa- tum sp. n. (N-India), L. spiculatum sp. n. (C-Nepal), L. spiiiosissimum sp. n. (C-Nepal), L. umbhakense sp. n. (NE-Ne- pal).Theexternalandsexualcharacters ofall thespeciesaredescribedandillustrated. Onegenus-groupandtwo species- groupsynonymiesareproposed: Lathrobium Gravenhorst, 1802 = Glyptomerodoschema Scheerpeltz, 1976,syn. n.;Lath- robium lassallei Coiffait, 1981 = L. sherpa Coiffait, 1982, syn. n.; L. emodense Coiffait, 1975 = L. goropanense Coif- fait, 1983, syn. n. Medonjalja/ensis (Coiffait, 1984) comb. n. is transferred from Lathrobium to the subtribe Medonina, redescribed, and illustrated. The Himalayan Lathrobium species are most unlikely to fonn a monophylum. Ten species groups are identified and characterised. In the Himalaya, Lathrobium species have been recorded only from the region between Kashmir in the west and Darjeeling (West Bengal) in the east. The altitudes range from 2400 to 5000 m. Most species have become known from central Nepal (25 species) and eastern Nepal (14 species), probably because collect- ingactivitywashighestintheseregions.Thedistributionsofthe individual speciesandofthespeciesgroupsaremapped. Acatalogue and a key to the HimalayanLathrobium species are provided. Key words. Taxonomy, rove beetles, Lathrobium, Medon, Himalaya, Nepal, India, new species, new synonymies, new combination, key to species, catalogue, diversity, vertical distribution INTRODUCTION The speciose paederine genus Lathrobium Gravenhorst, 1905, Tetartopeus Czwalina, 1888, or Pseudobium Mul- & 1802 is represented in the Palaearctic region by approxi- sant Rey, 1878 (Assing 2012a-d, in press). mately 350 described species in three subgenera (Assing Twenty-one species in two subgenera had previously 2010b; Smetana 2004; Schiilke unpubl.). However, new been recorded from the Himalaya (Coiffait 1975, 1981, species are being described every year, particularly from 1982a, b, 1983, 1984, 1987; Scheerpeltz 1976; Smetana China and Japan, suggesting that the known inventory of 2004); one ofthem is the type species ofthe monotypi- the East Palaearctic Lathrobium fauna is far from com- cal subgenus Glyptomerodoschema Scheerpeltz, 1976. plete. An accurate estimate oftotal species number is dif- Coiffait (1982b) provided a keytothe 21 HimalayanLath- ficult, since numerous species from regions otherthan the robium species known to him at thattime, butthis key in- Palaearctic have not been revised and may belong to oth- cluded several species ofLobrathium, as well as one of er genera ofLathrobiina. unknown generic and subtribal affiliations (Assing in The available biogeographic data suggestthatthe genus prep.). A revision ofthe Himalayan Lathrobium species has a Holarctic distribution. In the East Palaearctic, its has never been attempted. range extends toTaiwan in the southeast, where theLath- Unlike other lathrobiine genera such as Lobrathium, robium species are confined to high-altitude habitats Pseudolathra, and Pseudobium, Lathrobium species (Assing 2010b). In India, the genus is known only from and/or species groups are often distinguished also by the high-altitude habitats in the Himalaya.All the species pre- female secondary sexual characters (Assing 2010b, viously recorded from India (including Sri Lanka) and 2012a). The morphology ofthe female stemite VIII and Myanmar as Lathrobium by Cameron (1931) refertoLo- the female tergite IX are ofparticular taxonomic signifi- & brathium Mulsant Rey, 1878, Pseudolathra Casey, cance. The female sexual characters ofHimalayan Lath- robium species were previously unknown. Received: 14.04.2012 Corresponding editor: D. Ahrens Accepted: 10.10.2012 Revision ofHimalayan Lathrobium 143 While intheWest Palaearctic region aconsiderablepro- SNSD Senckenberg Naturkundliche Sammlungen portion ofthe species is either macropterous or wing-di- Dresden, Germany (O. Jager) moiphic, more or less wide-spread, and often found at low ZII Zoologisches Institut Innsbruck, Austria (W. elevations, the vast majoritiy of species in the East Schedl) Palaearctic is micropterous and more or less locally en- cAss author's private collection demic to individual mountain ranges. This is particularly cFel private collectionBenediktFeldmann, Miinster, true ofthe fauna ofregions such as the Himalaya, China, Germany and Japan. Not a single winged true Lathrobium species cKle private collection Andreas Kleeberg, Berlin, has been recorded from the Himalaya. Gennany The present study was inspired primarily by material cSme private collectionAles Smetana, Ottawa, Cana- made available to me by Benedikt Feldmann (Miinster) da and Andreas Kleeberg (Berlin). In order to identify the species represented in this material, types had to be ex- amined. Eventually, the typematerial ofall the species de- RESULTS scribed from the Himalaya and additional material from various public andprivate collections were studied, yield- Taxonomic changes. Therevision yieldedthree new syn- ing anunexpectednumberofundescribed species, as well onymies of genus-group and species-group names. The as several new synonymies and new combinations. subgeneric name Glyptomerodoschema Scheerpeltz, 1976 is placed in synonymy with Lathrobium Gravenhorst, 1802. In the two cases of species-group synonymy, the MATERIALAND METHODS juniorsynonym(Z. sherpa Coiffait, 1982, L. goropanense Coiffait, 1983) had been described from the same locali- The morphological studies were conducted using a Ste- ty as the senior name (L. lassallei Coiffait, 1981 and L. mi SV 11 microscope (Zeiss Germany) andaJenalab com- emodenseCoiffait, 1975, respectively)by the same author, pound microscope (Carl Zeiss Jena). A digital camera andthemale sexual characters ofeitherthejunior(L. sher- (Nikon Coolpix 995) was used for the photographs. pa) or the senior synonym {L. emodense) had been un- Head length was measured from the anterior margin of known. the fi"ons to theposteriormargin ofthe head, elytral length The type series ofthreepreviously described species {L. at the suture fi^om the apex ofthe scutellum to the poste- nepalense Coiffait, 1975, L. franzi Coiffait, 1975, L. rior margin ofthe elytra, and the length ofthe aedeagus nepalorientis Coiffait, 1984) are composed ofspecimens fi^om the apex of the ventral process to the base of the ofat least two species. The paratypes that had erroneous- aedeagal capsule. The "parameral" side (i.e., the side lybeen assumedto be conspecific with the holotype were where the sperm duct enters) is referred to as the ventral, either females or had been mis-sexed as females. the opposite side as the dorsal aspect. LathrobiumjaljalenseCoiffait, 1984 is transferredtothe The maps were created using MapCreator 2.0 (primap) genusMedon Stephens ofthe subtribe Medonina. Anoth- software. The coordinates ofsome localities were obtained erspecies originally described inLathrobium,L.perpusil- fi-omAhrens (2004). lum Coiffait, 1982, belongs to a genus ofMedonina, too; it will be treated elsewhere (Assing 2012d). Both species had erroneously been attributed to Lathrobium probably COLLECTION MATERIAL DEPOSITORIES because of their faint resemblance in colour and body shape with small Lathrobium species. FSF Forschungsinstitut Senckenberg, Frankfiirt, Ger- many (A. Hastenpflug-Vesmanis) In all, 32 Lathrobium species are newly described, 30 MHNG Museum d'Histoirenaturelle, Geneve, Switzer- ofthem from Nepal, one from West Bengal (India), and one from eastern Nepal and West Bengal. land (G. Cuccodoro) MNHNP Museum national d'Histoire naturelle, Paris, France (A. Taghavian) NHMW Species groups. The Himalayan Lathrobium species are Naturhistorisches Museum Wien, Austria (H. most unlikely to represent a monophyletic group. Prima- Schillhammer) NME rily based on the male and female sexual characters, but Naturkundemuseum Erfurt, Germany (M. Hart- also on external characters, ten species groups are iden- mann, assisted by W. Apfel) tified. SDEI Senckenberg Deutsches Entomologisches Insti- The most speciose group, theL. nepalensegroup, is dis- tut, Miincheberg, Germany (L. Behne) SMNS Staatliches Museum fur Naturkunde Stuttgart, tributed in central and eastern Nepal (Figs 2-3) and in- Germany (W. Schawaller, K. Wolf-Schwennin- cludes 15 species (L. bibarbatum sp. n., L. compressum sp. n., L. diremptum sp. n., L. exsertum sp. n., L. ignora- ger) Bonn zoological Bulletin 61 (2): 142-209 ©ZFMK 144 VolkerAssing turn sp. n., L. inexcisum sp. n., L. infractum sp. n., L. TheL. deuveigroup isrepresentedby eight species from janetscheki Scheerpeltz, 1976, L. khumbuense Coiffat, westem andcentral Nepal (Fig. 126), all ofthem confined 1982, L. kleebergi sp. n., L. lassallei, L. milkeense sp. n., to high-altitude habitats above 4000 m: L. aciforme, L. L. nepalense, L. nepalorientis, L. umbhakense sp. n.). apalatum sp. n.,L. bartheiCoiffait, 1987, L. deuvei Coif- Species ofthis group are characterisedbyusuallydarkcol- fait, 1981,L. ganeshense Coiffait, 1983,L. lamjunense sp. oration, small to moderately large body size, moderately n., L. palatum sp. n., L. rupinaicum sp. n. The species of small eyes composedofapproximately20-30 ommatidia, this group are characterised by moderately large to large a relatively weakly oblong (approximately 1.15 times as body size (length offorebody > 3.0 mm), conspicuously long as broad) and - in cross-section - strongly convex slender habitus with relatively long legs and antennae, pronotum with more or less pronounced microsculpture minute eyes composed ofvery few ommatidia, a strong- (e.g.. Figs 5,18, 28, 43), a male stemite VII with orwith- ly transverse and posteriorly excavate male stemite VII out weakly modified pubescence (e.g. Figs 6, 11, 19, 23, (e.g.. Figs 127, 132), a distinctly transverse male stemite 29, 35, 39), the shape ofthe male stemite VIII (posterior VIII with a deep (exception: L. ganeshense) and broadly margin with orwithout very small median excision, as in V-shapedposteriorexcision andmostlywith conspicuous Figs 7, 12-13, 20, 24, sometimes even convex inthe mid- fringes orclusterofdense dark setae (e.g.. Figs 128, 133), dle, as in Figs 30, 66), a distinctly sclerotised aedeagus a relatively large (1.3-2.0 mm) aedeagus with a long, with a lamellate dorsal plate (e.g.. Figs 8, 14-15, 21, strongly sclerotised, and apically acute ventral process, 25-26), and the presence ofmicropubescence in the pos- with a dorsal plate composed ofa lamellate basal portion terior portion of the female stemite VIII (e.g. Figs 10, and a distinctly sclerotised (not lamellate), apically 16-17, 22, 28). In addition, the female tergite IX is sep- hooked apical portion, and with long, slender, distinctly arated by a nan^ow membranous suture in the middle in sclerotised intemal stmctures (e.g.. Figs 129, 134-135); most species. the female stemite VIII is often strongly producedposte- TheL. mugiiicum group includes onlyone species from riorly (e.g.. Fig. 136). northwestern Nepal (Fig. 96), L. muguicum sp. n. It is The L. gladiator group includes only a single species, somewhat similarto the species oftheL. nepalensegroup, L. gladiatorfrom Kashmir, and is characterisedby apos- but has slightly larger eyes (composed ofapproximately teriorly laterally compressed, stronglybulging female ter- 30 ommatidia), a slightly more slender and less convex giteX (Figs 123-124), anaedeaguswithalongandstrong- pronotum without microsculpture (Fig. 87), a distinctlyV- ly sclerotised ventral process, with a dorsal plate com- shaped posterior excision ofthe male stemite VIII (Fig. posed ofa lamellate basal and a strongly sclerotised api- 89), and a different morphology ofthe aedeagus (dorso- calportion (Figs 120-121), a stronglytransversemale ster- ventrally flattenedventral process, lamellate dorsalplate, nite VII with an extensive cluster ofmodified setae (Fig. internal sac with long membranous tube, but without scle- 118), and by the shape ofthe male stemite VIII (posteri- rotised stmctures) (Figs 90-91). or margin produced on either side ofthe posterior exci- The L. emodense group comprises four species from sion) (Fig. 119). central Nepal (Fig. 96): L. annapurnense sp. n., L. emod- TheL. discissum group isrepresentedbytwo described ense, L. cwvum sp. n., and L. spinosissimum sp. n. They (and one undescribed) species from West Bengal (North are characterisedby relatively small body size, reddishto India) and the extreme east ofNepal (Figs 3, 126): L. dis- dark-brown coloration, moderately small eyes composed cissum sp. n. and L. separatum sp. n. It is distinguished ofapproximately 20-25 ommatidia, the presence ofmod- from other species groups particularly by the stmcture of ifiedsetae on the male stemiteVIII (Figs 93, 99, 104, 109) the female tergite IX (Fig. 174), which is divided into two X and often also on stemite VII (Figs 92, 98), the shape of hemi-tergites, so that the anterior margin oftergite al- themale stemiteVIII (posteriormargin sometimes point- most reaches the anterior margin of tergite IX (unique ed on either side of the median excision, as in Figs 93, among HimalayanLathrobium), and alsobythe morphol- 104), the morphology ofthe aedeagus (dorsal plate api- ogy of the aedeagus (ventral process apically narrowly cally at least weakly hooked, long, and stout, not lamel- tmncate inventralview; dorsalplatereduced; intemal sac late; ventralprocess not flat; intemal stmctures with scle- withmembranous tube,butwithout sclerotised spines; see rotised intemal stmctures) (Figs 94, 100, 105), andanan- Figs 171-172, 178-179) and by the unmodified pubes- teriorly undivided female tergite IX. cence ofthe male stemites VII and VIII (Figs 176-177, TheL. excisum group includes only a single species, L. 169-170). excisum sp. n. from eastem Nepal (Fig. 96). It is similar TheL.jumlensegroup includesthree species fromwest- to the species oftheL. emodense group, but distinguished em Nepal (Fig. 96), L.jumlense Coiffait, 1982, L. inus- primarily by aedeagal characters (dorsal plate lamellate; tum Coiffait, 1982, and L. planissimum sp. n., which are intemal structures without sclerotised spines, as in Figs characterised by small body size, uniformly reddish col- 115-116). oration, a conspicuously small (< 0.7 mm), weakly scle- rotised, and - in ventral view - broad aedeagus (Figs Bonn zoological Bulletin 61 (2): 142-209 ©ZFMK ^^^^^^^ FmOuRSsCHeUuNGmS ^^mKOENIG Bonn ndex 2012 Volume 61 zoological 1-2 Issue Bulletin formerly: Bonner zoologische Beitrage Bonn Volume61 zoological '^ou Bulletin Editor-in-Chief Fabian Herder Editorial Board Dirk Alirens Wolfgang Bohme Bonn Volume61 Netta Dorchin zoological '^2*0,2 Bulletin Renate van den Elzen Bernhard A. Huber Rainer Hutterer Gustav Peters Bradley Sinclair Dieter Stuning Topfer Till Wagner Philipp An open access journal of organismal zoology, published by Zoologisches Forschungsmuseum Alexander Koenig, Bonn Contents Volume 61 Assing, Volker: 49--128 A revision of East Palaearctic Lobrathium (Coleoptera: Staphylinidae: Paederinae) Assing, Volker: 142-209 A revision of tine Lathrobium species of the Himalaya (Coleoptera: Staphylinidae: Paederinae) Assing, Volker: 210-215 The genus Trisunius in the Himalaya (Coleoptera: Staphylinidae: Paederinae: Medonina) Auliya, Mark, Philipp Wagner & Wolfgang Bohme: 255-281 The herpetofauna of the Bijagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist Faucheux, Michel Joel: 129-134 The urticating apparatus in the larva of the Lappet Moth, Streblote panda Hubner, 1820 (Lepidoptera: Lasiocampidae) Koerber, Stefan: 29-30 Mercediella nom. nov., a replacement name for Camposichthys Figueiredo & Silva Santos, 1991 (Pisces: Pycnodontiformes) Nadein, Konstantin & Chi-Feng Lee: 41-48 New data about some Alticini from Taiwan with descriptions of two new species (Coleoptera: Chrysomelidae) Rajaei Sh., Hossein & Dieter Stuning 135-•139 Scotopteryx ^uznetzow (Wardikian, 1957) (Lepidoptera: Geometridae: Larentiinae), a new species for the fauna of Iran and Turkey Rosier, Herbert, Ivan Ineich, Thomas M. Wilms & Wolfgang Bohme: 241-254 Studies on the taxonomy of the Gekko vittatus Houttuyn, 1782 complex (Squamata: Gekkonidae) 1. On the variability of G. vittatus Houttuyn, 1782 sensu lato, with the description of a new species from Palau Islands, Micronesia Rosa, Gongalo M., Paolo Eusebio Bergo, Angelica Crottini & Franco Andreone: 31-34 Report of the life colouration of the enigmatic burrowing skink Voeltzkowia rubrocaudata (Grandidier, 1869) from southwestern Madagascar Sonnenberg, Rainer & Eckhard Busch: 13-28 Description of Scriptaphyosemion wieseae (Cyprinodontiformes: Nothobranchiidae), a new species from northern Sierra Leone Sonnenberg, Rainer & Jouke R. Van der Zee: 3-12 Aphyosemion pseudoelegans (Cyprinodontiformes: Nothobranchiidae), a new killifish species from the Cuvette centrale in the Congo Basin (Democratic Republic of Congo) Wagner, Philipp, Aaron M. Bauer & Wolfgang Bohme: 216-240 Amphibians and reptiles collected by Moritz Wagner, with a focus on the ZFMK collection Wagner, Philipp, Dennis Rodder & Thomas M. Wilms: 35-40 New data on the morphology and natural history of Tetradactylus ellenbergeri (Angel, 1922) (Sauria: Gerrhosauridae) and Trachylepis ivensii (Socage, 1879) (Sauria: Scincidae) in northeastern Zambia Publication dates: Vol. 61 Issue 1: July 2012 Vol. 61 Issue 2: December 2012 Revision ofHimalayan Lathrobiiim 145 184-185, 191-192), unmodified pubescence ofthe male westernmost representative is from Kashmir, the eastern- stemites VII and VIII (Figs 182-183, 189-190), and a most species from Darjeeling district in West Bengal, symmetric posterior excision ofthe male stemite VIII. northern India. The region with the highest diversity of The two species ofthe L. aculeatum group, L. aculea- described species is central Nepal (25 species), followed tiim Coiffait, 1982 and L. spiculatum sp. n., are distrib- by eastern Nepal (14 species) and westem Nepal (seven uted in central Nepal (Fig. 96). They are similar to the species). Only three species are known from northem In- species ofthe L.jumleme group, but distinguished by a dia, one from Kashmir and two from Darjeeling district long (>1.0 mm) and slenderaedeagus witha conspicuous- in West Bengal. These figures, however, are probably ly long and thin, needle-shaped ventral process (Figs strongly biased as a result ofthe different collecting ac- 196-197, 203-212), and a somewhat asymmetric poste- tivity in these regions. As far as the Staphylinidae is con- rior excision ofthe male stemite VIII (Figs 195, 202). cemed, central Nepal is without doubt the most frequent- The speciose L. pectinatum group includes eleven ly visited and best studied region in the Himalaya. Not a species from central and eastern Nepal (Fig. 214): L. at- single species has been recorded from the Himalaya east tritiim sp. n., L. barbatiim sp. n., L. barbulatum sp. n., L. ofWest Bengal, a poorly studied region from which on- calcaratiim, L. cassagnaui Coiffait, 1982, L. cavicrus sp. ly very little material ofStaphylinidae has become avail- n.,L. compressicnis sp. n.,L.fodens sp. n.,L.franziCoif- able. There is no evidence suggesting that the Lathrobi- fait, 1975, L. pectinatum Coiffait, 1981, L. privum sp. n. um fauna ofthe eastem Himalaya should be any less di- They are readily distinguished from all other Himalayan verse than that ofNepal. Moreover, the only Lathrobium species groups by the presence ofone or more transverse species known from the westem Himalaya west ofNepal rows ofpectinate setae onthe male stemite VII (e.g., Figs isL. gladiatorfrom Kashmir. Not a single specieshasbeen 215, 219), undoubtedly a synapomorphy constituting the recorded from the region between the type locality ofL. monophyly of this species group. The males of some gladiator and Nepal, a distance ofnearly 800 km! species, in some cases even the females, have conspicu- Without exception, all the currently known Himalayan ously modified metafemora, mesotibiae, and/or metatib- Lathrobium species are micropterous, microphthalmous, iae (Figs 225, 230, 237, 244, 251). and locally endemic. Many ofthem are known only from a single locality. Interestingly, the species groups have rather restricted distributions, too. None of the ten Diversity and biogeography. In all, 48 described species species groups (see preceding section) is distributedacross ofLathrobium are now known from the Himalaya. The all ofNepal (Figs 2, 3, 96, 126, 214). 2500 3000 3500 4000 4500 5000 altitude(m) Fig. 1. Pooled vertical distribution ofHimalayanLathrobium species. Bonn zoological Bulletin 61 (2): 142-209 ©ZFMK 146 VolkerAssing Catalogue ofthe Lathrohiiim species ofthe Himalaya In the catalogue, the species are listed alphabetically. The references in the catalogue are abbreviated as follows: App = present paper; C75 = Coiffait (1975), C81 = Coiffait (1981), C82a = Coiffait (1982a), C82b = Coiffait (1982b), C83 = Coiffait (1983), C84 = Coiffait (1984), C87 = Coiffait (1987), S76 = Scheerpeltz (1976). Species Distribution References Altitude aciforme sp. n. C-Nepal: Annapuma App 4500-4700 m m aculeatum Coiffait, 1982 C-Nepal: N-Dhaulagiri, App, C82a 3300-3400 annapiiniense sp. n. C-Nepal: Annapuma App 4900 m apalatum sp. n. C-Nepal: S-Manaslu: Meme Pokhari App 4300^400 m attritiim sp. n. C-Nepal: Bagmati prov.: Yardang ridge App 3250 m barbatum sp. n. C-Nepal: N-Annapuma App 3000-3500 m barbulatum sp. n. C-Nepal: N-Annapuma App 3050 m ^arr/je; Coiffait, 1987 W-Nepal: Jumla: Mt. Mahidoela App, C75, C87 5000 m = alticola Coiffait, 1975 bibarbatum sp. n. NE-Nepal: Taplejung district App 3400-3600 m calcaratum sp. n. C-Nepal: Kali-Gandaki valley App 7 cassagnaui Co'i{{ait, 1982 eastem C-Nepal: Mt. Kalinchock App, C82b 3000 m cavicrus sp. n. C-Nepal: Manaslu App 2800-3300 m compressicrus sp. n. C-Nepal: Manaslu App 3800^100 m compressum sp. n. eastem central Nepal, NNE Kathmandu, App 2900 m Bagmati province, Shermathang in Bairavkund Lekh cun'wn sp. n. C-Nepal: Dhaulagiri App 2700-2900 m m (/ewve/ Coiffait, 1981 C-Nepal: Manaslu: Himal Chuli App, C81 4300 diremptum sp. n. NE-Nepal: Taplejung district App 3250-3500 m discissum sp. n. N-India: W-Bengal: Darjeeling; App 2700-3100 m E-Nepal: Panchthar district W m emodense Coiffait, 1975 C-Nepal: Pokhara, Ghorepani env. App, C75, C83 3050-3100 =goropanense Coiffait, 1983; syn. n. exciswn sp. n. E-Nepal: Rolwaling Himal App 3300-3800 m exsertum sp. n. C-Nepal: Bagmati province: App 3100^800 m N Kathmandu, Yangri ridge and environs ofThare Pati fodens sp. n. W-Nepal: Jumla district App 3550^100 m NNW /ra«zz Coiffait, 1975 C-Nepal: Kathmandu App, C75 7 m ganeshense Coiffait, 1983 C-Nepal: Ganesh Himal App, C83 4300-4500 m gladiator Co\ifdi\U 1982 Kashmir: Pir Panjal pass App, C82b 3200-3700 ignoratum sp. n. C-Nepal: N Kathmandu, App 3600-3800 m western Langtang region m inexcisiim sp. n. NE-Nepal: Taplejung district App 3250-3700 infractiim sp. n. E-Nepal: Solukhumbu district App 3000 m m inustum Coiffait, 1982 W-Nepal: Jumla: Maharigaon env. C82a 3000-4000 janetscheki Scheei-peltz, 1976 E-Nepal: Kliumbu: Pangpoche env. App, S76 7 jumlense Coiffait, 1982 W-Nepal: Jumla: Dampa pass near ChantaApp, C82a 3500 m khumbuense Coiffait, 1982 E-Nepal: Khumbu: Lukla App, C82a, C83 3000^000 m m kleebergi sp. n. E-Nepal: Rolwaling Himal App 2400-3300 m lamjunense sp. n. C-Nepal: Lamjun Himal App 4300-4600 /flS5a//e/ Coiffait, 1981 eastem central Nepal, Mt. Kalinchock = sherpa Coiffait, 1982, syn. n. environs, region NE Barahbise App, C81, C82b 3000-3250 m milkeense sp. n. E-Nepal: Milke Himal App 2500 m mitguicum sp. n. NW-Nepal: Mugu district App 3200 m m nepalense Co'iiiaW, 1975 C-Nepal: Bagmati province App, C75 2800-3500 nepalorientis Co\fia\{, 1984 E-Nepal: Jaljale Himal App, C84 4000 m palatum sp. n. C-Nepal: Annapuma App 5000 m m pectinatum Coiffait, 1981 E-Nepal: Manaslu App, C81 3000-3500 planissimiim sp. n. NW-Nepal: Mahakali App 3450 m privum sp. n. W-Nepal: Jumla App 3500 m rupinaicum sp. n. C-Nepal: S-Manaslu App 4100-4500 m separatum sp. n. N-India: W-Bengal: Darjeeling App 2700-3100 m spiculatum sp. n. eastem C-Nepal: Langtang App 2900-4800 m spinosissimum sp. n. C-Nepal: Lamjun Himal App 3700 m umbhakense sp. n. E-Nepal: Sankhua Sabha and App 3450^200 m Taplejung districts Bonn zoological Bulletin 61 (2): 142 -209 ®ZFMK Revision ofHimalayanLathrobium 147 Natural history. As was to be expected based on the hy- long, shaped as in Figs 178-179; posteriormargin of pothesised Holarctic distribution of the genus, the Hi- stemite VII very weakly concave in the middle (Fig. malayan Lathrobium species are absent from lower ele- 176); stemite VIII with shallow posteriorexcision and vations. The altitudes range from 2400 to 5000 m, with with sparse pubescence (Fig. 177). N-India: West m most species occurring between 3000 and 4500 (Fig. Bengal: Darjeeling district (Fig. 3) 1). Species ofthe L. deiivei group appear to be specially L. separatum sp. n. adapted and confined to high-altitude habitats above 4000 3. Species of usually dark coloration; at least head, m. As far as can be inferred from the data specified from pronotum, and abdomen dark-brown to blackish- the labels, fi-om Franz' unpublished collection notes, as brown. Head andpronotum with microsculpture. Eyes well as from unpublished reports ofcolleagues who col- moderately small, composed ofat least approximate- lectedLathrobium inNepal, the species live in the leaflit- ly 20 ommatidia. Pronotum ratherweakly oblong, ap- ter of montane shrubland and forests (alder, birch, fir, proximately 1.15 times as longasbroad, broaderthan, spruce, rhododendron, etc.) (Figs 295-297). Species ofthe or approximately asbroad as head, and strongly con- L. deuveigroup have also been found in unforested, alpine vex in cross-section (e.g.. Figs 5, 18). Body size small habitats. On several occasions, two or more Lathrobium to moderately large; length offorebody 2.3-3.8 mm. species, sometimes ofthe same species group, were col- (5*: stemite VII with weakly modified pubescence at lected in the same localities. Teneral adults were found in most; stemite VIII often with modified setae, poste- April (three species). May (one species), June (two riorly at most with very shallow posterior excision species), September (one species), and October (one (e.g. Figs 20, 24), sometimes even produced in the species). middle (e.g.. Fig. 30); aedeagus usually rather com- pact, though often with long and acute ventral process, andwithmoderately sclerotised dorsal plate KEY TO THE HIMALAYAN LATHROBIUM at most. $ tergite IX mostly with membranous su- : SPECIES ture inthemiddle; stemiteVIII withmicropubescence posteriorly. Central and eastemNepal(Figs 2-3). The 1. S'- aedeagus without sclerotised dorsal plate, with L. nepalense group 4 long, dark membranous tube, but without distinctly - Pronotrmi without microsculpture, less convex in sclerotised structures in internal sac; ventral process cross-section, and almost always more slender. Eyes of aedeagus slender, apically narrowly truncate in mostly smallerand composed ofless than 20 omma- ventral view (Figs 171-172, 178-179); stemites VII tidia. Coloration mostly paler. S'- stemite VIII pos- andVIII withoutdistinctlymodifiedpubescence (Figs teriorly with deeper excision, never produced in the 169-170, 176-177). $: tergite IX distinctly divided middle; dorsal plate of aedeagus in dark-coloured into two hemi-tergites (Fig. 174); tergite X almost species strongly sclerotised. $ tergite IX neverwith : reaching anterior margin oftergite IX. Species from median suture; stemite VIII usually without micro- the extreme east ofNepal and from Darjeeling (In- pubescence posteriorly 18 dia). The Lathrobium discissum group 2 4. Smaller species; length offorebody at most 3.0 mm. - S'- aedeagus mostly with at least weakly sclerodsed Species from eastemNepal 5 dorsal plate and often with slerotised internal struc- - Larger species; length of forebody at least 3.0 mm. tures; ventral process of aedeagus apically not nar- Species from central and eastemNepal 11 rowly truncate in ventral view; stemites VII and/or 5. S'- stemiteVIII without modifiedsetae,posteriormar- VIII oftenwith distinctlymodified chaetotaxy. $ ter- gin broadly and weakly concave (Fig. 40); aedeagus : gite IX anteriorly atmostwith fine, membranous me- as in Fig. 41. 9 stemite VIII as in Fig. 42. Khumbu : dian suture; anterior portion oftergite X clearly dis- (Fig. 2) L. khumbuenseCoiffait tant from anterior margin oftergite IX 3 - S'- stemite VIII with modified setae, posterior mar- 2. Larger species: body length 7.5-9.0 mm; length of gin either with small and shallow median concavity forebody at least 3.1 mm. S'- aedeagus approximate- orproduced inthe middle; aedeagus ofdifferentmor- mm ly 1.5 long, shaped as in Figs 171-172; posteri- phology 6 or margin of stemite VII distinctly concave in the 6. (J : posterior margin of stemite VIII at least weakly middle (Fig. 169); stemiteVIII withmoderately deep convex in the middle 7 posterior excision andwith moderately dense pubes- - S: posterior margin ofstemite VIII weakly concave cence (Fig. 170). E-Nepal: Panchthar district; N-In- in the middle 8 dia: West Bengal: Darjeeling district (Fig. 126) 7. Smaller species; length offorebody 2.3-2.6 mm. <S: L. discissum sp. n. stemite VIII approximately as long as broad, on ei- Smaller species: body length 5.5-6.5 mm; length of ther side of middle with weakly defined cluster of mm mm forebody 3.0 at most. <S: aedeagus 0.9-1.0 dense setae, andwith distinctly convexposteriormar- Bonn zoological Bulletin 61 (2): 142-209 ©ZFMK : 148 VolkerAssing gin (Fig. 76); aedeagus smaller, approximately 0.8 lydefinedclusters ofmoderately dense setae andwith mm long, ventral process symmetric in ventral view less strongly convex posterior margin (Fig. 71); mm andweakly bent in lateral view (Figs 77-79). $: ster- aedeagus approximately 1.0 long and with less nite VIII strongly produced posteriorly (Fig. 81). Ta- slender ventral process in lateral and in ventral view plejung district (Fig. 3) L. inexcisum sp. n. (Figs 72-73). $: stemiteVIII as in Fig. 74. Taplejung - Larger species; length of forebody 2.6-3.0 mm. <S: district (Fig. 2) L. diremptum sp. n. stemite VIII distinctly transverse, with median clus- - S'- stemite VII with posteriormargin broadl and less ter ofmoderately dense black setae, and with weak- distinctly concave (Fig. 65); stemiteVIII with distinct ly convex posterior margin (Fig. 83); aedeagus larg- cluster ofvery dense setae on either side ofmiddle mm er, approximately 1.0 long, ventral process and with strongly convex posteriormargin (Fig. 66); mm strongly asymmetric in ventral view and stronglybent aedeagus 1.1 long and with more slender ven- in lateral view (Figs 84-85). $: sexual characters un- tral process both in lateral and in ventral view (Figs known. Solukhumbu district (Fig. 3) 67-68). $: stemite VIII as in Fig. 69. Taplejung dis- L. infractum sp. n. trict (Fig. 3) L. bibarbatum sp. n. 8. S'- ventral process of aedeagus strongly bent in lat- 14. S'- middle ofposterior margin ofstemite VIII weak- eral view (Figs 50, 54 9 ly concave, without additional modifications 15 - S'- ventral process ofaedeagus weakly bent and api- - (S: middle ofposterior margin ofstemite VIII either callyvery acute in lateral view (e.g.. Figs 60-63) ...10 convex, or somewhat elevated and with pair ofclus- mm 9 S'- aedeagus larger, approximately 1.05 long; ters ofdense black setae 17 ventral process ofaedeagus symmetric inventral view 15. S'- apical portion ofventral process ofaedeagus dis- and apically more acute in lateral view; dorsal plate tinctly elongate, slender, symmetric, and dorso-ven- strongly sclerotised and longer (Figs 54-55). Milke trally compressed (Figs 37-38). Species ofrelative- Himal (Fig. 2) L. milkeense sp. n. ly large size; length of forebody 3.4-3.6 mm. Bag- - <S'- aedeagus slightly smaller, 0.95 mm long; ventral mati province: Bairavkund Lekh (Fig. 2) process ofaedeagus distinctly asymmetric in ventral L. compressum sp. n. view and apically less acute in lateral view; dorsal - S'- apicalportion ofventralprocess ofaedeagus short- plate less strongly sclerotised and shorter (Figs er, less slender, and not dorso-ventrally compressed 50-51). Mt. Everest range (Fig. 2) 16 L.janetscheki Scheerpeltz 16. S'- apex ofventral process ofaedeagus very acute in 10. S'- stemite VIII approximately as long as wide (Fig. lateral view and slightly asymmetric in ventral view; 45); aedeagus shaped as in Figs 46-47. Jaljale Himal dorsal plate longer (Figs 25-27). $: stemite VIII as (Fig. 2) L. nepalorientis Coiffait in Fig. 28. Bagmati province (Fig. 2) - S'- stemite VIII transverse (Fig. 59); aedeagus L. ignoratum sp. n. shaped as in Figs 60-63. $: stemite VIII as in Fig. - S'- apex ofventral process stouter; dorsal plate short- 64. Sankhua Sabha and Taplejung districts (Fig. 3) er(Figs 14-15). $ stemiteVIII as in Figs 16-17. En- : L. umbhakense sp. n. virons ofKalinchok (Fig. 2) ... L. lassallei Coiffait 11. Species from eastern Nepal (east of 86°10'E longi- 17. S'- stemite VIII on either side ofmiddle with rather tude) 12 extensive clusterofdense black setae, posteriormar- - Species from central Nepal (west of 86°10'E longi- gin convex in the middle (Fig. 30); aedeagus with tude) 14 apex ofventral process shorter and stouter in lateral 12. S'- stemiteVIII with extensive median clusterofmod- view; dorsal plate shorter (Figs 31-32). $: stemite ified setae andwith broadly andweakly concavepos- VIII more strongly produced posteriorly (Fig. 33). teriormargin (Fig. 20); aedeaguswithrelatively short Bagmati province (Fig. 3) L. exsertum sp. n. and apically not very acute ventral process in lateral - S: stemite VIII with median cluster ofdense setae, view (Fig. 21). $: stemite VIII as in Fig. 22; tergite posteriormargin somewhatproduced and elevated in IX without median suture. Rolwaling Himal (Fig. 2) the middle, this elevation with pair of clusters of L. kleebergi sp. n. dense setae (Fig. 7); aedeagus with apex of ventral - S: stemite VIII with a cluster ofless distinctly mod- process longer, more slender, and more acute; dorsal ified setae on either side ofmiddle and with distinct- plate longer (Figs 8-9). $: stemite VIII less strong- lyconvexposteriormargin; aedeagus with longerand ly produced posteriorly (Fig. 10). Bagmati province apicallyvery acute ventralprocess in lateral view. $ (Fig. 2) L. nepalenseCoiffait tergite IX with median suture 13 18. Eyes moderately small, composed ofat least approx- 13. S'- stemite VII with posterior margin distinctly con- imately 20 ommatidia. Coloration ofbody brown to cave in the middle (Fig. 70); stemiteVIII withweak- blackish-brown, rarely uniformly reddish {L. emod- Bonn zoological Bulletin 61 (2); 142-209 ®ZFMK Revision ofHimalayan Lathrobium 149 mm ense, L. excisum). Mostly species of intermediate as in Fig. 92; aedeagus 1.2 long; ventral process body size (length offorebody < 3.1 mm). Nepal east- symmetric and shaped as in Figs 94-95; dorsal plate wards to Rolwaling Himal 19 only indistinctly hooked apically; internal structures - Eyes smaller, composed ofapproximatelyten omma- long andweakly curved in lateral view. Region to the tidia atmost. Eitherlarge and slender species (length west ofPokhara (Fig. 96) of forebody > 3.0 mm) of dark-reddish to reddish- L. emodense Coiffait brown coloration or small species ofmore or less uni- - S'- stemiteVIII with denserand more numerous mod- fonnly reddish coloration 24 ified setae in the middle; aedeagus with ventral 19. Largerspecies; length offorebody approximately 3.4 process ofdifferent shape and sometimes asymmet- mm. Eyes larger, composedofapproximately 30 om- ric in ventral view; dorsal plate more distinctly mm matidia. S: aedeagus (Figs 90-91) larger, 1.5 hooked in lateral view; intemal stmctures shorter, long, with slender, dorso-ventrally compressed ven- more strongly curved in lateral view, and ofsimilar tral process; internal sac with longmembranoustube, length 23 but without distinctly sclerotised internal structures; 23. S: stemite VII with more numerous stouterand short- dorsal plate flat and shorter; pubescence of stemite er modified setae in median portion (Fig. 98); stem- VIII not distinctly modified (Fig. 89). Northwestern ite VIII with more strongly modified (shorter and Nepal (Fig. 96). The L. muguicum group stouter) setae in posterior median portion, posterior L. muguicum sp. n. excision shallower and broader (Fig. 99); aedeagus - Smaller species; length offorebody < 3.1 mm. Eyes larger, approximately 1.3 mm long, shaped as inFigs smaller, composed of approximately 20-25 omma- 100-101; ventral process of aedeagus somewhat mm tidia. S'- aedeagus smaller, 0.8-1.3 long; ventral asymmetric in ventral view; dorsal plate longer; in- process less slenderin lateral view and dorso-ventral- temal stmcturesregularlycurved. Dhaulagiri (Fig. 96) ly not compressed; stemite VIII and mostly also VII L. curvum sp. n. with modified setae in the middle; posterior margin - <S: stemiteVII with fewerandless strongly modified ofstemite VIII oftenpointed on either side ofposte- setae in median portion (Fig. 103); stemite VIII with rior excision. Central Nepal and eastem Nepal ... 20 few and less stronglymodified setae in posteriorme- 20. S'- aedeagus with dark membranous tube, but with- dian portion, posterior excision much narrower (Fig. mm out sclerotised spines in intemal sac and with lamel- 104); aedeagus smaller, approximately 1.0 long, late dorsal plate, ventral process of characteristic shaped as in Figs 105-106; dorsal plate shorter; in- shape (Figs 115-116); stemite VII with modified, temal sac with differently shaped stmctures ofdiffer- stout black setae in median portion (Fig. 113); ster- ent lengths. $: stemite VIII as in Fig. 125. Annapur- nite VIII weakly transverse and with stout black se- na range: southem Lamjun Himal (Fig. 96) tae inmedianportion, posteriormarginpointed on ei- L. spinosissimum sp. n. ther side ofmiddle (Fig. 114). $ stemite VIII weak- 24. Moderately large to large species (length offorebody : ly oblong (Fig. 114). Eastem Nepal: Rolwaling Hi- > 3.0 mm) ofreddish to dark-brown coloration and mal (Fig. 96). The L. excisum group conspicuously slenderhabitus. Eyes minute and com- L. excisum sp. n. posed ofvery few ommatidia. S: stemiteVII strong- - S'- aedeagus with sclerotised intemal stmctures in in- ly transverse and strongly excavate posteriorly (e.g.. temal sac; dorsal plate stout (not flat), long, and api- Figs 127, 132); stemiteVIII transverse, with deep (ex- cally more or less distinctly hooked in lateral view. ception: L. ganeshense) and broadly V-shaped pos- CentralNepal. TheL. emodense group 21 terior excision and mostly with conspicuous fringes 21. Smaller species, length offorebody 2.4-2.5 mm. S'- or a cluster ofdense dark setae (e.g.. Figs 128, 133); mm mm aedeagus smaller, 0.83 long, shaped as in Fig. aedeagus (e.g.. Figs 129, 134-135) large, 1.3-1.9 110; intemal sac with two weakly sclerotised, not long; ventral process long and apically acute; dorsal spine-shaped stmctures; pubescence ofstemites VII plate composed ofa lamellate basal and a distinctly and VIII rather sparse (Figs 108-109). $: stemite sclerotised and apically hooked apical portion; inter- VIII as in Fig. 111. Annapuma (Fig. 96) nal sac with long, slender, and distinctly sclerotised L. annapurnense sp. n. stmctures. 9^ stemite VIII often strongly produced - Onaverage largerspecies; lengthofforebody2.4-2.9 posteriorly (e.g., Fig. 136). WestemandcentralNepal mm. S'- aedeagus > 1.0 mm long and of different (Fig. 126). Known only from high elevations above shape; intemal sac with at least four strongly sclero- 4000 m. The L. deuvei group 25 tised, more or less curved, spine-like stmctures ... 22 - Small species (length offorebody < 3.0 mm). Body 22. S- stemiteVIII with sparserand less numerous mod- of more or less uniformly reddish coloration. Male ified setae in the middle, posteriormargin pointed on sexual characters different 32 eitherside ofposteriorexcision (Fig. 93); stemite VII 25. Largest Himalayan representative of the genus, Bonn zoological Bulletin 61 (2): 142-209 ©ZFMK

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