Bonn zoological Bulletin formerly: Bonner zoologische Beitrage Volume 61 Issue 2 ZO An open access journal of organismal zoology, published by Zoologisches Forschungsmuseum Alexander Koenig, Bonn Bonn zoological Bulletin (BzB), formerly “Bonner zoologische Beitrage’, is published by the Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn. Two regular issues of BzB are published per year; supplements on focus topics are produced in irregular succession. Subscription price is 46 € per volume (year), including shipping costs. For subscription, back issues and institutional exchange, please contact the ZFMK library (ZFMK, Bibliothek, Frau Mareike Bollen, Adenauerallee 160, 53113 Bonn, Germany, tel. +49 228—-9122-216, fax: +49 228—-9122—212; m.bollen@zfmk.de). The online version of BzB is available free of charge at the BzB homepage: http://www.zoologicalbulletin.de. © 2012 Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany. ISSN 2190-7307 Produced by Eva-Maria Levermann, Kaiserstr. 129, 53113 Bonn, Germany; emlevermann@netcologne.de. Printed and bound by DCM, Werner-von-Siemens-Str. 13, 53340 Meckenheim, Germany. Bonn zoological Bulletin EDITOR-IN-CHIEF Fabian Herder, Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Ichthyology Section, Adenauerallee 160, 53113 Bonn, Germany, tel. +49 228-9122-255, fax: +49 228-9122—212; f.herder.zfmk@uni-bonn.de MANAGING EDITOR SUPPLEMENT SERIES Thomas Wesener, ZFMK, tel. +49 228-9122-425, fax: +49 228-9122-212; twesener@uni-bonn.de EDITORIAL BOARD Dirk Ahrens, Insects: Coleoptera, ZFMK, tel. +49 228-9122-286, fax: +49 228—-9122-332; d.ahrens.zfmk@uni-bonn.de Wolfgang Béhme, Amphibians and Reptiles, ZFMK, tel. +49 228-9122—250, fax: +49 228-9122-212; w.boehme.zfmk@uni-bonn.de Netta Dorchin, Insects: Diptera, Department of Zoology, Tel Aviv University, Tel Aviv 69978, Israel, tel. +972-3- 6409808, fax: +972-3-6409403; ndorchin@post.tau.ac.11 Renate van den Elzen, Birds, ZFMK, tel. +49 228-9122—231, fax: +49 228-9122-212; r.elzen.zfmk@uni-bonn.de Bernhard Huber, Invertebrates except Insects, ZFMK, tel. +49 228-9122-294, fax: +49 228-9122-212; b.huber.zfmk@uni-bonn.de Rainer Hutterer, Mammals, ZFMK, tel. +49 228-9122—261, fax: +49 228-9122-212; r.hutterer.zfmk@uni-bonn.de Gustav Peters, Mammals, ZFMK, tel. +49 228-9122—262, fax: +49 228-9122-212; g.peters.zfmk@uni-bonn.de Bradley Sinclair, Canadian National Collection of Insects, Ottawa Plant Laboratory — Entomology, CFIA, K.W. Neat- by Bldg., C.E.F., 960 Carling Ave., Ottawa, ON, Canada K1A 0C6, tel. + 1 613-759-1787, fax: + 1 613-759-1927; bradley.sinclair@inspection.ge.ca Dieter Stiining, Insects except Coleoptera and Diptera, ZFMK, tel. +49 228—9122—220, fax: +49 228-9122-212; d.stuening.zfmk@uni-bonn.de Till Tépfer, Birds, ZFMK, tel. + 49 228-9122-246, fax: +49 228-9122-212; t.toepfer@zfmk.de Philipp Wagner, Villanova University, Department of Biolo- gy, 800 Lancaster Avenue, Villanova, PA 19085-1699, USA; philipp.wagner.zfmk(@uni-bonn.de ADVISORY BOARD Theo C. M. Bakker, Rheinische Friedrich-Wilhelms-Univer- sitat, Institut fir Evolutionsbiologie & Okologie, 53113 Bonn, Germany, tel. +49 228—73—5130, fax: +49 228—73— 2321; t.bakker@uni-bonn.de Aaron M. Bauer, Villanova University, Department of Biolo- gy, 800 Lancaster Avenue, Villanova, PA 19085-1699, USA, tel. +1 610-519-4857, fax: +1 610-519-7863; aaron.bauer@villanova.edu Wieslaw Bogdanowicz, Museum and Institute of Zoology, Polish Academy of Sciences, Wilcza 64, 00-679 Warszawa, Poland, tel. +48 22—628—7304, fax: +48 22—629-6302; wieslawb@miiz.waw.pl Matthias Glaubrecht, Museum fiir Naturkunde Berlin, Leib- niz-Institut ftir Evolutions- und Biodiversitatsforschung an der Humboldt-Universitat zu Berlin, Invalidenstrasse 43, 10115 Berlin, Germany, tel. +49 30—2093—8504/ 8400, fax: +49 030—2093-—8565; matthias.glaubrecht@mfn- berlin.de Jeremy D. Holloway, The Natural History Museum, Depart- ment of Entomology, Cromwell Road, London, SW7 5BD, U.K.; j.holloway@nhm.ac.uk Tan Heok Hui, Raffles Museum of Biodiversity Research, National University of Singapore, Department of Biological Sciences, 6 Science Drive 2, #03-01, Singapore 117546, tel. +65-6516 1662, heokhui@nus.edu.sg Boris Krystufek, Slovenian Museum of Natural History, P. O. Box 290, Ljubljana, Slovenia; boris.krystufek@zrs.upr.si Wolfgang Schawaller, Staatliches Museum fir Naturkunde, Rosenstein 1, 70191 Stuttgart, Germany, tel. +49 711-8936—221, fax: +49 711—8936—100; schawaller.smns@naturkundemuseum-bw.de Ulrich K. Schliewen, Department of Ichthyology, Bavarian State Collection of Zoology, Miinchhausenstr. 21, 81247 Miinchen, Germany, tel. + 49 89-8107—110; schliewen@zsm.mwn.de Michael Schmitt, Ernst-Moritz-Arndt-Universitat, Allge- meine & Systematische Zoologie, Anklamer Str. 20, 17489 Greifswald, Germany, tel. +49 3834864242, fax: +49 3834864098; michael.schmitt@uni-greifswald.de W. David Sissom, Dept. of Life, Earth and Environmental Sciences, W. Texas A. & M. University, WTAMU Box 60808, Canyon, Texas 79016, USA, tel. +1 806-651-2578, fax: +1 806-651-2928; dsissom@mail.wtamu.edu Miguel Vences, Technical University of Braunschweig, Zoo- logical Institute, Spielmannstr. 8, 38106 Braunschweig, Germany, tel. + 49 531-391-3237, fax: + 49 31-391-8198; m.vences(@tu-bs.de Erich Weber, Eberhard-Karls-Universitat, Zoologische Schausammlung, Sigwartstr. 3, 72076 Tubingen, tel. +49 70712972616, fax +49 7071-295170; erich.weber@uni-tuebingen.de Editorial The present issue of the Bonn zoological Bulletin contains some quite comprehensive contributions. In the first two papers, Volker Assing studies the taxonomy of Himalayan rove beetles, including a revision of the genus Lathrobi- um. In this paper, Assing recognizes 48 species of these small staphylinids between Kashmir and West Bengal, including 32 species new to science. This work sheds light on the value of taxonomic work for recognizing diversi- ty in small insects, and the conspicuous correlation between species richness and collection activity highlights the importance of assessments and field explorations in remote areas. A total of 297 figures, a summarizing cat- alogue and an extensive key help making the results accessible. In his second paper in the present issue, Ass- ing likewise focuses on staphylinid beetles in the Himalaya, here on the genus 7risunius in Nepal. He pro- vides the first record of the genus for the country and adds the description of three new species, again including a key and distribution maps. Mark Auliya and co-authors also contribute to the ex- ploration of the fauna of a remote area. Their annotated check-list of the herpetofauna of the Bijagos archipelago (Guinea-Bissau, West Africa) provides several new records, and their check-list of the herpetofauna of the whole country of Guinea-Bissau sets a baseline for the recognition of anuran and reptile diversity in the area. Two additional publications also come from the field of herpetology. Herbert Résler et al. examined allopatric Gekko vittatus populations from Pacific islands that show distinct colour patterns, argue that one of these constitutes anew species, and formally describe a new taxon. Philipp Wagner et al. contribute an overview on expeditions and resulting collections of his namesake Moritz Wagner, an important German explorer of the 19" century. M. Wag- ner’s collections contain substantial amounts of amphib- ians and reptiles, including type material; much of the material has recently been transferred to the collection of Museum Koenig. The present paper is the first one in a series of contributions summarizing the knowledge on M. Wagner’s material, and provides a commented list of spec- imens valuable especially to museum curators and taxonomists. The portrait of an important new viper species collected by him and dedicated to him by two Swedish colleagues in 1984, also in BzB, constitutes the front cover image of this issue. As usual, all pdfs are online free of charge at www.zoologicalbulletin.de. Migration of the archive to the new website is nearly completed — thanks to Sebastian Hillen for his efforts! Finally, I am delighted to welcome Till Topfer as new board member. Till Topfer is ornithologist with special interest in evolutionary avian biology and systematics. In his studies, he combines collection-based morphological approaches with molecular and geographic analyses. He has recently joined the ZFMK team as curator for ornithol- ogy. A warm “welcome on board”! With best wishes for the New Year to all readers, authors, referees and “BzB-team” members, Tibi fluc Fabian Herder (Editor-in-Chief, Fish Curator at ZFMK) Bonn, December 2012 FEB 13 2013 LIBRARIES Bonn zoological Bulletin 61 (2): 142-209 A revision of the Lathrobium species of the Himalaya (Coleoptera: Staphylinidae: Paederinae) Volker Assing Gabelsbergerstr. 2, D-30163 Hannover, Germany; E-mail: vassing.hann@t-online.de. Abstract. In all, 48 named Lathrobium species, all of them micropterous, microphthalmous, and locally endemic, are re- cognised in the Himalaya, among them 32 species new to science: L. aciforme sp. n. (C-Nepal), L. annapurnense 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), LZ. 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. curvum sp. n. (C-Nepal), L. diremptum sp. n. (NE-Nepal), L. discissum sp. n. (N-India; E-Nepal), LZ. excisum sp. n. (E-Nepal), L. exsertum sp. n. (C-Nepal), L. fodens sp. n. (W-Ne- pal), L. ignoratum sp. n. (C-Nepal), L. inexcisum sp. n. (NE-Nepal), L. infractum sp. n. (E-Nepal), L. kleebergi sp. n. (E- Nepal), L. Jamjunense sp. n. (C-Nepal), L. milkeense sp. n. (E-Nepal), L. muguicum sp. n. (NW-Nepal), LZ. 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. spinosissimum sp. n. (C-Nepal), L. umbhakense sp. n. (NE-Ne- pal). The external and sexual characters of all the species are described and illustrated. One genus-group and two species- group synonymies are proposed: 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. Medon jaljalensis (Coiffait, 1984) comb. n. is transferred from Lathrobium to the subtribe Medonina, redescribed, and illustrated. The Himalayan Lathrobium species are most unlikely to form 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- ing activity was highest in these regions. The distributions of the individual species and of the species groups are mapped. A catalogue and a key to the Himalayan Lathrobium species are provided. Key words. Taxonomy, rove beetles, Lathrobium, Medon, Himalaya, Nepal, India, new species, new synonymies, new December 2012 combination, key to species, catalogue, diversity, vertical distribution INTRODUCTION The speciose paederine genus Lathrobium Gravenhorst, 1802 is represented in the Palaearctic region by approx1- mately 350 described species in three subgenera (Assing 2010b; Smetana 2004; Schiilke unpubl.). However, new species are being described every year, particularly from China and Japan, suggesting that the known inventory of the East Palaearctic Lathrobium fauna is far from com- plete. An accurate estimate of total species number is dif- ficult, since numerous species from regions other than the Palaearctic have not been revised and may belong to oth- er genera of Lathrobiina. The available biogeographic data suggest that the genus has a Holarctic distribution. In the East Palaearctic, its range extends to Taiwan in the southeast, where the Lath- robium species are confined to high-altitude habitats (Assing 2010b). In India, the genus is known only from high-altitude habitats in the Himalaya. All the species pre- viously recorded from India (including Sri Lanka) and Myanmar as Lathrobium by Cameron (1931) refer to Lo- brathium Mulsant & Rey, 1878, Pseudolathra Casey, Received: 14.04.2012 Accepted: 10.10.2012 1905, Tetartopeus Czwalina, 1888, or Pseudobium Mul- sant & Rey, 1878 (Assing 2012a-—d, in press). Twenty-one species in two subgenera had previously been recorded from the Himalaya (Coiffait 1975, 1981, 1982a, b, 1983, 1984, 1987; Scheerpeltz 1976; Smetana 2004); one of them is the type species of the monotypi- cal subgenus Glyptomerodoschema Scheerpeltz, 1976. Coiffait (1982b) provided a key to the 21 Himalayan Lath- robium species known to him at that time, but this key in- cluded several species of Lobrathium, as well as one of unknown generic and subtribal affiliations (Assing in prep.). A revision of the Himalayan Lathrobium species has never been attempted. Unlike other lathrobiine genera such as Lobrathium, Pseudolathra, and Pseudobium, Lathrobium species and/or species groups are often distinguished also by the female secondary sexual characters (Assing 2010b, 2012a). The morphology of the female sternite VIII and the female tergite [IX are of particular taxonomic signifi- cance. The female sexual characters of Himalayan Lath- robium species were previously unknown. Corresponding editor: D. Ahrens Revision of Himalayan Lathrobium 143 While in the West Palaearctic region a considerable pro- portion of the species is either macropterous or wing-di- morphic, more or less wide-spread, and often found at low elevations, the vast majoritiy of species in the East Palaearctic is micropterous and more or less locally en- demic to individual mountain ranges. This is particularly true of the fauna of regions such as the Himalaya, China, and Japan. Not a single winged true Lathrobium species has been recorded from the Himalaya. The present study was inspired primarily by material made available to me by Benedikt Feldmann (Munster) and Andreas Kleeberg (Berlin). In order to identify the species represented in this material, types had to be ex- amined. Eventually, the type material of all the species de- scribed from the Himalaya and additional material from various public and private collections were studied, yield- ing an unexpected number of undescribed species, as well as several new synonymies and new combinations. MATERIAL AND METHODS The morphological studies were conducted using a Ste- mi SV 11 microscope (Zeiss Germany) and a Jenalab com- pound microscope (Carl Zeiss Jena). A digital camera (Nikon Coolpix 995) was used for the photographs. Head length was measured from the anterior margin of the frons to the posterior margin of the head, elytral length at the suture from the apex of the scutellum to the poste- rior margin of the elytra, and the length of the aedeagus from the apex of the ventral process to the base of the aedeagal capsule. The “parameral” side (1.e., the side where the sperm duct enters) is referred to as the ventral, the opposite side as the dorsal aspect. The maps were created using MapCreator 2.0 (primap) software. The coordinates of some localities were obtained from Ahrens (2004). COLLECTION MATERIAL DEPOSITORIES FSFE Forschungsinstitut Senckenberg, Frankfurt, Ger- many (A. Hastenpflug-Vesmanis) Muséum d’ Histoire naturelle, Geneve, Switzer- land (G. Cuccodoro) MNHNP Muséum national d’Histoire naturelle, Paris, France (A. Taghavian) NHMW Naturhistorisches Museum Wien, Austria (H. Schillhammer) MHNG NME Naturkundemuseum Erfurt, Germany (M. Hart- mann, assisted by W. Apfel) SDEI Senckenberg Deutsches Entomologisches Insti- tut, Muincheberg, Germany (L. Behne) SMNS | Staatliches Museum fir Naturkunde Stuttgart, Germany (W. Schawaller, K. Wolf-Schwennin- ger) Bonn zoological Bulletin 61 (2): 142-209 SNSD_ Senckenberg Naturkundliche Sammlungen Dresden, Germany (O. Jager) ZI Zoologisches Institut Innsbruck, Austria (W. Schedl) cAss author’s private collection cFel private collection Benedikt Feldmann, Miinster, Germany cKle private collection Andreas Kleeberg, Berlin, Germany cSme __ private collection Ales Smetana, Ottawa, Cana- da RESULTS Taxonomic changes. The revision yielded three new syn- onymies of genus-group and species-group names. The subgeneric name Glyptomerodoschema Scheerpeltz, 1976 is placed in synonymy with Lathrobium Gravenhorst, 1802. In the two cases of species-group synonymy, the junior synonym (L. sherpa Coiffait, 1982, L. goropanense Coiffait, 1983) had been described from the same locali- ty as the senior name (L. Jassallei Coiffait, 1981 and L. emodense Coiffait, 1975, respectively) by the same author, and the male sexual characters of either the junior (L. sher- pa) or the senior synonym (L. emodense) had been un- known. The type series of three previously described species (L. nepalense Coiffait, 1975, L. franzi Coiffait, 1975, L. nepalorientis Coiffait, 1984) are composed of specimens of at least two species. The paratypes that had erroneous- ly been assumed to be conspecific with the holotype were either females or had been mis-sexed as females. Lathrobium jaljalense Coiffait, 1984 is transferred to the genus Medon Stephens of the subtribe Medonina. Anoth- er species originally described in Lathrobium, L. perpusil- lum Coiffait, 1982, belongs to a genus of Medonina, too; it will be treated elsewhere (Assing 2012d). Both species had erroneously been attributed to Lathrobium probably because of their faint resemblance in colour and body shape with small Lathrobium species. In all, 32 Lathrobium species are newly described, 30 of them from Nepal, one from West Bengal (India), and one from eastern Nepal and West Bengal. Species groups. The Himalayan Lathrobium species are most unlikely to represent a monophyletic group. Prima- rily based on the male and female sexual characters, but also on external characters, ten species groups are iden- tified. The most speciose group, the L. nepalense group, is dis- tributed in central and eastern Nepal (Figs 2—3) and in- cludes 15 species (L. bibarbatum sp. n., L. compressum sp. n., L. diremptum sp. n., L. exsertum sp. n., L. ignora- OZFMK 144 Volker Assing tum sp. n., L. inexcisum sp. n., L. infractum sp. n., L. Janetscheki Scheerpeltz, 1976, L. khumbuense Coiffat, 1982, L. kleebergi sp. n., L. lassallei, L. milkeense sp. n., L. nepalense, L. nepalorientis, L. umbhakense sp. n.). Species of this group are characterised by usually dark col- oration, small to moderately large body size, moderately small eyes composed of approximately 20-30 ommatidia, a relatively weakly oblong (approximately 1.15 times as long as broad) and — in cross-section — strongly convex pronotum with more or less pronounced microsculpture (e.g., Figs 5, 18, 28, 43), a male sternite VI with or with- out weakly modified pubescence (e.g. Figs 6, 11, 19, 23, 29, 35, 39), the shape of the male sternite VIII (posterior margin with or without very small median excision, as in Figs 7, 12-13, 20, 24, sometimes even convex in the mid- dle, as in Figs 30, 66), a distinctly sclerotised aedeagus with a lamellate dorsal plate (e.g., Figs 8, 14-15, 21, 25-26), and the presence of micropubescence in the pos- terior portion of the female sternite VII (e.g. Figs 10, 16-17, 22, 28). In addition, the female tergite [X is sep- arated by a narrow membranous suture in the middle in most species. The L. muguicum group includes only one species from northwestern Nepal (Fig. 96), L. muguicum sp. n. It 1s somewhat similar to the species of the L. nepalense group, but has slightly larger eyes (composed of approximately 30 ommatidia), a slightly more slender and less convex pronotum without microsculpture (Fig. 87), a distinctly V- shaped posterior excision of the male sternite VIII (Fig. 89), and a different morphology of the aedeagus (dorso- ventrally flattened ventral process, lamellate dorsal plate, internal sac with long membranous tube, but without scle- rotised structures) (Figs 90-91). The L. emodense group comprises four species from central Nepal (Fig. 96): L. annapurnense sp. n., L. emod- ense, L. curvum sp. n., and L. spinosissimum sp. n. They are characterised by relatively small body size, reddish to dark-brown coloration, moderately small eyes composed of approximately 20-25 ommatidia, the presence of mod- ified setae on the male sternite VIII (Figs 93, 99, 104, 109) and often also on sternite VII (Figs 92, 98), the shape of the male sternite VIII (posterior margin sometimes point- ed on either side of the median excision, as in Figs 93, 104), the morphology of the aedeagus (dorsal plate api- cally at least weakly hooked, long, and stout, not lamel- late; ventral process not flat; internal structures with scle- rotised internal structures) (Figs 94, 100, 105), and an an- teriorly undivided female tergite [X. The L. excisum group includes only a single species, L. excisum sp. n. from eastern Nepal (Fig. 96). It is similar to the species of the L. emodense group, but distinguished primarily by aedeagal characters (dorsal plate lamellate; internal structures without sclerotised spines, as in Figs 115-116). Bonn zoological Bulletin 61 (2): 142—209 The L. deuvei group is represented by eight species from western and central Nepal (Fig. 126), all of them confined to high-altitude habitats above 4000 m: L. aciforme, L. apalatum sp. n., L. barthei Coiffait, 1987, L. deuvei Coif- fait, 1981, L. ganeshense Coiffait, 1983, L. lamjunense sp. n., L. palatum sp. n., L. rupinaicum sp. n. The species of this group are characterised by moderately large to large body size (length of forebody > 3.0 mm), conspicuously slender habitus with relatively long legs and antennae, minute eyes composed of very few ommatidia, a strong- ly transverse and posteriorly excavate male sternite VII (e.g., Figs 127, 132), a distinctly transverse male sternite VII with a deep (exception: L. ganeshense) and broadly V-shaped posterior excision and mostly with conspicuous fringes or cluster of dense dark setae (e.g., Figs 128, 133), a relatively large (1.3-2.0 mm) aedeagus with a long, strongly sclerotised, and apically acute ventral process, with a dorsal plate composed of a lamellate basal portion and a distinctly sclerotised (not lamellate), apically hooked apical portion, and with long, slender, distinctly sclerotised internal structures (e.g., Figs 129, 134-135); the female sternite VIII is often strongly produced poste- riorly (e.g., Fig. 136). The L. gladiator group includes only a single species, L. gladiator from Kashmir, and is characterised by a pos- teriorly laterally compressed, strongly bulging female ter- gite X (Figs 123-124), an aedeagus with a long and strong- ly sclerotised ventral process, with a dorsal plate com- posed of a lamellate basal and a strongly sclerotised api- cal portion (Figs 120-121), a strongly transverse male ster- nite VII with an extensive cluster of modified setae (Fig. 118), and by the shape of the male sternite VIII (posteri- or margin produced on either side of the posterior exci- sion) (Fig. 119). The L. discissum group is represented by two described (and one undescribed) species from West Bengal (North India) and the extreme east of Nepal (Figs 3, 126): L. dis- cissum sp. n. and L. separatum sp. n. It is distinguished from other species groups particularly by the structure of the female tergite IX (Fig. 174), which is divided into two hemi-tergites, so that the anterior margin of tergite X al- most reaches the anterior margin of tergite IX (unique among Himalayan Lathrobium), and also by the morphol- ogy of the aedeagus (ventral process apically narrowly truncate in ventral view; dorsal plate reduced; internal sac with membranous tube, but without sclerotised spines; see Figs 171-172, 178-179) and by the unmodified pubes- cence of the male sternites VII and VIII (Figs 176-177, 169-170). The L. jumlense group includes three species from west- ern Nepal (Fig. 96), L. jumlense Coiffait, 1982, L. inus- tum Coiffait, 1982, and L. planissimum sp. n., which are characterised by small body size, uniformly reddish col- oration, a conspicuously small (< 0.7 mm), weakly scle- rotised, and — in ventral view — broad aedeagus (Figs ©ZFMK Bonn zoological Bulletin formerly: Bonner zoologische Beitrage Bonn zoological Bulletin An open access journal of organismal zoology, published by Zoologisches Forschungsmuseum Alexander Koenig, Bonn A iGeiic (3 ope acs paral el crpntaral solos pubes by Zeeloraths loectergsrmaers Aad Coen, Lean Bonn zoological Bulletin ANG ENic Volume 61 Issue 2 2012 Index. 2012 Volume 61 Issue 1-2 Editor-in-Chief Fabian Herder Editorial Board Dirk Ahrens Wolfgang BOhme Netta Dorchin Renate van den Elzen Bernhard A. Huber Rainer Hutterer Gustav Peters Bradley Sinclair Dieter StUning Till T6pfer Philipp Wagner Contents Volume 61 Assing, Volker: 49-128 A revision of East Palaearctic Lobrathium (Coleoptera: Staphylinidae: Paederinae) Assing, Volker: A revision of the Lathrobium species of the Himalaya (Coleoptera: Staphylinidae: Paederinae) 142-209 Assing, Volker: The genus Trisunius in the Himalaya (Coleoptera: Staphylinidae: Paederinae: Medonina) 210-215 Auliya, Mark, Philipp Wagner & Wolfgang Bohme: The herpetofauna of the Bijagés archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist 255-281 Faucheux, Michel Joél: The urticating apparatus in the larva of the Lappet Moth, Streblote panda Hubner, 1820 (Lepidoptera: Lasiocampidae) 129-134 Koerber, Stefan: Mercediella nom. nov., a replacement name for Camposichthys Figueiredo & Silva Santos, 1991 (Pisces: Pycnodontiformes) 29-30 Nadein, Konstantin & Chi-Feng Lee: New data about some Alticini from Taiwan with descriptions of two new species (Coleoptera: Chrysomelidae) 41-48 Rajaei Sh., Hossein & Dieter StUning Scotopteryx kuznetzovi (Wardikian, 1957) (Lepidoptera: Geometridae: Larentiinae), a new species for the fauna of Iran and Turkey 135-139 Rosler, Herbert, Ivan Ineich, Thomas M. Wilms & Wolfgang BOhme: Studies on the taxonomy of the Gekko vittatus Houttuyn, 1782 complex (Squamata: Gekkonidae) |. On the variability of G. vittatus Houttuyn, 1782 sensu lato, with the description of a new species from Palau Islands, Micronesia 241-254 Rosa, Gongalo M., Paolo Eusebio Bergo, Angelica Crottini & Franco Andreone: Report of the life colouration of the enigmatic burrowing skink Voeltzkowia rubrocaudata (Grandidier, 1869) from southwestern Madagascar 31-34 Sonnenberg, Rainer & Eckhard Busch: Description of Scriptaphyosemion wieseae (Cyprinodontiformes: Nothobranchiidae), a new species from northern Sierra Leone 13-28 Sonnenberg, Rainer & Jouke R. Van der Zee: Aphyosemion pseudoelegans (Cyprinodontiformes: Nothobranchiidae), a new killifish species 3-12 Wagner, Philipp, Aaron M. Bauer & Wolfgang Bohme: Amphibians and reptiles collected by Moritz Wagner, with a focus on the ZFMK collection 216-240 Wagner, Philipp, Dennis Rodder & Thomas M. Wilms: New data on the morphology and natural history of Tetradactylus ellenbergeri (Angel, 1922) (Sauria: Gerrhosauridae) and Trachylepis ivensii (Bocage, 1879) (Sauria: Scincidae) in northeastern Zambia Publication dates: Vol. 61 Issue 1: July 2012 Vol. 61 Issue 2: December 2012 35-40 Revision of Himalayan Lathrobium 145 184-185, 191-192), unmodified pubescence of the male sternites VII and VIII (Figs 182-183, 189-190), and a symmetric posterior excision of the male sternite VIII. The two species of the LZ. aculeatum group, L. aculea- tum Coiffait, 1982 and L. spiculatum sp. n., are distrib- uted in central Nepal (Fig. 96). They are similar to the species of the L. jumlense group, but distinguished by a long (>1.0 mm) and slender aedeagus with a conspicuous- ly long and thin, needle-shaped ventral process (Figs 196-197, 203-212), and a somewhat asymmetric poste- rior excision of the male sternite VIII (Figs 195, 202). The speciose L. pectinatum group includes eleven species from central and eastern Nepal (Fig. 214): L. at- tritum sp. n., L. barbatum sp. n., L. barbulatum sp. n., L. calcaratum, L. cassagnaui Coiffait, 1982, L. cavicrus sp. n., L. compressicrus sp. n., L. fodens sp. n., L. franzi Coif- fait, 1975, L. pectinatum Coiffait, 1981, L. privum sp. n. They are readily distinguished from all other Himalayan species groups by the presence of one or more transverse rows of pectinate setae on the male sternite VII (e.g., Figs 215, 219), undoubtedly a synapomorphy constituting the monophyly of this species group. The males of some species, in some cases even the females, have conspicu- ously modified metafemora, mesotibiae, and/or metatib- lae (Figs 225, 230, 237, 244, 251). Diversity and biogeography. In all, 48 described species of Lathrobium are now known from the Himalaya. The westernmost representative is from Kashmir, the eastern- most species from Darjeeling district in West Bengal, northern India. The region with the highest diversity of described species is central Nepal (25 species), followed by eastern Nepal (14 species) and western Nepal (seven species). Only three species are known from northern In- dia, one from Kashmir and two from Darjeeling district in West Bengal. These figures, however, are probably strongly biased as a result of the different collecting ac- tivity in these regions. As far as the Staphylinidae is con- cerned, central Nepal is without doubt the most frequent- ly visited and best studied region in the Himalaya. Not a single species has been recorded from the Himalaya east of West Bengal, a poorly studied region from which on- ly very little material of Staphylinidae has become avail- able. There is no evidence suggesting that the Lathrobi- um fauna of the eastern Himalaya should be any less di- verse than that of Nepal. Moreover, the only Lathrobium species known from the western Himalaya west of Nepal is L. gladiator from Kashmir. Not a single species has been recorded from the region between the type locality of L. gladiator and Nepal, a distance of nearly 800 km! Without exception, all the currently known Himalayan Lathrobium species are micropterous, microphthalmous, and locally endemic. Many of them are known only from a single locality. Interestingly, the species groups have rather restricted distributions, too. None of the ten species groups (see preceding section) 1s distributed across all of Nepal (Figs 2, 3, 96, 126, 214). n species 2500 3000 3500 4000 4500 5000 altitude (m) Fig. 1. Pooled vertical distribution of Himalayan Lathrobium species. Bonn zoological Bulletin 61 (2): 142-209 OZFMK 146 Volker Assing Catalogue of the Lathrobium species of the 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 aciforme sp. N. aculeatum Coiffait, 1982 annapurnense sp. n. apalatum sp. n. attritum sp. Nn. barbatum sp. n. barbulatum sp. n. barthei Coiffait, 1987 = alticola Coiffait, 1975 bibarbatum sp. n. calcaratum sp. n. cassagnaui Coiffait, 1982 cavicrus sp. n. compressicrus sp. 0. compressum sp. 0. curvum sp. n. deuvei Coiffait, 1981 diremptum sp. n. discissum sp. n. emodense Coiffait, 1975 = goropanense Coiffait, 1983; syn. n. excisum sp. N. exsertum sp. n. fodens sp. n. franzi Coiffait, 1975 ganeshense Coiffait, 1983 gladiator Coiffait, 1982 ignoratum sp. n. inexcisum sp. N. infractum sp. n. inustum Corffait, 1982 Janetscheki Scheerpeltz, 1976 jumlense Coiffait, 1982 khumbuense Coiffait, 1982 kleebergi sp. n. lamjunense sp. n. lassallei Coiffait, 1981 = sherpa Coiffait, 1982, syn. n. milkeense sp. n. muguicum sp. n. nepalense Coiffait, 1975 nepalorientis Coiffait, 1984 palatum sp. n. pectinatum Coiffait, 1981 planissimum sp. n. privum sp. n. rupinaicum sp. n. separatum sp. n. spiculatum sp. n. Spinosissimum sp. 1. umbhakense sp. n. Distribution C-Nepal: Annapurna C-Nepal: N-Dhaulagiri, C-Nepal: Annapurna C-Nepal: S-Manaslu: Meme Pokhari C-Nepal: Bagmati prov.: Yardang ridge C-Nepal: N-Annapurna C-Nepal: N-Annapurna W-Nepal: Jumla: Mt. Mahidoela NE-Nepal: Taplejung district C-Nepal: Kali-Gandaki valley eastern C-Nepal: Mt. Kalinchock C-Nepal: Manaslu C-Nepal: Manaslu eastern central Nepal, NNE Kathmandu, Bagmati province, Shermathang in Bairavkund Lekh C-Nepal: Dhaulagiri C-Nepal: Manaslu: Himal Chuli NE-Nepal: Taplejung district N-India: W-Bengal: Darjeeling; E-Nepal: Panchthar district C-Nepal: W Pokhara, Ghorepani env. E-Nepal: Rolwaling Himal C-Nepal: Bagmati province: N Kathmandu, Yangri ridge and environs of Thare Pati W-Nepal: Jumla district C-Nepal: NNW Kathmandu C-Nepal: Ganesh Himal Kashmir: Pir Panjal pass C-Nepal: N Kathmandu, western Langtang region NE-Nepal: Taplejung district E-Nepal: Solukhumbu district W-Nepal: Jumla: Maharigaon env. E-Nepal: Khumbu: Pangpoche env. References A App, C75, C83 App App App App, C75 App, C83 App, C82b App App App C82a App, S76 W-Nepal: Jumla: Dampa pass near Chanta App, C82a E-Nepal: Khumbu: Lukla E-Nepal: Rolwaling Himal C-Nepal: Lamjun Himal eastern central Nepal, Mt. Kalinchock environs, region NE Barahbise E-Nepal: Milke Himal NW-Nepal: Mugu district C-Nepal: Bagmati province E-Nepal: Jaljale Himal C-Nepal: Annapurna E-Nepal: Manaslu NW-Nepal: Mahakali W-Nepal: Jumla C-Nepal: S-Manaslu N-India: W-Bengal: Darjeeling eastern C-Nepal: Langtang C-Nepal: Lamjun Himal E-Nepal: Sankhua Sabha and Taplejung districts Bonn zoological Bulletin 61 (2): 142—209 App, C82a, C83 App App App, C81, C82b Altitude 4500-4700 m 3300-3400 m 4900 m 4300-4400 m 3250 m 3000-3500 m 3050 m 5000 m 3400-3600 m z 3000 m 2800-3300 m 3800-4100 m 2900 m 2700-2900 m 4300 m 3250-3500 m 2700-3100 m 3050-3100 m 3300-3800 m 3100-4800 m 3550-4100 m 9 4300-4500 m 3200-3700 m 3600-3800 m 3250-3700 m 3000 m 3000-4000 m 9 3500 m 3000-4000 m 2400-3300 m 4300-4600 m 3000-3250 m 2500 m 3200 m 2800-3500 m 4000 m 5000 m 3000-3500 m 3450 m 3500 m 4100-4500 m 2700-3100 m 2900-4800 m 3700 m 3450-4200 m ©ZFMK Revision of Himalayan Lathrobium 147 Natural history. As was to be expected based on the hy- pothesised Holarctic distribution of the genus, the Hi- malayan Lathrobium species are absent from lower ele- vations. The altitudes range from 2400 to 5000 m, with most species occurring between 3000 and 4500 m (Fig. 1). Species of the L. dewvei group appear to be specially adapted and confined to high-altitude habitats above 4000 m. As far as can be inferred from the data specified from the labels, from Franz’ unpublished collection notes, as well as from unpublished reports of colleagues who col- lected Lathrobium in Nepal, the species live in the leaf lit- ter of montane shrubland and forests (alder, birch, fir, spruce, rhododendron, etc.) (Figs 295-297). Species of the L. deuvei group have also been found in unforested, alpine habitats. On several occasions, two or more Lathrobium species, sometimes of the same species group, were col- lected in the same localities. Teneral adults were found in April (three species), May (one species), June (two species), September (one species), and October (one species). KEY TO THE HIMALAYAN LATHROBIUM SPECIES 1. : aedeagus without sclerotised dorsal plate, with long, dark membranous tube, but without distinctly sclerotised structures in internal sac; ventral process of aedeagus slender, apically narrowly truncate in ventral view (Figs 171—172, 178-179); sternites VII and VIII without distinctly modified pubescence (Figs 169-170, 176-177). &: tergite IX distinctly divided into two hemi-tergites (Fig. 174); tergite X almost reaching anterior margin of tergite [X. Species from the extreme east of Nepal and from Darjeeling (In- dia). The Lathrobium discissum group ............ 7) — 4: aedeagus mostly with at least weakly sclerotised dorsal plate and often with slerotised internal struc- tures; ventral process of aedeagus apically not nar- rowly truncate in ventral view; sternites VII and/or VIII often with distinctly modified chaetotaxy. 9: ter- gite [X anteriorly at most with fine, membranous me- dian suture; anterior portion of tergite X clearly dis- tant from anterior margin of tergite IX ............ 3) 2. Larger species: body length 7.5—9.0 mm; length of forebody at least 3.1 mm. ¢: aedeagus approximate- ly 1.5 mm long, shaped as in Figs 171-172; posteri- or margin of sternite VII distinctly concave in the middle (Fig. 169); sternite VIII with moderately deep posterior excision and with moderately dense pubes- cence (Fig. 170). E-Nepal: Panchthar district; N-In- dia: West Bengal: Darjeeling district (Fig. 126) SSE RNSE OA a ROR TU EE L. discissum sp. n. — Smaller species: body length 5.5—6.5 mm; length of forebody 3.0 mm at most. ¢: aedeagus 0.9-1.0 mm Bonn zoological Bulletin 61 (2): 142-209 long, shaped as in Figs 178-179; posterior margin of sternite VII very weakly concave in the middle (Fig. 176); sternite VII with shallow posterior excision and with sparse pubescence (Fig. 177). N-India: West Bengal: Darjeeling district (Fig. 3) EEE Serge S hese ee ahs sekcleceleenins L. separatum sp. n. Species of usually dark coloration; at least head, pronotum, and abdomen dark-brown to blackish- brown. Head and pronotum with microsculpture. Eyes moderately small, composed of at least approximate- ly 20 ommatidia. Pronotum rather weakly oblong, ap- proximately 1.15 times as long as broad, broader than, or approximately as broad as head, and strongly con- vex in cross-section (e.g., Figs 5, 18). Body size small to moderately large; length of forebody 2.3—3.8 mm. &: sternite VII with weakly modified pubescence at most; sternite VIII often with modified setae, poste- riorly at most with very shallow posterior excision (e.g. Figs 20, 24), sometimes even produced in the middle (e.g., Fig. 30); aedeagus usually rather com- pact, though often with long and acute ventral process, and with moderately sclerotised dorsal plate at most. 9: tergite IX mostly with membranous su- ture in the middle; sternite VIII with micropubescence posteriorly. Central and eastern Nepal (Figs 2—3). The Jb PG DOHNA PSE (3 OW 0) wracacddovacoobpacdce sicbaconasosnes 4 Pronotum without microsculpture, less convex in cross-section, and almost always more slender. Eyes mostly smaller and composed of less than 20 omma- tidia. Coloration mostly paler. @: sternite VIII pos- teriorly with deeper excision, never produced in the middle; dorsal plate of aedeagus in dark-coloured species strongly sclerotised. 9: tergite [IX never with median suture; sternite VHI usually without micro- PUbESCENCeIpPOStetionlyarweesdeacecte ese eee seen 18 Smaller species; length of forebody at most 3.0 mm. Species irommeastemNepal-.ss.ccs400-4-0cmeseee oe 3) Larger species; length of forebody at least 3.0 mm. Species from central and eastern Nepal ............ 11 &: sternite VIII without modified setae, posterior mar- gin broadly and weakly concave (Fig. 40); aedeagus as in Fig. 41. 9: sternite VIII as in Fig. 42. Khumbu CTD nO hee Sen L. khumbuense Coiffait &: sternite VIII with modified setae, posterior mar- gin either with small and shallow median concavity or produced in the middle; aedeagus of different mor- JOLOO RAY os ccc ct neue aanG Lon eRe ace Rn eA oceans anene 6 S: posterior margin of sternite VIII at least weakly Cconvexsinuhemiddlcrasyasscaseneeeeessa rece: 7 S: posterior margin of sternite VIII weakly concave imetinve middle aaa anya eee pole cneie ern tones: 8 Smaller species; length of forebody 2.3-2.6 mm. 3: sternite VIII approximately as long as broad, on ei- ther side of middle with weakly defined cluster of dense setae, and with distinctly convex posterior mar- ©OZFMK 148 10. Volker Assing gin (Fig. 76); aedeagus smaller, approximately 0.8 mm long, ventral process symmetric in ventral view and weakly bent in lateral view (Figs 77-79). 9: ster- nite VII strongly produced posteriorly (Fig. 81). Ta- plejung district (Fig. 3) ......... L. inexcisum sp. 0. Larger species; length of forebody 2.6-3.0 mm. ¢: sternite VIII distinctly transverse, with median clus- ter of moderately dense black setae, and with weak- ly convex posterior margin (Fig. 83); aedeagus larg- er, approximately 1.0 mm long, ventral process strongly asymmetric in ventral view and strongly bent in lateral view (Figs 84-85). 2: sexual characters un- known. Solukhumbu district (Fig. 3) at Sh ear aer pact a en Pass arian mtg L. infractum sp. n. 3: ventral process of aedeagus strongly bent in lat- eral views (igs s ONS 4 ress e cee easeee oer eemceiacencre 9 3: ventral process of aedeagus weakly bent and api- cally very acute in lateral view (e.g., Figs 60-63) ...10 S: aedeagus larger, approximately 1.05 mm long; ventral process of aedeagus symmetric in ventral view and apically more acute in lateral view; dorsal plate strongly sclerotised and longer (Figs 54—55). Milke imal (Figg ies sseseeseernceee L. milkeense sp. n. &: aedeagus slightly smaller, 0.95 mm long; ventral process of aedeagus distinctly asymmetric in ventral view and apically less acute in lateral view; dorsal plate less strongly sclerotised and shorter (Figs 50-51). Mt. Everest range (Fig. 2) ssbteia cues atdatbaasseia tte cease peace L. janetscheki Scheerpeltz &: sternite VIII approximately as long as wide (Fig. 45); aedeagus shaped as in Figs 46—47. Jaljale Himal (Eig 2) ch ca eee ee eee L. nepalorientis Coiffait S: sternite VIII transverse (Fig. 59); aedeagus shaped as in Figs 60-63. @: sternite VIII as in Fig. 64. Sankhua Sabha and Taplejung districts (Fig. 3) Te EP at MAAS ae nA L. umbhakense sp. n. . Species from eastern Nepal (east of 86°10’E longi- CUT AS renee as Mee OOS Pan A RE 12 Species from central Nepal (west of 86°10’E longi- [HG <)) ERR MOR Ae aa OP tN ham earns 14 &: sternite VII with extensive median cluster of mod- ified setae and with broadly and weakly concave pos- terior margin (Fig. 20); aedeagus with relatively short and apically not very acute ventral process in lateral view (Fig. 21). 2: sternite VIII as in Fig. 22; tergite IX without median suture. Rolwaling Himal (Fig. 2) ct Aura sil yatta nae SA eee L. kleebergi sp. n. &: sternite VIII with a cluster of less distinctly mod- ified setae on either side of middle and with distinct- ly convex posterior margin; aedeagus with longer and apically very acute ventral process in lateral view. 9: tergite IX with median suture -..-.-s ase 13 . Q: sternite VII with posterior margin distinctly con- cave in the middle (Fig. 70); sternite VIII with weak- Bonn zoological Bulletin 61 (2): 142-209 14. 1S: 16. Ws ly defined clusters of moderately dense setae and with less strongly convex posterior margin (Fig. 71); aedeagus approximately 1.0 mm long and with less slender ventral process in lateral and in ventral view (Figs 72-73). 2: sternite VIII as in Fig. 74. Taplejung NACE (LN, A) secogavesc50000008 L. diremptum sp. n. 3: sternite VII with posterior margin broad| and less distinctly concave (Fig. 65); sternite VII with distinct cluster of very dense setae on either side of middle and with strongly convex posterior margin (Fig. 66); aedeagus 1.1 mm long and with more slender ven- tral process both in lateral and in ventral view (Figs 67-68). 2: sternite VIII as in Fig. 69. Taplejung dis- trict(Fige Sie moe cues L. bibarbatum sp. n. 3: middle of posterior margin of sternite VIII weak- ly concave, without additional modifications...... 15 3: middle of posterior margin of sternite VIII either convex, or somewhat elevated and with pair of clus- ters ofidense blackisetaee feat. seer eee eee eee 7 3: apical portion of ventral process of aedeagus dis- tinctly elongate, slender, symmetric, and dorso-ven- trally compressed (Figs 37-38). Species of relative- ly large size; length of forebody 3.4-3.6 mm. Bag- mati province: Bairavkund Lekh (Fig. 2) POE g te ern RES IGE SS iO sanigerch L. compressum sp. 0. 3: apical portion of ventral process of aedeagus short- er, less slender, and not dorso-ventrally compressed 3: apex of ventral process of aedeagus very acute in lateral view and slightly asymmetric in ventral view; dorsal plate longer (Figs 25-27). 9: sternite VIII as in Fig. 28. Bagmati province (Fig. 2) LS Woe RCSA ae eran ee L. ignoratum sp. n. 3: apex of ventral process stouter; dorsal plate short- er (Figs 14-15). 2: sternite VIII as in Figs 16-17. En- virons of Kalinchok (Fig. 2) ... L. lassallei Coiffait &: sternite VIII on either side of middle with rather extensive cluster of dense black setae, posterior mar- gin convex in the middle (Fig. 30); aedeagus with apex of ventral process shorter and stouter in lateral view; dorsal plate shorter (Figs 31-32). 9: sternite VII more strongly produced posteriorly (Fig. 33). Bagmati province (Fig. 3)......... L. exsertum sp. n. &: sternite VIII with median cluster of dense setae, posterior margin somewhat produced and elevated in the middle, this elevation with pair of clusters of dense setae (Fig. 7); aedeagus with apex of ventral process longer, more slender, and more acute; dorsal plate longer (Figs 8-9). 2: sternite VIII less strong- ly produced posteriorly (Fig. 10). Bagmati province (CRA) Petree attain SonGesyadasecs L. nepalense Coiffait 18. Eyes moderately small, composed of at least approx- imately 20 ommatidia. Coloration of body brown to blackish-brown, rarely uniformly reddish (L. emod- ©ZFMK 19. 20. Pe 22. Revision of Himalayan Lathrobium ense, L. excisum). Mostly species of intermediate body size (length of forebody < 3.1 mm). Nepal east- Wards tOmRolwalinovklimal sees. qeccescececeee eee ee 19 Eyes smaller, composed of approximately ten omma- tidia at most. Either large and slender species (length of forebody > 3.0 mm) of dark-reddish to reddish- brown coloration or small species of more or less uni- formalyared dishyc olorationtey-naeeac--ecssecae sec 24 Larger species; length of forebody approximately 3.4 mm. Eyes larger, composed of approximately 30 om- matidia. @: aedeagus (Figs 90-91) larger, 1.5 mm long, with slender, dorso-ventrally compressed ven- tral process; internal sac with long membranous tube, but without distinctly sclerotised internal structures; dorsal plate flat and shorter; pubescence of sternite VIII not distinctly modified (Fig. 89). Northwestern Nepal (Fig. 96). The L. muguicum group BEE Reacts siieiiossis Pulctaitionmnetaly's L. muguicum sp. n. Smaller species; length of forebody < 3.1 mm. Eyes smaller, composed of approximately 20—25 omma- tidia. J: aedeagus smaller, 0.8—1.3 mm long; ventral process less slender in lateral view and dorso-ventral- ly not compressed; sternite VIII and mostly also VII with modified setae in the middle; posterior margin of sternite VIII often pointed on either side of poste- rior excision. Central Nepal and eastern Nepal ... 20 3: aedeagus with dark membranous tube, but with- out sclerotised spines in internal sac and with lamel- late dorsal plate, ventral process of characteristic shape (Figs 115-116); sternite VI with modified, stout black setae in median portion (Fig. 113); ster- nite VIII weakly transverse and with stout black se- tae in median portion, posterior margin pointed on ei- ther side of middle (Fig. 114). 2: sternite VIII weak- ly oblong (Fig. 114). Eastern Nepal: Rolwaling Hi- mal (Fig. 96). The L. excisum group a pBpOaC USO ARH BH Ect Eee REEe Tere ee ee L. excisum sp. n. &: aedeagus with sclerotised internal structures in in- ternal sac; dorsal plate stout (not flat), long, and api- cally more or less distinctly hooked in lateral view. Central Nepal. The L. emodense group ............ 21 Smaller species, length of forebody 2.4-2.5 mm. ¢: aedeagus smaller, 0.83 mm long, shaped as in Fig. 110; internal sac with two weakly sclerotised, not spine-shaped structures; pubescence of sternites VII and VIII rather sparse (Figs 108-109). 9: sternite VII as in Fig. 111. Annapurna (Fig. 96) dhe: Qe etieist saa a OE Reto eue sane L. annapurnense sp. n. On average larger species; length of forebody 2.4-2.9 mm. @: aedeagus > 1.0 mm long and of different shape; internal sac with at least four strongly sclero- tised, more or less curved, spine-like structures ... 22 ¢: sternite VIII with sparser and less numerous mod- ified setae in the middle, posterior margin pointed on either side of posterior excision (Fig. 93); sternite VII Bonn zoological Bulletin 61 (2): 142-209 24. Ie 149 as in Fig. 92; aedeagus 1.2 mm long; ventral process symmetric and shaped as in Figs 94—95; dorsal plate only indistinctly hooked apically; internal structures long and weakly curved in lateral view. Region to the west of Pokhara (Fig. 96) aie ciel pa chicacn sae I aa Re ee L. emodense Coiffait : sternite VIII with denser and more numerous mod- ified setae in the middle; aedeagus with ventral process of different shape and sometimes asymmet- ric in ventral view; dorsal plate more distinctly hooked in lateral view; internal structures shorter, more strongly curved in lateral view, and of similar Tena tla eet yer pa ere eerste aE he ee NE ee 23 . 6: sternite VII with more numerous stouter and short- er modified setae in median portion (Fig. 98); stern- ite VIII with more strongly modified (shorter and stouter) setae in posterior median portion, posterior excision shallower and broader (Fig. 99); aedeagus larger, approximately 1.3 mm long, shaped as in Figs 100-101; ventral process of aedeagus somewhat asymmetric in ventral view; dorsal plate longer; in- ternal structures regularly curved. Dhaulagiri (Fig. 96) BSR aeeicineteierten sda cross vo anneemnaseees L. curvum sp. n. &: sternite VII with fewer and less strongly modified setae in median portion (Fig. 103); sternite VIN with few and less strongly modified setae in posterior me- dian portion, posterior excision much narrower (Fig. 104); aedeagus smaller, approximately 1.0 mm long, shaped as in Figs 105-106; dorsal plate shorter; in- ternal sac with differently shaped structures of differ- ent lengths. 2: sternite VII as in Fig. 125. Annapur- na range: southern Lamjun Himal (Fig. 96) shah ota ia denis GonaHaNeac oe taamees L. spinosissimum sp. n. Moderately large to large species (length of forebody > 3.0 mm) of reddish to dark-brown coloration and conspicuously slender habitus. Eyes minute and com- posed of very few ommatidia. @: sternite VII strong- ly transverse and strongly excavate posteriorly (e.g., Figs 127, 132); sternite VII transverse, with deep (ex- ception: L. ganeshense) and broadly V-shaped pos- terior excision and mostly with conspicuous fringes or a cluster of dense dark setae (e.g., Figs 128, 133); aedeagus (e.g., Figs 129, 134-135) large, 1.3—1.9 mm long; ventral process long and apically acute; dorsal plate composed of a lamellate basal and a distinctly sclerotised and apically hooked apical portion; inter- nal sac with long, slender, and distinctly sclerotised structures. 9: sternite VIII often strongly produced posteriorly (e.g., Fig. 136). Western and central Nepal (Fig. 126). Known only from high elevations above 40 00imaiithewsdeuver eroupiees.casecs-ceeeeaeeeeeee 25 Small species (length of forebody < 3.0 mm). Body of more or less uniformly reddish coloration. Male Sexvalicharactensidittenemths-en. esac eee eels 32 Largest Himalayan representative of the genus, ©OZFMK 150 Volker Assing length of forebody > 4.0 mm. ¢@: sternite VII broad- ly and rather deeply excised posteriorly (Fig. 157); sternite VIII with deep posterior excision (Fig. 158); aedeagus shaped as in Figs 159-160. Western Nepal: Jumla region (Fig. 126) ......... L. barthei Coiffait Smaller species; length of forebody < 4.0 mm. Male sexual characters different. Central Nepal ......... 26 Body smaller; length of forebody 3.4 mm at most. 3: aedeagus smaller, 1.3—1.4 mm long, and with short- er dorsal plate (Fig. 155); posterior margin of stern- ite VII distinctly excavate and weakly bisinuate in the middle (Fig. 153); sternite VIII with deep and broad- ly V-shaped posterior excision (Fig. 154). Southern Manaslu Himal (Fig. 126) ... L. rupinaicum sp. n. Body larger; length of forebody at least 3.4 mm. 3: aedeagus larger, > 1.5 mm long, and with longer dor- sal plate. Sternite VI and/or VII of different shape andichactotaxy ss aas eee enee eee eee Asi] 27. &: posterior excision of sternite VIII relatively shal- low (Fig. 163); posterior margin of sternite VII broad- ly concave, without distinct median excavation (Fig. 162); aedeagus 1.6 mm long; ventral process apical- ly very acute and straight (Figs 164-165). 2: stern- ite VHI relatively weakly produced posteriorly (Fig. 166). Ganesh Himal (Fig. 126) Pas ea aos ee eee L. ganeshense Coiffait 3: posterior excision of sternite VIII deep; posterior margin of sternite VII with distinct median excava- tion; aedeagus of different shape. 2: sternite VIII more strongly produced posteriorly (exceptions: L. apalatum and L. deuvei). Distribution different ... 28 . GO: aedeagus with ventral process much stouter in api- cal portion (Fig. 150); sternite VII with conspicuous- ly deep and broad posterior excision, its depth dis- tinctly more than half the length of sternite (Fig. 149); sternite VII with distinctly bisinuate posterior exci- sion (Fig. 148). 2: sternite VIII transverse and obtuse- ly pointed posteriorly (Fig. 151). Southern Manaslu Himali(Fig: 126) ete eee L. apalatum sp. n. S: aedeagus with ventral process longer and more slender in apical portion; sternite VIII with less deep and/or less broad posterior excision. @: sternite VIII oblong and distinctly, convexly produced pos- teniorly (exception: Zaidemver) es. varanasi 29 29. ©: sternite VIII obtusely pointed posteriorly (Fig. 130). 3: posterior excision of sternite VII broad, mod- erately deep, and not bisinuate (Fig. 127); posterior excision of sternite VIII very deep, its depth more than half the length of sternite, and symmetric (Fig. 128); aedeagus shaped as in Fig. 129. Manaslu: Hi- maliChulin(hig< 26) nee L. deuvei Coiffait °: sternite VIII strongly and convexly produced pos- teriorly. ¢: sternites VII and/or VIII of different shape and chaetotaxy; aedeagus of different morphology. ATINAP UE A TAN PS Acree tec Soccer eee eee 30 Bonn zoological Bulletin 61 (2): 142—209 30. Sills 33% 34. ¢: aedeagus larger, 1.8-2.0 mm long, shaped as in Figs 134-135; sternite VII with deeper posterior ex- cavation (Fig. 132); posterior excision of sternite VIII somewhat asymmetric (Fig. 133). 2: sternite VI- II as in Fig. 136. Northern Annapurna (Fig. 126) SE man cane ei ore dan oaRo ne L. lamjunense sp. n. 3: aedeagus smaller, < 1.8 mm long; sternite VII with slightly less deep posterior excavation. Distribution different: 2.1/2: say. Seeece Reece een eee 31 &: ventral process of aedeagus apically sharply bent in lateral view (Fig. 140); sternite VHI with deeper and symmetric posterior excision (Fig. 139); sternite Vilas in Fig. 138. 9: sternite VIII as in Fig. 141. An- napurna: Kang La pass (Fig. 126) SESS Ns PE eee Ee L. palatum sp. n. ¢: ventral process of aedeagus apically smoothly curved in lateral view (Fig. 145); sternite VIII with less deep and slightly asymmetric posterior excision (Fig. 144); sternite VII as in Fig. 143. Q: sternite VUI as in Fig. 146. Annapurna: Pisang (Fig. 126) fete vc Scr eM cl ee CE L. aciforme sp. n. . 9: tergite X laterally compressed, strongly bulging, almost keeled in posterior portion (Figs 123-124); sternite VIII as in Fig. 122. 4: aedeagus (Figs 120-121) 1.2 mm long, with long and strongly scle- rotised ventral process; dorsal plate of aedeagus with lamellate basal and strongly sclerotised apical portion; sternite VII strongly transverse and with extensive cluster of modified setae (Fig. 118); sternite VII pro- duced on either side of posterior excision (Fig. 119). Kashmir. The L. gladiator species group hs eRe i eee eee L. gladiator Coiffait ©: tergite X weakly convex in cross-section. <: aedeagus of different morphology, dorsal plate not strongly sclerotised; sternites VI and VUI of differ- ent shape and chaetotaxy. Nepal .................. 33 &: sternite VII without transverse combs of pectinate SETAC sisi decane Sue eee 34 3: sternite VII with transverse combs of pectinate se- tae (e.g., Figs 215, 219). Central and eastern Nepal (Fig. 214). The L. pectinatum group ............... 38 &: aedeagus weakly sclerotised and slender, > 1.0 mm long, with conspicuously long and thin, needle-shaped ventral process; sternite VII weakly oblong or ap- proximately as long as broad, without distinctly mod- ified pubescence and with somewhat asymmetric pos- terior excision. 9: sternite VIII distinctly oblong. The L. aculeatum group. Central Nepal (Fig. 96) ...... 35 S: aedeagus much smaller, < 0.7 mm long, and of much broader shape, ventral process not thin and nee- dle-shaped; sternite VII without modified setae; ster- nite VII oblong, without modified setae, and with symmetric posterior excision. 9: sternite VIII distinct- ly oblong. Western Nepal (Fig. 96). The L. jumlense (100) 1) Oe ee eR eet careers sodancacdiiahtadandooadodcs 36 36. Sus 38. Revision of Himalayan Lathrobium . G: aedeagus 1.2—1.3 mm long, ventral process ex- tremely long, longer than basal portion of aedeagus (Figs 203-212); posterior margin of sternite VII in- distinctly concave, almost truncate (Fig. 201); stern- ite VIII approximately as long as broad and with broader posterior excision (Fig. 202). 9: sternite VIII as in Fig. 213; tergite X shorter than tergite [X in the middle. Bagmati province: region ca. 40 km north of Kathmandu (Fig. 96) L. spiculatum sp. n. ¢: aedeagus approximately 1.1 mm long, ventral process shorter than basal portion of aedeagus (Figs 196-197); posterior margin of sternite VII more dis- tinctly concave (Fig. 194); sternite VIII oblong and with narrower posterior excision (Fig. 195). 9: ster- nite VIII as in Fig. 198; tergite X longer than tergite IX in the middle. Northern Dhaulagiri range (Fig. 96) > SOG ENO DEER RES ce OE Cane EES L. aculeatum Coiffait ¢: aedeagus dorso-ventrally distinctly flattened and apically less acute in ventral view (Figs 277-279); posterior excision of sternite VIII broadly V-shaped (Fig. 276). Extreme northwest of Nepal: Mahakali re- aon (WEI, SIS) soooossooonsasecds L. planissimum sp. n. S: aedeagus dorso-ventrally not distinctly flattened, ventral process apically acute in ventral view; poste- rior excision of sternite VIII not distinctly V-shaped. umlasre stoni (194196) eee erecece one ee vareee ee ee 37 S: sternite VII relatively weakly transverse and with weakly concave posterior margin (Fig. 182); poste- rior excision of sternite VIII deeper (Fig. 183); aedea- gus much larger, approximately 0.65 mm long and relatively narrower in ventral view (Figs 184-185). ©: sternite VIII slightly less slender (Fig. 186) 2 cogs OS SOOO HERE CHORE Seen enone L. jumlense Coiffait G: sternite VII distinctly transverse and with more strongly concave posterior margin (Fig. 189); poste- rior excision of sternite VII less deep and almost of semi-circular shape (Fig. 190); aedeagus minute (smaller than in any other Himalayan Lathrobium species), 0.33 mm long and relatively broader in ven- tral view (Figs 191-192). 9: sternite VIII slightly more slender (Fig. 193) ......... L. inustum Coiffait &: metatibia modified: either conspicuously flattened, or in basal third with pronounced postero-ventral tooth and in apical half distinctly excavate (note that females of most of the species are unknown and that this may not always be a secondary sexual character); ventral process of aedeagus shorter, not needle- shaped; sternite VII with at least two complete trans- verse combs of pectinate setae. Central Nepal: Man- aslurandAnnapumaman Cesereeepereecne er erenoee: 39 ¢: metatibia moderately flattened at most, otherwise unmodified; ventral process of aedeagus long, thin and mostly somewhat needle-shaped; sternite VII with one or two transverse rows of pectinate setae. si eigiiae Sect mieten gee Re es eran teeters Sah aicdrnctahciok 43 Bonn zoological Bulletin 61 (2): 142-209 39} 40. 41. 42. 43. 15] ¢: metatibia conspicuously flattened, without tooth; ventral process of aedeagus of broadly triangular shape and symmetric in ventral view. Central Nepal: Manasluinane creerraanacascnce nce eae neree eae 40 ¢: metatibia with distinct postero-ventral tooth in basal third and strongly excavate in apical half (Fig. 225); ventral process of aedeagus more slender and distinctly asymmetric in ventral view. Central Nepal: Northern Annapurna and Manaslu ranges ......... 41 Apical half of mesotibia conspicuously flattened in both sexes. @: ventral process of aedeagus broader and slightly longer (Figs 221—222); sternite VII with two uninterrupted and two interrupted transverse rows of pectinate setae (Fig. 219); sternite VIII as in Fig. 220. Y: sternite VIII as in Fig. 223. Barapokhari Lekh (Big@21A) rsh ee Sas L. compressicrus sp. n. Mesotibia not conspicuously flattened. @: ventral process of aedeagus less broad and slightly shorter (Figs 217-218); sternite VII with two complete and a rudimentary third transverse row of pectinate setae (Fig. 215); sternite VIII as in Fig. 216. Distribution: By fo24 7 Ca Sas Screen eon aca ae L. pectinatum Coiffait &: sternite VII with posterior margin and posterior transverse combs of pectinate setae almost straight (Fig. 284); ventral process of aedeagus almost sym- metric in ventral view and apically less acute in lat- eral view (Figs 286-287). 9: sternite VIII relatively longer (Fig. 289). Manaslu range (Fig. 214) A GS DHE ORG ODOR CROC AROC UC Aase aan L. cavicrus sp. 0. &: sternite VII with posterior margin and posterior transverse combs of pectinate setae distinctly con- cave; ventral process of aedeagus distinctly asymmet- ric in ventral view and apically more acute in later- al view. 2: sternite VIII relatively shorter (Fig. 231). INorthermAnnapumasrange’. sane cac-see eee neeenee 42 3: sternite VII with apical row of pectinate setae of flatly trapezoid shape (Fig. 226); aedeagus shaped as in Figs 228-229; sternite VIII as in Fig. 227. 9: ster- nite VII as in Fig. 231. Distribution: Fig. 214 AGAROSE SACS UNRATE CaN arerS L. barbatum sp. n. S: sternite VII with apical row of pectinate setae curved (Fig. 232); aedeagus shaped as in Figs 234-235; sternite VII as in Fig. 233. Distribution: OA Da Ge eres eychde eae sia erase sya sie L. barbulatum sp. n. ¢: metafemur distinctly modified, with postero-ven- tral tooth-like process approximately in the middle (e.g., Figs 237, 244); sternite VI with one or two transverse rows of pectinate setae; aedeagus with long and more or less needle-shaped ventral process ... 44 S: metafemur not conspicuously modified; sternite VII with one transverse row of — in one species rel- atively sparse — pectinate setae; sternite VIII without pubescence in the middle and with somewhat asym- INE Ih CIPOSLEMIOMEXCISIONE Ns ee ee seer eeeeseeeecceeeeL: 46 15 2 Volker Assing mA) 44. %: tooth-like process of metafemur apically acute (Fig. 237); sternite VI with two long transverse rows of pectinate setae (Fig. 238); sternite VIII with weak- ly asymmetric posterior excision (Fig. 239); aedea- gus approximately 1.0 mm long, ventral process somewhat spear-shaped and symmetric in ventral view (Figs 240-241). Central Nepal: Bagmati provin- ce (Fig. 214) L. franzi Coiffait &: tooth-like process of metafemur not distinctly acute apically; sternite VII with one transverse row of pectinate setae; aedeagus at least approximately 1.1 mm long; ventral process asymmetric and not spear- shaped in ventral view 45 45. 4: metafemur with symmetric and somewhat axe- shaped process (Fig. 251); sternite VII with straight transverse row of pectinate setae and with weakly concave posterior margin (Fig. 252); sternite VIII with asymmetric posterior excision (Fig. 253); aedea- gus with much longer and more slender ventral process somewhat resembling a golf club (Figs 254- 255). Q: sternite VIII strongly oblong (Fig. 256). West Nepal: Jumla district (Fig. 214) ... LZ. privum sp. n. 3: metafemur asymmetric, not axe-shaped (Fig. 244); sternite VII with flatly trapezoid transverse row of pectinate setae, posterior margin with trapezoid ex- cavation in the middle (Fig. 245); posterior excision of sternite VII almost symmetric (Fig. 246); ventral process shorter, less slender, and somewhat scoop- 46. 47. shaped (Figs 247-248). Central Nepal: Kali-Ganda- ki valley (valley separating the Annapurna and Dhaulagiri ranges) (Fig. 214) HE oe SSE DN Tae OO L. calcaratum sp. n. 3: ventral process of aedeagus gradually narrowed toward apex and not needle-shaped in ventral view (Figs 271—272); sternite VII posteriorly with few and not contiguous pectinate setae (Fig. 269); sternite VIII as in Fig. 270. Bagmati province, Yardang ridge west of Barahbise (Fig. 214) ...... L. attritum sp. n. 3: ventral process thin and slender, not gradually nar- rowed towards apex in ventral view; sternite VII pos- teriorly with transverse comb of numerous contigu- ous pectinate setae 47 3: aedeagus 1.3 mm long, ventral process longer and more slender, distinctly needle-shaped (Figs 266-267); sternite VII with transverse row of pecti- nate setae bisinuate (Fig. 264); posterior excision of sternite VIII smaller (Fig. 265). 9: sternite VIII shaped as in Fig. 268. West Nepal: Jumla region (Fig. 214) L. fodens sp. n. ¢S: aedeagus 1.15 mm long, ventral process shorter and stouter (Figs 260-261); sternite VII with almost straight transverse row of pectinate setae (Fig. 258); posterior excision of sternite VIII larger (Fig. 259). ®: sternite VIII as in Fig. 262. Central Nepal: Kalin- chok (Fig. 214) L. cassagnaui Coiffait THE HIMALAYAN LATHROBIUM SPECIES The Lathrobium nepalense group Fig. 2. Distributions of species of the L. nepalense group: L. ignoratum (open diamonds); L. nepalense (filled squares); L. com- pressum (open star); L. /assaillei (filled triangles); L. kleebergi (open circles); L. khumbuense (open triangle); L. janetscheki (filled diamond); L. nepalorientis (filled star); L. milkeense (open square); L. diremptum (filled circles). Bonn zoological Bulletin 61 (2): 142-209 ©ZFMK Revision of Himalayan Lathrobium 153 Fig. 3. Distributions of species of the L. nepalense (open symbols) and the L. discissum groups (filled circle): L. exsertum (open triangles); L. infractum (open star); L. umbhakense (open squares); L. bibarbatum (open diamond); L. inexcisum (open circles); L. separatum (filled circle). Lathrobium nepalense Coiffait, 1975 (Figs 2, 4-10) Lathrobium nepalensis [sic] Coiffait, 1975: 182. Type material examined. Holotype @: “Zw. Mulkharka u. Tare-Pati / Pa 158 [overleaf] / Zentral-Nepal, Sept.-Okt. 1971, lg. H. Franz / Holotype / Lathrobium nepalense H. Coiffait 1974 / Lathrobium nepalense Coiffait, det. V. Ass- ing 2011” (NHMW). Paratypes [see also type material of L. ignoratum]: 19: same data as holotype, but “Paratype” (NHMW). Comment. The original description is based on a male holotype and two female paratypes from “Entre Mulkhar- ka et Tare-Pati”, one female from “bois au-dessous de Fu- lung”, and one female from “chemin de Pokhara vers Goropani” (Coiffait 1975). The holotype and two paratypes from the Franz collection at the NHMW were examined; the “female” from Fulung proved to be a male of an undescribed species (see description of L. ignora- tum). In view of the generally restricted distributions of Himalayan Lathrobium species, there is little doubt that the female paratype from the environs of Pokhara, too, refers to a different, probably undescribed species. The aedeagus of the holotype is damaged: the apical portion of the ventral process is broken off. It was evidently un- damaged when Coiffait (1975) figured it. Additional material examined. Nepal: 22 exs. [partly tener- al], Bagmati province, below Thare Pati, 3300 m, 10.IV.1981, leg. Lobl & Smetana (cSme, cAss); 15 exs. [partly teneral], same data, but 11.IV.1981 (cSme, cAss); 1 ex., same data, but 9.1V.1981 (cSme); 10 exs. [partly teneral], same data, but 3400 m, 13.I1V.1981 (cSme, cAss); 26 exs., same data, but 3500 m, Bonn zoological Bulletin 61 (2): 142-209 12.1V.1981 (cSme, cAss); 11 exs., Bagmati province, Malem- chi, 2800 m, 18.IV.1981, leg. Lobl & Smetana (cSme, cAss); 10 exs. [partly teneral], Bagmati province, near Mere Dara, 3000 m, 7.1V.1981, leg. Lobl & Smetana (cSme, cAss); 4 exs. [par- tly teneral], Bagmati province, Mere Dara, 3200 m, 8.IV.1981, leg. Lobl & Smetana (cSme, cAss); 8 exs., Nuwakot district, be- tween Ghopte and Thare Pati, 3220 m, 23.1V.1985, leg. Smetana (cSme, cAss); 3 exs., same data, but 3250 m (cSme); 1 ex., same data, but 3150 m, 25.IV.1985 (cAss); 1 ex., same data, but 3200 m, 25.IV.1985 (cSme); 2 exs, same data, but 3200 m, 26.1V.1985 (cSme, cAss). Redescription. Body length 6.4—8.0 mm; length of fore- body 3.0—3.8 mm. Habitus as in Fig. 4. Coloration: body dark-brown to blackish-brown, with the abdominal apex, often the elytra, and sometimes also the head paler brown; legs pale-reddish to reddish-brown; antennae reddish, api- cal half often somewhat infuscate. Head (Fig. 5) approximately as long as broad or weak- ly transverse; punctation moderately coarse, sparse in me- dian dorsal portion, somewhat less sparse in posterior and lateral portions, but interstices on average broader than di- ameter of punctures; interstices with fine microreticula- tion. Eyes weakly projecting from lateral contours of head, small, 1/4—1/3 times as long as postocular region in dor- sal view. Pronotum (Fig. 5) 1.10—1.15 times as long as broad and 1.05—-1.10 times as broad as head, strongly convex in cross-section; punctation similar to that of head, but slight- ly finer; interstices with microreticulation. ©ZFMK 154 Volker Assing 7 Figs 4-10. Lathrobium nepalense. 4: habitus; 5: forebody; 6: male sternite VII; 7: male sternite VIII; 8: aedeagus in lateral view; 9: apical portion of aedeagus in ventral view; 10: female sternite VIII. Scale bars: 4—5: 1.0 mm; 6—10: 0.5 mm. Elytra short, 0.55—0.60 times as long as pronotum (Fig. 5); humeral angles weakly marked; punctation shallow and ill-defined; interstices without distinct microsculpture. Hind wings completely reduced. Abdomen broader than elytra; punctation fine and mod- erately dense, sparser on posterior tergites; posterior mar- gin of tergite VII without palisade fringe. &: protarsomeres I-IV strongly dilated; sternite VII im- pressed along the middle, posterior margin broadly and weakly concave (Fig. 6); sternite VIII impressed along the middle, the impression with numerous weakly modified setae, at posterior margin with pair of elevations with con- spicuously dense setae, posterior margin weakly bisinu- ate in the middle (Fig. 7); aedeagus (Figs 8—9) approxi- mately 1.2 mm long, with slender and apically acute ven- tral process, with lamellate, but distinctly sclerotised dor- sal plate, and with long, sclerotised internal structures. Bonn zoological Bulletin 61 (2): 142—209 ©: protarsomeres I-IV dilated, but somewhat less so than in male; sternite VIII much longer than tergite VIII, distinctly produced and finely pubescent posteriorly (Fig. 10); tergite LX with fine median suture; tergite X slightly longer than tergite IX in the middle. Comparative notes. This species is most similar and ev- idently closely related to the following 14 species, from which it is distinguished particularly by the different shape of the ventral process of the aedeagus, as well as by the conspicuous modifications of the male sternite VIII. Distribution and natural history. The type locality is sit- uated between Mulkharka [27°46’N, 85°26’E] and Thare Pati [28°02’N, 85°29’E] to the northeast of Kathmandu. The additional material was found near Thare Pati, Mere Dara (ca. 5 km south of Thare Pati), and Malemchi ©ZFMK Revision of Himalayan Lathrobium 155 [28°01 N, 85°31’E] at altitudes of 2800-3500 m (Fig. 2). Part of the material collected in April is slightly teneral. Between Ghopte and Thare Pati, L. nepalense was found together with the closely related and similar L. exsertum. According to Franz’ diary, the type specimens were col- lected in a rhododendron forest above “Bulumje” [?] on PPXGlOTAe Lathrobium lassallei Coiffait, 1981 (Figs 2, 11—17) Lathrobium lassallei Coiffait, 1981: 331 f. Lathrobium sherpa Coiffait, 1982b: 290 f.; syn. n. Type material examined. L. Jassallei: Holotype 3: “Nepal IX 80, Barabinse [sic] / Mt Kalingchok, 3000 D. L / Holotype / Lathrobium lassallei H. Coiffait 1981 / Lathrobium lassallei Coiffait, det. V. Assing 2011” (MNHNP). L. sherpa: Holotype @ [aedeagus and abdominal seg- ments [IX—X missing]: “Nepal X.81, Kalingchok, 3.100 PC / Holotype / Lathrobium sherpa H. Coiffait 1982 / Lath- robium lassallei Coiffait, det. V. Assing 2011” (MNHNP). Paratype 2: same data as holotype (MNHNP). Comment. The original description of L. /assallei is based on a unique holotype male from “Barabinse [sic], Mt Kalingchock, 3000 m” (Coiffait 1981). The specimen is deposited in the Coiffait collection at the MNHNP. Lathrobium sherpa was described from a male holotype and a female paratype from “Massif du Kalingchok, pres de Barbabise [sic], 3100 m” (Coiffait 1982b). The aedea- gus of the holotype is missing. According to a footnote (Coiffait 1982b: 291), it was lost before the drawings were finished. Remarkably, there is no reference to L. lassallei in the original description of L. sherpa, although the type localities of both species are practically identical and de- spite their similar external appearance. Since convincing evidence was found neither in the ex- ternal nor in the male sexual characters that L. sherpa should represent a distinct species, the name is placed in synonymy with L. /assallei. 12 13 Figs 11-17. Lathrobium lassallei (12, 14, 16: Kalinchok; 11, 13, 15, 17: Ting Sang La). 11: male sternite VII; 12-13: male ster- nite VIII; 14-15: aedeagus in lateral view; 16-17: female sternite VIII. Scale bars: 0.5 mm. Bonn zoological Bulletin 61 (2): 142-209 ©ZFMK 156 Volker Assing Additional material examined. Nepal: 10 exs., Dolakha Di- strict, E Ting Sang La [27°49°N, 86°03’E], 3100 m, 12.-13.V1.2000, leg. Schawaller [628] (SMNS, cAss); 13 exs., Dolakha District, SW Kalinchok Mt., 3100 m, 19.-23.IV.1995, leg. Martens & Schawaller [421] (SMNS, cAss); 9 exs., Bag- mati province, NE Barahbise, Yardang ridge, 3250 m, 5.V.1981, leg. Lobl & Smetana (cSme, cAss); 2 exs., Bagmati province, NE Barahbise, Pokhare, 3000 m, 7.V.1981, leg. L6bl & Smeta- na (cSme, cAss). Redescription. Body length 7—8 mm; length of forebody 3.0-3.5 mm. External characters similar to those of L. nepalense; reliably distinguished only by the sexual char- acters. ¢: posterior margin of tergite VIII truncate to weakly convex; sternite VII weakly impressed in postero-medi- an portion, posterior margin broadly and weakly concave (Fig. 11); sternite VHI impressed along the middle, the im- pression with numerous weakly modified setae, posteri- or margin weakly concave in the middle (Figs 12-13); aedeagus (Figs 14-15) 1.0—1.1 mm long, with slender and apically acute ventral process, and with lamellate dorsal plate. ©: posterior margin of tergite VIII weakly angled in the middle; sternite VII much longer than tergite VIII, dis- tinctly produced posteriorly, posteriorly very finely pubes- cent (Figs 16-17); tergite [IX with fine median suture; ter- gite X approximately as long as tergite [IX in the middle, or nearly so. Comparative notes. This species is highly similar and ev- idently closely related to L. nepalense, from which it is distinguished particularly by the different shape of the ven- tral process of the aedeagus and by the male secondary sexual characters. Intraspecific variation. The material from Ting Sang La is distinguished from that collected on or near Mt. Kalin- chok by slight differences in the shape of the ventral process of the aedeagus (Fig. 15), as well as in the shapes of the male and female sternites VIII (Figs 13, 17). Since no additional evidence was found suggesting that the two samples should belong to different species, these differ- ences are attributed to intra- rather than interspecific vari- ation. Distribution and natural history. The known distribu- tion of L. /assallei is confined to the Kalinchok range and the region to the northeast of Barahbise, some 60 km ENE of Kathmandu (Fig. 2), where it has been collected at al- titudes of 3000-3250 m. Lathrobium kleebergi sp. n. (Figs 2, 18-22, 296-297) Type material. Holotype ¢: “Ost-Nepal, Rolwaling Hi- mal / Rolwaling Ufer zw. Simigaon u. Nyimare 2700 m, Bonn zoological Bulletin 61 (2): 142-209 17.05.2000 leg. A. Kleeberg / Holotypus ¢ Lathrobium kleebergi sp. n. det. V. Assing 2011” (cAss). Paratypes: 1 ex.: same data as holotype (cKle); 9 exs.: “Ost-Nepal, Rol- waling Himal / Rolwaling Tal, Nyimare, 3300 m, 19.05.2000, leg. A. Kleeberg” (cKle, cAss); 3 exs.: “Ost- Nepal, Rolwaling Himal / Nyimare, 3000 m, 18.05.2000, leg. A. Kleeberg” (cKle); 3 exs.: “Ost-Nepal, Rolwaling Himal / oberh. Simigaon, 2700-2800 m, 31.05.2000, leg. A. Kleeberg” (cKle); 15 exs.: same data, but “01.06.2000” (cKle, cAss); 14 exs.: “Ost-Nepal, Rolwaling Himal / westl. Daldung La Pass, 3300 m, 28.05.2000, leg. A. Klee- berg” (cKle, cAss); 1 ex.: same data, but “29.05.2000” (cKle); 13 exs.: same data, but 30.05.2000” (cKle, cAss); 1 ex.: “Nepal, Rolwaling Himal, upp. Simigaon vill. 2400- 2700 m, 01.06.2000, leg. J. Schmidt” (cKle); 14 exs.: “upp. Simigaon vill., ca. 3300 m, 28.05.2000, leg. J. Schmidt” (cKle, cAss); 8 exs.: “Nepal, Rolwaling Himal, upp. Simigaon vill. 2700-2800 m, 01.06.2000, leg. J. Schmidt” (cKle, cAss); 4 exs.: “Nepal, Rolwaling vall. Dugong Kharka, 2700-2800 m, 17.05.2000, leg. J. Schmidt” (cKle, cAss); 8 exs.: “Nepal, Rolwaling vall., bef. Beding vill., 3300 m, 19.05.2000, leg. J. Schmidt” (cKle, cAss). Description. Body length 7-8 mm; length of forebody 3.0-3.5 mm. Coloration: body dark-brown to blackish- brown, with the abdominal apex, often the elytra, and sometimes also the head paler brown; legs pale-reddish to reddish-brown; antennae reddish, apical half often somewhat infuscate. Head (Fig. 18) approximately as long as broad or weak- ly transverse; punctation moderately coarse, sparse in me- dian dorsal portion, somewhat less sparse in posterior and lateral portions, but interstices on average broader than di- ameter of punctures; interstices with fine microreticula- tion. Eyes weakly projecting from lateral contours of head, small, 1/4—1/3 times as long as postocular region in dor- sal view. Pronotum (Fig. 18) 1.10—1.15 times as long as broad and 1.05—1.10 times as broad as head, strongly convex in cross-section; punctation and microsculpture similar to those of head. Elytra short, 0.55—0.60 times as long as pronotum (Fig. 18); humeral angles weakly marked; punctation shallow and ill-defined; interstices without distinct microsculpture. Hind wings completely reduced. Protarsomeres I-IV di- lated in both sexes, but more so in male than in female. Abdomen broader than elytra; punctation fine and mod- erately dense; posterior margin of tergite VII without pal- isade fringe. G: posterior margin of tergite VIII truncate to weakly convex; sternite VII weakly impressed in postero-medi- an portion, posterior margin broadly and weakly concave (Fig. 19); sternite VII broadly impressed along the mid- dle, the impression with numerous modified setae, poste- ©ZFMK Revision of Himalayan Lathrobium 157 Figs 18-28. Lathrobium kleebergi (18-22) and L. ignoratum (23-28). 18: forebody; 19, 23: male sternite VII; 20, 24: male ster- nite VIII; 21, 25-26: aedeagus in lateral view; 22, 28: female sternite VIII; 27: apical portion of aedeagus in ventral view. Scale bars: 18: 1.0 mm; 19-28: 0.5 mm. Bonn zoological Bulletin 61 (2): 142-209 ©Z7FMK 158 Volker Assing rior margin weakly concave, without distinct median in- cision (Fig. 20); aedeagus (Fig. 21) approximately 1.0—1.1 mm long, with slender and apically acute ventral process, and with lamellate dorsal plate. ©: posterior margin of tergite VIII weakly angled in the middle; sternite VIII much longer than tergite VIII, dis- tinctly produced posteriorly, posteriorly finely pubescent (Fig. 22); tergite [X not divided in the middle; tergite X approximately as long as tergite [X in the middle, or near- ly so. Comparative notes. Lathrobium kleebergi is highly sim- ilar and closely related to L. nepalense, L. lassallei, and other species of the L. nepalense group. It is reliably dis- tinguished from them by the male primary and secondary sexual characters, from most of them also by the absence of a median suture of the female tergite IX. Etymology. The species is dedicated to Andreas Kleeberg, who, together with Joachim Schmidt, collected the type specimens. Distribution and natural history. Lathrobium kleeber- gi is currently known from several localities in the Rol- waling Himal in eastern Nepal (Fig. 2), where the speci- mens were collected at altitudes of 2400-3300 m (Figs 296-297). In one locality, the species was found togeth- er with L. excisum. One of the dissected females had a ma- ture egg in the ovaries. Lathrobium ignoratum sp. n. (Figs 2, 23-28) Type material. Holotype: “Nepal Rasuwa Dis., north slope above Syabru 3600 m, 18.1V.85 A. Smetana / Holo- typus ¢ Lathrobium ignoratum sp. n., det. V. Assing 2012” (cSme). Paratypes: 5 exs.: same data as holotype (cSme, cAss); 5 exs.: same data, but “19.IV.85” (cSme); 5 exs.: same data, but “3650 m, 17.1V.85” (cSme, cAss); 2 exs.: same data, but 3800 m, 18.IV.85” (cSme); 1: ‘““Wald un- terhalb Fulung / Pa 175 [overleaf] / Zentral-Nepal, Sept—Okt. 1971, lg. H. Franz / Paratype [of L. nepalense]” (NHMW). Description. Length of forebody 3.35 mm. External char- acters highly similar to those of L. nepalense and allied species. 3: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VIII very weakly convex; sternite VII weak- ly impressed in postero-median portion, this impression with moderately dense black setae, posterior margin broadly and weakly concave (Fig. 23); sternite VIII im- pressed along the middle, this impression with numerous long black setae, posterior margin shallowly concave in the middle (Fig. 24); aedeagus 1.0-1.1 mm long, with Bonn zoological Bulletin 61 (2): 142-209 slender and apically acute ventral process, and with dis- tinctly sclerotised, lamellate dorsal plate (Figs 25—27). ©: protarsomeres I-IV dilated, but less so than in male; posterior margin of tergite VII weakly, obtusely angled in the middle; sternite VIII convexly produced and fine- ly pubescent posteriorly (Fig. 28); tergite [IX with fine me- dian suture; tergite X distinctly longer than tergite IX in the middle. Comparative notes. The similar external and sexual char- acters suggest that L. ignoratum is closely related to L. nepalense and allied species, from which it is distin- guished by the shape and chaetotaxy of the male sternites VU and VIII, as well as by the morphology of the aedea- gus. From L. nepalense, with which it was previously con- founded, it is separated by slightly smaller body size, the less transverse male sternite VII, the less dense and finer setae in the posterior impression of the male sternite VII, the absence of setae in the middle of the impression of the male sternite VIII, the absence of dense clusters of setae near the middle of the posterior margin of sternite VIII, as well as by the smaller aedeagus with a differently shaped ventral process and a differently shaped dorsal plate. Etymology. The specific epithet (Latin, past participle of ignorare: to mistake, to confound) alludes to the fact that the paratype of L. nepalense from Fulung, now paratype of L. ignoratum, was previously both missexed and misidentified. Distribution and natural history. The species was found near Syabru [28°10’N, 85°20’E] and near the Fulung Monastery [28°07’N, 85°20’E], to the north of Kathman- du, central Nepal (Fig. 2). The specimens from Syabru were collected at an altitude of 3600-3800 m. According to Franz’ diary, the specimen from Fulung was sifted from dry and slightly mouldy leaf litter in an old oak forest with rhododendron forest with scattered pine and fir on IEXETOF Ae Lathrobium exsertum sp. n. (Figs 3, 29-33) Type material. Holotype 3: “Nepal (Prov. Bagmati), Yan- gri Ridge, 4200 m, 21.1V.81, Lobl & Smetana / Holoty- pus ¢ Lathrobium exsertum sp. n., det. V. Assing 2012” (cSme). Paratypes: 344, 19 [1 slightly teneral]: same da- ta as holotype (cSme, cAss); 34, 19: “Nepal (Prov. Bag- mati), Yangri Ridge, 4350 m, 22.1V.81, Lobl & Smetana” (cSme, cAss); 12, 39 9: “Nepal (Prov. Bagmati), Yangri Ridge, 4700-4800 m, 22.IV.81, Lobl & Smetana” (cSme, cAss); 34 [1 teneral]: “Nepal Newakot Di., betw. Ghopte and Thare Pati, 3220 m, 23.IV.85 A. Smetana” (cSme, cAss); 34.4: same data, but “3150 m, 24.IV.85” ©ZFMK Revision of Himalayan Lathrobium 159 ies Sea Figs 29-38. Lathrobium exsertum (29-33) and L. compressum (34-38). 29, 35: male sternite VI; 30, 36: male sternite VIII; 31, 37: aedeagus in lateral view; 32, 38: aedeagus in ventral view; 33: female sternite VIII; 34: forebody. Scale bars: 34: 1.0 mm; 29-33, 35-38: 0.5 mm. Bonn zoological Bulletin 61 (2): 142-209 ©ZFMK 160 Volker Assing (cSme, cAss); 1d, 299: same data, but “3100 m, 24.1V.85” (cSme); 1: same data, but “3150 m, 25.IV.85” (cSme); 14, 19: same data, but “3200 m, 26.IV.85” (cSme, cAss). Description. Body length 6.3-8.5 mm; length of forebody 3.1-3.7 mm. External characters as in L. nepalense. &: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VIII weakly convex in the middle; sternite VI impressed along middle, this impression with mod- erately dense black setae, posterior margin broadly and weakly concave (Fig. 29); sternite VII impressed along the middle, this impression with moderately dense black setae, on either side of median impression with extensive cluster of conspicuously dense black setae, posterior mar- gin produced in the middle (Fig. 30); aedeagus (Figs 31-32) approximately 1.3 mm long, ventral process mod- erately slender in lateral view and broad in ventral view; dorsal plate lamellate, but distinctly sclerotised. ©: protarsomeres I-IV dilated, but less so than in male; posterior margin of tergite VIII obtusely angled in the mid- dle; sternite VIII distinctly produced and finely pubescent posteriorly (Fig. 33); tergite [IX with fine median suture; tergite X much longer than tergite [X in the middle. Comparative notes. Lathrobium exsertum is distin- guished from the similar, geographically close, and part- ly even syntopic L. nepalense by the completely differ- ent shape and chaetotaxy of the male sternite VIII, the shorter, less acute (lateral view), a much broader ventral process of the aedeagus (ventral view), and by the poste- riorly more produced female sternite VIII. Etymology. The specific epithet (Latin, past participle of exserere: to extend, to stick out) refers to the posteriorly distinctly produced female sternite VIII, the only charac- ter distinguishing female L. exsertum from female L. nepalense. Distribution and natural history. The specimens were collected in the Yangri ridge, some 25 km to the east of Thare Pati, and between Ghopte and Thare Pati (Fig. 3) at altitudes of 3100-4800 m. Some of the beetles are no- ticeably teneral. Between Ghopte and Thare Pati, L. exser- tum was found together with L. nepalense. Lathrobium compressum sp. n. (Figs 2, 34-38) Type material. Holotype ¢: “Nepal (Prov. Bagmati), above Shermathang, 2900 m, 26.IV.81, Lobl & Smetana / Holotypus 3 Lathrobium compressum sp. n., det. V. Ass- ing 2011” (MHNG). Paratypes: 14, 19: same data as holotype (MHNG, cAss). Bonn zoological Bulletin 61 (2): 142-209 Description. Body length 7.0—7.7 mm; length of forebody 3.4-3.6 mm. External characters highly similar to those of L. nepalense, except for the larger average body size and the very shallow, almost obsolete microreticulation on the head and pronotum (Fig. 34). S: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VIII truncate; sternite VII weakly impressed in postero-median portion, posterior margin broadly and weakly concave (Fig. 35); sternite VIII impressed along the middle, this impression with numerous weakly mod- ified setae, posteriorly with patch of dense dark setae on either side of middle, posterior margin weakly concave, with very small and shallow median incision (Fig. 36); aedeagus approximately 1.3 mm long, with very slender and apically acute ventral process, and with distinctly scle- rotised, lamellate dorsal plate (Figs 37-38). ©: protarsomeres strongly dilated, only slightly less so than in male; posterior margin of tergite VIII weakly con- vex in the middle; sternite VIII much longer than tergite VII, distinctly produced and finely pubescent posterior- ly, of similar shape as that of L. kleebergi; tergite [X with fine median suture; tergite X distinctly longer than tergite IX in the middle. Comparative notes. The similar external and sexual char- acters suggest that L. compressum is closely related to L. nepalense, L. lassallei, and allied species, from which it is distinguished by slightly larger average body size, shal- lower microreticulation of the head and pronotum, and particularly by the male sexual characters. Etymology. The specific epithet (Latin, adjective) alludes to the dorso-ventrally strongly compressed ventral process of the aedeagus. Distribution and natural history. The type locality is sit- uated near Shermathang in the Bairavkund Lekh, some 55 km NNE of Kathmandu, in Bagmati province (Fig. 2). The specimens were collected at an altitude of 2900 m. Lathrobium khumbuense Coiffait, 1982 (Figs 2, 39-42) Lathrobium khumbuensis [sic] Coiffait, 1982a: 90. Type material examined. Holotype 9: “Umg. Lughla, Khumbu, Nepal, lg. H. Franz / Pa 260 [overleaf] / Type / Lathrobium khumbuensis H. Coiffait 1979 / Lathrobium khumbuense Coiffait, det. V. Assing 2011” (NHMW). Paratype [teneral]: same data as holotype (MNHNP). Comment. The original description is based on a holo- type and a paratype, both females, from “Népal, environs de Lughla, Khumbu” (Coiffait 1982a) deposited in the col- lections at the NHMW and in the Coiffait collection at the MNHNP, respectively. The male sexual characters were ©ZFMK Revision of Himalayan Lathrobium 161 described by Coiffait (1983), based on a specimen that was evidently collected together with the types, as can be in- ferred from Franz’ identical sample number (“Pa 260”). This specimen was erroneously labelled by Coiffait as the allotype; it does not have type status, since it was not in- cluded in the original description. Additional material examined. Nepal: 14, Khumbu, Lughla env., “Alm Dugdinma, 3000-4000 m, leg. Franz, “Pa 260”, “Al- lotype” (MNHNP). Redescription. Species of moderate size; body length 5.6—6.5 mm; length of forebody 2.5—2.8 mm. Coloration: head and pronotum dark-brown; elytra reddish to brown; abdomen reddish-brown to brown; legs and antennae red- dish. Head approximately as broad as long; punctation coarse and rather sparse, even sparser in median dorsal portion; interstices with distinct microreticulation, almost matt, on average broader than diameter of punctures. Eyes mod- erately small and not distinctly projecting from lateral con- Figs 39-47. Lathrobium khumbuense (39-42) and L. nepalorientis (43-47). 39, 44: male sternite VII; 40, 45: male sternite VI- II; 41, 46: aedeagus in lateral view; 42: female sternite VIII; 43: forebody; 47: aedeagus in ventral view. Scale bars: 43: 1.0 mm; 39-42, 44-47: 0.5 mm. Bonn zoological Bulletin 61 (2): 142-209 ©ZFMK 162 Volker Assing tours of head, approximately 1/3 the length of postocular region in dorsal view and composed of numerous (> 30) ommatidia. Pronotum broad, short, and strongly convex 1n cross- section, approximately 1.15 times as long as broad and ap- proximately as broad as head; lateral margins straight and almost subparallel, only indistinctly converging posteri- ad; punctation similar to that of head; interstices with pro- nounced microreticulation and almost matt. Elytra short, approximately 0.55 times as long as prono- tum; humeral angles weakly marked; punctation shallow and ill-defined; interstices without distinct microsculpture. Hind wings completely reduced. Metatibia weakly com- pressed. Abdomen broader than elytra, widest at segment VI; punctation dense and not particularly fine; interstices with finely transverse microsculpture; posterior margin of ter- gite VII without palisade fringe. ¢: protarsomeres I-IV moderately dilated; posterior margin of tergite VII indistinctly angled in the middle; sternite VII moderately transverse, with shallow median impression, without modified pubescence, and with weakly concave posterior margin (Fig. 39); sternite VIII weakly transverse, with long black pubescence in poste- rior half, posterior margin very weakly concave, almost truncate, and without median excision (Fig. 40); aedea- gus approximately 0.9 mm long, with long, slender, and apically acute ventral process, with a lamellate, weakly sclerotised dorsal plate, and with dark membranous inter- nal structures (Fig. 41). ©: protarsomeres I-IV almost as dilated as in male; ter- gite VII of similar shape as in male; sternite VIII distinct- ly longer than tergite VIII, with convex posterior margin, at posterior margin with fine pubescence (Fig. 42); tergite IX with fine median suture; tergite X somewhat longer than tergite LX in the middle. Comparative notes. As can be inferred from external (head and pronotum with pronounced microreticulation; eyes with numerous ommatidia; short and convex prono- tum) and sexual characters (sternite VII without distinct- ly modified pubescence and posterior margin; sternite VI- II without posterior excision and without distinctly mod- ified setae; general morphology of the aedeagus without sclerotised internal structures and with weakly sclerotised dorsal plate), L. khumbuense is closely related to L. nepalense and allied species. It is distinguished from them by smaller size and by the male primary and secondary sexual characters. Distribution and natural history. Lathrobium khum- buense is currently known only from the type locality near “Lughly”, today probably Lukla (27°41°N, 86°43’E) in Khumbu, eastern Nepal (Fig. 2), where the type specimens Bonn zoological Bulletin 61 (2): 142-209 were collected at an altitude of 3000-4000 m. The paratype is teneral. Lathrobium nepalorientis Coitfait, 1984 (Figs 2, 43-47) Lathrobium nepalorientis Coiffait, 1984: 381 f. Type material examined. Holotype ¢: “Nepal or. 7.83, Jaljale Himal 4000 m / Type / Lathrobium nepalorientis H. Coiffait / Lathrobium nepalorientis Coiffait, det. V. Ass- ing 2011” (MNHNP). Paratype <: see holotype of L. mil- keense. Comment. The original description is based on a male holotype from “Neépal oriental, Jaljale Himal 4000 m” and a paratype (“une femelle’) from “Milke Himal, 2500 m” deposited in the Coiffait collection at the MNHNP (Coif- fait 1984). An examination of the two type specimens re- vealed that the paratype is in fact a male and that it is not conspecific with the holotype. Redescription. Species of moderate size; body length 6.1 mm; length of forebody 2.9 mm. Coloration: forebody brown; abdomen blackish-brown to brown; legs and an- tennae reddish. Head (Fig. 43) approximately as broad as long; punc- tation coarse and rather sparse, even sparser in median dor- sal portion; interstices with distinct microreticulation, dis- tinctly broader than diameter of punctures. Other external characters as in L. khumbuense. 3: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VIII indistinctly angled in the middle; ster- nite VII moderately transverse, with shallow median im- pression, without modified pubescence, and with weakly concave posterior margin (Fig. 44); sternite VIII as long as wide, with long median impression, on either side of middle with dense black setae, posterior margin with very shallow, almost indistinct median excision (Fig. 45); aedeagus | mm long, with slender, weakly curved (later- al view), and apically acute ventral process and with weak- ly sclerotised, lamellate dorsal plate (Figs 46-47). ©: unknown. Comparative notes. Lathrobium nepalorientis is reliably distinguished from the similar L. khumbuense only by the male sexual characters, particularly the shape and chaeto- taxy of the male sternite VIII and the shape of the ventral process of the slightly larger aedeagus. Distribution and natural history. This species is current- ly known only from the type locality, the Jaljale Himal in eastern Nepal (Fig. 2), where the holotype was collected at an altitude of 4000 m. OZFMK Revision of Himalayan Lathrobium 163 49 48 52 Figs 48-55. Lathrobium janetscheki (48-51) and L. milkeense (52-55). 48, 52: male sternite VII; 49, 53: male sternite VII; 50, 54: aedeagus in lateral view; 51: aedeagus in ventral view; 55: apical portion of aedeagus in ventral view. Scale bars: 0.5 mm. Lathrobium janetscheki Scheerpeltz, 1976 (Figs 2, 48— 51) Lathrobium (Glyptomerodoschema) janetscheki Scheer- peltz, 1976: 29 ff. Type material examined. Holotype ~ [somewhat dam- aged]: “¢ / Himalaya-Expedition Prof. Dr. Janetschek 1961 nach Nepal / loc. 51. / Lathrobium (Glyptomero- doschema n. subg.) Janetscheki n. sp. / Holotypus / Ty- pus Lathrobium (Glyptomerodoschema) Janetscheki O. Scheerpeltz / Lathrobium (Glyptomerodoschema n. subg.) Janetscheki n. sp., det. Scheerpeltz / Lathrobium janetsche- ki Scheerpeltz, det. V. Assing 2011” (ZII). Bonn zoological Bulletin 61 (2): 142-209 Comment. The original description is based on a unique male holotype from “Basislager Yaral (Pangpoche)” (Scheerpeltz 1976). The specimen, which 1s in rather poor condition, is deposited in the collections of the ZII. Scheerpeltz (1976) described the subgenus Glyp- tomerodoschema to accommodate the type species Lath- robium janetscheki Scheerpeltz, 1976, stating that the sub- genus was distinguished from other species groups of the Lathrobium s. str. by the short elytra, reduced hind wings and small eyes, and from Glyptomerus Miller, 1856 by the presense of ommatidia. Coiffait (1982b) already ar- gued that attributing the depigmented, micropterous, and microphthalmous Himalayan Lathrobium species to a sub- genus of their own was unjustified. Such adaptive reduc- tions are known from several species groups in Lathrobi- ©OZFMK 164 Volker Assing um both in the West and in the East Palaearctic regions. Moreover, attributing L. janetscheki to a subgenus of its own would undoubtedly render the nominate subgenus pa- raphyletic. Therefore, Glyptomerodoschema is placed in synonymy with Lathrobium. Redescription. External characters as in L. nepalorientis and L. nepalense. ¢: protarsomeres I-IV strongly dilated; sternite VII strongly transverse, with median impression of triangu- lar shape and with relatively long dark setae, posterior margin weakly concave (Fig. 48); sternite VIII moderate- ly transverse and with pronounced, sparsely pubescent me- dian impression, on either side of this impression with dense dark setae, posterior margin weakly concave in the middle (Fig. 49); aedeagus 1 mm long, ventral process strongly asymmetric in ventral view (Figs 50-51). ©: unknown. Comparative notes. Lathrobium janetscheki is evident- ly closely related to L. nepalorientis and L. khumbuense, as can be inferred particularly from the similar modifica- tions of the male sternites VII and VIII, as well as from the similar morphology of the aedeagus and similar ex- ternal characters. It is readily distinguished from its close relatives especially by the strongly asymmetric ventral process of the aedeagus. Distribution and natural history. The type locality is sit- uated near Pangpoche [27°51°N, 86°48’E] in the Mount Everest region, eastern Nepal (Fig. 2). The holotype was sifted from rhododendron litter and grass in April. Lathrobium milkeense sp. n. (Figs 2, 52—55) Type material. Holotype @: “Népal or. 7-83, Milke Hi- mal, 2500 m / Allotype [of L. nepalorientis] / Holotypus 3 Lathrobium milkeense sp. n., det. V. Assing 2011” (MNHNP). Description. External characters as in L. nepalorientis. ¢: protarsomeres I-IV strongly dilated; sternite VII strongly transverse, with median impression with relative- ly long dark setae, posterior margin weakly concave (Fig. 52); sternite VIII moderately transverse, median impres- sion oblong and with sparse, weakly modified dark setae, on either side of impression with dense dark setae, pos- terior margin weakly concave in the middle (Fig. 53); aedeagus 1.1 mm long, ventral process strongly curved in lateral view, dorsal plate relatively massive and strongly sclerotised, apically very acute and in the middle broad in dorsal view (Figs 54-55). ©: unknown. Bonn zoological Bulletin 61 (2): 142-209 Comparative notes. As can be inferred from the similar external and male sexual characters, L. milkeense is close- ly allied to the L. nepalorientis, L. khumbuense, and L. Janetscheki, but distinguished from them particularly by the morphology of the aedeagus, especially the shape of the ventral process and of the strongly sclerotised dorsal plate. Etymology. The specific epithet (adjective) is derived from the name of the mountain range where the species was discovered. Distribution and natural history. The type locality is sit- uated in the Milke Himal in eastern Nepal (Fig. 2). The holotype was collected at an altitude of 2500 m. Lathrobium umbhakense sp. n. (Figs 3, 56—64) Type material. Holotype 3: “Nepal-Expeditionen Jochen Martens / 390 Sankhua Sabha Distr., Thudam, mixed fo- rest mainly Betula/Rhododendron, 3550-3650 m, 25—27 May 88 J. Martens & W. Schawaller / Holotypus ¢ La- throbium umbhakense sp. n., det. V. Assing 2011” (SMNS). Paratypes: 44.2, 22 9: same data as holotype (SMNS, cAss); 14, 19: “Nepal-Expeditionen Jochen Martens / 362 Taplejung Distr., upper Simbua Khola Val- ley, near Yalung, 3450-3700 m, mature Abies-Rhododen- dron-Juniperus forest, 13 May 1988 J. Martens & W. Schawaller leg.” (SMNS); 2¢'4, 19: “Nepal-Expeditio- nen Jochen Martens / 387 Sankhua Sabha Distr., Kangla Khola E Thudam, dwarf Rhododendron, rock debris, 4100-4200 m, 24—-25May88 Martens & Schawaller” (SMNS, cAss). Description. Body length 5.3—-6.7 mm; length of forebody 2.6-2.9 mm. Habitus and forebody as in Figs 56-57. Ex- ternally highly similar to L. nepalorientis. ¢: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VIII convex to indistinctly pointed in the middle; sternite VII strongly transverse, median impres- sion with relatively long dark setae, posterior margin weakly concave (Fig. 58); sternite VIII moderately trans- verse, median impression oblong, not very deep, and with moderately sparse modified dark setae, on either side of impression with dense dark setae, posterior margin weak- ly concave in the middle (Fig. 59); aedeagus approximate- ly 0.9-1.0 mm long, ventral process evenly curved and slender in lateral view, dorsal plate relatively large, but not massive and relatively weakly sclerotised (Figs 60-63). ©: protarsomeres I-IV dilated, but distinctly less so than in male; posterior margin of tergite VII obtusely point- ed in the middle; sternite VIII oblong, longer than tergite VIII, produced and with fine setae posteriorly (Fig. 64); ©ZFMK Revision of Himalayan Lathrobium 165 60 61 62 63 Figs 56-64. Lathrobium umbhakense. 56: habitus; 57: forebody; 58: male sternite VII; 59: male sternite VII; 60-62: aedeagus in lateral view; 63: aedeagus in ventral view; 64: female sternite VIII. Scale bars: 56-57: 1.0 mm; 58—64: 0.5 mm. tergite IX not divided in the middle; tergite X slightly longer than tergite [IX in the middle. Comparative notes. As can be inferred from the similar external and male sexual characters, L. umbhakense is closely related to L. nepalorientis and allied species, but distinguished from them particularly by the morphology Bonn zoological Bulletin 61 (2): 142-209 of the aedeagus, especially the shape of the ventral process and the dorsal plate (L. nepalorientis: male sternite VIII not transverse; aedeagus with ventral process of slightly different shape in lateral view and dorsal plate much small- er). It is additionally separated from the syntopic L. inex- cisum by the anteriorly undivided female tergite IX. ©ZFMK 166 Volker Assing < bane a Sy SS ALPES rr 70 Figs 65-74. Lathrobium bibarbatum (65-69) and L. diremptum (70-74). 65, 70: male sternite VU; 66, 71: male sternite VIL; 67, 72: aedeagus in lateral view; 68: aedeagus in ventral view; 69, 74: female sternite VIII; 73: apical portion of aedeagus in ventral view. Scale bars: 0.5 mm. Etymology. The specific epithet (adjective) is derived from the name of the mountain range where the species is probably endemic. Distribution and natural history. The species was found in three localities near Thudam and Yalung in the very northeast of Nepal, close to the border with Tibet (Fig. 3). The specimens were collected in a mixed forest with rho- dodendron, from rock debris, and litter of dwarf rhodo- Bonn zoological Bulletin 61 (2): 142-209 dendron at altitudes of 3450-4200 m, in one locality to- gether with L. inexcisum. Lathrobium bibarbatum sp. n. (Figs 3, 65-69) Type material. Holotype ¢: “Nepal-Expeditionen Jochen Martens / 380 Taplejung Distr., above Walungchung Go- la, open Abies-Betula forest, 3400-3600 m, 21 May 1988 ©ZFMK Revision of Himalayan Lathrobium 167 J. Martens & W. Schawaller / Holotypus ¢ Lathrobium bibarbatum sp. n., det. V. Assing 2011” (SMNS). Paratypes: 14, 12: same data as holotype (SMNS, cAss). Description. Body length 7.0—7.5 mm; length of forebody 3.1-3.2 mm. Whole body dark-reddish. Externally high- ly similar to L. nepalorientis, but eyes less convex, slight- ly smaller, and composed of < 30 ommatidia. ¢: protarsomeres I-IV strongly dilated; sternite VII strongly transverse, median impression with relatively long dark setae, posterior margin very weakly concave (Fig. 65); sternite VII weakly transverse and with pro- nounced, deep, wide, long median impression with sparse black setae, on either side of this impression with conspicuous cluster of dense dark setae, posterior margin produced in the middle, not concave or incised (Fig. 66); aedeagus 1.1 mm long, ventral process evenly curved, slender, and apically acute in lateral view, dorsal plate long, lamellate, and weakly sclerotised (Figs 67—68). ©: protarsomeres I-IV dilated, but less so than in male; sternite VIII oblong, longer than tergite VIII, convexly produced and with micropubescence posteriorly (Fig. 69); tergite [X with median suture; tergite X distinctly longer than tergite IX in the middle. Comparative notes. Based on the external and sexual characters, L. bibarbatum is closely related to L. nepalo- rientis and allied species. It is distinguished from them par- ticularly by the conspicuous shape and chaetotaxy of the male sternite VII and by the morphology of the aedea- gus (shapes of ventral process and of dorsal plate). Etymology. The specific epithet (Latin, adjective: with two beards) alludes to the conspicuous clusters of dark se- tae on the male sternite VIII. Distribution and natural history. The type locality is sit- uated in Taplejung district, in the very northeast of Nepal (Fig. 3). The specimens were collected in an open fir and birch forest at an altitude of 3400-3600 m. Lathrobium diremptum sp. n. (Figs 2, 70-74) Type material. Holotype @: “Nepal-Expeditionen Jochen Martens / 361 Taplejung Distr., upper Simbua Khola Val- ley, near Tseram, 3250-3350 m, mature Abies-Rhododen- dron forest, 10-15 May 88 J. Martens & W. Schawaller leg. / Holotypus ¢ Lathrobium diremptum sp. n., det. V. Assing 2011” (SMNS). Paratype @: “Nepal-Expeditionen Jochen Martens / 359 Taplejung Distr., pasture Lassetham NW Yamputhin, 3300-3500 m, mature Abies-Rhododen- dron forest, 6-9 May 1988 J. Martens & W. Schawaller leg.” (cAss). Bonn zoological Bulletin 61 (2): 142-209 Description. Body length 7.0—7.2 mm; length of forebody 3.1-3.3 mm. Externally indistinguishable from L. bibar- batum. ¢: protarsomeres I-IV strongly dilated; sternite VII strongly transverse, median impression with relatively sparse, long dark setae, posterior margin very weakly con- cave in the middle (Fig. 70); sternite VIII weakly trans- verse and with moderately deep, wide, long median im- pression with sparse black setae, on either side of this im- pression with moderately dense dark setae, posterior mar- gin convex, not concave or incised (Fig. 71); aedeagus ap- proximately | mm long, ventral process evenly curved in lateral view, somewhat asymmetric, relatively broad, and apically acute in ventral view, dorsal plate moderately long, lamellate, and distinctly sclerotised (Figs 72—73). ©: protarsomeres I-IV dilated, but less so than in male; sternite VHI oblong, longer than tergite VIII, convexly produced and with micropubescence posteriorly (Fig. 74); tergite [X with fine median suture; tergite X distinctly longer than tergite [X in the middle. Comparative notes. Lathrobium diremptum is evidently most closely related to L. bibarbatum, as is suggested by the similar external morphology and particularly by the male and female sexual characters: the posteriorly con- vex male sternite VII, the morphology of the aedeagus, and the anteriorly divided, but contiguous female tergite IX. It 1s distinguished from L. bibarbatum by the shape and chaetotaxy of the male sternite VIII (without pro- nounced cluster of dense dark setae on either side of mid- dle; posterior margin less strongly produced), the shape of the male sternite VII (posterior margin concave in the middle), and by morphology of the aedeagus (ventral process broader and distinctly asymmetric in ventral view; dorsal plate shorter and more strongly sclerotised). Etymology. The specific epithet (Latin, past participle of dirimere: to separate) alludes to the anteriorly divided fe- male tergite IX. Distribution and natural history. The type specimens were found in two localities near Tseram and near Yam- puthin in Taplejung district, northeastern Nepal (Fig. 2). They were collected in mature fir and rhododendron forests at an altitude of 3250-3500 m. Lathrobium inexcisum sp. n. (Figs 3, 75-81) Type material. Holotype ¢: “Nepal-Expeditionen Jochen Martens / 361 Taplejung Distr., upper Simbua Khola Val- ley, near Tseram, 3250-3350 m, mature Abies-Rhododen- dron forest, 10-15 May 88 J. Martens & W. Schawaller leg. / Holotypus ¢ Lathrobium inexcisum sp. n., det. V. ©ZFMK 168 Volker Assing 84 85 Ay Wy Ax ARMA sees : ch WY PP eS 82 Figs 75-85. Lathrobium inexcisum (75-81) and L. infractum (82-85). 75, 82: male sternite VII; 76, 83: male sternite VII; 77-78, 84: aedeagus in lateral view; 79, 85: aedeagus in ventral view; 80: female tergite VIH; 81: female sternite VIII. Scale bars: 0.5 mm. Assing 2011” (SMNS). Paratypes: 44'¢, 49. 9: same da- ta as holotype (SMNS, cAss); 24: “Nepal-Expeditionen Jochen Martens / 362 Taplejung Distr., upper Simbua Kho- la Valley, near Yalung. 3450-3700 m, mature Abies-Rho- dodendron-Juniperus forest, 13 May 1988 J. Martens & W. Schawaller leg.” (SMNS, cAss); 299 [1 teneral]: “Nepal-Expeditionen Jochen Martens / 359 Taplejung Dis- tr., pasture Lassetham NW Yamputhin, 3300-3500 m, ma- ture Abies-Rhododendron forest, 6-9 May 1988 J. Martens & W. Schawaller leg.” (SMNS); 1¢: ““Nepal-Ex- peditionen Jochen Martens / 275 Taplejung Dist., Alm Las- Bonn zoological Bulletin 61 (2): 142-209 setham, Abies-Rhododendron-Wald, 3500 m, Berlese, 6—7 Sep83 Martens & Daams” (SMNS); 1: ““Nepal-Expedi- tionen Jochen Martens / 277 Taplejung Dist. Khola, Yalung, 3450 m, 8Sep83 J. Martens” (SMNS); 1: ‘“Nepal-Expeditionen Jochen Martens / 275 Taplejung Dist., Alm Lassetham, Abies-Rhododendron-Wald, 3500 m, Berlese, 6-7 Sep 83 Martens & Daams” (SMNS); 19: ‘“Nepal-Expeditionen Jochen Martens / 367 Tapleyung Dis- tr., pass Deorali W Yamputhin, 3400 m, Abies-Rhododen- dron forest, 17 May 1988 J. Martens & W. Schawaller leg.” (SMNS). ©OZFMK Revision of Himalayan Lathrobium 169 Description. Body length 5.3—6.2 mm; length of forebody 2.32.6 mm. Coloration: head and pronotum reddish- brown to dark-brown; elytra reddish to reddish-brown; ab- domen brown to blackish; legs reddish to brown; anten- nae reddish. Apart from the slightly smaller body size, ex- ternally highly similar to the preceding species. ¢: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VIII weakly convex to indistinctly angled in the middle; sternite VII distinctly transverse, median impression with long dark setae, posterior margin very weakly concave (Fig. 75); sternite VIII approximately as long as broad and with moderately deep, wide, long me- dian impression with sparse black setae, on either side of this impression with moderately dense dark setae, poste- rior margin convex, not concave or incised (Fig. 76); aedeagus approximately 0.8 mm long, ventral process sub- apically bent in lateral view, relatively broad and apical- ly acute in ventral view, dorsal plate lamellate, very thin, moderately long, and weakly sclerotised (Figs 77—79). ©: protarsomeres I-IV dilated, but less so than in male; posterior margin of tergite VII obtusely pointed in the middle (Fig. 80); sternite VIII oblong, longer than tergite VII, strongly convexly produced and with micropubes- cence posteriorly (Fig. 81); tergite [X with median suture; tergite X distinctly longer than tergite [X in the middle. Comparative notes. As can be inferred particularly from the similar shape of the male sternite VIII and the anteri- orly divided female tergite IX, L. inexcisum is closely al- lied to L. diremptum and L. bibarbatum. It is distinguished from both species particularly by smaller body size, by the shape and chaetotaxy of the male sternite VII, and by the distinctly smaller aedeagus with a ventral process and dor- sal plate of different shapes. From the syntopic L. umb- hakense, it 1s additionally separated by the anteriorly di- vided female tergite IX. Etymology. The specific epithet (Latin, adjective) alludes to the absence of a posterior excision of the male stern- ite VIII. Distribution and natural history. The type specimens were found in several localities near Tseram, Yalung, and Yamputhin in Taplejung district, northeastern Nepal (Fig. 3). They were collected in mature fir, rhododendron, and juniper forests at an altitude of 3250-3700 m, in one lo- cality together with L. umbhakense. One female taken in early May is teneral. Lathrobium infractum sp. n. (Figs 3, 82-85) Type material. Holotype ¢@: “523 Nepal: Solukhumbu Distr., E Pangkongma La, 3000 m, 17.V.1997, leg. W. Schawaller / Holotypus ¢ Lathrobium infractum sp. n., Bonn zoological Bulletin 61 (2): 142-209 det. V. Assing 2011” (SMNS). Paratypes: 19 [without ab- dominal apex]: same data as holotype (cAss). Description. Body length 5.8 mm; length of forebody 2.6—3.0 mm. Whole body reddish. Externally highly sim- ilar to L. umbhakense and L. bibarbatum, except for the slightly larger eyes. S: protarsomeres I-IV strongly dilated; sternite VII strongly transverse, median impression with weakly mod- ifed setae, posterior margin weakly concave (Fig. 82); ster- nite VII weakly transverse and with extensive median im- pression with sparse black setae, without lateral clusters of dense dark setae, posterior margin convex, in the mid- dle indistinctly concave (Fig. 83); aedeagus | mm long, ventral process long, sharply bent in lateral view, strong- ly asymmetric in ventral view, dorsal plate lamellate and weakly sclerotised (Figs 84-85). ©: protarsomeres I-IV dilated, but less so than in male; other sexual characters not available (missing in paratype). Comparative notes. Based on the external and sexual characters, L. infractum is closely related to L. nepalori- entis and allied species. It is distinguished from them par- ticularly by the weakly modified pubescence of the male sternites VII and VIII, the extensive impression on the male sternite VIII, as well as by the conspicuous shape of the ventral process of the aedeagus. From many of the re- lated species, it is additionally separated by the slightly larger eyes. Etymology. The specific epithet (Latin, adjective: flexed, broken) alludes to the conspicuous shape of the ventral process of the aedeagus. Distribution and natural history. The type locality is sit- uated in Solukhumbu district in eastern Nepal (Fig. 3). The specimens were collected at an altitude of 3000 m. The Lathrobium muguicum group Lathrobium muguicum sp. n. (Figs 86—91, 96) Type material. Holotype @: “576 Nepal: Mugu Distr., SW Rara Lake, 3200 m, 12.V1I.1998, leg. W. Schawaller / Holotypus ¢ Lathrobium muguicum sp. n., det. V. Assing 2011” (SMNS). Description. Body length 7 mm; length of forebody 3.4 mm. Habitus as in Fig. 86. Coloration: body blackish- brown; legs and antennae reddish. Head (Fig. 87) as long as broad; punctation moderate- ly coarse and not very dense; interstices with distinct mi- croreticulation, on average somewhat broader than diam- eter of punctures, slightly sparser in median dorsal por- ©ZFMK 170 Volker Assing 95 yh 92 Figs 86-95. Lathrobium muguicum (86-91) and L. emodense (92-95). 86: habitus; 87: forebody; 88, 92: male sternite VI; 89, 93: male sternite VIII; 90, 94: aedeagus in lateral view; 91, 95: aedeagus in ventral view. Scale bars: 86-87: 1.0 mm; 88—95: 0.5 mm. tion. Eyes weakly projecting from lateral contours of head, approximately 1/3 the length of postocular region in dor- sal view and composed of approximately 30 ommatidia. Pronotum (Fig. 87) 1.18 times as long as broad and 1.04 times as broad as head; punctation similar to that of head, but slightly coarser; interstices without microsculpture. Elytra short, approximately 0.6 times as long as prono- tum (Fig. 87); humeral angles weakly marked; punctation shallow and weakly defined; interstices without distinct microsculpture. Hind wings completely reduced. Metati- bia slightly compressed. Abdomen broader than elytra; punctation moderately fine and dense, somewhat sparser on posterior than on an- terior tergites; posterior margin of tergite VII without pal- Bonn zoological Bulletin 61 (2): 142-209 isade fringe; posterior margin of tergite VIII weakly con- vex. 3: protarsomeres I-IV strongly dilated; sternite VII ex- tensively impressed and with sparse long black setae in postero-median portion, posterior margin weakly concave (Fig. 88); sternite VIII moderately transverse, with shal- low median impression, this impression with weakly mod- ified black setae, posterior excision small and almost V- shaped (Fig. 89); aedeagus 1.5 mm long, ventral process thin and blade-shaped, dorsal plate lamellate and moder- ately sclerotised, internal sac with a long membranous tube (Figs 90-91). ©: unknown. ©ZFMK Revision of Himalayan Lathrobium 171 Fig. 96. Distributions of the species of the L. muguicum (open square), the L. emodense (stars, circles), the L. jumlense (dia- monds), the L. excisum (filled square), and the L. aculeatum groups (triangles): L. planissimum (grey diamond); L. muguicum (open square); L. jumlense (black diamond); L. inustum (white diamonds); L. aculeatum (black triangle); L. curvum (black star); L. emod- ense (white circles); L. spinosissimum (white star); L. annapurnense (black circle); L. spiculatum (white triangles); L. excisum (black squares). Comparative notes. Neither the external nor the male sex- ual characters suggest a closer relationship to any of the other species recorded from Nepal. In general external ap- pearance, L. muguicum is somewhat similar to the species of the L. nepalense group, but at once distin- guished from them by absence of microsculpture on the pronotum and by the completely different male sexual characters. Etymology. The specific epithet (adjective) is derived from the name of the district where the type locality is sit- uated. Distribution and natural history. The species is known only from one locality in Mugu district, northwestern Nepal (Fig. 96). The holotype was collected at an altitude of 3200 m. The Lathrobium emodense species group Lathrobium emodense Coiffait, 1975 (Figs 92-96) Lathrobium emodense Coiffait, 1975: 183 f. Lathrobium goropanensis [sic] Coiffait, 1983: 172; syn. n. Type material examined. L. emodense: Holotype 9: “Umg. Goropani, W. Pokhara / Pa 143 [overleaf] / Zen- tral-Nepal, Sept.-Okt. 1971, lg. H. Franz / Holotype / La- throbium emodense H. Coiffait 1975 / Lathrobium emo- dense Coiffait, det. V. Assing 2011” (NHMW). Bonn zoological Bulletin 61 (2): 142-209 L. goropanense: Holotype °: “Umg. Goropani, W. Pok- hara / Pa 111 [overleaf] / Holotype / Lathrobium goropa- nensis H. Coiffait 1983 / Lathrobium emodense Coiffait, det. V. Assing 2011” (MNHNP). Comment. Lathrobium emodense was described from a single female from “Environs de Goropani a |’ouest de Pokhara” (Coiffait 1975) deposited in the Franz collection at the NHMW. In the original description of L. goropa- nense, which is based on a unique female holotype from “Népal, environs de Goropani” (Coiffait 1983) deposited in the Coiffait collection at the MNHNP, there is no ref- erence whatsoever to L. emodense, although the type lo- calities of both species are practically the same. Also, the locality labels of both specimens are identical, except for the overleaf sample number. A comparison revealed that the holotype of L. emodense is slightly smaller than that of L. goropanense, but otherwise they are indistinguish- able. In consequence, there 1s little doubt that both names refer to the same species, so that L. goropanense is placed in synonymy with L. emodense. Fortunately, a male from the vicinity of the the environs of Ghorepani has become available, so that an interpre- tation of this species 1s now possible. Additional material examined. Nepal: |, 1°, Parbat district, Punhill at Ghoropani Pass, 3050-3100 m, 8.X.1983, leg. Sme- tana & L6bl (MHNG, cAss). Redescription. Body length 5.5—6.7 mm; length of fore- body 2.4—2.9 mm. Coloration: body more or less uniform- ly reddish. ©ZFMK 172 Volker Assing Head approximately as long as broad or weakly trans- verse; punctation coarse and moderately dense; interstices without distinct microsculpture (some shallow traces may be visible on frons), at least as broad as diameter of punc- tures in median dorsal portion, narrower than diameter of punctures in lateral and posterior dorsal portions. Eyes not projecting from lateral contours of head, small, composed of approximately 20 ommatidia. Pronotum 1|.20—1.25 times as long as broad and as broad as head; punctation similar to that of head; interstices with- out microsculpture. Elytra short, 0.55—0.60 times as long as pronotum; humeral angles weakly marked; punctation shallow; in- terstices without distinct microsculpture. Hind wings com- pletely reduced. Metatibia somewhat compressed. Abdomen broader than elytra; punctation moderately fine and dense; posterior margin of tergite VII without pal- isade fringe; posterior margin of tergite VII truncate in both sexes. 3: protarsomeres I-IV strongly dilated; sternite VII ex- tensively impressed and with moderately dense modified setae in postero-median portion, posterior margin broad- ly and weakly concave (Fig. 92); sternite VII broadly im- pressed along the middle and with moderately numerous modified setae, posterior margin with small posterior ex- cision, on either side of this excision acutely produced (Fig. 93); aedeagus 1.2 mm long, with straight and api- cally acute ventral process (lateral view), a long, straight (lateral view), and distinctly sclerotised dorsal plate, and with several strongly sclerotised internal structures of char- acteristic shape (Figs 94—95). ©: protarsomeres I-IV dilated, but slightly less so than in male; sternite VIII oblong and convexly produced pos- teriorly; tergite [IX undivided; tergite X distinctly longer than tergite [X in the middle. Comparative notes. As can be inferred from the similar external morphology, the similar shapes and chaetotaxy of the male sternites VII and VHI, and particularly by the similarly derived morphology of the aedeagus (shape of ventral process, long and straight dorsal plate, numerous long sclerotised structures in internal sac), L. emodense is closely related to the three following species. It is dis- tinguished from them by larger body size and especially by the shape of the male sternite VII and by the morphol- ogy of the aedeagus. Distribution and natural history. The species is current- ly known only from the environs of Ghorepani in the southwestern extension of the Annapurna range (Fig. 96). The additional specimens were collected at an altitude of 3050-3100 m. According to Franz’ diary, the holotype of L. goropanense was sifted from leaf litter in a montane forest on a southwestern slope on 19.1X.1971, and the Bonn zoological Bulletin 61 (2): 142-209 holotype of L. emodense was collected near Goropani by sifting leaf litter on 27.1X.1971. Lathrobium curvum sp. n. (Figs 96-101) Type material. Holotype @: “Nepal Himalaya, Dhawala- giri, 2004, Baglung Lekh / 30 km west Baglung, N Tara Khola [28°22’°N, 83°20’E], 27-2900 m, 20.05.04, leg. J. Schmidt / Holotypus 4 Lathrobium curvum sp. n., det. V. Assing 2011” (cAss). Description. Body length 6.3 mm; length of forebody 2.9 mm. Coloration: body brown with somewhat paler legs and abdominal apex (apparently darkened post-mortem, coloration of body in freshly collected material probably reddish). Head (Fig. 97) approximately as long as broad; punc- tation coarse and rather dense; interstices without distinct microsculpture, approximately as broad as diameter of punctures in median dorsal portion. Eyes not projecting from lateral contours of head, small, composed of approx- imately 20 ommatidia. Pronotum (Fig. 97) 1.25 times as long as broad and as broad as head; punctation similar to that of head, but slightly less coarse; interstices without microsculpture. Elytra short, 0.6 times as long as pronotum (Fig. 97); humeral angles weakly marked; punctation shallow and ill-defined; interstices without distinct microsculpture. Hind wings completely reduced. Metatibia slightly com- pressed. Abdomen broader than elytra; punctation fine and mod- erately dense; posterior margin of tergite VII without pal- isade fringe. 3: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VIII truncate; sternite VII impressed and with numerous modified setae in postero-median portion, pos- terior margin concave in the middle (Fig. 98); sternite VI- II broadly impressed along the middle and with numer- ous modified setae, posterior margin with broadly concave posterior excision in the middle (Fig. 99); aedeagus large, 1.3 mm long, with apically acute ventral process, strong- ly sclerotised and apically curved dorsal plate, and with large, strongly curved, and strongly sclerotised internal structures (Figs 100-101). ©: unknown. Comparative notes. Based on the similarly derived shape and chaetotaxy of the male sternite VII and VIII, as well as on the similar general morphology of the aedeagus (strongly sclerotised dorsal plate; distinctly sclerotised in- ternal structures; shape of ventral process), L. curvum and the two following species form a well-defined species group confined to the Annapurna and Dhaulagiri ranges OZFMK Revision of Himalayan Lathrobium 173 104 Figs 97-106. Lathrobium curvum (97-101) and L. spinosissimum (102-106). 97, 102: forebody; 98, 103: male sternite VII; 99, 104: male sternite VIII; 100, 105-106: aedeagus in lateral view; 101: aedeagus in ventral view. Scale bars: 97, 102: 1.0 mm; 98-101, 103-106: 0.5 mm. in central Nepal. Among the species of this group, L. curvum and L. spinosissimum are evidently most closely related, as is suggested by the similarly derived morphol- ogy of the aedeagus (shape of internal structures and of the dorsal plate). From L. spinosissimum and L. annapur- Bonn zoological Bulletin 61 (2): 142-209 nense, both from the Annapurna, L. curvum is distin- guished particularly by the male primary and secondary sexual characters (especially the shape and chaetotaxy of the male sternite VIII and the morphology of the aedea- gus), from L. annapurnense additionally by larger size. ©ZFMK 174 Volker Assing Etymology. The name (Latin, adjective) alludes to the sickle-shaped internal structures of the aedeagus. Distribution and natural history. The type locality is sit- uated in the Dhaulagiri range, central Nepal (Fig. 96), at an altitude of 2700-2900 m. Lathrobium spinosissimum sp. n. (Figs 96, 102-106, 125) Type material. Holotype @: ““S Lamjun Himal, W-slope Taunja Danda [28°25’N, 84°12’E], 3700 m, 9.8.95 / Nepal Annapurna-Mts., leg. Frabrizi, Jager, Schmidt / Holoty- pus ¢ Lathrobium spinosissimum sp. n., det. V. Assing 2011” (SDEI). Paratypes: 14, 12: same data as holotype (SDEI, cAss). Description. Body length 5.8—6.1 mm; length of forebody 2.7 mm. Coloration: body brown to dark-brown; legs and antennae reddish-brown. Head (Fig. 102) approximately as long as broad or weakly oblong; punctation coarse and rather dense; inter- stices without distinct microsculpture (some shallow traces may be visible on frons), at least as broad as diameter of punctures in median dorsal portion, narrower than diam- eter of punctures in lateral and posterior dorsal portions. Eyes not projecting from lateral contours of head, small, composed of approximately 20 ommatidia. Pronotum (Fig. 102) approximately 1.25 times as long as broad and as broad as head; punctation similar to that of head, but slightly less coarse; interstices without mi- crosculpture. Elytra short, approximately 0.55 times as long as prono- tum (Fig. 102); humeral angles weakly marked; puncta- tion shallow and ill-defined; interstices without distinct mi- crosculpture. Hind wings completely reduced. Metatibia somewhat compressed. Abdomen broader than elytra; punctation moderately fine and dense; posterior margin of tergite VII without pal- isade fringe; posterior margin of tergite VHI weakly con- vex in both sexes. 3: protarsomeres I-IV strongly dilated; sternite VII shallowly impressed and with moderately dense modified setae in postero-median portion, posterior margin broad- ly concave (Fig. 103); sternite VHI broadly impressed along the middle and with moderately numerous modified setae, posterior margin with small posterior excision in the middle (Fig. 104); aedeagus 1.0 mm long, with straight and apically acute ventral process (lateral view), a long, distinctly sclerotised, and apically hooked dorsal plate, and with several strongly sclerotised internal structures of char- acteristic shape (Figs 105—106). 2: protarsomeres I-IV dilated, but slightly less so than in male; sternite VII oblong and convexly produced pos- teriorly (Fig. 125); tergite [X undivided; tergite X slight- ly longer than tergite [X in the middle. Bonn zoological Bulletin 61 (2): 142-209 Comparative notes. Lathrobium spinosissimum 1s char- acterised particularly by the morphology of the aedeagus (shape of ventral process and dorsal plate, internal struc- tures), as well as by the shape and chaetotaxy of the male sternites VII and VIII. The similar primary and second- ary sexual characters suggest that the species is closely related to the smaller L. annapurnense. Etymology. The specific epithet (adjective, superlative of spinosus) alludes to the conspicuous assortment of scle- rotised internal structures. Distribution and natural history. The type locality is sit- uated in the southern Lamjun (or Lamjung) Himal in cen- tral Nepal (Fig. 96). The specimens were collected at an altitude of 3700 m. Lathrobium annapurnense sp. n. (Figs 96, 107-111) Type material. Holotype @: “Nepal-Himalaya, Annapur- na Mts., S Lamjun Himal, Namun Pass, S Hang, 4900 m, 28.5.1993, leg. J. Schmidt / Holotypus ¢ Lathrobium an- napurnense sp. n., det. V. Assing 2011” (cAss). Paratypes: 14, 19: same data as holotype (cKle). Description. Body length 5.0—5.5 mm; length of forebody 2.42.5 mm. Coloration: body dark-brown; legs reddish- brown; antennae reddish. Head (Fig. 107) approximately as long as broad; punc- tation coarse and rather dense; interstices without mi- crosculpture, approximately as broad as diameter of punc- tures in median dorsal portion, narrower than diameter of punctures in lateral and posterior dorsal portions. Eyes not projecting from lateral contours of head, small, composed of approximately 25 ommatidia. Pronotum (Fig. 107) 1.20—1.25 times as long as broad and as broad as head; punctation similar to that of head, but slightly less coarse; interstices without microsculpture. Elytra short, 0.6 times as long as pronotum (Fig. 107); humeral angles weakly marked; punctation shallow and ill-defined; interstices without distinct microsculpture. Hind wings completely reduced. Metatibia weakly com- pressed. Abdomen broader than elytra; punctation fine and dense; posterior margin of tergite VII without palisade fringe. ¢: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VIII truncate; sternite VII shallowly im- pressed and with relatively sparse, weakly modified se- tae in postero-median portion, posterior margin broadly concave (Fig. 108); sternite VII broadly impressed along the middle and with moderately numerous modified se- tae, posterior margin with concave posterior excision in the middle (Fig. 109); aedeagus 0.83 mm long, with rel- ©ZFMK Revision of Himalayan Lathrobium 175 ee 111 114 Figs 107-117. Lathrobium annapurnense (107-111) and L. excisum (112-117). 107, 112: forebody; 108, 113: male sternite VII; 109, 114: male sternite VIII; 110, 115-116: aedeagus in lateral view; 111, 117: female sternite VIII; 116: aedeagus in ventral view. Scale bars: 107, 112: 1.0 mm; 108-111, 113-117: 0.5 mm. atively broad ventral process (lateral view), a long, strong- _—_ sternite VIII weakly elongated, convexly produced pos- ly sclerotised, and apically hooked dorsal plate, and with __ teriorly (Fig. 111); tergite [IX undivided; tergite X approx- membranous internal structures (Fig. 110). imately as long as tergite IX in the middle. ©: protarsomeres I-IV dilated, but slightly less so than in male; posterior margin of tergite VII weakly convex; Bonn zoological Bulletin 61 (2): 142-209 OZFMK 176 Volker Assing Comparative notes. Among the species of the L. emod- ense group, L. annapurnense is characterised particular- ly by the morphology of the aedeagus (shape of ventral process and dorsal plate, internal sac without spine-like sclerotised structures), the shape and chaetotaxy of the male sternites VII and VIII, as well as by its small size. From most Himalayan Lathrobium species of similarly small size, it is additionally distinguished by the darker coloration. Etymology. The specific epithet (adjective) is derived from the name of the mountain range where the type lo- cality is situated. Distribution and natural history. The species is known from only one locality in the Annapurna range (Fig. 96), where the specimens were collected at an altitude of 4900 m. The Lathrobium excisum group Lathrobium excisum sp. n. (Figs 96, 112-117, 295) Type material. Holotype @: “Ost-Nepal, Rolwaling Hi- mal / nordéstl. unterh. d. Daldung La Pass, 3800 m, 27.05.2000 leg. A. Kleeberg / Holotypus 4 Lathrobium excisum sp. n., det. V. Assing 2011” (cAss). Paratypes: 22°: same data as holotype (cKle); 329: “Ost-Nepal, Rolwaling Himal / Rolwaling Tal, Nyimare, 3300 m, 19.05.2000 leg. A. Kleeberg” (cKle, cAss). Description. Body length 5.5—6.7 mm; length of forebody 2.5—3.0 mm. Coloration: body reddish, with slightly dark- er abdominal segments HI—VI and with slightly paler legs. Head (Fig. 112) weakly oblong, 1.05—1.10 times as long as broad; punctation relatively coarse and moderately dense; interstices with very indistinct traces of microretic- ulation, glossy. Eyes composed of approximately 20 om- matidia, with pigmentation. Pronotum (Fig. 112) approximately 1.3 times as long as broad and slightly narrower than head; punctation approx- imately as dense as that of head, but less coarse; interstices without microsculpture. Elytra short, approximately 0.6 times as long as prono- tum (Fig. 112). Hind wings completely reduced. Legs not compressed. Abdomen broader than elytra; punctation rather fine and dense; interstices with distinct microsculpture; posterior margin of tergite VII without palisade fringe. &: protarsomeres I-IV more strongly dilated than in fe- male; posterior margin of tergite VIII truncate; sternite VII extensively, but not very deeply impressed in posterior me- dian portion, this impression with numerous modified se- Bonn zoological Bulletin 61 (2): 142-209 tae, posterior margin weakly concave (Fig. 113); sternite VII broadly impressed along the middle, this impression with pair of clusters of numerous modified setae, poste- rior margin with distinct median excision (Fig. 114); aedeagus 1.15 mm long, with apically weakly curved ven- tral process (lateral view) and with lamellate dorsal plate (Figs 115-116). ®: posterior margin of tergite VIII unmodified; stern- ite VIII (Fig. 117) convexly produced posteriorly; tergite IX not divided in the middle; tergite X distinctly longer than tergite IX in the middle. Comparative notes. Lathrobium excisum is distinguished from other microphthalmous and depigmented represen- tatives of the genus particularly by the shapes and chaeto- taxy of the male sternites VII and VIII, as well as by the morphology of the aedeagus. Based on the similar shape and chaetotaxy of the male sternite VIII, this species is probably most closely affiliated with the species of the L. emodense group, from which it is separated by the mor- phology of the aedeagus (absence of sclerotised spines in internal sac, lamellate dorsal plate, shape of ventral process). Etymology. The specific epithet (Latin, adjective) refers to the posterior excision of the male sternite VIII, one of the characters distinguishing L. excisum from the geo- graphically close and externally similar L. janetscheki. Distribution and natural history. Lathrobium excisum was collected in two localities in the Rolwaling Himal, eastern Nepal (Fig. 96), at altitudes of 3300 and 3800 m (Fig. 295). In one of the localities, the species was found together with L. kleebergi. The Lathrobium gladiator group Lathrobium gladiator Coiffait, 1982 (Figs 118—124) Lathrobium gladiator Coiffait, 1982b: 286 f. Type material examined. Holotype @: “Inde Samum, Bhadarwak, 3200-3700 T.D / Holotype / Lathrobium glad- iator H. Coiffait 1982 / Lathrobium gladiator Corffait, det. V. Assing 2011” (MNHNP). Paratype 2: same data as holotype (MNHNP). Comment. The original description is based on a male holotype and a female paratype from “Inde, Pir Panyal, Ghadarwak, Samum, 3200 m” (Coiffait 1982b), both de- posited in the Coiffait collection at the MNHNP. Redescription. Body length 5.0—5.2 mm; length of fore- body 2.5 mm. Coloration: body uniformly reddish-yellow. ©ZFMK Revision of Himalayan Lathrobium 177 118 124 Figs 118-125. Lathrobium gladiator (118-124) and L. spinosissimum (125). 118: male sternite VII; 119: male sternite VIII; 120: aedeagus in lateral view; 121: aedeagus in ventral view; 122, 125: female sternite VIII; 123: female tergites [X—X in dorsal view. 124: female tergites [x—X in lateral view. Scale bars: 0.5 mm. Head approximately as long as broad or weakly oblong; punctation coarse and relatively sparse; interstices with distinct microreticulation, on average broader than diam- eter of punctures. Eyes not projecting from lateral contours of head, small, composed of approximately ten ommatidia. Pronotum 1.25—1.30 times as long as broad and slight- ly narrower than head, weakly tapering posteriad; punc- tation finer than, and of similar density as that of head; interstices without microsculpture. Elytra short, 0.55—0.60 times as long as pronotum; humeral angles weakly marked; punctation shallow; in- terstices without distinct microsculpture. Hind wings com- pletely reduced. Metatibia somewhat compressed. Abdomen broader than elytra; punctation moderately fine and dense; posterior margin of tergite VII without pal- isade fringe; posterior margin of tergite VIII weakly con- vex in both sexes. Bonn zoological Bulletin 61 (2): 142-209 &: protarsomeres I-IV strongly dilated; sternite VII with extensive and distinct impression, this impression with nu- merous modified setae, posterior margin broadly concave, in the middle with additional concavity (Fig. 118); stern- ite VII narrowly and distinctly impressed along the mid- dle, this impression with numerous modified, short and stout setae, posterior excision V-shaped and moderately deep, on either side of this incision with acute projections (Fig. 119); aedeagus 1.2 mm long, ventral process long and apically hooked, dorsal plate strongly sclerotised, in- ternal sac with sclerotised internal structure (Figs 120-121). ®: protarsomeres I-IV dilated, but distinctly less so than in male; sternite VII oblong, longer than tergite VIII, and rather narrowly produced posteriorly (Fig. 122); tergite [X undivided; tergite X distinctly longer than tergite [X in the middle, strongly bulging, almost keeled in posterior half in cross-section (Figs 123—124). ©ZFMK 178 Volker Assing Fig. 126. Distributions of the species of the L. discissum (stars) and the L. deuvei groups (other symbols): L. barthei (filled square); L. palatum (filled triangle); L. aciforme (open circle); L. lamjunense (filled diamonds); L. apalatum (filled circles); L. deuvei (open square); L. rupinaicum (open diamonds); L. ganeshense (open triangles); L. discissum (open stars). Comparative notes. Based on the male primary and sec- ondary sexual characters, this species is most closely al- lied to the species of the L. emodense group. It is distin- guished from them particularly by the distinctive shape and chaetotaxy of the male sternite VII (shape somewhat similar to that of L. emodense), the morphology of the aedeagus, as well as by the conspicuous shape of the fe- male tergite X. Distribution and natural history. The species is current- ly known only from the environs of the Pir Panjal pass [ca. 33°38°N, 74°31°E] in Kashmir. According to the la- bels attached to the type specimens they were collected between 3200 and 3700 m. The Lathrobium deuvei group Lathrobium deuvei Coiffait, 1981 (Figs 126—130) Lathrobium deuvei Coiffait, 1981: 329. Type material examined. Holotype @: “Nepal VIII.80, Dudh Pokari / 4300 Himal Chuli D.L / Holotype / Lath- robium deuvei H. Coiffait 1981 / Lathrobium deuvei Coif- fait, det. V. Assing 2011” (MNHNP). Paratypes: 14, 19: same data as holotype (but labelled “Paratype” and “Al- lotype”, respectively) (MNHNP). Comment. The original description is based on the male holotype and nine paratypes from ““Duth [sic] Pokhari, Hi- mal Chuli 4300 m” deposited in the Coiffait collection. Bonn zoological Bulletin 61 (2): 142-209 Redescription. Body length 7.5—8.5 mm; length of fore- body 3.7-3.9 mm. Coloration: body reddish-brown to dark-brown. Head approximately as broad as long; punctation coarse and dense; interstices in lateral and posterior dorsal por- tions on average narrower than diameter of punctures; in- terstices without microreticulation, glossy. Eyes very small, composed of approximately ten ommatidia. Pronotum relatively slender, 1.25—1.30 times as long as broad and approximately 0.95 times as broad as head, no- ticeably tapering posteriad; punctation similar to that of head, but slightly finer; interstices on average as broad as, or slightly wider than diameter of punctures, without mi- crosculpture and glossy. Elytra short, approximately 0.6 times as long as prono- tum; humeral angles weakly marked; punctation shallow and ill-defined; interstices without distinct microsculpture. Hind wings completely reduced. Metatibia unmodified. Abdomen broader than elytra, widest at segment VI; punctation dense and fine; interstices with shallow fine- ly transverse microsculpture; posterior margin of tergite VII without palisade fringe. ¢: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VII] weakly convex; sternite VII strongly transverse, with shallow and rather extensive impression posteriorly, this impression with numerous black setae, posterior margin with broadly and rather deeply concave posterior excavation, middle of this excavation not dis- tinctly bisinuate (Fig. 127); sternite VIII with numerous short black setae, particularly at margin of posterior ex- cision, posterior excision large, acute, and very deep, reaching clearly beyond middle of sternite (Fig. 128); ©ZFMK Revision of Himalayan Lathrobium 179 Figs 127-136. Lathrobium deuvei (127-130) and L. lamjunense (131-136). 127, 132: male sternite VII; 128, 133: male sternite VIII; 129, 134-135: aedeagus in lateral view; 130, 136: female sternite VHI; 131: forebody. Scale bars: 131: 1.0 mm; 127-130, 132-136: 0.5 mm. aedeagus large, approximately 1.9 mm long; ventral process long, slender, and apically hooked; dorsal plate with lamellate basal portion and with long, slender, strong- ly sclerotised, and apically hooked apical portion; inter- nal sac with very long, slender, and apically acute sclero- tised structures (Fig. 129). Bonn zoological Bulletin 61 (2): 142-209 ©: protarsomeres I-IV dilated, but somewhat less so than in male; tergite VIII of similar shape as in male; ster- nite VIII posteriorly convex, not conspicuously produced (Fig. 130); tergite [X not divided in the middle; tergite X much shorter than tergite [X in the middle. ©ZFMK 180 Volker Assing Comparative notes. Lathrobium deuvei and the six fol- lowing species form a group of highly similar and un- doubtedly closely related species, all of which are distrib- uted in central and western Nepal. The representatives of this group, hereafter referred to as the L. deuvei group, are practically indistinguishable based on external characters and characterised by moderate to large body size, minute eyes with very few ommatidia, a slender and posteriorly tapering pronotum, a strongly transverse and posteriorly distinctly excavated male sternite VII, a transverse stern- ite VII with a deep (exception: L. ganeshense) and broad- ly V-shaped posterior excision and often conspicuous fringes or cluster of dense dark setae, a large and slender aedeagus with an apically acute, either hooked or straight ventral process, with a dorsal plate composed of a lamel- late basal portion and a slender, distinctly sclerotised and apically hooked apical portion, and with long, distinctly sclerotised, and apically acute internal structures. In ad- dition, some species have a posteriorly distinctly produced female sternite VII. Lathrobium deuvei is distinguished from other species of the L. deuvei group particularly by the shape of the ven- tral process of the aedeagus, the shape of the male stern- ite VII (strongly transverse, posterior excavation regular- ly concave and without bisinuate anterior margin), the shape of the male sternite VII (posterior excision conspic- uously deep, its depth much more than half the length of sternite), and by the weakly modified female sternite VII. Distribution and natural history. Lathrobium deuvei is known only from the type locality, the Himal Chul in the Manaslu range (Fig. 126). The type specimens were col- lected at an altitude of 4300 m. Lathrobium lamjunense sp. n. (Figs 126, 131—136) Type material. Holotype ¢: “Lamjun Himal, Dudh Pokhari [28°26’N, 84°16’E], 444600 m [sic], 15.8.95 / Nepal Annapurna-Mts., leg. Fabrizi, Jager, Schmidt / Holotypus ¢ Lathrobium lamjunense sp. n., det. V. Ass- ing 2011” (SDEI). Paratypes: 15 exs.: same data as holo- type (SDEL, cAss); 3 exs.: “S Lamjun Himal, Wasserschei- de Khudi-Miyardi-Chhar Khola [28°25’N, 84°15’E], 15.8.95, 4300 m / Nepal Annapurna-Mts., leg. Fabrizi, Jager, Schmidt” (SDEI, cAss); 2 exs.: “S Lamjun Himal, E Taunja Danda, Quellgeb. Chhar Khola [28°25’N, 84°14°E], 4300 m, 14.8.95 / Nepal Annapurna-Mts., leg. Fabrizi, Jager, Schmidt” (SDEI). Description. Rather large species; body length 7.5—9.0 mm; length of forebody 3.5—-4.0 mm. Coloration: body reddish-brown to dark-brown. Forebody as in Fig. 131. External characters as in L. deuvei. Bonn zoological Bulletin 61 (2): 142-209 &: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VIII indistinctly angled in the middle; ster- nite VII weakly impressed and with numerous blackish se- tae posteriorly, posterior margin strongly excavated in the middle, anterior margin of this excavation weakly bisin- uate (Fig. 132); sternite VIUI weakly impressed and with numerous short black setae, posterior excision pro- nounced, deeply and broadly V-shaped (Fig. 133); aedea- gus large, 1.8—2.0 mm long; ventral process long, slen- der, and apically hooked; dorsal plate with lamellate basal portion and with long, slender, strongly sclerotised, and apically hooked apical portion; internal sac with very long, slender, and apically acute sclerotised structures (Figs 134-135). ©: protarsomeres I-IV dilated, but somewhat less so than in male; tergite VIII of similar shape as in male; ster- nite VIII posteriorly conspicuously produced in the mid- dle (Fig. 136); tergite [IX not divided in the middle; ter- gite X shorter than tergite [X in the middle. Comparative notes. Lathrobium lamjunense 1s reliably distinguished from other species of the L. deuvei group only by the male primary and secondary sexual charac- ters. Etymology. The specific epithet (adjective) is derived from the name of the mountain range where the species was discovered. Distribution and natural history. Lathrobium lamjunense may be endemic to the Lamjun Himal in cen- tral Nepal (Fig. 126). The species has been collected at altitudes of 4300-4600 m. Lathrobium palatum sp. n. (Figs 126, 137-141) Type material. Holotype <: “Nepal c.: Annapurna Mts.: Kang La-Pass, N-/ slope, E Manang, 5000 m, 5.V1I.1994, leg. J. Schmidt / Holotypus ¢ Lathrobium palatum sp. n., det. V. Assing 2011” (SDEI). Paratypes: 292 9: same da- ta as holotype (SDEI, cAss). Description. Body length 6.5—8.0 mm; length of forebody 3.4-3.8 mm. Coloration: whole body uniformly reddish. Head weakly oblong (Fig. 137). Other external characters as in L. deuvei. 3: posterior margin of tergite VIII weakly convex in the middle; sternite VII similar to that of ZL. deuvei, but pos- terior excavation shallower and with more distinctly bis- inuate anterior margin (Fig. 138); sternite VII similar to that of L. deuvei (Fig. 139); aedeagus similar to that of L. deuvei, but smaller, 1.7 mm long, ventral process apical- ly angled, not curved (Fig. 140). ©OZFMK Revision of Himalayan Lathrobium 181 138 143 Figs 137-146. Lathrobium palatum (137-141) and L. aciforme (142-146). 137, 142: forebody; 138, 143: male sternite VI; 139, 144: male sternite VIII; 140, 145: aedeagus in lateral view; 141, 146: female sternite VIII. Scale bars: 137, 142: 1.0 mm; 138-141, 143-146: 0.5 mm. ©: secondary sexual characters similar to those of L. deuvei, but sternite VIII strongly produced posteriorly (Fig. 141) Comparative notes. Lathrobium palatum is distin- guished from the similar L. dewvei particularly by the no- ticeably oblong head, smaller average size, paler col- oration, the shape of the posterior excavation of the male Bonn zoological Bulletin 61 (2): 142-209 sternite VII, by the morphology of the smaller aedeagus (shape of the apex of the ventral process and of the inter- nal structures), and by the shape of the female sternite Vill. Etymology. The name is an adjective is derived from the Latin noun pala (shovel, spade) and alludes to the shape of the female sternite VIII. ©ZFMK 182 Volker Assing Distribution and natural history. The type locality is sit- uated in the northern Annapurna Himal to the east of Man- ang, central Nepal (Fig. 126). The specimens were col- lected at an altitude of 5000 m. Lathrobium aciforme sp. n. (Figs 126, 142-146) Type material. Holotype @: “Nepal: N-Annapurna Mts.: Pisang Peak, / N-slope, 4500-4700 m, 6.V1.1994, leg. J. Schmidt / Holotypus ¢ Lathrobium aciforme sp. n., det. V. Assing 2011” (SDEI). Paratypes: 5434, 199 9: same data as holotype (SDEI, cAss). Description. Body length 7.0—8.5 mm; length of forebody 3.4-3.9 mm. Coloration: whole body reddish to reddish- brown. Head often noticeably oblong and posteriorly ta- pering (Fig. 142). Other external characters as in L. deu- vel. &: posterior margin of tergite VIII weakly convex; ster- nite VII similar to that of L. deuvei, but posterior excava- tion shallower and with more distinctly bisinuate anteri- or margin (Fig. 143); sternite VII similar to that of L. deu- vei, but posterior excision shallower and somewhat asym- metric (Fig. 144); aedeagus similar to that of L. deuvei, but smaller, 1.7—1.8 mm long, apex of ventral process of slightly different shape, and internal structures subapical- ly abruptly narrowed and apically needle-shaped (Fig. 145). ©: sternite VIII similar to that of L. palatum, but pos- terior projection shorter and basally broader; other second- ary sexual characters as in L. deuvei. Comparative notes. Lathrobium aciforme is reliably dis- tinguished from other representatives of the L. deuvei group by the shape of the aedeagus (especially the shape of the apex of the ventral process and the shape of the in- ternal structures), the shape and chaetotaxy of the male sternite VII, the shallower and somewhat asymmetric pos- terior excision of the male sternite VII, as well as by the shape of the female sternite VII. Etymology. The specific epithet (Latin, adjective) alludes to the needle-shaped apex of the internal structures of the aedeagus, one of the characters separating this species from its closest relatives. Distribution and natural history. The type locality, the Pisang Himal, is situated in the northern Annapurna Hi- mal (Fig. 126), not far from the type locality of L. pala- tum. The specimens were collected at an altitude of 4500-4700 m. One of the females is teneral. Bonn zoological Bulletin 61 (2): 142—209 Lathrobium apalatum sp. n. (Figs 126, 147-151) Type material. Holotype @: “Meme Pokhari Lekh, Meme Pokhari [28°22’N, 84°35’E], 4300 m, 8.9. / Nepal Hi- malaya, Manaslu-Mts., lg. Schmidt 1995 / Holotypus 2 Lathrobium apalatum sp. n., det. V. Assing 2011” (SDEI). Paratypes: 34.4, 19: same data as holotype (SDEI, cAss); 733, 529: “Himal Chuli, Meme Pokhari Lekh: Dudh Pokhari [28°23’N, 84°35’E], 4400 m, 9.9. / Nepal Hi- malaya, Manaslu-Mts., lg. Schmidt 1995” (SDEI, cAss). Description. Body length 7.0—-8.5 mm; length of forebody 3.4-3.9 mm. Forebody as in Fig. 147. Coloration: whole body reddish to reddish-brown, with the abdominal seg- ments III-VI or IH-VII often darker. Other external char- acters as in L. deuvei. G: posterior margin of tergite VIII weakly convex or truncate; sternite VII similar to that of L. deuvei, but pos- terior excavation shallower and with more distinctly bi- sinuate anterior margin (Fig. 148); sternite VIII similar to that of L. deuvei, but posteriorly with denser blackish pu- bescence and posterior excision even deeper and broad- er (Fig. 149); aedeagus similar to that of L. dewvei, but smaller, 1.7—1.8 mm long, ventral process apically stouter, apical portion of dorsal plate shorter, and internal structures of slightly different shape (Fig. 150). ©: sternite VIII not conspicuously produced posterior- ly, posterior margin obtusely angled (Fig. 151); other sec- ondary sexual characters as in L. deuvei. Comparative notes. Lathrobium apalatum 1s reliably dis- tinguished from other representatives of the L. deuvei group by the shape of the aedeagus (especially the stouter apex of the ventral process), the shape and chaetotaxy of the male sternite VII, the deeper and broader posterior ex- cision of the male sternite VIII, as well as by the complete- ly different shape of the female sternite VIII. Etymology. The specific epithet (Latin, adjective) signi- fies that, in contrast to other species of the L. deuvei group, the female sternite VIII is not conspicuously produced pos- teriorly. Distribution and natural history. The species is only known from the Meme (or Mimi) Pokhari in the south- ern Manaslu Himal (Fig. 126). The specimens were col- lected at an altitude of 4300-4400 m; one of them is ten- eral. Lathrobium rupinaicum sp. n. (Figs 126, 152-155) Type material. Holotype ¢ [dissected prior to present study]: “Nepal-Expeditionen Jochen Martens / 241 Gorkha Dist., Rupina La, S-Seite, 4500-4100 m, 9 Aug ©ZFMK Revision of Himalayan Lathrobium 183 149 Figs 147-155. Lathrobium apalatum (147-151) and L. rupinaicum (152-155). 147, 152: forebody; 148, 153: male sternite VI; 149, 154: male sternite VIII; 150, 155: aedeagus in lateral view; 151: female sternite VIII. Scale bars: 147, 152: 1.0 mm; 148-151, 153-155: 0.5 mm. 83 Martens & Schawaller leg. / Lathrobium n. sp. cf. deu- vei Coiff., det. 198 [sic] G. de Rougemont / Holotypus ¢ Lathrobium rupinaicum sp. n., det. V. Assing 2011” (SMNS). Paratypes: 1 [dissected prior to present study]: “Nepal-Expeditionen Jochen Martens / 239 Gorkha Dist., zw. Tabruk und Rupina La, 4400-4500 m, 9Aug83 Martens & Schawaller” (SMNS); 1¢ [dissected prior to present study]: “Nepal-Expeditionen Jochen Martens / 238 Bonn zoological Bulletin 61 (2): 142-209 Gorkha Dist., zw. Tabruk und Rupina La, 4100-4400 m, 9 Aug 83 Martens & Schawaller” (cAss). Comment. Two of the type specimens have evidently been subject to post-mortem darkening, most likely as a result of the application of a chemical of unknown iden- tity. ©ZFMK 184 Volker Assing Description. Body length 6.8—7.5 mm; length of forebody 3.2—3.4 mm. Coloration: whole body reddish. Externally highly similar to the preceding species, except for the smaller size (Fig. 152). 3: posterior margin of tergite VIII truncate; sternite VII with shallow and weakly bisinuate posterior excavation, anterior margin of this excavation with longer black se- tae (Fig. 153); sternite VII in the middle with extensive cluster of dense blackish setae, posterior excision broad and moderately deep (Fig. 154); aedeagus (Fig. 155) 1.3—1.4 mm long, of similar general morphology as in L. deuvei. @: unknown. Comparative notes. Lathrobium rupinaicum 1s distin- guished from other representatives of the L. deuvei group by the smaller average size (see measurements of length of forebody), by the shape and chaetotaxy of the male ster- nites VII and VII, as well as by the distinctly smaller aedeagus with a much shorter dorsal plate. Etymology. The specific epithet (adjective) is derived from the name of the pass where this species was discov- ered. Distribution and natural history. The type specimens were collected in three localities near Rupina La, a pass in the southern Manaslu Himal (Fig. 126), central Nepal, at altitudes of 4100-4500 m. Lathrobium barthei Coiffait, 1987 (Figs 126, 156—160) Lathrobium alticola Coiffait, 1975: 331 f.; preoccupied. Lathrobium barthei Coiffait, 1987: 497; replacement name. Type material examined. Holotype ¢: “Umg. Mahi- doela-Pal}, S000 m, b. Maharigaon / Pa 208 [overleaf] / Gebiet von Jumla, Westnepal, lg. H. Franz / Holotype / Lathrobium alticola H. Coiffait 1975 / Lathrobium barthei Coiffait, det. V. Assing 2011” (NHMW). Comment. The original description of L. alticola is ba- sed on a unique male holotype from “Environs du Col de Mahidoela, 5000 m, pres de Maharigaon, région de Jula, Népal occidental” (Coiffait 1975). The preoccupied name L. alticola was subsequently replaced with L. barthei by Coiffait (1987). Redescription. Body length 8.2 mm; length of forebody 4.3 mm. Forebody as in Fig. 156. Coloration: whole body reddish. Other external characters similar to those of L. deuvei. 3: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VIII weakly convex; sternite VII shallowly, but extensively impressed and with numerous blackish se- Bonn zoological Bulletin 61 (2): 142—209 tae posteriorly, posterior margin strongly excavated in the middle, anterior margin of this excavation not bisinuate (Fig. 157); sternite VIII weakly impressed and with dense blackish pubescence in the middle, posterior excision pro- nounced, deep, acute, and slightly asymmetric (Fig. 158); aedeagus large, approximately 1.6 mm long; ventral process long and slender; dorsal plate with lamellate basal portion and with long, slender, strongly sclerotised, and apically hooked apical portion; internal sac with very long, slender, and apically acute sclerotised structures (Figs 159-160). (Note that the apex of the ventral process is ev- idently broken off in the holotype.) ©: unknown. Comparative notes. As can be inferred from the similar- ly derived shape and chaetotaxy of the male sternites VII and VII, as well as from the similarly derived morphol- ogy of the aedeagus, L. barthei 1s evidently closely relat- ed to L. deuvei and its allies. It is reliably distinguished from other representatives of this group only by the male primary and secondary sexual characters. Distribution and natural history. Lathrobium barthei is the only representative of the L. deuvei group in West Nepal; all other species of this group are distributed in cen- tral Nepal. The holotype was collected at the Mahidoela pass [29°20’N, 82°23’E] near Maharigaon, Jumla region (Fig. 126), at an altitude of 5000 m. According to Franz’ diary, the holotype was sifted from moss at rocks near snow on 23.V.1972. In contrast to the altitude stated on the label, the altitude specified in the diary is 5200 m. Lathrobium ganeshense Coiffait, 1983 (Figs 126, 161-166) Lathrobium ganeshensis [sic] Coiffait, 1983: 170. Type material examined. Holotype ¢: “Népal VII.82, Ganesh Himal, Lari, 4500 TD / Holotype / Lathrobium ganeshensis [sic] H. Coiffait 1983 / Lathrobium ganesh- ense Coiffait, det. V. Assing 2011” (MNHNP). Paratypes: 14, 19: same data as holotype (MNHNP). Comment. The original description is based on a male holotype and five paratypes (four males and one female) from “Ganesh Himal, Lari, 4500 m” (Coiffait 1983), all of them deposited in the Coiffait collection. Additional material examined. Nepal: 20 exs., Ganesh Himal, Jaisuli Kund env., 4300-4500 m, 13.—16.VI.2000 (SNSD, cAss). Redescription. Body length 6.7—8.0 mm; length of fore- body 3.2—-3.9 mm. Forebody as in Fig. 161. Apart from the slightly smaller body size, externally highly similar to L. barthei (including coloration). ©OZFMK Revision of Himalayan Lathrobium 185 158 163 166 Figs 156-166. Lathrobium barthei (156-160) and L. ganeshense (161-166). 156, 161: forebody; 157, 162: male sternite VII; 158, 163: male sternite VIII; 159, 164: aedeagus in lateral view; 160, 165: aedeagus in ventral view; 166: female sternite VIII. Scale bars: 156, 161: 1.0 mm; 157—160, 162—166: 0.5 mm. S: protarsomeres I-IV moderately dilated; posterior margin of tergite VIII weakly convex; sternite VII strongly transverse, with shallow, extensive median im- pression posteriorly, this impression with numerous blackish setae, posterior margin broadly concave (Fig. 162); sternite VII transverse, with shallow impression posteriorly, this impression with cluster of very dense black setae, posterior excision broadly V-shaped, but not Bonn zoological Bulletin 61 (2): 142-209 very deep, its depth approximately 1/4 the length of ster- nite (Fig. 163); aedeagus 1.6 mm long, with long, almost straight, slender, and apically acute ventral process, a bi- sinuate (lateral view) and apically strongly hooked dor- sal plate composed of a lamellate basal and a strongly scle- rotised apical portion, internal sac with two long sclero- tised structures (Figs 164-165). OZFMK 186 Volker Assing ©: protarsomeres I-IV dilated, but slightly less so than in male; posterior margin of tergite VIII weakly convex; sternite VIII distinctly longer than tergite VIII, its poste- rior margin moderately convexly produced in the middle (Fig. 166); tergite LX undivided in the middle; tergite X shorter than tergite [X in the middle. Comparative notes. Lathrobium ganeshense is distin- guished from other species of the L. dewvei group by slightly smaller body size, the shape of the posterior mar- gin of the strongly transverse male sternite VI, the shape (posteriorly less deeply excised) and chaetotaxy (presence of a cluster of dense black setae in the middle) of the male sternite VIL, the morphology of the aedeagus, as well as by the shape of the female sternite VII. Distribution and natural history. The type locality, Lari (a zinc mine; approximately 28°14’N, 85°11’), is situat- ed to the southeast of the Paldol peak in the Ganesh Hi- mal, central Nepal. The additional material was collect- ed near Jaisuli Kund [28°13’N, 85°13’E] (Fig. 126). The specimens were collected at an altitude of 4300-4500 m. The Lathrobium discissum group Lathrobium discissum sp. n. (Figs 126, 167-174) Type material. Holotype @: “India W. Bengal, Darjee- ling distr., Tonglu 3100 m 16.X.1978, Besuchet-Lobl / Holotypus 4 Lathrobium discissum sp. n., det. V. Assing 2012” (MHNG). Paratypes: 443, 499 [4 exs. at least slightly teneral]: same data as holotype (MHNG, cAss); 13: same data, but 2700 m (cAss); 243, 19: “Nepal-Ex- peditionen Jochen Martens / 324 Panchthar Distr., Dhor- par Kharka, mature Rhododendron-Lithocarpus forest, 2700 m, 13-16 Apr 88 Martens & Schawaller” (SMNS, cAss). Description. Species of moderately large size; body length 7.5—9.0 mm; length of forebody 3.1—3.8 mm. Habitus as in Fig. 167. Coloration: head and pronotum reddish-brown to blackish-brown; elytra reddish to reddish-brown; ab- domen dark-brown to blackish; legs reddish to dark- brown; antennae reddish. Head (Fig. 168) approximately as broad as long or weakly oblong; punctation moderately coarse and rather sparse; interstices with distinct fine microreticulation, al- most matt, distinctly broader than diameter of punctures. Eyes small, but composed of more than 20 ommatidia, not projecting from lateral contours of head, and 1/5—1/4 the length of postocular region in dorsal view. Pronotum (Fig. 168) relatively slender, 1.25—1.30 times as long as broad and approximately as broad as head, or nearly so; lateral margins almost subparallel in dorsal Bonn zoological Bulletin 61 (2): 142-209 view; punctation similar to that of head; interstices with- out microsculpture and glossy. Elytra short, approximately 0.55 times as long as prono- tum (Fig. 168) ; humeral angles weakly marked; puncta- tion fine, shallow, and ill-defined; interstices with barely noticeable traces of microsculpture. Hind wings complete- ly reduced. Metatibia somewhat compressed, but other- wise unmodified. Abdomen broader than elytra, widest at segment VI; punctation dense and moderately fine; interstices with shallow, finely transverse microsculpture; posterior mar- gin of tergite VII without palisade fringe. &: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VIII weakly convex; sternite VII with shal- low median impression posteriorly, posterior margin dis- tinctly concave in the middle, this concavity with fringe of longer setae (Fig. 169); sternite VHI with impunctate median impression posteriorly, posterior excision moder- ately deep and broad (Fig. 170); aedeagus approximate- ly 1.5 mm long and slender, without distinct dorsal plate; ventral process long, slender, and apically truncate in ven- tral view; internal sac with long, dark, membranous tube (Figs 171-172). ©: protarsomeres I-IV dilated, but somewhat less so than in male; tergite VIII of similar shape as in male; sternite VII only slightly longer than tergite VIII and with strong- ly convex posterior margin (Fig. 173); tergite IX almost completely divided in the middle, tergite X almost reach- ing anterior margin of tergite [X (Fig. 174). Comparative notes. Lathrobium discissum is undoubted- ly most closely related to the similar, but distinctly small- er, syntopic L. separatum; for characters distinguishing these species see the comparative notes in the following section. Both species are separated from all other Hi- malayan congeners by the modifications of the male ster- nites VII and VIII, by the morphology of the aedeagus (slender, ventral process apically truncate in ventral view, long dark membranous tube in the internal sac, absence of a distinct dorsal plate), as well as by the almost com- pletely divided female tergite IX. Etymology. The specific epithet (Latin, past participle of discindere: to tear or cut apart) refers to the divided fe- male tergite IX. Distribution and natural history. Lathrobium discissum was found in two localities near Tonglu [27°02’N, 88°05’E] in Darjeeling district, West Bengal, North India, and in one locality [27°05’N, 87°55’E] in the very east of Nepal (Fig. 126). The specimens were collected at al- titudes of 2700 and 3100 m, in two localities together with L. separatum. Some of the type specimens from India are teneral. ©ZFMK Revision of Himalayan Lathrobium 187 peu ae Re eds ee —— « Q. 4 179 175 178 174 Figs 167-180. Lathrobium discissum (167-174) and L. separatum (175-180). 167: habitus; 168, 175: forebody; 169, 176: male sternite VII; 170, 177: male sternite VIII; 171, 178: aedeagus in lateral view; 172, 179: aedeagus in ventral view; 173, 180: fe- male sternite VIII; 174: female abdominal segments IX—X. Scale bars: 167-168, 175: 1.0 mm; 169-174, 176-180: 0.5 mm. Lathrobium separatum sp. n. (Figs 3, 175-180) Type material. Holotype @: “India W. Bengal, Darjee- ling distr., Tonglu 3100 m 16.X.1978, Besuchet-Lobl / Holotypus 4 Lathrobium separatum sp. n., det. V. Ass- ing 2011” (MHNG). Paratypes: 23:4, 29 9: same data as holotype (MHNG, cAss); 529: same data, but 2700 m (MHNG, cAss). Bonn zoological Bulletin 61 (2): 142-209 Description. Rather small species; body length 5.5—6.5 mm; length of forebody 2.7—3.0 mm. Coloration: head and pronotum reddish to brown; elytra reddish to reddish- brown; abdomen dark-brown; legs and antennae reddish. Head (Fig. 175) weakly oblong; punctation moderate- ly coarse and rather sparse; interstices with distinct fine microsculpture, almost matt, distinctly broader than diam- eter of punctures. Eyes small, but composed of more than ©ZFMK 188 Volker Assing 20 ommatidia, not projecting from lateral contours of head, and approximately 1/5 the length of postocular region in dorsal view. Pronotum (Fig. 175) relatively slender, approximately 1.3 times as long as broad, or nearly so, and approximate- ly as broad as head; lateral margins almost subparallel in dorsal view; punctation similar to that of head; interstices without microsculpture and glossy. Elytra short, 0.55—0.60 times as long as pronotum (Fig. 175); humeral angles weakly marked; punctation fine, shallow, and ill-defined; interstices with barely noticeable traces of microsculpture. Hind wings completely reduced. Metatibia somewhat compressed, but otherwise unmodi- fied. Abdomen broader than elytra, widest at segment VI; punctation dense and moderately fine; interstices with fine transverse microsculpture; posterior margin of tergite VII without palisade fringe. &: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VIII weakly convex; sternite VII with shal- low median impression posteriorly and with sparse, un- modified pubescence, posterior margin broadly and weakly concave (Fig. 176); sternite VIII with narrow im- punctate median impression posteriorly, pubescence un- modified, posterior excision very shallow (Fig. 177); aedeagus 0.9—1.0 mm long and slender, without distinct dorsal plate; ventral process long, slender, subapically curved (lateral view), and apically truncate (ventral view); internal sac with long, dark, membranous tube (Figs 178-179). ©: protarsomeres I-IV dilated, but somewhat less so than in male; tergite VII of similar shape as in male; ster- nite VIL only slightly longer than tergite VIII and with convex posterior margin (Fig. 180); tergite LX almost com- pletely divided in the middle, tergite X almost reaching anterior margin of tergite IX. Comparative notes. Based on the similar morphology of the male sexual characters, as well as on the almost com- pletely separated female tergite IX, clearly a synapomor- phic character, L. separatum is apparently the sister species of the syntopic L. discissum. It is distinguished from this species by distinctly smaller body size (no overlap), the shape and chaetotaxy of the male sternite VII and VIII, and the much smaller and more slender aedeagus. Etymology. The specific epithet (Latin, adjective) refers to the divided female tergite IX. Distribution and natural history. Lathrobium separatum was found in two localities near Tonglu in Darjeeling dis- trict, West Bengal, North India (Fig. 3). The specimens were collected at altitudes of 2700 and 3100 m, together with L. discissum. Bonn zoological Bulletin 61 (2): 142-209 The Lathrobium jumlense group Lathrobium jumlense Coiffait, 1982 (Figs 96, 181-186) Lathrobium jumlensis [sic] Coiffait, 1982a: 89 f. Type material examined. Holotype <: ‘““Dampa-PaB, ge- gen Chauta / Gebiet von Jumla, Westnepal, lg. H. Franz / Pa 229 [overleaf] / Type / Lathrobium jumlensis [sic] H. Coiffait 1979 / Lathrobium jumlense Coiffait, det. V. Assing 2011” (NHMW). Paratype 2: same data as holo- type (NHMW). Comment. The original description is based on a male holotype and three paratypes, a male and two females, from ““Népal, Dampa Pass, prés Chanta, région de Jum- la” (Coiffait 1982a). The holotype and a female paratype were located in the Franz collection at the NHMW. Redescription. Small species, body length 4.14.5 mm; length of forebody 1.8—1.9 mm. Coloration: body uniform- ly yellowish-red. Head (Fig. 181) weakly oblong, nearly 1.1 times as long as broad, dilated posteriad; punctation moderately coarse and very sparse; interstices much broader than diameter of punctures and with shallow microreticulation. Eyes not projecting from lateral contours of head, very small, com- posed of approximately ten ommatidia. Pronotum (Fig. 181) rather broad, approximately 1.2 times as long as broad and as broad as head; posterior mar- gin truncate to weakly concave; punctation similar to that of head, but somewhat less sparse; interstices without m1- crosculpture. Elytra short, approximately 0.6 times as long as prono- tum (Fig. 181); humeral angles weakly marked; puncta- tion shallow, fine, and sparse; interstices without distinct microsculpture. Hind wings completely reduced. Metati- bia slightly compressed. Abdomen slender and with subparallel margins, approx- imately as broad as elytra; punctation moderately fine and dense; posterior margin of tergite VII without palisade fringe; posterior margin of tergite VIII truncate to weak- ly convex in both sexes. 3: protarsomeres I-IV strongly dilated; sternite VII rel- atively weakly transverse, without modified setae, and with very weakly concave posterior margin (Fig. 182); sternite VIII distinctly oblong, without modified setae, and with rather broad, moderately deep, and almost sem1-cir- cular posterior excision (Fig. 183); aedeagus small, ap- proximately 0.65 mm long, ventral process broad and api- cally very acute in ventral view, thin and sinuate in later- al view; dorsal plate weakly sclerotised (Figs 184-185). ©: protarsomeres I-IV dilated, but slightly less so than in male; sternite VHI oblong, only slightly longer than ter- gite VIII, and convexly produced posteriorly (Fig. 186); ©ZFMK Revision of Himalayan Lathrobium 189 187 189 Figs 181-193. Lathrobium jumlense (181-186) and L. inustum (187-193). 181, 188: forebody; 182, 189: male sternite VII; 183, 190: male sternite VIII; 184, 191: aedeagus in lateral view; 185, 192: aedeagus in ventral view; 186, 193: female sternite VIII; 187: habitus. Scale bars: 187: 1.0 mm; 181, 188: 0.5 mm; 182-186, 189-193: 0.2 mm. tergite IX undivided; tergite X approximately as long as tergite [X in the middle. Comparative notes. The similar external characters and particularly the similar general morphology of the aedea- gus (ventral process broad and apically acute in ventral view, sclerotised internal structures absent) suggest that Bonn zoological Bulletin 61 (2): 142-209 L. jumlense 1s closely related to L. inustum. For charac- ters separating these two species see the following sec- tion. From the externally similar species of the L. pecti- natum and L. aculeatum groups, L. jumlense is readily dis- tinguished by the absence of pectinate setae on the male sternite VII and by the completely different morphology of the aedeagus, respectively. ©OZFMK 190 Volker Assing Distribution and natural history. The type locality is sit- uated in Jumla district, western Nepal (Fig. 96). Accord- ing to Franz’ diary, the type specimens were sifted from leaf litter in a birch and fir forest with bamboo at an alti- tude of 3500 m on 29.1X.1972. Lathrobium inustum Coiffait, 1982 (Figs 96, 187—193) Lathrobium inustum Coiffait, 1982a: 88 f. Type material examined. Holotype ¢: “Umg. Mahari- gaon, 3000-3500 m / Pa 200 [overleaf] / Gebiet von Jum- la, Westnepal, lg. H. Franz / Type / Lathrobium inustum H. Coiffait 1979 / Lathrobium inustum Corffait, det. V. Assing 2011” (NHMW). Paratypes: 24.4, 19 [9 without head and pronotum]: same data as holotype (NHMW); 19: “Dzunda Khola-Tal b. Talphi 3000-3500 m / Pa 194 [over- leaf] / Gebiet von Jumla, Westnepal, lg. H. Franz / Para- type” (NHMW). Comment. The original description is based on a male holotype and five paratypes, two males and three females, from ““Népal, environs de Maharigaon, région de Jumla”, one male paratype from “Népal, Environs de Alm Darghari, 4000 m’”, and four paratypes, one male and three females, from, “Népal, Dzunda Khola Tal, pres Talphi” (Coiffait 1982a). The holotype and four paratypes were located in the Franz collection at the NHMW. The spec- imens listed as additional material below were evidently collected together with the types, but not included in the type series. Additional material examined. Nepal: 6 exs., “Dzunda Kho- la-Tal b. Talphi 3000-3500 m / Pa 194 [overleaf] / Gebiet von Jumla, Westnepal, lg. H. Franz” (NHMW, cAss); 3 exs., “Umg. Alm Darghari b. Maharigaon, 4000 m / Pa 211 [overleaf] / Ge- biet von Jumla, Westnepal, lg. H. Franz” (NHMW, cAss). Redescription. Small species, body length 3.6—4.8 mm; length of forebody 1.7—2.1 mm. Habitus and forebody as in Figs 187-188. Coloration: body uniformly yellowish- red. External characters as in L. jumlense. 3: protarsomeres I-IV strongly dilated; sternite VII moderately transverse, without modified setae, and with broadly concave posterior margin (Fig. 189); sternite VII distinctly oblong, without modified setae, and with rather broad, moderately deep, and almost semi-circular posterior excision (Fig. 190); aedeagus minute, approxi- mately 0.30—0.35 mm long, subapical portion broad in ventral view; ventral process weakly sclerotised, almost membranous, and apically acute in ventral view; dorsal plate weakly sclerotised (Figs 191-192). 2: protarsomeres I-IV dilated, but slightly less so than in male; sternite VIII oblong, only slightly longer than ter- gite VIII, and with convex posterior margin (Fig. 193); ter- Bonn zoological Bulletin 61 (2): 142-209 gite [IX undivided; tergite X approximately as long as ter- gite IX in the middle. Comparative notes. As can be inferred from the similar external characters and particularly the similar sexual char- acters (male sternite VII without modified setae; male ster- nite VIII without modified setae and with almost semi-cir- cular posterior excision; aedeagus small, subapically broad in ventral view, with weakly sclerotised, short, and api- cally acute ventral process, and with weakly sclerotised dorsal plate; shape of female sternite VHI), L. inustum is most closely related to L. jumlense. It is readily distin- guished from this species by the more transverse male ster- nite VII, as well as by the much smaller and differently shaped aedeagus. In fact, the aedeagus is distinctly small- er than in any other Himalayan representative of the genus. Distribution and natural history. This species 1s current- ly known from three geographically close localities near Maharigaon [29°20’N, 82°23’E] and Talphi [29°18’N, 82°22’E] in Jumla district, western Nepal (Fig. 96). The specimens were collected at altitudes of 3000-4000 m. Ac- cording to Franz’ diary, the holotype was sifted near “Sinamoro [?]” from leaf litter and moss in a forest on 21.1X.1972, the specimens from “Alm Darghari” near “Sinamoro [?]” from leaf litter, and the specimens from the environs of Talphi were collected in a forest near “Alm Kharana [?]” on 20.[X.1972. Lathrobium planissimum sp. n. (Figs 96, 274-281) Type material. Holotype @: “Nepal P. Mahakali, D Darchula, 14 km NNE, Ghusa, 3450 m, plateau before Api / 29°56’06”N, 80°56°36”E, leg. M. Hartmann, 8./9.V1.2005, sieving in deciduous forest / Holotypus 3 Lathrobium planissimum sp. n., det. V. Assing 2012” (NME). Paratypes: 234, 599: same data as holotype (NME, cAss). Description. Small species, body length 4.24.6 mm; length of forebody 1.9—2.1 mm. Habitus as in Fig. 274. Coloration: body uniformly reddish. External characters as in L. jumlense and L. inustum. S: protarsomeres I-IV strongly dilated; sternite VII moderately transverse, without modified setae, and with broadly concave posterior margin (Fig. 275); sternite VIII oblong, without modified setae, and with broadly V- shaped posterior excision (Fig. 276); aedeagus small, ap- proximately 0.60 mm long, dorso-ventrally conspicuous- ly flattened; ventral process broad; dorsal plate weakly sclerotised (Figs 277-279). ©: protarsomeres I-IV dilated, but slightly less so than in male; sternite VIII oblong, only somewhat longer than tergite VIII, and with convex posterior margin (Figs ©ZFMK Revision of Himalayan Lathrobium 19] 280-281); tergite [IX undivided; tergite X approximately as long as tergite IX in the middle. Comparative notes. As can be inferred from the similar external and male sexual characters, L. planissimum be- longs to the L. jumlense group. It is distinguished from the other two species of this group only by the different shape of the posterior excision of the male sternite VII, as well as by the different morphology of the aedeagus. Distribution and natural history. The type locality is sit- uated in the southern slopes of the Byasrikh Himal, Ma- hakali province, in the very northwest of Nepal. The spec- imens were sifted from leaf litter in a deciduous forest at an altitude of 3450 m. The Lathrobium aculeatum group Lathrobium aculeatum Coiffait, 1982 (Figs 96, 194-198) Lathrobium aculeatum Coiffait, 1982a: 88. Type material examined. Holotype ¢: “nérdl. Dhaula- giri, Gompa/Tarakot [ca. 28°51°N, 83°00’E], 3300-3400 m, 2.-6.VI.1973 / Type / Holotypus / Lathrobium aculea- tum H. Coiffait 1979” (FSF). Paratype 2: same data as holotype, but “Paratypus” (FSF). Comment. The original description is based on a male holotype and five paratypes from “Népal, nordl. Dhaula- girl, Gompa/Tarakot, 3300-3400 m” (Coiffait 1982a). The holotype and a female paratype were examined. The for- mer is somewhat teneral, which explains the misleading illustration of the aedeagus in Coiffait (1982a). Redescription. Body length 5.0—5.3 mm; length of fore- body 2.3—2.4 mm. Coloration: body more or less uniform- ly reddish. Head distinctly oblong, approximately 1.1 times as long as broad; punctation moderately coarse and rather sparse in median dorsal portion, somewhat less sparse in poste- rior and lateral portions, but interstices on average broad- er than diameter of punctures; interstices with fine, shal- low microreticulation. Eyes very small, with fewer than ten ommatidia. Pronotum approximately 1.25 times as long as broad and slightly broader than head; punctation similar to that of head, but slightly finer; interstices without microsculpture. Elytra short, approximately 0.6 times as long as prono- tum; humeral angles weakly marked; punctation shallow and ill-defined; interstices without distinct microsculpture. Hind wings completely reduced. Metatibia unmodified. Abdomen broader than elytra; punctation relatively coarse and dense; posterior margin of tergite VII without palisade fringe. Bonn zoological Bulletin 61 (2): 142-209 3: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VIII truncate; sternite VII weakly impressed posteriorly, pubescence unmodified, posterior margin broadly and weakly concave (Fig. 194); sternite VIII ob- long, pubescence unmodified, posterior excision relative- ly narrow and moderately deep (Fig. 195); aedeagus 1.0 mm long and slender, with very long and thin ventral process (but ventral process shorter than basal portion of aedeagus), lamellate dorsal plate, and without sclerotised internal structures (Figs 196-197). ©: protarsomeres I-IV distinctly dilated, but slightly less so than in male; tergite VII with weakly convex poster1- or margin; sternite VIII oblong, gradually tapering poste- riad, posterior margin convex (Fig. 198); tergite [X not di- vided in the middle; tergite X slightly longer than tergite IX in the middle. Comparative notes. Lathrobium aculeatum is charac- terised particularly by the conspicuously slender shape of the aedeagus and the ventral process, as well as by the re- duced eyes, oblong head, and the shape and chaetotaxy of the male sternites VII and VIII. It is evidently most closely related to L. spiculatum, from which it is distin- guished particularly by the distinctly shorter ventral process of the aedeagus and by the narrower posterior ex- cision of the male sternite VIII. Distribution and natural history. L. aculeatum 1s prob- ably endemic to the Dhaulagiri range (Fig. 96). The type specimens were collected at an altitude of 3300-3400 m. Lathrobium spiculatum sp. n. (Figs 96, 199-213) Type material: Holotype 3: “Nepal (Prov. Bagmati), Yan- gri Ridge, 4200 m, 21.1V.81, Lébl & Smetana / Holoty- pus ¢ Lathrobium spiculatum sp. n., det. V. Assing 2011” (cSme). Paratypes: 4¢°3, 729: same data as holotype (MHNG, cSme, cAss); 599: “Nepal (Prov. Bagmati), Yangri Ridge, 4200 m, 23.IV.81, Lobl & Smetana” (MH- NG, cAss); 73.3, 229: “Nepal (Prov. Bagmati), Yangri Ridge, 4150 m, 24.IV.81, Lobl & Smetana” (cSme, cAss); 734, 82° [2 exs. teneral]: “Nepal (Prov. Bagmati), Yan- gri Ridge, 4350 m, 22.IV.81, L6bl & Smetana” (MHNG, cSme, cAss); 29.4, 229: “Nepal (Prov. Bagmati), Yan- gri Ridge, 4700-4800 m, 22.IV.81, Lobl & Smetana” (MHNG, cSme, cAss); 963, 829 [partly teneral]: “Nepal (Prov. Bagmati), Mere Dara, 3200 m, 8.IV.81, Lobl & Smetana” (MHNG, cSme, cAss); 19: “Nepal (Prov. Bagmati), Mere Dara, 3100-3300 m, 7.IV.81, Lobl & Sme- tana” (cSme); 14, 22 9: “Nepal (Prov. Bagmati), near Me- re Dara, 3000 m, 7.I1V.81, Lobl & Smetana” (MHNG, cSme); 744, 599 [partly slightly teneral]: “Nepal (Prov. Bagmati), below Thare Pati, 3300 m, 11.IV.81, Lobl & Smetana” (MHNG, cSme. cAss); 244: same data, but ©ZFMK 192 Volker Assing 203 204 205 206 207 38 208 209 211 212 Figs 194-213. Lathrobium aculeatum (194-198) and L. spiculatum (199-213). 194, 201: male sternite VII; 195, 202: male ster- nite VIII; 196, 203, 205, 207, 209, 211: aedeagus in lateral view; 197, 204, 206, 208, 210, 212: aedeagus in ventral view; 198, 213: female sternite VII; 199: habitus; 200: forebody. Scale bars: 199-200: 1.0 mm; 194-198, 201-213: 0.5 mm. Bonn zoological Bulletin 61 (2): 142-209 ©OZFMK Revision of Himalayan Lathrobium 193 “10.1V.81” (cSme, cAss); 14, 62 9: “Nepal (Prov. Bag- mati), below Thare Pati, 3500 m, 12.1V.81, Lobl & Sme- tana” (MHNG, cSme, cAss); 29 9: “Nepal (Prov. Bagma- ti), below Thare Pati, 3400 m, 13.IV.81, Lébl & Smeta- na” (cSme); 499: “Nepal (Prov. Bagmati), Malemchi, 2900 m, 14.1V.81, Lobl & Smetana” (MHNG, cSme, cAss); 14: “Nepal Nuwakot Di., betw. Ghopte and Thare Pati 3150 m, 24.IV.85 A. Smetana” (cAss); 24 [1 ten- eral]: same data, but “25.IV.85” (cAss); 19: same data, but “3200 m 23.IV.85” (cSme); 14: same data, but “3200 m, 26.1V.85” (cSme); 29 9: “Zw. Mulkharka u. Tare-Pati / Pa 158* [overleaf] / Zentral-Nepal, Sept—Okt. 1971, lg. H. Franz / Paratype [of L. franzi]” (NHMW). Description. Body length 4.3—5.3 mm; length of forebody 2.0—2.5 mm. Habitus as in Fig. 199. Coloration: body uni- formly reddish. Head (Fig. 200) distinctly oblong, approximately 1.1 times as long as broad; punctation moderately coarse and rather sparse in median dorsal portion, somewhat less Sparse in posterior and lateral portions, but interstices on average broader than diameter of punctures; interstices with fine, shallow microreticulation. Eyes very small, with fewer than ten ommatidia. Pronotum (Fig. 200) approximately 1.20—1.25 times as long as broad and slightly broader than head; posterior margin weakly concave; punctation similar to that of head; interstices without microsculpture. Elytra short, approximately 0.6 times as long as prono- tum (Fig. 200); humeral angles weakly marked; puncta- tion shallow and ill-defined; interstices without distinct mi- crosculpture. Hind wings completely reduced. Metatibia somewhat flattened in both sexes. Abdomen broader than elytra; punctation relatively coarse and dense; posterior margin of tergite VII without palisade fringe. ¢: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VII weakly convex; sternite VII weakly im- pressed posteriorly, pubescence unmodified, posterior margin broadly and weakly concave (Fig. 201); sternite VIII oblong, posterior excision almost V-shaped and mod- erately deep, margin of excision with very fine and short setae; tergites and sternite of segments [X—X extremely elongated (Fig. 202); aedeagus |.2—1.3 mm long and slen- der, ventral process of somewhat variable shape and vari- able length, long, thin, and more or less strongly bent ven- trad in lateral view, dorsal plate lamellate and weakly scle- rotised, internal sac without distinctly sclerotised struc- tures (Figs 203-212). ©: protarsomeres I-IV dilated, but less so than in male; tergite VIII with truncate to weakly convex posterior mar- gin; sternite VIII oblong, gradually tapering posteriad, pos- terior margin strongly convex (Fig. 213); tergite IX not divided in the middle; tergite X slightly longer than ter- gite [IX in the middle. Bonn zoological Bulletin 61 (2): 142-209 Intraspecific variation. The ventral process of the aedeagus is of remarkably variable length and shape (Figs 203-212). The relative length ranges from slightly short- er to somewhat longer than the basal portion of the aedea- gus. The ventral keel in the basal half of the ventral process may be pronounced to practically obsolete. Moreover, the base of the ventral process (ventral view) may range from very thin (much thinner than the basal portion of the aedea- gus) to relatively broad and apically gradually tapering. These character conditions are linked by transitional states, even in material from the same locality, so that the ob- served differences are attributed to intra- rather than in- terspecific variation. Comparative notes. This species 1s characterised partic- ularly by the extremely long ventral process of the aedea- gus (distinctly longer even than that of L. aculeatum), as well as by the reduced eyes, oblong head, the shape and chaetotaxy of the male sternites VII and VIII, the conspic- uously elongated sclerites of the male segments IX and X, and the long female tergite [X (longer in the middle than tergite X). Etymology. The specific epithet is an adjective derived from the Latin noun spiculum (spine, sting) and alludes to the conspicuously long and thin ventral process of the aedeagus. Distribution and natural history. Lathrobium spicula- tum is probably endemic to the Langtang region to the north and northeast of Kathmandu (Fig. 96). The — part- ly teneral — specimens were collected at altitudes of 2900- 4800 m. In some localities, this species was collected to- gether with L. nepalense and/or L. exsertum. According to Franz’ diary, his specimens were collected in a rhodo- dendron forest above “Bulumje” [?] on 7.X.1971. The Lathrobium pectinatum group Lathrobium pectinatum Coiffait, 1981 (Figs 214-218) Lathrobium pectinatum Coiffait, 1981: 329 ff. Type material examined. Holotype ¢: “Nepal VIII80, Dudh Pokari / 3000 Himal Chuli D.L / Holotype / Lath- robium pectinatum H. Coiffait 1981 / Lathrobium pecti- natum Coiffait, det. V. Assing 2011” (MNHNP). Paratype S: same data as holotype (but labelled “Paratype”) (MNHNP). Comment. The original description is based on a male holotype and three paratypes from “Dudh Pokhari, Himal Chuli 3000 m” (Coiffait 1981) deposited in the Coiffait collection. ©OZFMK 194 Volker Assing Fig. 214. Distributions of the species of the L. pectinatum group: L. privum (open star); L. fodens (open circles); L. calcaratum (filled triangle); L. barbulatum (open diamond); L. barbatum (filled circle); L. compressicrus (filled diamond); L. pectinatum (open triangles); L. cavicrus (westernmost open triangle); L. attritum (filled star); L. cassagnaui (open squares). Additional material examined. Nepal: 13, Manaslu, Dudh Pokhari Lekh, between Kharka and Malemchi Kharka [28°18°N, 84°35’E], 3300-3500 m, 12.-13.1X.1995, leg. Schmidt (cAss); 13, 1129, Manaslu, SW Merne Pokhari, affluent of Ngadi Kho- la, spring area, 28°22’N, 84°31°E, 3200-3300 m, 11.V.2005, leg. Schmidt (NME, cAss); 14, 229, Manaslu, E-slope of Ngadi Khola Valley, 28°22’N, 84°30’E, 2800-3000 m, 13.V.2005, leg. Schmidt (NME, cAss). Redescription. Body length 5.3—6.0 mm; length of fore- body 2.5—2.6 mm. Coloration: body uniformly reddish. Head approximately as long as wide; punctation coarse and rather sparse; interstices with distinct microreticula- tion, broader than diameter of punctures in median dor- sal portion. Eyes not projecting from lateral contours of head, very small, composed of fewer than ten ommatidia. Pronotum approximately 1.25 times as long as broad and as broad as head; punctation similar to that of head, but slightly less coarse; interstices without microsculpture. Elytra short, 0.55—0.60 times as long as pronotum; humeral angles weakly marked; punctation shallow and ill-defined; interstices without distinct microsculpture. Hind wings completely reduced. Metatibia distinctly com- pressed and somewhat bent, sexually dimorphic. Abdomen broader than elytra; punctation distinct and dense; posterior margin of tergite VII without palisade fringe. 3: protarsomeres I-IV strongly dilated; metatibia con- spicuously flattened; posterior margin of tergite VIII con- vex in the middle; sternite VII posteriorly impressed and Bonn zoological Bulletin 61 (2): 142-209 with two transverse rows each composed of numerous pal- isade setae, anterior to these rows with additional long black setae, posterior margin distinctly concave (Fig. 215); sternite VIII in the middle impressed and with very sparse pubescence, posterior margin with rather deep and asym- metric excision (Fig. 216); aedeagus 0.85 mm long, with symmetric ventral process of roughly triangular shape and with lamellate dorsal plate (Figs 217-218). ©: protarsomeres I-IV dilated, but somewhat less so than in male; metatibia noticeably flattened in apical half; tergite VIII with convex posterior margin; sternite VIII ob- long, slightly longer than tergite VIII, and convexly pro- duced posteriorly; tergite X slightly longer than tergite IX in the middle. Comparative notes. Lathrobium pectinatum is readily distinguished from other small-sized and subanophthal- mous Himalayan Lathrobium species particularly by the modifications of the male sternites VII and VIII, as well as by the morphology of the aedeagus. Females are sep- arated from those of the syntopic L. cavicrus by the less strongly flattened metatibiae and by the posteriorly less broadly produced sternite VIII. Distribution and natural history. The species is known from four localities in the Manaslu Himal, in central Nepal (Fig. 214). It has been collected at altitudes of 2800-3500 m, in two localities together with the similar and closely related L. cavicrus. ©OZFMK Revision of Himalayan Lathrobium 195 2014 292 220 215 219 Figs 215-223. Lathrobium pectinatum (215-218) and L. compressicrus (219-223). 215, 219: male sternite VII; 216, 220: male sternite VIII; 217, 221: aedeagus in lateral view; 218, 222: aedeagus in ventral view; 223: female sternite VIII. Scale bars: 0.5 mm. Lathrobium compressicrus sp. n. (Figs 214, 219-223, 2713) Type material. Holotype @: “C-Nepal, Manaslu massif, Barapokhari Lekh, 23 km NE Besisahar Vill., 28°21°N, 84°33’E, 14.1X.2000, leg. A. Hetzel / 3800-4100 m, sieved from moss and Rhododendron leaf litter / Holotypus ¢ Lathrobium compressicrus sp. n. det. V. Assing 2011” (cAss). Paratypes: 23:3, 429: same data as holotype (cFel, cAss). Description. Body length 5.0—6.2 mm; length of forebody 2.12.6 mm; males on average larger than females. Eyes composed of fewer than ten ommatidia. Bonn zoological Bulletin 61 (2): 142-209 Apical half of mesotibiae and all of metatibiae strong- ly flattened in both sexes. Other external characters high- ly similar to those of L. barbatum (see the following sec- tion). S: protarsomeres I-IV moderately dilated; posterior margin of tergite VIII convex; sternite VII strongly trans- verse and with concave posterior margin, in posterior me- dian portion with four conspicuous transverse rows of pectinate setae, the two anterior rows interrupted in the middle and shorter than the two posterior rows (Fig. 219); sternite VIII with relatively broad and deep posterior ex- cision (Fig. 220); aedeagus stout and approximately 0.9 mm long, ventral process subapically strongly narrowed and apically acute in ventral view (Figs 221—222). ©ZFMK 196 Volker Assing ®: protarsomeres I-IV slightly less dilated than in male; posterior margin of tergite VIII strongly convex in the mid- dle (Fig. 273); sternite VII strongly narrowed posteriad (Fig. 223); tergite [X not divided in the middle; tergite X distinctly longer than tergite IX in the middle. Comparative notes. Lathrobium compressicrus is evi- dently most closely related to the geographically close L. pectinatum, as is suggested by the similarly derived mor- phology of the metatibiae, the similar morphology of the aedeagus, and the similar modifications of the male ster- nites VI and VIII, but distinguished from this species by the flattened apical half of the mesotibia, the chaetotaxy of the male sternite VII, and the broader and slightly longer ventral process of the aedeagus. Etymology. The specific epithet is a noun in apposition composed of the Latin adjective compressus and the Lat- in noun crus (shin); it refers to the conspicuous shape of the meso- and metatibiae. Distribution and natural history. As can be inferred from the restricted distributions of other Lathrobium species in the Himalaya, as well as from the adaptive reductions of the pigmentation, the eyes, and the wings, the species is probably endemic to the Manaslu range, where it was found in the Barapokhari Lekh (Fig. 214). The type spec- imens were sifted from moss and rhododendron litter at an altitude of 3800-4100 m. Lathrobium barbatum sp. n. (Figs 214, 224-231) Type material. Holotype ¢: “Nepal, Annapurna Mts., above Temang, 3000-3500 m, 28°30’44N, 83°18°37E, 06.V.2007, leg. J. Schmidt / Holotypus 3 Lathrobium bar- batum sp. n. det. V. Assing 2011” (NME). Paratypes: 1<, 29: same data as holotype (NME, cAss). Description. Body length 4.8—5.5 mm; length of forebody 2.3—2.5 mm. Coloration: body uniformly reddish. Head (Fig. 224) weakly oblong; punctation coarse and rather sparse; interstices with distinct microreticulation, broader than diameter of punctures in median dorsal por- tion, on average at least as broad as diameter of punctures in lateral and posterior dorsal portions. Eyes not project- ing from lateral contours of head, very small, composed of approximately ten ommatidia. Pronotum (Fig. 224) approximately 1.25 times as long as broad and as broad as head; punctation similar to that of head, but slightly less coarse; interstices without mi- crosculpture. Elytra short, 0.55—0.60 times as long as pronotum (Fig. 224); humeral angles weakly marked; punctation shallow and mostly ill-defined; interstices without distinct mi- Bonn zoological Bulletin 61 (2): 142-209 crosculpture. Hind wings completely reduced. Mesotibia abruptly dilated in basal half; metatibia modified and with conspicuous sexual dimorphism. Abdomen broader than elytra; punctation distinct and dense; posterior margin of tergite VII without palisade fringe. G: protarsomeres I-IV strongly dilated; mesotibia di- lated in basal third; metatibia in basal third with pro- nounced toothlike projection and in apical two thirds with pronounced concavity (Fig. 225); posterior margin of ter- gite VII convex; sternite VII posteriorly impressed and with three transverse rows each composed of numerous palisade setae (Fig. 226); sternite VIII in the middle im- pressed and with very sparse pubescence, posterior mar- gin with rather deep and asymmetric excision (Fig. 227); aedeagus approximately 0.9 mm long, with asymmetric ventral process and with lamellate dorsal plate (Figs 228-229). ©: protarsomeres I-IV dilated, but less so than in male; modifications of metatibia much less pronounced than in male (Fig. 230); posterior margin of tergite VII weakly convex; sternite VII barely longer than broad, posterior margin obtusely and almost triangularly produced (Fig. 231); tergite [IX undivided; tergite X slightly longer than tergite IX in the middle. Comparative notes. Lathrobium barbatum is readily dis- tinguished from other species of the L. pectinatum group by the modifications of the sexually dimorphic metatibi- ae, the conspicuous chaetotaxy of the male sternite VII, the asymmetric posterior excision of the male sternite VIII, as well as by the asymmetric aedeagus. Etymology. The specific epithet (adjective: bearded) al- ludes to the conspicuous chaetotaxy of the male sternite VU. Distribution and natural history. The type locality 1s sit- uated near Temang [28°32’N, 84°19’E] in the northern slopes of the Annapurna range (Fig. 214) at an altitude of 3000-3500 m. Lathrobium barbulatum sp. n. (Figs 214, 232-235) Type material. Holotype 3: “Nepal Manang Distr., 4 km SE Pisang 3050 m 26.1X.83 Smetana & Lobl/ Holotypus 3S Lathrobium barbulatum sp. n. det. V. Assing 2011” (MHNG). Description. Body length 5.2 mm; length of forebody 2.4 mm. External characters as in L. barbatum, distinguished only by the male sexual characters: 3: protarsomeres I-IV strongly dilated; modifications of meso- and metatibia as in L. barbatum; posterior mar- ©ZFMK Revision of Himalayan Lathrobium 197 234 235 Py yth 232 231 233 Figs 224-235. Lathrobium barbatum (224-231) and L. barbatulum (232-235). 224: forebody; 225: male metatibia; 226, 232: male sternite VII; 227, 233: male sternite VIII; 228, 234: aedeagus in lateral view; 229, 235: aedeagus in ventral view; 230: fe- male metatibia; 231: female sternite VII. Scale bars: 224: 1.0 mm; 225—235: 0.2 mm. gin of tergite VIII convex; sternite VII posteriorly im- pressed and with three transverse rows each composed of numerous palisade setae (Fig. 232); sternite VIII in the middle impressed and with very sparse pubescence, pos- terior margin with rather deep and weakly asymmetric ex- cision (Fig. 233); aedeagus approximately 0.95 mm long, with asymmetric ventral process and with lamellate dor- sal plate (Figs 234-235). ©: unknown. Bonn zoological Bulletin 61 (2): 142-209 Comparative notes. Lathrobium barbulatum is distin- guished from the highly similar and undoubtedly closely related L. barbatum by the shape of the apical row of pal- isade setae of the male sternite VII, the less strongly asym- metric posterior excision of the male sternite VHI, and by the morphology of the aedeagus (shape of the apex of the ventral process, transversely rectangular dorsal plate). From other species of the L. pectinatum group, it is read- ily distinguished also by the modifications of the male meso- and metatibiae. ©ZFMK 198 Volker Assing Etymology. The specific epithet (Latin, adjective derived from barbula: small beard) alludes to the conspicuous chaetotaxy of the male sternite VII and to the close rela- tionship with L. barbatum. Distribution and natural history. The type locality is sit- uated near Pisang [28°38’N, 84°06’E] in the northern slopes of the Annapurna range (Fig. 214), not far from the type locality of L. barbatum, at an altitude of 3050 m. Lathrobium cavicrus sp. n. (Figs 214, 282-289) Type material. Holotype @: “Nepal Manaslu Mts., SW Merne Pokhari, 3200-3300 m, leg. Schmidt, 11.V.2005 / 28°21°41N 84°30°42E, Quellgebiet, mittl. NebenfluB des Ngadi Khola / Holotypus ¢ Lathrobium cavicrus sp. n. det. V. Assing 2012” (NME). Paratypes: 73.0, 1199: same data as holotype (NME, cAss); 79 9: “Nepal Man- aslu Mts., 28°21°36N 84°30°04E, E slope of Ngali Kho- la Vall., 2800-3000 m, leg. Schmidt, 13.V.2005” (NME). Description. Body length 5.0-6.0 mm; length of forebody 2.3—2.6 mm. Forebody as in Fig. 282. Externally highly similar to the syntopic L. pectinatum; reliably distinguished only by the primary and secondary sexual characters. 3: protarsomeres I-IV strongly dilated; metafemur en- larged, but without posterior tooth; metatibia with ventral tooth-like projection in basal third and and in apical two thirds with pronounced concavity (Fig. 283); posterior margin of tergite VIII weakly convex; sternite VII strongly transverse and with weakly concave posterior margin, in posterior median portion with four conspicu- ous, transverse, almost straight, and uninterrupted rows of pectinate setae (Fig. 284); sternite VII with relatively broad, moderately deep, and somewhat asymmetric pos- terior excision (Fig. 285); aedeagus stout and 0.90—0.95 mm long, ventral process subapically strongly narrowed, apically acute, and almost symmetric in ventral view (Figs 286-287). ©: protarsomeres I-IV dilated, but somewhat less so than in male; posterior margin of tergite VIII convexly pointed in the middle (Fig. 288); sternite VHI oblong and convexly produced posteriorly (Fig. 289); tergite [X not divided in the middle; tergite X distinctly longer than ter- gite IX in the middle. Comparative notes. As can be inferred from the similar- ly derived modifications of the male metatibiae and of the male sternites VII and VIII, as well as from the similar general morphology of the aedeagus, L. cavicrus is most closely related to L. barbatus and L. barbatulus from the Annapurna range. It is distinguished from them by the less distinctly concave posterior margin and the almost Bonn zoological Bulletin 61 (2): 142-209 straight transverse rows of pectinate setae of the male ster- nite VII, as well as by the differently shaped, almost sym- metric ventral process of the aedeagus. From the syntopic and externally similar L. pectinatum, L. cavicrus is dis- tinguished by the male primary and secondary sexual char- acters, by the more strongly flattened female metatibiae, as well as by the posteriorly more broadly produced fe- male sternite VIII. Etymology. The specific epithet is a noun in apposition composed of the Latin adjective cavus (excavate) and the Latin noun crus (shin); it refers to the conspicuous shape of the male metatibiae. Distribution and natural history. This species is known only from two adjacent localities in the Manaslu range in central Nepal (Fig. 214), where it was collected at altitudes of 2800-3300 m, together with numerous specimens of L. pectinatum. Lathrobium franzi Coiffait, 1975 (Figs 214, 236-241) Lathrobium franzi Coiffait, 1975: 185. Type material examined. Holotype <: “Zentral-Nepal, Sept._Okt. 1971, lg. H. Franz / Wald unterhalb Fulung / Pa 174 [overleaf] / Holotype / Lathrobium franzi H. Coif- fait 1974 / Lathrobium franzi Coiffait, det. V. Assing 2011” (NHMW). Paratypes: see type material of L. spiculatum. Comment. The original description is based on a male holotype from ‘“Fulung, Népal central”, one female paratype from “‘chemin entre Gosatkunde et le monastere de Fulung”, and three female paratypes from “entre Mulkarka [sic] et Tare Pati” (Coiffait 1975). Two exam- ined female paratypes from between Mulkharka and Thare Pati are not conspecific with the holotype; they refer to L. spiculatum (see below). Redescription. Moderately small species, body length 5.7 mm; length of forebody 2.5 mm. Coloration: body uni- formly yellowish-red. Head (Fig. 236) distinctly oblong, approximately 1.1 times as long as broad; punctation coarse and moderate- ly dense; interstices with distinct microreticulation, broader than diameter of punctures in median dorsal por- tion, on average at least as broad as diameter of punctures in lateral and posterior dorsal portions. Eyes not project- ing from lateral contours of head, very small, composed of approximately 10 ommatidia. Pronotum (Fig. 236) 1.25 times as long as broad and as broad as head; punctation similar to that of head, but slightly less coarse; interstices without microsculpture. Elytra short, 0.55 times as long as pronotum (Fig. 236); humeral angles weakly marked; punctation shallow and ©ZFMK Revision of Himalayan Lathrobium 199 242 243 247 248 245 Figs 236-248. Lathrobium franzi (236-241) and L. calcaratum (242-248). 236, 243: forebody; 237, 244: male metafemur; 238, 245: male sternite VII; 239, 246: male sternite VIII; 240, 247: aedeagus in lateral view; 241, 248: aedeagus in ventral view; 242: habitus. Scale bars: 236, 242-243: 1.0 mm; 238-239, 245-246: 0.5 mm; 237, 240-241, 244, 247-248: 0.2 mm. ill-defined; interstices without distinct microsculpture. Hind wings completely reduced. Abdomen broader than elytra; punctation moderately fine and dense; posterior margin of tergite VII without pal- isade fringe. ¢: protarsomeres I-IV strongly dilated; metafemur with distinct ventral tooth in the middle (Fig. 237); metatibia slightly sinuate and strongly dilated apically; sternite VII strongly transverse, posteriorly extensively impressed in the middle and with three transverse rows each composed Bonn zoological Bulletin 61 (2): 142-209 of numerous palisade setae, posterior margin weakly con- cave (Fig. 238); sternite VIII moderately transverse, pos- teriorly with long median impression, with sparse, unmod- ified pubescence, and with weakly asymmetric, moderate- ly deep posterior excision (Fig. 239); aedeagus 1.0 mm long and of highly distinctive morphology; basal portion large, ventral process long, slender, and apically spear- shaped (Figs 240-241). Q: not examined. ©ZFMK 200 Volker Assing Comparative notes. This species is distinguished from other microphthalmous and depigmented representatives of the genus particularly by the modifications of the metafemur and metatibia, by the conspicuous chaetotaxy of the male sternite VII, as well as by the conspicuous shape of the aedeagus. Distribution and natural history. The type locality is sit- uated near the Fulung Monastery, to the northwest of the Gosainkund lakes, to the north-northwest of Kathmandu, in central Nepal (Fig. 214). Lathrobium ignoratum was collected in the same locality. According to Franz’ diary, the holotype was collected by sifting forest and rhododen- dron litter on 11.X.1971. Lathrobium calcaratum sp. n. (Figs 214, 242—248) Type material. Holotype 3: “Kali-Gandaki-Tal, zw. Gha- sa u. Lete / Pa 138 [overleaf] / Holotypus 4 Lathrobium calcaratum sp. n., det. V. Assing 2011” (NHMW). Description. Body length 5.7 mm; length of forebody 2.85 mm. Habitus as in Fig. 242. Coloration: body uniformly reddish. Head (Fig. 243) weakly oblong, 1.04 times as long as broad; punctation coarse and rather sparse, even sparser in median dorsal portion; interstices with distinct, but shal- low microreticulation, somewhat glossy, much broader than diameter of punctures in median dorsal portion, on average at least as broad as diameter of punctures in lat- eral and posterior dorsal portions. Eyes not projecting from lateral contours of head, very small, composed of approx- imately ten ommatidia. Pronotum (Fig. 243) 1.25 times as long as broad and as broad as head; punctation similar to that of head; inter- stices without microsculpture. Elytra short, 0.57 times as long as pronotum (Fig. 243); humeral angles weakly marked; punctation shallow, sparse, and fine; interstices without distinct microsculp- ture, much broader than diameter of punctures. Hind wings completely reduced. Metafemur with conspicuous sexu- al dimorphism. Abdomen broader than elytra; punctation distinct, moderately fine, and dense; posterior margin of tergite VII without palisade fringe. &: protarsomeres I-IV strongly dilated; metafemur with pronounced, apically rounded ventral tooth in the middle (Fig. 244); metatibia in apical two thirds somewhat dilat- ed; sternite VII distinctly transverse, with pronounced me- dian impression of triangular shape, this impression lat- erally delimited by a fringe of conspicuously dense pecti- nate setae, posterior margin in the middle with rather broad and not very deep excavation of trapezoid shape, anteri- or margin of this excavation with transverse row of nu- Bonn zoological Bulletin 61 (2): 142-209 merous pectinate setae (Fig. 245); sternite VII with dis- tinct median impression without pubescence, posterior ex- cision relatively deep and V-shaped (Fig. 246); aedeagus 1.1 mm long, ventral process strongly asymmetric and of highly distinctive shape (Figs 247-248). 2: unknown. Comparative notes. As can be inferred from the shape and chaetotaxy of the male sternites VII and VIII, as well as from the similar general morphology of the aedeagus, L. calcaratum belongs to the L. pectinatum group. The similarly derived modifications (presence of a pro- nounced ventral tooth in the middle) of the male metafe- mur, a unique character among Himalayan representatives of the genus, suggests that L. calcaratum is most closely related to L. franzi. It is additionally separated from all its congeners particularly by the conspicuous shape and chaetotaxy of the male sternite VIL as well as by the shape of the strongly asymmetric ventral process of the aedea- gus. Etymology. The specific epithet (Latin, adjective: with spurs) alludes to the tooth-like ventral projections of the male metafemora. Distribution and natural history. The type locality is sit- uated in the Kali-Gandaki valley, the valley separating the Dhaulagiri and Annapurna ranges, between Lete and Ghasa [approximately 28°37’N, 83°38’E] in central Nepal (Fig. 214). According to Franz’ diary, the holotype was collected in a stand of alder, in the vicinity of a rho- dodendron forest, on 26.1X.1971. Lathrobium privum sp. n. (Figs 214, 249-257) Type material. Holotype @: “581 Nepal: Jumla Distr., Khali-Lagna pass, 3500 m, 16.—17.VI.1998, leg. W. Scha- waller / Holotypus 4 Lathrobium privum sp. n., det. V. Assing 2011” (SMNS). Paratypes: 19: same data as holo- type (cAss). Description. Body length 5.8—6.2 mm; length of forebody 2.7—3.1 mm. Habitus and forebody as in Figs 249-250. Coloration: body uniformly reddish. Posterior margin of abdominal tergite VIII distinctly convexly produced in the middle. Other external characters as in L. calcaratum. 3S: protarsomeres I-IV strongly dilated; metafemur strongly dilated in the midde and with pronounced, api- cally broadly truncate ventral tooth in the middle (Fig. 251); metatibia somewhat compressed, gradually dilated apically; sternite VII distinctly transverse, with shallow median impression, on either side of this impression with a cluster of dense dark setae posteriorly, posterior margin weakly concave and with transverse row of approximate- OZFMK Revision of Himalayan Lathrobium 201 Figs 249-262. Lathrobium privum (249-257) and L. cassagnaui (258-262). 249: habitus; 250: forebody; 251: male metafemur; 252, 258: male sternite VII; 253, 259: male sternite VIII; 254, 260: aedeagus in lateral view; 255, 261: aedeagus in ventral view; 256, 262: female sternite VIII; 257: female abdominal segments IX and X in dorsal view. Scale bars: 249-250: 1.0 mm; 252-262: 0.5 mm; 251: 0.2 mm. ly 30 pectinate setae (Fig. 252); sternite VIII with distinct (Fig. 253); aedeagus 1.15 mm long, ventral process long median impression without pubescence posteriorly, pos- and thin, somewhat shaped like a golf club (Figs terior excision relatively deep and somewhat asymmetric 254-255). Bonn zoological Bulletin 61 (2): 142-209 ©ZFMK 202 Volker Assing ©: protarsomeres distinctly dilated, but less so than in male; sternite VII much longer than broad and longer than tergite VIII, distinctly tapering posteriad (Fig. 256); ter- gite [IX undivided anteriorly; tergite X reduced, apparent- ly absent (Fig. 257). Comparative notes. As can be inferred from the shape and chaetotaxy of the male sternites VII and VIII, as well as from the similar general morphology of the aedeagus, L. privum belongs to the L. pectinatum group. The simi- larly derived modifications (presence of a pronounced ven- tral tooth in the middle) of the male metafemur suggests that L. privum is most closely related to L. franzi and L. calcaratum. It is distinguished from these species partic- ularly by the strongly dilated male metafemur, the shape of the tooth on the male metafemur, by the modifications of the male sternites VII and VIII, as well as by the shape of the ventral process of the aedeagus, which somewhat resembles a golf club. Etymology. The specific epithet (Latin, adjective: free of) alludes to the reduced female tergite X. Distribution and natural history. The type locality is sit- uated in Jumla district, western Nepal (Fig. 214), at an al- titude of 3500 m. Lathrobium cassagnaui Coiffait, 1982 (Figs 214, 258-262) Lathrobium cassagnaui Coiffait, 1982b: 287 ff. Type material examined. Holotype ¢: “Nepal X.81, Kalingchok, 3000 m PC / Holotype / Lathrobium cassag- naui H. Coiffait 1982 / Lathrobium cassagnaui Coiffait, det. V. Assing 2011” (MNHNP). Comment. The original description is based on a unique male holotype from “Népal, Massif du Kalingchok, prés de Barbabise [sic]” (Coiffait 1982b). Additional material examined. Nepal: 19, Dolakha district, SW Kalinchok, 3100 m, 19.—23.IV.1995, leg. Martens & Scha- waller (SMNS). Redescription. Moderately small species, body length 4.8-5.3 mm; length of forebody 2.2—2.3 mm. Coloration: body uniformly yellowish-red. Head oblong, almost 1.1 times as long as broad; punc- tation coarse and rather sparse, even sparser in median dor- sal portion; interstices with distinct microreticulation, broader than diameter of punctures. Eyes not projecting from lateral contours of head, very small, composed of ap- proximately eight ommatidia. Bonn zoological Bulletin 61 (2): 142-209 Pronotum 1.28 times as long as broad and as broad as head; punctation similar to that of head, but slightly denser; interstices without microsculpture, except for some shallow traces in posterior half. Elytra short, 0.58 times as long as pronotum; humeral angles weakly marked; punctation shallow and ill-defined; interstices without distinct microsculpture. Hind wings completely reduced. Abdomen broader than elytra; punctation moderately fine and dense; posterior margin of tergite VII without pal- isade fringe. 3: protarsomeres I-IV strongly dilated; hind legs with- out modifications, except for the slightly compressed metatibia; sternite VII strongly transverse, with shallow median impression posteriorly, posterior margin broadly and weakly concave, in the middle with one transverse row of numerous palisade setae (Fig. 258); sternite VIII weak- ly transverse, posteriorly with shallow median impression without pubescence, posterior excision rather deep and somewhat asymmetric (Fig. 259); aedeagus rather large in relation to body size, 1.2 mm long, ventral process straight, long, and spine-shaped (Figs 260-261). Q: protarsomeres I-IV moderately dilated; posterior margin of tergite VII very obtusely angled in the middle; sternite VII weakly oblong and with broadly convex pos- terior margin (Fig. 262); tergite [X not divided in the mid- dle; tergite X slightly longer than tergite [X in the mid- dle. Comparative notes. This species is distinguished from other representatives of the L. pectinatum group particu- larly by the modifications of the male sternite VII and by the distinctive shape of the aedeagus. Distribution and natural history. The species is current- ly known only from the type locality, the Kalinchok, a mountain some 13 km to the east of Barahbise and almost 70 km to east-northeast of Kathmandu in central Nepal (Fig. 214). Apart from the altitude (3000 m), bionomic da- ta are not available. Lathrobium fodens sp. n. (Figs 214, 263-268) Type material. Holotype 3: “Nepal, Prov. Karnali, D: Jumla, Churta, E Jagdula valley, 3800-4100 m NN, 03.VI1.2007 / 29°09’49N, 82°31°09E, alpine mats, leg. M. Hartmann / Holotypus ¢ Lathrobium fodens sp. n., det. V. Assing 2011” (NME). Paratypes: 14: same data as holotype (cAss); 14, 49 9: “Nepal, P: Karnali, D: Jum- la, Churta, E below Mori La, 3550-3800 m NN, 01.VI.2007 / 29°09°58”N, 82°29712”E, leg. M. Hart- mann, mixed pine forest” (NME, cAss). ©ZFMK Revision of Himalayan Lathrobium 203 265 268 270 273 Figs 263-273. Lathrobium fodens (263-268), L. attritum (269-272), and L. compressicrus (273). 263: forebody; 264, 269: male sternite VII; 265, 270: male sternite VIII; 266, 271: aedeagus in lateral view; 267, 272: aedeagus in ventral view; 268: female ster- nite VIII; 273: female tergite VHI. Scale bars: 263: 1.0 mm; 264-273: 0.5 mm. Description. Body length 5-6 mm; length of forebody 2.42.7 mm. Coloration: body more or less uniformly red- dish. Head (Fig. 263) approximately as long as broad or weakly oblong; punctation coarse and rather sparse; in- terstices with distinct microreticulation, much broader than diameter of punctures in median dorsal portion. Eyes not projecting from lateral contours of head, very small, com- posed of fewer than 10 ommatidia. Pronotum (Fig. 263) approximately 1.25 times as long as broad and as broad as head; punctation similar to that of head; midline broadly impunctate; interstices without microsculpture. Elytra short, approximately 0.6 times as long as prono- tum (Fig. 263); humeral angles weakly marked; puncta- tion shallow and rather ill-defined; interstices without dis- Bonn zoological Bulletin 61 (2): 142-209 tinct microsculpture. Hind wings completely reduced. Metatibia slightly compressed. Abdomen broader than elytra; punctation shallow and moderately dense; posterior margin of tergite VI without palisade fringe. &: protarsomeres I-IV strongly dilated; posterior mar- gin of tergite VII truncate; sternite VII weakly impressed, posteriorly with a pair of clusters of blackish setae, pos- terior margin bisinuate in the middle and with fringe of numerous black palisade setae (Fig. 264); sternite VIII weakly transverse, median impression without setae, pos- terior excision somewhat U-shaped and in somewhat asymmetric position (Fig. 265); aedeagus 1.2—1.3 mm long; ventral process conspicuously long and thin; dorsal plate broad and lamellate (Figs 266-267). ©ZFMK 204 Volker Assing ©: protarsomeres I-IV dilated, but somewhat less so than in male; posterior margin of tergite VIII weakly con- vex in the middle; sternite VII oblong and convexly pro- duced posteriorly (Fig. 268); tergite [X not divided in the middle; tergite X slightly longer than tergite IX in the mid- dle. Comparative notes. Among the species of the L. pecti- natum group, L. fodens 1s particularly characterised by the conspicuously long and thin ventral process of the aedea- gus, as well as by the shape and chaetotaxy of the male sternites VII and VUI. Etymology. The specific epithet is the present participle of the Latin verb fodere (to sting) and refers to the con- spicuous shape of the ventral process of the aedeagus. Distribution and natural history. The species was col- lected in two localities in Karnali province, Jumla district, West Nepal (Fig. 214), at altitudes of 3550-4100 m. Lathrobium attritum sp. n. (Figs 214, 269-272) Type material. Holotype 3: “Nepal (Prov. Bagmati), Yar- dang ridge NE Barahbise, 3250 m, 5.V.81, Lobl & Sme- tana / Holotypus ¢ Lathrobium attritum sp. n., det. V. Ass- ing 2012” (cAss). Description. Body length 4.6 mm; length of forebody 2.2 mm. Coloration: body pale reddish. External characters as in L. cassagnaui. ¢: protarsomeres I-IV moderately dilated; hind legs without modifications; sternite VII rather strongly trans- verse, with shallow median impression posteriorly, pos- terior margin broadly and weakly conave, in the middle with one transverse row of approximately 8 stout, but not very dense palisade setae (Fig. 269); sternite VII weak- ly transverse, posteriorly with shallow median impression without pubescence, posterior excision weakly asymmet- ric and rather deep (Fig. 270); aedeagus 1.15 mm long, slender, with long, acute, and in lateral view almost straight ventral process (Figs 271—272). ©: unknown. Comparative notes. Based on the modifications of the male sternite VIII and on the similar general morpholo- gy of the aedeagus, L. attritum is undoubtedly most close- ly related to the geographically close L. cassagnaui, from which it is distinguished particularly by the less numer- ous pectinate setae at the posterior margin of the male ster- nite VII, as well as as by the shape of the aedeagus. Etymology. The specific epithet (Latin, adjective: rubbed off, worn out) refers to the relatively sparse pectinate se- tae at the posterior margin of the male sternite VII. Bonn zoological Bulletin 61 (2): 142-209 Distribution and natural history. The type locality is sit- uated in the Yardang ridge to the west of Barahbise, Bag- mati province, central Nepal (Fig. 214). The holotype was collected at an altitude of 3250 m. UNIDENTIFIED AND UNDESCRIBED SPECIES Lathrobium sp. 1 Material examined. Nepal: 12, Manaslu, Barapokhari Lekh, 23 km NE Besisahar, 28°21’N; 84°33’E, 3800-4100 m, sifted from moss and rhododendron lit- ter, 14.1X.2000, leg. Hetzel (cAss). Comment. The above female was collected in the same locality as the types of L. compressicrus. It is smaller than L. compressicrus and does not have flattened femora and tibiae. Lathrobium sp. 2 Material examined. Nepal: 29 9: “Kali-Gandaki-Tal, zw. Ghasa u. Lete / Pa 138 [overleaf]” (NHMW, cAss). Comment. This species is characterised by conspicuous- ly small body size (length of forebody: 1.7—1.8 mm; much smaller than the syntopic L. calcaratum), a uniformly pale- reddish body, and eyes composed of fewer than ten om- matidia. Lathrobium sp. 3 Material examined. Nepal: 19: “576 Nepal: Mugu Distr., SW Rara Lake, 3200 m, 12.VI.1998, leg. W. Scha- waller “ (SMNS). Comment. This species is characterised by small body size (length of forebody: 2.9 mm), a uniformly pale-red- dish body, and eyes composed of fewer than ten omma- tidia. Lathrobium sp. 4 Material examined. Nepal: 19: “628 Nepal: Dolakha Distr., E Ting Sang La, 3100 m, 12.—13.VI.2000, leg. W. Schawaller “ (SMNS). Comment. This species is characterised by small body size (length of forebody: 2.2 mm), a uniformly pale-red- dish body, and eyes composed of fewer than ten omma- tidia. ©ZFMK Revision of Himalayan Lathrobium 205 Figs 274-289. Lathrobium planissimum (274-281) and L. cavicrus (282-289). 274: habitus; 275, 284: male sternite VII; 276, 285: male sternite VIII; 277, 286: aedeagus in lateral view; 278-279, 287: aedeagus in ventral view; 280, 288: female tergite VI- II; 281, 289: female sternite VIII; 283: male metatibia. Scale bars: 274, 282: 1.0 mm; 283: 0.5 mm; 275-281, 284-289: 0.2 mm. Bonn zoological Bulletin 61 (2): 142-209 ©ZFMK 206 Volker Assing Lathrobium sp. 5 Material examined. Nepal: 19: “215 Dhading Dist., Ankhu Khola Tal, Ankhu Sangi, 650 m, Kulturland, Wald- reste, 24.-25. Jul 83 Martens & Schawaller “ (SMNS). Comment. This species is characterised by small body size (length of forebody: 2.5 mm) and small eyes com- posed of fewer than ten ommatidia. The teneral specimen has evidently been subject to post-mortem darkening. Lathrobium sp. 6 Material examined. Nepal: 1 [teneral; dissected prior to present study; aedeagus damaged]: “282 Taplejung Dist. S. Gunsa, 3900-3600 m, Abies/Rhododend., 10 Sep 83 Martens & Daams |. “ (SMNS). Comment. This species is characterised by small body size (length of forebody: 2.9 mm) and moderately small eyes. The teneral specimen has evidently been subject to post-mortem darkening. Lathrobium sp. 7 Material examined. Nepal: 29 9: “Nepal-Expeditionen Jochen Martens / 361 Taplejung Distr., upper Simbua Kho- la Valley, near Tseram, 3250-3350 m, mature Abies-Rho- 290 293 Figs 290-294. dodendron forest, 10-15 May 88 J. Martens & W. Schawaller leg.” (SMNS, cAss). Comment. This species is belongs to the L. discissum group. It is distinguished from L. discissum by even larg- ensize: Lathrobium sp. 8 Material examined. Nepal: 299: “Nepal Khandbari Dis., Goru Dzure Dara, W slope 3600 m, 9.IV.84 Smetana & Lébl” (SMNS, cAss). 399: “Nepal (Prov. Bagmati), below Thare Pati, 3500 m, 12.IV.81, Lobl & Smetana” (MHNG, cAss). Comment. This species belongs to the L. nepalense group. It is distinguished from most species of this group by its small size (length of forebody: 2.5—2.7 mm). Lathrobium sp. 9 Material examined. Nepal: 49 9: “Nepal (Prov. Bag- mati), below Thare Pati, 3500 m, 12.IV.81, Lobl & Smetana” (MHNG, cSme, cAss). Comment. This species is highly similar to, and was found together with L. spiculatum. It is distinguished from this species by on average slightly larger body size (length of forebody: 2.32.6 mm), a more oblong head and prono- 291 294 292 Medon jaljalensis. 290: habitus; 291: male sternite VII; 292: male sternite VIII; 293: aedeagus in lateral view; 294: aedeagus in ventral view. Scale bars: 290: 1.0 mm; 291-294: 0.2 mm. Bonn zoological Bulletin 61 (2): 142-209 ©ZFMK 207 Revision of Himalayan Lathrobium type locality of L. excisum NE Dal- above Simigaon, 2700-2800 m (26 specimens of L. kleebergi); 297 Figs 295-297. Lathrobium habitats in the Rolwaling Himal; photos: Andreas Kleeberg. 295 dung La pass, 3800 m; 296 ©ZFMK W Daldung La pass, 3300 m (28 Bonn zoological Bulletin 61 (2): 142-209 specimens of L. kleebergi). 208 Volker Assing tum, larger eyes (composed of approximately 20 omma- tidia), coarser punctation of the forebody, and by a broad- er, less oblong, and posteriorly less strongly convex fe- male sternite VIII. SPECIES EXCLUDED FROM LATHROBIUM In the course of the revision it was discovered that two species from Nepal that had originally been assigned to Lathrobium, L. jaljalense Coiffait, 1984 and L. perpusil- lum Coiffait, 1982, in fact belong to genera of the sub- tribe Medonina. The latter species is dealt with by Assing (2012d). Medon jaljalensis (Coiffait, 1984), comb. n. (Figs 290-294) Lathrobium jaljalensis [sic] Coiffait, 1984: 382. Type material examined. Holotype ¢: “Nepal XI.78 / Jaljale Himal, 2950 m PC31 / Holotype / Lathrobium jal- jalensis H. Coiffait / Medon jaljalensis (Coiffait), det. V. Assing 2011” (MNHNP). Paratype 3: “Nepal XI.78.3 / Jaljale Himal, 2920 m PC-3 / Paratype / Lathrobium jal- jalensis H. Coiffait / Medon jaljalensis (Corffait), det. V. Assing 2011” (MNHNP). Comment. The original description is based on a male holotype and three paratypes, a male and two females, from “Népal oriental, Jaljale Himal 2950 m” deposited in the Coiffait collection (Coiffait 1984). An examination of the holotype and the male paratype revealed that the species belongs to the genus Medon Stephens, 1833. Redescription. Body length 3.7-3.8 mm; length of fore- body 2.0 mm. Habitus as in Fig. 290. Coloration: forebody uniformly reddish to dark-reddish; abdomen uniformly reddish to dark-brown with reddish apex; legs and anten- nae yellowish to reddish-yellow. Head almost as wide as long; lateral margins behind eyes straight and subparallel; punctation moderately coarse and moderately sparse, somewhat sparser in median dor- sal portion than elsewhere; interstices mostly broader than diameter of punctures and without microsculpture. Eyes moderately large and weakly convex, approximately 0.7 times as long as postocular region in dorsal view. Pronotum approximately as long as broad and slightly broader than head; punctation similar to that of head, but somewhat denser; interstices without distinct microsculp- ture, glossy. Elytra short, approximately 0.7 times as long as prono- tum, slightly widened posteriad; humeral angles weakly marked; punctation fine, shallow, and dense; interstices without distinct microsculpture. Hind wings completely reduced. Bonn zoological Bulletin 61 (2): 142-209 Abdomen broader than elytra; punctation very fine and moderately dense, barely noticeable in the pronounced mi- crosculpture; posterior margin of tergite VII without pal- isade fringe. 3: sternite VII weakly transverse, posterior margin with indistinct excision in the middle, on either side of middle with a row of approximately eight longer marginal setae (Fig. 291); sternite VIII oblong, with moderately deep pos- terior excision (Fig. 292); aedeagus approximately 0.5 mm long, shaped as in Figs 293-294. Comparative notes. As can be inferred from the shape and chaetotaxy of the male sternite VII and from the mor- phology of the aedeagus, this species undoubtedly belongs to the Medon apicalis group. It is distinguished from oth- er species of this group by the male sexual characters, from most of them additionally by the short elytra. For illus- trations of the micropterous representatives of the M. api- calis group known from Nepal see Assing (2010a). Distribution and natural history. Medon jaljalensis is currently known only from the type locality in eastern Nepal, where it was collected at an altitude of almost 3000 m. The reduced wings, the absence of a palisade fringe at the posterior margin of the abdominal tergite VII, and the absence of additinal records suggest that the species has a restricted distribution. Acknowledgements. I am indebted to the colleagues indicated in the material section for the loan of material under their care. Special thanks are due to Benedikt Feldmann, Andreas Kleeberg, and Ales Smetana for the generous permission to retain sever- al holotypes and single males needed for future reference pur- poses. Lee H. Herman (New York) cross-checked the names treated in this article against his unpublished catalogue of Paed- erinae. Thierry Deuve (MNHNP), Wolfgang Schawaller (SMNS), and Joachim Schmidt (Greifswald) kindly assisted in the identification of localities in Nepal, Irene Schatz (Innsbruck) in arranging the loan of the holotype of Lathrobium janetsche- ki. Harald Schillhammer (NHMW) made scans of excerpts from Franz’ diary available. Benedikt Feldmann proof-read the man- uscript. REFERENCES Ahrens D (2004) Monographie der Sericini des Himalaya (Co- leoptera: Scarabaeidae). Dissertation.de —Verlag im Internet GmbH, Berlin: 534 pp. Assing V (2010a) A revision of Palaearctic Medon. VIII. A new species from Nepal and additional records (Coleoptera: Staphylinidae: Paederinae). 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Linzer Biologische Beitrage 44 (1): 409-419 Assing V (2012d) The genus 7risunius in the Himalaya (Coleoptera: Staphylinidae: Paederinae: Medonina). Bonn Zo- ological Bulletin 61 (2): 210-215 Assing V (in press) The Pseudolathra species of the East Palaearctic and the Oriental regions (Coleoptera: Staphylin- idae: Paederinae). Beitrage zur Entomologie, Keltern 62 (2) (2012) Cameron M (1931) The Fauna of British India including Cey- lon and Burma. Coleoptera. Staphylinidae. Vol. 2. Taylor and Francis, London: viii + 257 pp. Coiffait H (1975) Xantholininae, Paederinae et Euaesthetinae ré- coltés au Népal par le professeur Franz (Col. Staphylinidae). Nouvelle Revue d’Entomologie 5 (2): 153-186 Coiffait H (1981) Staphylinides nouveaux du Népal. Nouvelle Revue d’Entomologie 11 (4): 323-335 Bonn zoological Bulletin 61 (2): 142-209 Coiffait H (1982a) Contribution a la connaissance des staphy- linides de l’Himalaya (Népal, Ladakh, Cachemir). Sencken- bergiana Biologica 62 (1981): 21-179 Coiffait H (1982b) Staphylinides (Col.) de la région himalayenne et de I’Inde (I. Xantholininae, Staphylininae et Paederinae). Entomologica Basiliensia 7: 231-302 Coiffait H (1983) Staphylinides du massif du Ganesh Himal (Né- pal Central) récoltés par Th. Deuve et E. Queinnec. Descrip- tions de nouvelles espéces et d’un nouveau genre himalayens. Nouvelle Revue d’Entomologie 13 (2): 161-179 Coiffait H (1984) Contribution a la connaissance des staphyli- nides de |’ Himalaya (Coleoptera, Staphylinidae). Annales de la Société Entomologique de France (N. S.) 20 (4): 373-387 Coiffait H (1987) Nouvelles corrections homonymiques (Col. Staphylinidae). Nouvelle Revue d’Entomologie (N. S.) 3 (4) (1986): 497-498 Scheerpeltz O (1976) Wissenschaftliche Ergebnisse der von Prof. Dr. H. Janetschek im Jahre 1961 in das Mt.-Everest-Gebiet Nepals unternommenen Studienreise (Col. Staphylinidae). Khumbu Himal. Ergebnisse des Forschungsunternehmens Ne- pal Himalaya 5: 1—75 Smetana A (2004) Subfamily Paederinae Fleming, 1821. In: Lobl I. & Smetana A (eds.) Catalogue of Palaearctic Coleoptera. Vol. 2. Hydrophiloidea — Histeroidea — Staphylinoidea. Apol- lo Books, Stenstrup: 579-624 ©ZFMK Bonn zoological Bulletin 61 (2): 210-215 December 2012 The genus 7risunius in the Himalaya (Coleoptera: Staphylinidae: Paederinae: Medonina) Volker Assing Gabelsbergerstr. 2, D-30163 Hannover, Germany; E-mail: vassing.hann@t-online.de. Abstract. The genus Jrisunius Assing, 2011 is reported from Nepal for the first time. Four species are (re-)de- scribed and illustrated: 7. perpusillus (Coiffait, 1982), comb. n. (ex Lathrobium Gravenhorst, 1802), 7: alesi sp. n. (northeastern Nepal), 7) opaciceps sp. n. (central Nepal: Manaslu), and 7? manasluensis sp. n. (central Nepal: Manaslu). Including these species, 7risunius is now represented in the Himalaya by five species and comprises a total of 14 species. A supplement to a recent key is provided. The distribution of the genus in the Himalaya is mapped. Key words. Taxonomy, rove beetles, Trisunius, Lathrobium, Himalaya, Nepal, new species, new combination, key to species INTRODUCTION The recently described medonine genus 7risunius previ- ously comprised ten species. Its known distribution ranges from the Himalaya, where it was represented only by 7. monticola (Cameron, 1932) from Uttaranchal, to eastern China and Thailand. Eight of the ten species have been recorded only from the Chinese province Yunnan. A cat- alogue and a key to species are provided by Assing (2011). A taxonomic revision of Himalayan Lathrobium Gravenhorst (Assing 2012) revealed that one of the species, L. perpusillum Coiffait, 1982, in fact belongs to Trisunius. A second, undescribed species was discovered among unidentified Lathrobium material from Nepal, and two additional species, both of them undescribed, were found in Paederinae material from Nepal deposited in the Naturkundemuseum Erfurt. These findings suggest that Trisunius 1s widespread in the Himalaya and that the genus is probably represented in this regions by numerous ad- ditional species that remain to be discovered. For a redescription and illustrations of 7: monticola see Assing (2011). MATERIAL AND METHODS The morphological studies were conducted using a Ste- mi SV 11 microscope (Zeiss Germany) and a Jenalab com- pound microscope (Carl Zeiss Jena). A digital camera (Nikon Coolpix 995) was used for the photographs. Head length was measured from the anterior margin of the frons to the posterior margin of the head, elytral length at the suture from the apex of the scutellum to the posteri Received: 14.04.2012 Accepted: 10.10.2012 or margin of the elytra, and the length of the aedeagus from the apex of the ventral process to the base of the aedea- gal capsule. The “parameral” side (1.e., the side where the sperm duct enters) is referred to as the ventral, the oppo- site side as the dorsal aspect. The map was created using MapCreator 2.0 (primap) software. COLLECTION MATERIAL DEPOSITORIES MNHNP Muséum national d’Histoire naturelle, Paris, France (A. Taghavian) NME _ Naturkundemuseum Erfurt, Germany (M. Hart- mann, assisted by W. Apfel) cAss author's private collection RESULTS Trisunius perpusillus (Coiffait, 1982), comb. n. (Figs 1-4, 21) Lathrobium perpusillum Coiffait, 1982: 289 f. Type material examined. Holotype @: “Nepal X.81, Kalingchok, 3100 m P.C / Holotype / Lathrobium perpusil- lum H. Coiffait 1982 / Trisunius perpusillus (Corffait), det. V. Assing 2011” (MNHNP). Comment. The original description is based on a unique male holotype from “Népal, Massif du Kalingchok, pres Corresponding editor: D. Ahrens The genus Zrisunius in the Himalaya an Figs 1-10. Trisunius perpusillus (1-4) and T: alesi (5-10). 1, 6: forebody; 2, 7: male sternite VII; 3, 8: male sternite VIII; 4, 9: aedeagus in lateral view; 5: habitus; 10: ventral process of aedeagus in ventral view. Scale bars: 5: 1.0 mm; 1, 6: 0.5 mm; 2-3, 7-8: 0.2 mm; 4, 9-10: 0.1 mm. de Barbabise [sic], 2370 m [sic]” (Coiffait 1982). An ex- amination of the holotype revealed that this species does not belong to Lathrobium, but to Trisunius. Redescription. Very small species, body length 2.8 mm; length of forebody 1.45 mm. Coloration: body uniform- ly reddish; legs and antennae yellowish. Head (Fig. 1) oblong, almost 1.1 times as long as broad; punctation very fine, barely noticeable in the pronounced microreticulation, except for a macropuncture on either side in median dorsal portion. Anterior margin of labrum with two tooth-like projections on either side of the U- shaped median incision. Eyes not projecting from lateral contours of head, somewhat less than half as long as pos- tocular region in dorsal view. Pronotum (Fig. 1) approximately 1.05 times as long as broad and as broad as head; punctation dense and more Bonn zoological Bulletin 61 (2): 210-215 distinct than that of head, interstices without distinct mi- crosculpture, glossy. Elytra short, approximately 0.7 times as long as prono- tum (Fig. 1); humeral angles weakly marked; punctation fine and dense; interstices without distinct microsculpture. Hind wings completely reduced. Abdomen broader than elytra; punctation very fine and dense, barely noticeable in the pronounced microsculpture; posterior margin of tergite VII without palisade fringe. &: protarsomeres I-IV moderately dilated; sternite VII moderately transverse, with broad and shallow posterior margin (Fig. 2); sternite VIII approximately as long as wide and with small V-shaped posterior excision (Fig. 3); aedeagus approximately 0.4 mm long, shaped as in Fig. 4. ©: unknown. ©ZFMK 212 Volker Assing Figs 11-20. Trisunius manasluensis (11-15) and T. opaciceps (16-20). 11, 16: habitus; 12, 17: male sternite VII; 13, 18: male sternite VIII; 14, 19: aedeagus in lateral view; 15, 20: aedeagus in ventral view. Scale bars: 11, 16: 1.0 mm; 12—13, 17—18: 0.2 mm; 14-15, 19-20: 0.1 mm. Comparative notes. Trisunius perpusillus is distin- guished from its congeners particularly by the morphol- ogy of the aedeagus. For additional characters distinguish- ing it from other Himalayan representatives of the genus see the key at the end of the article. Distribution and natural history. The species is current- ly known only from the type locality, the Kalinchok, a mountain some 13 km to the east of Barahbise and almost 70 km to east-northeast of Kathmandu in eastern central Nepal (Fig. 21). According to the label attached to the holotype, the specimen was collected at an altitude of 3100 m. Bonn zoological Bulletin 61 (2): 210-215 Trisunius alesi sp. n. (Figs 5—10, 21) Type material. Holotype @: “Nepal, Khandbari District / above Tashigaon, 3600 m 6.1V.1982, A. & Z. Smetana /Holotypus 3 Trisunius alesi sp. n., det. V. Assing 2012” (cAss). Etymology. This species is dedicated to Ales Smetana, Ot- tawa, who collected the holotype. His generous offer to retain single males for future reference purposes is much appreciated. Description. Body length 3.8 mm; length of forebody 2.0 mm. Habitus as in Fig. 5. Coloration: body uniformly red- dish. ©ZFMK The genus Jrisunius in the Himalaya 213 Fig. 21. Distribution of Zrisunius in the Himalaya: 7: monticola (square); T: manasluensis and T. opaciceps (circle); T: perpusil- lus (triangle); 7? alesi (diamond). Head (Fig. 6) weakly oblong, approximately 1.05 times as long as broad; punctation dense and fine; interstices with microreticulation and almost matt. Anterior margin of labrum with two tooth-like projections on either side of the U-shaped median incision. Eyes not projecting from lateral contours of head, barely one third as long as pos- tocular region in dorsal view, composed of approximate- ly 15 ommatidia. Pronotum 1.1 times as long as broad and approximate- ly 0.9 times as broad as head; punctation and microsculp- ture similar to that of head; midline with narrow shiny band reaching neither anterior nor posterior margins (Fig. 6). Elytra short, approximately 0.75 times as long as prono- tum, dilated posteriorly (Fig. 6); humeral angles weakly marked; punctation fine and dense; interstices without dis- tinct microsculpture. Hind wings completely reduced. Abdomen distinctly broader than elytra; punctation very fine and dense; interstices with microsculpture; posterior margin of tergite VII with narrow palisade fringe. 3: protarsomeres I-IV moderately dilated; sternite VII moderately transverse, in posterior half with extensive me- dian impression, this impression furnished with two rather extensive clusters of dense dark setae, posterior margin bisinuate, weakly concave in the middle (Fig. 7); sternite Bonn zoological Bulletin 61 (2): 210-215 VII weakly transverse, posteriorly with median impres- sion without modified pubescence, posterior excision rel- atively deep and almost V-shaped (Fig. 8); aedeagus 0.53 mm long, with relatively shout ventral process (Figs 9-10). ©: unknown. Comparative notes. 7risunius alesi is distinguished from other Himalayan congeners particularly by its larger size, the uniformly reddish coloration of the body, the small eyes, and by the male sexual characters. Distribution and natural history. The type locality is sit- uated near Tashigaon [27°36’N, 87°15’E] in Khandbari district, northeastern Nepal (Fig. 21). The holotype was collected at an altitude of 3600 m. Trisunius manasluensis sp. n. (Figs 11-15, 21) Type material. Holotype ¢@: “Nepal, Manaslu Mts., 28°22’N, 84°29’E, E slope of Ngali Khola Vall., 2000-2300 m, leg. Schmidt, 15.V.2005 / Holotypus ¢ Trisunius manasluensis sp. n., det. V. Assing 2012” (NME). ©ZFMK 214 Volker Assing Etymology. The specific epithet is a latinized adjective derived from the name of the mountain range where this species was discovered. Description. Body length 3.3 mm; length of forebody 1.7 mim. Habitus as in Fig. 11. Coloration: forebody dark-red- dish; abdomen dark-brown; legs and antennae reddish. Head 1.05 times as long as broad; dorsal surface with very dense microreticulation, with subdued shine; punc- tation dense and shallow. Anterior margin of labrum with two tooth-like projections on either side of the U-shaped median incision. Eyes not projecting from lateral contours of head, less than half as long as postocular region in dor- sal view, composed of approximately 30-40 ommatidia. Pronotum approximately 1.1 times as long as broad and 0.95 times as broad as head; punctation fine, dense, and shallow; microsculpture weakly pronounced, dorsal sur- face therefore more shiny than head; midline without im- punctate band. Elytra short, 0.8 times as long as pronotum, lateral mar- gins diverging posteriad in dorsal view; humeral angles weakly marked; punctation dense and shallow; interstices without distinct microsculpture. Hind wings completely reduced. Abdomen somewhat broader than elytra; punctation very fine, dense on anterior tergites, somwhat sparser on posterior tergites; interstices with shallow microsculpture; posterior margin of tergite VII without palisade fringe. S: protarsomeres I-IV weakly dilated; sternite VII strongly transverse, posteriorly with shallow median im- pression, this impression with slightly stouter setae, pos- terior margin broadly and weakly concave (Fig. 12); ster- nite VII moderately transverse, posterior excision almost V-shaped and not very deep (Fig. 13); aedeagus 0.45 mm long, ventral process slender and of distinctive shape (Figs 14-15). Comparative notes. This species is characterised partic- ularly by the reduced hind wings, the absence of a pal- isade fringe at the posterior margin of tergite VII, and by the distinctive shape of the aedeagus. Etymology. The specific epithet is a latinized adjective derived from the name of the mountain range where this species was discovered. Distribution and natural history. The type locality is sit- uated in the Manaslu range in central Nepal (Fig. 21), where the holotype was found at an altitude of 2000-2300 m, together with the following species. Bonn zoological Bulletin 61 (2): 210-215 Trisunius opaciceps sp. n. (Figs 16-21) Type material. Holotype ¢: “Nepal, Manaslu Mts., 28°22’N, 84°29’E, E slope of Ngali Khola Vall., 2000-2300 m, leg. Schmidt, 15.V.2005 / Holotypus 3 Trisunius opaciceps sp. n., det. V. Assing 2012” (NME). Paratype °: same data as holotype (cAss). Etymology. The specific epithet is a noun in apposition and alludes to the dull head. Description. Body length 2.8—3.0 mm; length of forebody 1.6—1.7 mm. Habitus as in Fig. 16. Coloration: head and pronotum dark-brown; elytra reddish-brown with paler posterior margins; abdomen blackish-brown; legs reddish- brown; antennae reddish. Head approximately as broad as long or weakly oblong; dorsal surface with pronounced, very dense microreticu- lation, matt; punctation moderately dense and shallow, barely noticeable in the microsculpture. Anterior margin of labrum with two tooth-like projections on either side of the U-shaped median incision. Eyes weakly projecting from lateral contours of head, little more than half as long as postocular region in dorsal view, composed of approx- imately 40-50 ommatidia. Pronotum approximately 1.1 times as long as broad and approximately 0.95 times as broad as head; punctation fine, dense, and shallow; microsculpture weakly pro- nounced, dorsal surface therefore much more shiny than head; impunctate band along midline indistinct or very narrow. Elytra approximately as long and 1.10—1.15 times as wide pronotum, lateral margins subparallel in dorsal view; humeral angles marked; punctation fine and dense; inter- stices with indistinct microsculpture. Hind wings appar- ently present, but possibly of reduced length. Abdomen approximately as broad as elytra; punctation very fine and dense; interstices with shallow microsculp- ture; posterior margin of tergite VII with palisade fringe. 3: protarsomeres I-IV weakly dilated; sternite VII mod- erately transverse, posteriorly with shallow median im- pression, pubescence unmodified, posterior margin weak- ly concave in the middle (Fig. 17); sternite VIII weakly transverse, posteriorly with narrow median impression without pubescence in the middle, posterior excision rel- atively deep and V-shaped (Fig. 18); aedeagus small, 0.30 mm long, and apically bifid in ventral view (Figs 19-20). Comparative notes. Based on the synapomorphically de- rived morphology of the aedeagus (small size, apically bi- fid), 7. opaciceps is undoubtedly most closely related to T. monticola, from which it is distinguished by smaller size, smaller eyes, more pronounced microreticulation of the head, shorter elytra, and by the shape of the aedeagus (particularly in lateral view). For illustrations of TZ mon- OZFMK The genus 7risunius in the Himalaya 215 ticola see Assing (2011). For characters distinguishing 7: opaciceps from other Himalayan congeners see the key at the end of the article. Distribution and natural history. The type locality is identical to that of 7) manasluensis and situated in the Manaslu range in central Nepal (Fig. 21) at an altitude of 2000-2300 m. KEY TO SPECIES In the key in Assing (2011), the species treated in the pres- ent paper would key out at couplet 3, together with 7? mon- ticola. In order to account for the additional species, the key is modified as follows: Bee Specics om the ldimalayares ns. scseseeseee sce. 3a SMES PECICS MOMMY UnMan Chinaeecn-nesceceeeesee ees -- 4 3a. Elytra at least approximately as long as pronotum. Eyes at least half as long as postocular region in dor- sal view. G: sternite VIII with relatively deep, V- shaped posterior excision (Fig. 18); aedeagus conspic- uously small, approximately 0.3 mm long, apically bi- iG! im TATE WEN (18, A) oosccossocessosenasscace 3b — Elytra distinctly shorter than pronotum. Eyes less than half as long as postocular region in dorsal view. ¢: sternite VIII mostly with less deep and broader pos- terior excision; aedeagus larger, distinctly longer than 0.3 mm, apically not bifid, and with slender ventral (UROCESS so cgooseadonsamoetacu noc c ona oecnr aac iad aes 3¢ 3b. Elytra longer than pronotum (Assing 2011: Fig. 58). Eyes distinctly more than half as long as postocular region in dorsal view. Dorsal surface of head with shallower microsculpture, with subdued shine. ¢: aedeagus shaped as in Assing (2011: Figs 65-66). In- dia: Uttaranchal (Fig. 21) ...... monticola (Cameron) — Elytra approximately as long as pronotum (Fig. 16). Eyes slightly more than half as long as postocular re- gion in dorsal view. Dorsal surface of head with pro- nounced microreticulation and matt. 4: aedeagus shaped as in Figs 19-20. Central Nepal (Fig. 21) 00000 0 COU CHOBE AERO ea Ee eae eae opaciceps sp. 0. 3c. Larger species; body length 3.8 mm; length of fore- body approximately 2.0 mm. Eyes smaller, composed of only approximately 15 ommatidia, barely one third as long as postocular region in dorsal view. Prono- tum with distinct microsculpture (Fig. 6). Abdominal Bonn zoological Bulletin 61 (2): 210-215 tergite VII with narrow palisade fringe posteriorly. ¢: sternite VII with extensive median impression pos- teriorly, this impression with two clusters of dense dark setae (Fig. 7); sternite VIII with deep posterior excision (Fig. 8); aedeagus larger, 0.53 mm long, shaped as in Figs 9-10. Eastern Nepal: Khandbari dis- CRC (ES DIN) ae rrecereine cers ane incense Smee alesi sp. n. — Smaller species; body length 3.3 mm at most; length of forebody 1.7 mm at most. Eyes relatively larger, composed of at least approximately 30 ommatidia, nearly half as long as postocular region in dorsal view. Pronotum without, or with very shallow microsculp- ture (Fig. 1). Abdominal tergite VI without palisade fringe posteriorly. @: sternite VII of different shape and chaetotaxy; sternite VIII with less deep posteri- or excision; aedeagus 0.45 mm long at most and of ditlerentashapemerce es come ere oe ceria 3d 3d. Larger species; body length 3.3 mm; length of fore- body 1.7 mm. ¢: sternite VII more strongly transverse and with somewhat stouter pubescence in posterior median impression (Fig. 12); sternite VHI with larg- er posterior excision (Fig. 13); aedeagus with ventral process of distinctive shape (Figs 14-15). Central Nepal: Manaslu (Fig. 21) ...... manasluensis sp. n. — Smaller species; body length 2.8 mm; length of fore- body 1.45 mm. @: sternite VII neither with distinct impression nor with modified pubescence (Fig. 2); sternite VIII with very small posterior excision (Fig. 3); aedeagus of different shape (Fig. 4). Nepal: Kalin- Choka(Big2il)) aeeecerncesc es T. perpusillus (Coiffait) Acknowledgements. I am indebted to the colleagues indicated in the material section for the loan of material under their care. Special thanks are due to Ales Smetana (Ottawa) for the gener- ous gift of the holotype of 7 alesi. Benedikt Feldmann (Mtin- ster) proof-read the manuscript. REFERENCES Assing V (2011) Trisunius gen.nov. from the southern East Palaearctic and the Oriental regions (Coleoptera: Staphylin- idae: Paederinae: Medonina). Linzer biologische Beitrage 43 (1): 195-220 Assing V (2012) A revision of the Lathrobium species of the Hi- malaya (Coleoptera: Staphylinidae: Paederinae). Bonn Zoo- logical Bulletin 61 (2): 142—209 Coiffait H (1982) Staphylinides (Col.) de la région himalayenne et de I’Inde (I. Xantholininae, Staphylininae et Paederinae). Entomologica Basiliensia 7: 231—302 ©ZFMK Bonn zoological Bulletin 61 (2): 216-240 December 2012 Amphibians and reptiles collected by Moritz Wagner, with a focus on the ZFMK collection Philipp Wagner” , Aaron M. Bauer & Wolfgang Béhme ‘Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA. ‘Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-53113 Bonn, Germany. “Corresponding author: E-mail: philipp.wagner.zfmk@uni-bonn.de. Abstract. Moritz Wagner (1813-1887) is one of the least poorly-known German explorers, geographers and biologists of the 19" century. Between 1836 and 1860, expeditions led him to Algeria, the Caucasus Region, as well as to North-, Central- and South-America. Beside his important scientific contributions to biology, geography and ethnogra- phy he also collected large numbers of plant and animal specimens. The collected material is scattered among several European museums and university collections because Wagner only obtained a permanent position after his last voyage. Prior to this he donated his material to experts, funding societies or the institutions where he was a student or in whose collections he worked. The present article is a first contribution towards a review of the herpetological collections made by Moritz Wagner, which includes type material of several amphibians and reptiles. Most specimens were found in the Zoologische Staatssammlung Miinchen and the Gottingen collection, which is now housed at the Museum Koenig in Bonn, but the collections in Berlin, Leiden, Paris and Vienna also hold material collected by Wagner. Key words. Moritz Wagner, historic collections, type material. INTRODUCTION Friedrich Johann Carl Moritz Wagner (1813-1887, Fig. 1) was a German scientist and naturalist, honorary pro- fessor of ethnology at the Ludwig-Maximilians Univer- sity and first curator of the Museum of Ethnology (at this time still named “k6niglich ethnographische Sammlung” [= the Royal Ethnographic Collection]) in Munich. He was a well-known explorer and many of his books where trans- lated to English, French or even Armenian (see Fig. 2) dur- ing his lifetime. As a scientist he worked in different sub- ject areas including geography, botany, zoology, ethnol- ogy, geology, politics, and journalism. Wagner produced notable results in many of these fields (Beck 1951, 1953, 1971; M. Wagner 1861, 1862; P. Wagner 2008). For ex- ample, he hypothesized the course of the Panama Canal decades before it was formally planned and built (M. Wag- ner 1861, Beck 1971) and was the discoverer of the west- ern source of the river Euphrates (Beck 1971). As a zo- ologist, he was one of the first scientists to work in the field of zoogeography, comparing distribution patterns, most importantly, of insects, fishes and mammals, but al- so of reptiles. These disciplines are also reflected in Wagner’s field collections. During his voyages, he collected large num- bers of insects, but only a few vertebrate vouchers, main- ly fishes. Parts of these collections were lost during the expeditions or during shipment (see below). Moreover, his collections are not stored in one institution, but were dis- persed to various important European natural history mu- seums (e.g., the museums of Berlin, Munich, Paris and Vi- enna). He often donated his material to taxonomists (e.g., Received: 02.05.2012 Accepted: 14.11.2012 Arnold A. Berthold, Hermann Schlegel, Franz Steindachn- er) who identified the specimens and sometimes added a chapter to his scientific books (e.g., Schlegel in M. Wag- ner 1841). Other parts of the collections were sold to mu- seums to finance his travels, or were donated to institutes where he studied or worked. It was only his last voyage to South America that was not privately organized, but rather funded by the Bavarian King Maximilian II. There- fore, the main parts of the zoological collection of this ex- pedition were inventoried in the collections of the Zool- ogische Staatssammlung in Munich (e.g., at least 72 fish- es, Neumann 2006), but partly later exchanged with oth- er institutions (e.g., NMW) or destroyed in the Second World War. Nevertheless, Moritz Wagner had a strong re- lationship to his brother, Rudolf Wagner, who was Pro- fessor and Director of the Institute of Physiology in Got- tingen, where Moritz Wagner studied geology and geog- raphy. It is, therefore, not surprising that Moritz Wagner donated many specimens from his collections from Alge- ria and Caucasus to the Institute of Physiology and the Zo- ological Museum at the University of Gottingen. I’ Expedition: Algeria (1836-38) In the introduction to his book about Algeria Wagner (1841) mentioned that he visited the North African coast in 1834 before his longer voyage to Algeria two years lat- er. Even though this was a short stay it may be assumed that Wagner collected at least insect material, as he also studied this group before his first longer expedition. Corresponding editor: F. Herder Amphibians and reptiles collected by Moritz Wagner 217 Fig. 1. Moritz Wagner (1813-1887), portrait from Franz Hanfstaeng! about 1860. Courtesy of the Stadtmuseum Miinchen. Wagner travelled (1836-38) to Algeria during the French occupation and war (1830-1847) as member of a scientific group within the French army. This was the same expedition during which Alphonse Guichenot, a well- known herpetologist from the Paris Museum, collected da- ta for his herpetological monograph (Guichenot 1850). The amphibians and reptiles collected by Wagner were lat- er identified by Hermann Schlegel (in M. Wagner 1841), a famous herpetologist of the time who was then the as- sistant of Coenraad Jacob Temminck, Director of the Riyksmuseum van Natuurlijke Historie [today Naturalis] in Leiden. As a result of Schlegel’s involvement many AI- gerian specimens are now part of the Leiden collection. However, Wagner also donated material from Algeria to the Géttingen collection, as he studied geology and ge- ography at this university after his return from Algeria (see P. Wagner 2008). The most important herpetological re- sults of this journey were a series of specimens of the then poorly-known, 7rogonophis wiegmanni (Fig. 3) and the description of Bufo mauritanicus by Schlegel (in M. Wag- ner 1841). Bonn zoological Bulletin 61 (2): 216-240 De Expedition: Caucasia (1842-44) During this expedition Wagner collected material mainly in the Caucasus region east of the Black Sea, today in the modern countries of Armenia, Azerbaijan, Georgia, northwestern Iran, southern Russia, Ukraine (Crimean Peninsula) and Turkey. Like the material from Algeria, parts of Wagner’s Caucasian collection were also exam- ined by specialists. The amphibians and reptiles were iden- tified by Arnold Adolph Berthold (see below, and M. Wag- ner 1850). Moritz Wagner donated his collected material to the Zoological Museum of the University of Géttingen, but also to the Institute of Physiology at the same univer- sity, where his brother, Rudolf Wagner, was director at this time. Rudolf mentioned a human skull collected by his brother with the collection number 507 from Caucasia (R. Wagner 1852). Moreover, the catalogue of the Zoologi- cal Museum lists vouchers collected by M. Wagner but obtained from the Institute of Physiology (see species ac- count). Berthold (in M. Wagner 1850) mentioned that Wagner collected 29 amphibian and reptile species rep- resenting 18 genera, but Berthold actually listed 30 species in total. However, several taxa (Amphibia: Hyla viridis; Sauria: Lacerta strigata, Lacerta viridis, Ophiops elegans, Anguis fragilis; Ophidia: Typhlops vermicularis, Coronel- la collaris, Coluber leopardinus, Coluber sauromates, Vipera berus) are mentioned by Berthold (in M. Wagner 1850), but these are neither found in the catalogue of the museum nor in the collection. Therefore, most probably Wagner also gave parts of this collection to other institutes, or Berthold exchanged these specimens before they were entered in the catalogue. To date it has not been possible to locate these missing vouchers. 3” expedition: North- and Central America (1852-55) and 4 expedition: Central- and South America (1857-60) Between 1852 and 1855 Wagner travelled together with the Austrian Carl Ritter von Scherzer through North Amer- ica to Central America (e.g. Honduras, Nicaragua, Costa Rica, El Salvador). Scherzer (Anonymus 1856) reported to the Austrian Academy of Science that only a few ver- tebrates were collected and donated to Austrian institutes and private collections. Only a small number of amphib- ians or reptiles have been identified in the Vienna collec- tion. Furthermore a donation of material to other specif- ic Austrian collections is not indicated in the publications about their travels (Scherzer & M. Wagner 1857; M. Wag- ner 1870; M. Wagner and Scherzer 1854, 1856) and, there- fore, the whereabouts of other material in Austria remains unresolved. The number of vertebrates collected was ad- ditionally reduced because some material was lost due to inadequate preservation during the expedition. Moreover, some material from Central America, held in storage pri- or to shipping, was destroyed during an earthquake on April 16 1854 in San Salvador (see M. Wagner, 1870). ©ZFMK 218 Philipp Wagner et al. UES tb ener wv Ue UCU h NUUCAWUL ANE PAL bb rk S£USUUSU bv LOLUPLN, PUTT CLES 2. PPT PAM ATUATUD UPPROULbuD NEMSEY IS \ we x mocis—igag § 2 8 icherel? a f i; Ub bv AUSAEW Oo Wo ur LPL md. 1851. TUTSA U- Ar menisile U bers Syorrs Vor SM + Wasnss ‘ yore Oo Mc nd ea avo? Ay pbb cifoeds Sagare padh bon Cyarom VU8B UAPS 4 Veme Peg gore. Fig. 2. Left: Lake Sewan and Mount Ararat (above) and an unknown area (below) in Armenia. Figured in the 1851 published Ar- menian translation of Reise nach dem Ararat und dem Hochland Armenien (Wagner 1848). Right: The title page of the Armenian edition (above) and handwritten information about the publisher (below). Courtesy of the V6lkerkundemuseum Minchen. Therefore, major parts of the third expedition , except those specimens he shipped before the earthquake, were destroyed. Later, on his fourth expedition (1858/1859), Wagner travelled alone to Central and northern South America and collected many insects and fishes (see Neu- mann 2006), but most probably only a small number of amphibians or reptiles. Most of these specimens were in- ventoried in the collection of the Zoologische Staatssamm- lung in Munich, because this journey was funded by the Bavarian King, Maximilian II, although later some were exchanged with the museum in Vienna or destroyed in the Second World War (nearly the entire ZSM fish collection and about half of the herpetological collection). Neumann (2006) mentioned that for the fish collected the location “Neu-Granada” was corrected by Wagner (1864) into “Panama” to indicate that he exclusively collected in this area, which was a province of New-Granada until 1903. Bonn zoological Bulletin 61 (2): 216-240 Moritz Wagner as biogeographer and darwinist Besides his relevance as a collector, Moritz Wagner also was a well-reputed biologist with a broad knowledge of many different groups, such as insects, fish and mammals, but also plants. Continuing from his first contributions, Moritz Wagner studied distribution patterns within differ- ent groups (e.g., Wagner 1841, mammals, birds, butter- flies, beetles, and land mollusks). This predates “The Ge- ographical Distribution of Animals” by Alfred Russell Wallace (1876) which is generally recognized as one of the early contributions to zoogeography. Therefore, Moritz Wagner must be recognized as one of the founders of this field in biology. Moreover, as he studied distribu- tion pattern in detail and firstly described rivers as a ge- ographic barrier between sibling species. Later, he was in- fluenced by e.g., Leopold von Buch and Alexander von Humboldt and therefore biogeography was a part of near- ly all of his contributions, but rarely he published works OZFMK Amphibians and reptiles collected by Moritz Wagner 219 exclusively in this field (e.g., Wagner 1866). However, it was the publication of Darwin’s epochal work “On the Origin of Species” which stimulated Moritz Wagner to de- velop his “Migrationstheorie” [= theory on the migration of species]. Here, he first described allopatric (geograph- ic) speciation, which was later adopted by Ernst Mayr. Even though Wagner (e.g., 1868) initially saw his theory as a good argument to support Darwin’s work, it was gen- erally recognized by others e.g., Haeckel and Weismann as anti-Darwinian. Moritz Wagner (1868) argued that the geographic isolation of two populations is the most im- portant requirement for speciation. Darwin, on the other hand, at first recognized Wagner’s theory, but considered isolation as a less important factor in speciation; later he totally ignored it (P. Wagner 2008). Remarks on the herpetological collection of the Zoolog- ical Museum at the University of Gottingen in respect to material collected by Moritz Wagner Apart from several historically important private collec- tions (which partly became state museums later on), the Zoological Museum of Gottingen is among the oldest in- stitutional German natural history museum. Since 1773 it had been a department of the Royal (1.e., Kingdom of Han- nover) Academic Museum and received its own building in 1793. The first director and curator of the zoological collection was, from 1776 onwards, the famous Johann Friedrich Blumenbach (1752-1840) whose work “Hand- buch der Naturgeschichte” [= Treatise of natural history] established his reputation as one of the founders of the modern discipline of comparative anatomy (see BOhme & Bischoff 1984). After Blumenbach’s death Arnold Adolph Berthold (1803-1861) became his successor. His fields of research were especially broad, one focus being the first experimen- tal studies on hormonal effects. However, of relevance for the present paper are his taxonomic works about crus- taceans, amphibians and reptiles. In this context, he con- tributed an important section to Moritz Wagner’s book “Reise nach Kolchis” [= Travel to Colchis] where he ex- amined the material donated by M. Wagner to the Zoo- logical Museum in Gottingen. Berthold’s connection to M. Wagner was most likely due to Wagner’s brother Rudolf Wagner, who was Professor and Director of the Institute of Physiology at the University of Gottingen and may have influenced Moritz to deposit his materials from Colchis as well as other voyages in this museum. After Berthold’s death, Wilhelm Moritz Keferstein (1833-1870) was able to continue the herpetological tradition at the G6ttingen museum with his work on amphibians from Costa Rica and Australia. But with his early death the phase of active herpetological research ended. Four years later, in 1874, the herpetology section improved when E. Ehlers became the curator and herpetologists like George Bonn zoological Bulletin 61 (2): 216-240 Albert Boulenger and Franz Werner studied the collection. In addition, important collections from South America, Australia and Asia were accessioned into the collection during this time. As a result, spaces for public exhibitions and scientific collections became limited and probably therefore, some collections were donated to other muse- ums (including a bird collection to Hannover, the fish col- lection to Hamburg). The herpetology collection was re- vised twice in Gottingen, in 1968 and 1970, and a third time during the inventory in Bonn, and several specimens must be regarded as lost. Unfortunately, it was not possi- ble to ascertain the reasons of these losses between the time of the inventories and the transfer to the Museum Koenig in Bonn, where the collection is stored today. It is the aim of this paper to summarize the herpetologi- cal collections of Moritz Wagner, with a special focus on the Gottingen material (now stored in Bonn), but also in- cluding known material from the museums in Berlin, Lei- den, Munich and Vienna. MATERIAL & METHODS Species accounts are headed with the current valid name of the identified species. Information of specimen records is referenced with “Material” which lists the specimens with their current collection number, old collection num- bers 1f available, locality data, collector and, if given, the collection date. The “Catalogue name” provides the his- torical museum catalogue number if the specimen was originally inventoried in a different collection, together with the species name given in the respective catalogue. If a specimen is mentioned in the literature, the “Citation” references the respective publication, accompanied by the mentioned species name and locality (if the locality is dif- ferent from that given with the specimen or in the muse- um catalogue). Is there no citation available the section was removed. Other information is provided under “Re- marks”. Museum abbreviations used within this publication: NMW, Naturhistorisches Museum, Wien, Austria; RMNH, Naturalis [formerly Rijksmuseum van Natuurlijke Historie], Leiden, Netherlands; ZFMK, Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany; ZMB, Museum fiir Naturkunde [formerly Zoologisches Museum Berlin], Berlin, Germany; ZMUG, Zoologisches Museum der Universitat Gottingen, Gottingen, Germany [sometimes incorrectly referred to as ZIUG= Zoological Institut of the University of Gottingen, which is a differ- ent collection]; ZSM, Zoologische Staatssammlung Munchen, Munich, Germany. OZFMK List of extant specimens of amphibians and reptiles col- lected by Moritz Wagner Amphibia, Gymnophiona [Caeciliidae| Caecilia albiventris (Daudin, 1803) Material: ZSM 2570/0 from “Ecuador” collected by “M. Wagner”. Catalogue name: “Caecilia albiventris”. Amphibia, Caudata [Ambystomidae] Ambystoma maculatum (Shaw, 1802) Material: ZSM 1259/0 [5 ex.] from “Wisconsin,” USA col- lected by “M. Wagner”. Catalogue name: “Ambystoma maculatum”. Ambystoma jeffersonianum (Green, 1827) Material: ZSM 1261/0 [8 ex.] from “Wisconsin,” USA col- lected by “M. Wagner”. Catalogue name: “Ambystoma jeffersonianum’. Remarks: This species does not occur in Wisconsin and it needs to be clarified if it is a misidentification or a wrong locality.. [Proteidae] Necturus maculosus Rafinesque, 1818 Material: ZFMK 27701 [ZMUG 26a] from “Nord Ameri- ka” [=North America, no further locality specified] donat- ed by the “Ph[ysiological] I[nstitute]”; ZFMK 27702 [ZMUG 26c] from “Nord Amerika” [=North America, no further locality specified] collected in 1853. Catalogue name: “Necturus maculatus” [ZMUG].Re- marks: There is no collector specified in the ZMUG cat- alogue but the locality is one collected by Wagner in 1853 and the donation was via the former the Physiological In- stitute, where Wagner’s brother was director. This strong- ly suggests that it was collected by Moritz Wagner. [Plethodontidae| Bolitoglossa palmata (Werner, 1897) Material: NMW 22862 [lectotype] from “Cordillera,” Ecuador collected by “M. Wagler [sic!].” Catalogue name: “Spelerpes palmatus”’. Citation: As “Spelerpes palmatus” by Werner (1897a); as “Spelerpes palmatus Werner” collected by “M. Wagler [sic!]” by Haupl et al. (1994). Remarks: Werner (1897a) mentioned that the type series consists of three specimens from Ecuador, two of which are housed at the ZSM, and one in the NMW collection. Moreover, he indicated that the specimens are from dif- ferent collectors ([Ludwig Karl]“Schmarda” [Austrian bio- geographer and collector, 1819-1908] and “M. Wagner”) and from the original source [‘(...) besitzt das zoolog. Mu- Bonn zoological Bulletin 61 (2): 216-240 220 Philipp Wagner et al. seum der Wiener Universitat ein junges, die zoolog. Sammlung des Staates in Miinchen zwei grofiere Exem- plare, welche alle aus Ecuador stammen (Schmarda bezw. M. Wagner coll.)*] it may be concluded that the Vienna specimen was collected by “Schmarda”, while the two Munich specimens by “M. Wagner”. In any case, the col- lector “M. Wagler” mentioned by Haupl et al. (1994) is obviously a misspelling, but refers rather to Moritz Wag- ner than to “Schmarda”’. This specimen (NMW 22862 [for- merly ZIUW Q43 fide Haupl et al. 1994]) was designat- ed as the lectotype by Brame & Wake (1962: 173). [Salamandridae] Notophthalmus viridescens (Rafinesque, 1820) Material: ZSM 1241/0 [8 ex.], ZSM 1243/0 [6 ex.] from “Wisconsin,” USA collected by “M. Wagner”. Catalogue name: “Triturus viridescens”’. Ommatotriton ophryticus (Berthold, 1846) Material: ZFMK 27793 [ZMUG 22a] [lectotype] from “Tiflis” [=Tbilisi], Georgia, collected by “M. Wagner” in 1846. Catalogue name: “Triton ophriticus”’ [sic!] [ZMUG]. Citation: “Triton ophryticus n. sp.“, from ,,bei Tiflis* [= near Tbilisi], (Berthold 1846); “Triton ophryticus n. sp.“, from ,,aus Grusien [Georgia] und von den Abhangen des Kaukasus,” Berthold in M. Wagner (1850, p. 335). Remarks: Mentioning two specimens, Berthold (in M. Wagner 1850) intended to describe “Triton ophryticus” in his checklist of the amphibians and reptiles collected by Moritz Wagner. However, the publishing of this book dragged out and Berthold published the description in ad- vance (Berthold 1846) in identical form to the later-pub- lished chapter (in Wagner 1850). One of the two syntypes seems to be lost and therefore ZFMK 27793 was desig- nated as the lectotype by Bohme & Bischoff (1984). Triturus karelini Strauch, 1870 Material: ZFMK 27779 [ZMUG 16m, ZMUG 18] from “Tiflis” [=Tbilisi], Georgia, collected by “M. Wagner” in 1846. Catalogue name: “Triton cristatus” [ZMUG 16m]; “7. vul- garis” [ZMUG 18]. Remarks: Berthold (in M. Wagner 1850) mentioned three species of the genus Triturus: T. ophryticus, T. taeniatus and 7. nycthemerus. While the first was collected in “Ti- flis”, the latter two were collected at the “tiirkisch-pon- tischen Kiiste [southern shore of the Black Sea]” and are therefore not identical with the given locality of this spec- imens. The first of these species was described by Berthold (1846, in Wagner 1850) and so that obviously this spec- imen was not mentioned by him (Berthold in Wagner 1850). ©ZFMK Amphibians and reptiles collected by Moritz Wagner 22] Amphibia, Anura [Bufonidae|] Atelopus longirostris (Cope, 1868) Material: ZSM 1017/0 from “Ecuador” [no further local- ity specified] collected by “M. Wagner”. Catalogue name: “Atelopus longirostris”’. Atelopus varius (Lichtenstein & von Martens, 1856) Material: NMW 3875.14 [4 ex., syntypes] from “Costa Rica” [no further locality specified] collected by “M. Wag- nene Catalogue name: “Hylaemorphus pluto”. Citation: As “Hylaemorphus pluto” by Schmidt (1858), Savage (1972). Remarks: According to Savage (1972), these specimens are the syntypes of Hylaemorphus pluto Schmidt, 1858. The type locality “Costa Rica” was later restricted to “Cartago, Canton Cartago, Provincia Cartago, Costa R1- ca” by Savage (1974). Bufo mauritanicus Schlegel, 1841 Material: ZFMK 27979 [ZMUG 56a] from “Algier,” Al- geria [no further locality specified] collected by “M. Wag- ner’; RMNH 2122 [holotype] from “Umgebungen von Al- gier” [=vicinities of Algier] collected by “M.F. Wagner”. Catalogue name: “Bufo mauritanicus ”’. Citation: “Bufo mauritanicus,’ Schlegel (in M. Wagner 1841, part HI, 134). Remarks: Even though this species was described by Schlegel (in M. Wagner 1841) the ZFMK voucher is not a type specimen, because Schlegel clearly mentioned a sin- gle voucher [“Das vorliegende Exemplar wurde in den Umgebungen von Algier gefunden.”, page 137 in Wagn- er 1841] and this holotype is present in the herpetology collection of Naturalis in Leiden. However, Schlegel ex- plicitly mentioned that it was not his intention to descri- be Bufo mauritanicus as a new species (“Wenn wir der in Algerien vorkommenden groBen Kroéte diesen neuen Na- men beilegen, so geschieht dies nur, um ihre Herkunft an- zudeuten, und wir wollen dieselbe keineswegs schon als neue Art angesehen wissen.”), but he gave a detailed des- cription and diagnosis. Therefore according to the Inter- national Code of zoological Nomenclature (ICZN 1999) this 1s a valid description. Schlegel is widely accepted as the sole author, but the entire description is in the plural, giving the impression of more than one author. This was a common style of formal writing at the time, but the in- tention of the usage of the German ‘wir’ cannot be final- ly resolved. Wagner (1841) mentioned that the species is common in the lowlands of Metidscha and rare near Bona. Dendrobates tinctorius (Cuvier, 1797) Material: ZSM 1012/0 from “Central Amerika” [=Central America, no further locality specified] collected by “M. Wagner”. Catalogue name: “Dendrobates tinctorius”’. Bonn zoological Bulletin 61 (2): 216-240 [Discoglossidae| Discoglossus pictus pictus Otth, 1837 Material: ZFMK 27894 [ZMUG 35a] from ,,Algier,* Al- geria [no further locality specified] collected by ““M. Wag- ner’; RMNH 1955 from “Algeria” [no further locality specified] collected by “M. F. Wagner”. Catalogue name: “Discoglossus pictus” [ZMUG]. Citation: “Rana picta,” Schlegel (in M. Wagner 1841, part I, 134). Remarks: Even though Schlegel (in M. Wagner 1841) mentioned the genus Discoglossus he did not accept this taxonomic assignation of ‘modern authors’ and still re- ferred the species to the genus Rana. [Hemiphractidae| Gastrotheca marsupiata (Duméril & Bibron, 1841) Material: ZSM 1188/0 [4 ex.] from “Pichincha,” Ecuador collected by “M. Wagner”. Catalogue name: “Gastrotheca marsupiata’’. Hemiphractus fasciatus Peters, 1862 Material: ZSM 36/0 [holotype] from “Catassatal” [sic!] (““Pastassa-Thal an der Ostseite der Anden [=Pastaza val- ley on the eastern slope of the Andes] in Ecuador” accord- ing to the description) collected by “M. Wagner”. Catalogue name: “Ceratohyla fasciata”. Citation: As “Hemiphractus fasciatus” by Trueb (1974) and Glaw & Franzen (2006). Remarks: According to Trueb (1974) the locality is erro- neous. [Hylidae] Duellmanohyla uranochroa (Cope, 1875) Material: NMW 6175 from “Costa Rica,” [no further lo- cality specified] collected by “M. Wagner”. Catalogue name: “Hyla uranochroa”. Hyla meridionalis Boettger, 1874 Material: RMNH 1699 from “Algeria” [no further local- ity specified] collected by M. Wagner. Catalogue name: “Hyla arborea”. Citation: As “Hyla arborea” by Schlegel (in M. Wagner 1841, part HI, 133). Remarks: Although Schlegel (in M. Wagner 1841) men- tioned several specimens, this is the only known vouch- er and was, at this time, the first record for Algeria. Hyla versicolor (LeConte, 1825) Material: ZSM 1162/0 [3 ex.] from “Wisconsin,” USA col- lected by “M. Wagner”. Catalogue name: “Hyla versicolor”. [Ranidae| Lithobates catesbeianus (Shaw, 1802) Material: NMW 2926 from “Costa Rica” [no further lo- OZFMK 222 Philipp Wagner et al. cality specified] collected by “M. Wagner”. Catalogue name: “Rana pipiens”. Remarks: This species does not does not occur in Costa Rica and was probably collected during Wagner’s North America travels. Pelophylax saharicus (Boulenger, 1913) Material: RMNH 1942 from “Algeria” [no further local- ity specified] collected by “M. Wagner”. Catalogue name: “Rana esculenta”. Citation: As “Rana esculenta” by Schlegel (in M. Wagn- er 1841, part IH, 133). Remarks: Schlegel (in M. Wagner 1841) mentioned sev- eral specimens, but this is the only known surviving voucher of this species from Wagner’s travels. [Strabomantidae] Pristimantis unistrigatus (Ginther, 1859) Material: ZSM 1052/0 from “Stid Amerika” [South Amer- ica, no further locality specified] collected by “M. Wag- ner Catalogue name: “E/eutherodactylus unistrigatus”. Reptilia, Testudinae [Chelydridae] Chelydra_ rossignoni (Bocourt, 1868) Material: ZMB 125 from “Mittelamerica” [=Central America, no further locality specified] collected by “Wag- ner”. Catalogue name: “Chelydra (Emysaurus) serpentina””. Chelydra_ serpentina (Linnaeus, 1758) Material: ZFMK 73776 [ZMUG 29b]_ from “Nordamer[ika]” [=North America, no further locality specified] collected by “Mor. Wagner” in 1853. Catalogue name: “Cinosternum pennsylvanicum” [ZMUG]. Remarks: Specimens of the genera Kinosternon and Chelydra are of course very distinct and specimens were likely mixed up earlier in the Géttingen (ZMUG) collec- tion. [Geoemydidae] Rhinoclemmys punctularia (Daudin, 1801) Material: ZSM 88/0 from “Centralamerika” [=Central America, no further locality specified] collected by “M. Wagner”. Catalogue name: Nicoria punctularia. [Kinosternidae| Kinosternon subrubrum Lacépeéde, 1788 Material: ZFMK 73778 [ZMUG 29a] _ from “Nordamer[ika]” [=North America, no further locality specified] collected by “Mor. Wagner” in 1853. Bonn zoological Bulletin 61 (2): 216-240 Catalogue name: “Cinosternum pennsylvanicum” [ZMUG]. [Testudinidae} Testudo graeca graeca Linnaeus, 1758 Material: RMNH 3234 from “Algeria” [no further local- ity specified] collected by “M. Wagner”. Catalogue name: “7estudo graeca”’. Citation: As “Zestudo graeca var. mauritanica” by Schlegel (in M. Wagner 1841, part II, 106). Remarks: Schlegel (in M. Wagner 1841) only mentioned T. graeca var. mauritanica but obviously the collected se- ries contained more than one species (see next species ac- count). Wagner (1841) mentioned that he found this species at all localities he visited in Algeria, and refers to areas with the Mediterranean dwarf Palm (Chamaerops humilis) as typical habitat where the specimens hide un- der leaves of this palm. Testudo graeca Linnaeus, 1758 Material: ZFMK 73732-733 [ZMUG 2e] from “Algier,” [=Algeria, no further locality specified] collected by “M. Wagner’, donated by “Phys[iologisches] Inst[itut]*. Catalogue name: “7estudo ibera” |ZMUG]. Citation: As “Zestudo graeca var. mauritanica” by Schlegel (in M. Wagner 1841, part III, 106). Remarks: The specimens were donated to the Zoological Museum in Gottingen from the Institute of Physiology of the same university. The head of the institute at this time was Rudolf Wagner, the older brother of Moritz Wagner. M. Wagner thus gave material directly to his brother as well as to the museum and its director, A. Berthold. Reptilia, Amphisbaenia [Trogonophiidae] Trogonophis wiegmanni Kaup, 1830 Material: RMNH 3584 from “Algeria” [no further local- ity specified] collected by “M. Wagner”. Catalogue name: “Amphisbaena wiegmannii”’. Citation: As “Amphisbaena wiegmannii” by Schlegel (in M. Wagner 1841, part III, 122, tab. VI). Remarks: Schlegel (in M. Wagner 1841) mentioned that Wagner collected several specimens but to date this is the single known preserved specimen in a reference collec- tion. He gave a very detailed description and figured the at this time largest specimen of the series (see Fig. 2). Moritz Wagner (1841) mentioned it as a rare species with a disjunct distribution and even restricted to small areas in the habitats. He collected three specimens in March 1837 at “Cap Matifu” (“eight hours” east of Algiers) un- der stones and mentioned this locality as the easternmost in Algeria. In May 1838 he collected a series under stones near Mostaganem and mentioned that this is the sole lo- cality in Algeria where the species is common. He de- scribed the habitat as a vegetationless sandy area and men- tioned May as mating season. ©ZFMK Amphibians and reptiles collected by Moritz Wagner 223 é “7, A Fis ae : “AY. COG Ti a Z J s , 4 ; ‘ 7 ¥ a 9 5 a 455 >. ce ; cist =o ees eS ae “fend > wapoense? 3 ‘ 1 4 a 3 F 7 ; mh % i bi Ly 4 . z 1 f Y Vv ¥ a $ / s } es Login / Lf VIA Aftsl Ei? SLL f leryz WtttP tl. Ciel at mal dof Mf Lovech Fig. 3. The specimen of Zrogonophis wiegmanni Kaup, 1830 figured in the rare atlas volume of Wagner (1841). Bonn zoological Bulletin 61 (2): 216-240 OZFMK 224 Philipp Wagner et al. Reptilia, Sauria [Agamidae] Paralaudakia caucasia (Kichwald, 1831) Material: ZFMK 20840-841 [ZMUG 414c] from “Tiflis” [=Tbilissi], Georgia collected by “M. Wagner” in 1846. Catalogue name:“Agama stellio’”’[ZMUG]. Citation: As “Stellio caucasius Eichwald” from “aus Grusien [Georgia] und von den Abhangen des Kaukasus [and the mounatin slopes of the Caucasus], by Berthold (in M. Wagner 1850, p. 329). Remarks: Surprisingly, Berthold’s (in M. Wagner 1850) diagnosis of the voucher was initially right but later ig- nored in the ZMUG catalogue and inventoried there as A. stellio. Berthold (in M. Wagner 1850) also only mentioned one voucher, whereas there are two specimens with the same locality in the collection. After 1890 the vouchers were stored in formalin and lost coloration, but Berthold (in M. Wagner 1850) described the relatively freshly pre- served specimens as having a ground color of dirty green to yellow with a yellow vertebral stripe, head gray and bel- ly olive-green. Lateral parts of the body with black mark- ings. Phrynocephalus persicus De Filippi, 1863 Material: ZFMK 26381 [ZMUG 154a] from “Urmia’, Iran collected by “M. Wagner” in 1846. Catalogue name: “Phrynocephalus helioscopus” |[ZMUG]. Citation: As “Phrynocephalus helioscopus Kaup” from “aus Grusien [Georgia] und von den Abhangen des Kau- kasus“ by Berthold (in M. Wagner 1850, p. 330). Remarks: Berthold (in M. Wagner 1850) mentioned two collected specimens with the remark that the species is very widespread along the coast of the Caspian Sea. [Anguidae] Pseudopus apodus (Pallas, 1775) Material: ZFMK 26394 [ZMUG 36d] from “Tiflis” [=Tbilissi], Georgia collected by “M. Wagner” in 1846. Catalogue name: “Ophisaurus apus”’ [ZMUG]. Citation: As “Pseudopus serpentinus Merrem” from “aus Grusien [Georgia] und von den Abhangen des Kaukasus” by Berthold (in M. Wagner 1850, p. 331). Remarks: Berthold (in M. Wagner 1850) mentioned two vouchers of ,Pseudopus serpentinus Merrem’ and assigned a juvenile specimen to ,Pseudopus fischeri Ménet.’. [Chamaeleonidae] Chamaeleo chamaeleon (Linnaeus, 1758) Material: RMNH 2990 from “Algeria” [no further local- ity specified] collected by “M. Wagner”. Catalogue name: “Chamaeleon africanus”. Citation: As “Chamaeleo africanus” by Schlegel (in M. Wagner 1841, part IH, 111). Remarks: Wagner (1841) mentioned that he found the chameleon only in the vicinities of Algiers, in the west- Bonn zoological Bulletin 61 (2): 216-240 erm parts of the country, where it is common in Arzew, Mostaganem and Oran, and in inland places (Mascara, Tlemsan). In the lowlands of Metidscha he found the species on shrubs, mainly on oleander (Nerium oleander). [Corytophanidae] Basiliscus basiliscus (Linnaeus, 1758) Material: ZSM 494/0 from ‘“Mittelamerika” [=Central America, no further locality specified] collected by “M. Wagner”; ZSM 495/0 from “Mittelamerika” [=Central America, no further locality specified] collected by “M. Wagner”; ZSM 507/0 from “Panama” [no further locali- ty specified] collected by “M. Wagner”. Catalogue name: “Basiliscus mitratus” [ZSM 494/0]; “Basiliscus americanus” [ZSM 495/0; ZSM 507/0]. Basiliscus vittatus Wiegmann, 1828 Material: ZSM 496/0 [3 ex.] from “Central-Amerika” [=Central America, no further locality specified] collect- ed by “? M. Wagner”. Catalogue name: “Basiliscus vittatus ”. Remarks: Even though the collector is mentioned as ques- tionable, the locality is consistent with other ZSM mate- rial collected by M. Wagner. [Dactyloidae] Anolis auratus (Daudin, 1802) Material: ZSM 484/0 from “Chiriqui,” Panama collected by “M. Wagner”; ZSM 485/0 from ‘“Mittelamerika” [= Central America, no further locality specified] collected by “M. Wagner”. Catalogue name: “Norops auratus ”’. Anolis carolinensis Voigt, 1832 Material: ZSM 467/0 [2 ex.], ZSM 468/0 [5 ex.], ZSM 469/0 [14 ex.] from “Louisiana,” USA collected by “M. Wagner”. Catalogue name: “Anolis carolinensis”. Anolis porcatus Gray, 1840 Material: ZMB 517 [3 ex.] from “Centralam[erca]” [= Central America, no further locality specified] collected by “Moritz Wagner”. Catalogue name: “Anolis porcatus ”’. Remarks: This is a Cuban endemic species and not found in Central America. However, on their sea passage back from Honduras to New York, Wagner and Scherzer trav- elled via Jamaica, Haiti, Puerto Rico, St. Thomas and Ha- vana. Therefore, it seems obvious that the locality men- tioned in the catalogue 1s incorrect. Anolis sagrei Duméril & Bibron, 1837 Material: NUW 12714 from “Costa Rica” [no further lo- cality specified] collected by “M. Wagner”. Catalogue name: “Anolis sagrei”’. ©ZFMK Amphibians and reptiles collected by Moritz Wagner 225 [Gymnophthalmidae] Bachia heteropa (Lichtenstein & von Martens, 1856) Material: ZMB 52679 [formerly ZMB 1175, paralecto- type] from “Palmas de St. Matias“ [=San Matias, Antio- quia, Colombia? fide Bauer & Giinther 1994] (label on the jar only reads “Venezuela’) collected by “Wagner”. Catalogue name: “Chalcides heteropus’’, “Cophias hetero- pus”. Citation: As “Chalcides heteropus”’ by Lichtenstein & von Martens (1856). Pholidobolus affinis (Peters, 1863) Material: ZSM 644/0/1 [holotype] from “Pichincha,” Ecuador, collected by “M. Wagner”; ZSM 644/0/2-5 [3 adults, | subadult, ?paratypes], all with the same data as the holotype. Catalogue name: “Ecpleopus affinis (Ptrs.) — aspidolae- mus Pet.”’. Citation: As “Ecpleopus affinis”’ by Peters (1863). Remarks: According to Franzen & Glaw (2007) ZSM 644/0/1 agrees with Peters’ description of Ecpleopus affi- nis and is without doubt the holotype. The complete se- ries consists of five specimens (see above) and Franzen & Glaw (2007) recognized these specimens as question- able paratypes. [Hoplocercidae] Enyalioides laticeps (Guichenot, 1855) Material: ZSM 499/0 from “Pastassathal” [=Pastaza val- ley], Ecuador collected by “M. Wagner”. Catalogue name: “Enyalioides laticeps”’. Enyalioides praestabilis (O’Shaughnessy, 1881) Material: ZSM 500/0 from “Pastassathal Tal” [=Pastaza valley], Ecuador collected by “M. Wagner”. Catalogue name: “Enyalioides praestabilis ”. [Iguanidae] Ctenosaura similis (Gray, 1831) Material: NMW 13090 from “Nicoya, Costa Rica” col- lected by “M. Wagner”. Catalogue name: “Ctenosaura similis”. Iguana iguana (Linnaeus, 1758) Material: ZSM 539/0 from “Panama” [no further locali- ty specified] collected by “M. Wagner”; ZSM 541/0 from “Mittelamerika” [=Central America, no further locality specified] collected by “M. Wagner”. Catalogue name: “/guana tuberculata”’. [Lacertidae] Acanthodactylus savignyi Audouin, 1809 Material: RMNH 3496 from “Algeria” [no further local- ity specified] collected by “M. Wagner”. Catalogue name: “Lacerta savignyi”’. Bonn zoological Bulletin 61 (2): 216-240 Citation: None. Remarks: This species is not mentioned by Schlegel (in Wagner 1841), but the specimen probably refer to the men- tioned “Lacerta pardalis Lichtenstein 1823”. Especially as Schlegel (in Wagner 1841, page 115) mentioned that he had many problems to identify this species and refer to it only because of its uncertain identidy. However, Schlegel (in Wagner 1841) also mentioned a series of spec- imens collected by Moritz Wagner but the whereabouts of other specimens remains unclear. Darevskia praticola (Eversmann, 1834) Material: ZFMK 21330 [ZMUG 21f] from “Tiflis” [=Tbilissi], Georgia collected by “M. Wagner” in 1847. Catalogue name: “Lacerta muralis var. chalybdea” [ZMUG]. Citation: As “Lacerta muralis Latr.” from “aus Grusien [Georgia] und von den Abhangen des Kaukasus” by Berthold (in M. Wagner 1850, p. 331). Remarks: Berthold (in M. Wagner 1850) specified two specimens of the species but did not recognize a subspe- cific level. Most probably this is one of the mentioned vouchers. There are only two other “Lacerta” species men- tioned: “Lacerta oxycephala” (see below) and Lacerta stri- gata, which are very different from Darevskia. Mesalina guttulata (Lichtenstein, 1823) Material: RMNH 3436 from “Oran, Algeria” collected by “M. Wagner”. Catalogue name: “Lacerta guttata’’. Citation: As “Lacerta guttulata” by Schlegel (in M. Wag- ner 1841, part IH, 113). Remarks: Schlegel (in M. Wagner 1841) mentioned two specimens collected by Wagner which was the first record of this species for Algeria. The other specimen could be the G6ttingen specimen, which is apparently lost (see be- low). Parvilacerta parva (Boulenger, 1887) Material: ZFMK 21229 [ZMUG 21g] from “Urmia,” Iran collected by “M. Wagner” in 1847. Catalogue name: “Lacerta muralis” [ZMUG]. Citation: As “Lacerta muralis Latr.” from “aus Grusien [Georgia] und von den Abhangen des Kaukasus” by Berthold (in M. Wagner 1850, p. 331). Remarks: Berthold (in M. Wagner 1850) mentioned two vouchers, but only one was located by us (see also remarks in Darevskia praticola). However, this species is not known from Lake Urmia today, but occurs in the Ararat region in Armenia which was also visited by Wagner. Podarcis tauricus (Pallas, 1814) Material: ZFMK 26521 [ZMUG 239a], ZFMK 21225 [ZMUG 21b], ZFMK 26522 [ZMUG 247d] from “Con- stantinople” [=Istanbul], Turkey collected by “M. Wag- ©ZFMK 226 Philipp Wagner et al. ner” collected by “M. Wagner” in “1847 [ZFMK 21225, ZFMK 26522)”. Catalogue name: “Lacerta taurica” [ZMUG 239a]; “Lac- erta muralis’ [ZMUG 21b]; “Lacerta oxycephala” [ZMUG 247d]. Citation: As “Lacerta oxycephala Schlegel” from “aus Grusien [Georgia] und von den Abhangen des Kaukasus” by Berthold (in Wagner 1850, p. 331); As “Lacerta tau- rica Pallas” from “von der tiirkisch-pontischen Kiiste” by Berthold (in Wagner 1850, p. 339). Remarks: Berthold (in Wagner 1850) mentioned two vouchers of this species from the ‘tiirkisch-pontischer Kiiste’ (= southern coastal parts of the Black Sea), which likely resemble these specimens. Berthold (in Wagner 1850) pointed out that the voucher of L. oxycephala 1s a juvenile specimen with a snout-vent length of “3.6 zoll” [=inch] and a tail length of “2.4 zoll”. Podarcis vaucheri (Boulenger, 1905) Material: RMNH 3465 from “Algeria” [no further local- ity specified] collected by “M. Wagner”. Catalogue name: “Lacerta muralis”. Citation: As “Lacerta muralis” by Schlegel (in M. Wag- ner 1841, part III, 113). Remarks: Schlegel (in M. Wagner 1841) mentioned the species as common in coastal areas, but Wagner also en- countered it in inland Algeria. Scelarcis perspicillata (Duméril & Bibron, 1839) Material: RMNH 3827 from “Algeria” [no further local- ity specified] collected by “M. Wagner”. Catalogue name: “Lacerta perspicillata”’. Remarks: This species is not mentioned by Schlegel (in M. Wagner 1841). Timon pater (Lataste, 1880) Material: RMNH 3501 from “Algeria” [no further local- ity specified] collected by “M. Wagner”’. Catalogue name: “Lacerta ocellata”’. Citation: As “Lacerta ocellata” by Schlegel (in M. Wag- ner 1841, part HI, 112). Remarks: Wagner (1841) mentioned “Lacerta ocellata” as common along the coast between Oran and Bona, but also mentioned it from Belida, Constantine and Mascara in inland Algeria. [Phrynosomatidae] Sceloporus acanthinus Bocourt, 1873 Material: ZSM 551/0 from “Mittel-Amerika” [=Central America, no further locality specified] collected by “M. Wagner”. Catalogue name: “Sceloporus acanthinus ”’. Sceloporus grammicus Wiegmann, 1828 Material: ZFMK 26788 [ZMUG 107a] from “Nord Ameri- Bonn zoological Bulletin 61 (2): 216-240 ka” [=North America, no further locality specified] col- lected by “M. Wagner” in 1853. Catalogue name: “Sceloporus grammicus” [ZMUG]. Sceloporus undulatus (Bosc & Daudin, 1801) Material: ZSM 554/0 [2 ex.] from “Louisiana,” USA col- lected by “M. Wagner”. Catalogue name: “Sceloporus undulatus”. [Phyllodactylidae] Tarentola mauritanica (Linnaeus, 1758) Material: RMNH 2742 from “Algeria” [no further local- ity specified] collected by “M. Wagner”. Catalogue name: “Platydactylus fascicularis ”’. Citation: As “Platydactylus fascicularis” by Schlegel (in M. Wagner 1841, part III, 108). Remarks: Wagner (1841) mentioned the gecko as common between Bona and Oran. He describes olive trees as the preferred habitat where the specimens are found on the bark of old trees. [Scincidae] Chalcides ocellatus (Forsskal, 1775) Material: RMNH 2500 from “Algeria” [no further local- ity specified] collected by “M. Wagner”. Catalogue name: “Scincus ocellatus”. Citation: As “Scincus ocellatus” by Schlegel (in M. Wag- ner 1841, part II, 120). Remarks: Wagner (1841) mentioned the species was com- mon in coastal areas between Bona and Oran (most com- monly in the area around Mostaganem), but he found the specimens only between April and October. He described the habitat as a sandy area and found the specimens un- der stones and rocks. Chalcides chalcides (Linnaeus, 1758) Material: RMNH 3667 from “Algeria” [no further local- ity specified] collected by “M. Wagner”. Catalogue name: “Scincus chalcides”. Citation: As “Seps chalcides” by Schlegel (in M. Wagn- er 1841, part IH, 121). Remarks: Wagner (1841) mentioned the species as com- mon in the vicinities of Algier, Bona and Oran, where he found the individuals in hollows but not under stones. Plestiodon fasciatus (Linnaeus, 1758) Material: ZSM 831/0 [5 ex.] from “Louisiana,” USA col- lected by “M. Wagner”; ZSM 2564/0 from “Florida,” USA collected by “M. Wagner”. Catalogue name: “Eumeces fasciatus”. Scincella lateralis (Say, 1823) Material: ZSM 805/0 [4 ex.] from “Florida,” USA collect- ed by “Wagner”. Catalogue name: “Leiolepisma laterale”. ©ZFMK Amphibians and reptiles collected by Moritz Wagner [Tropiduridae] Stenocercus guentheri (Boulenger, 1885) Material: ZSM 522/0 [3 ex.] from “Pichincha” [=Pichin- cha Mountain near Quito], Ecuador collected by “M. Wag- Neher. Catalogue name: “Liocephalus guentheri”’. Reptilia, Serpentes [Boidae| Corallus hortulanus (Linnaeus, 1758) Material: ZSM 1362/0 from “Mittel-Amerika” [=Central America, no further locality specified] collected by “M. Wagner’. Catalogue name: ,,Corallus cookei” [Colubridae| Atractus guentheri (Wucherer, 1861) Material: ZSM 1949/0 from “Ecuador” [no further local- ity specified] collected by “M. Wagner”. Catalogue name: ,,Atractus guentheri” Coluber constrictor Linnaeus, 1758 Material: ZFMK 31702 [ZMUG 110b] from “Nord Ameri- ka” [=North America, no further locality specified] col- lected by “M. Wagner” in 1854. Catalogue name: “Zamenis constrictor” [ZMUG]. Remarks: In contrast to the other material of Wagner from North America, this voucher was collected or catalogued in 1854. Coluber flagellum Shaw, 1802 Material: NUW 26579 from “Costa Rica” [no further lo- cality specified] collected by “M. Wagner”. Catalogue name: “Masticophis flagellum”. Remarks: If the specimen is correctly identified, it would be the first record for Costa Rica, which is very unlikely. Therefore, the specimen should re-identified. Coniophanes fissidens (Ginther, 1858) Material: NMW 27357 from “Costa Rica” [no further lo- cality specified] collected by “M. Wagner”. Catalogue name: “Coniophanes fissidens ”’. Dendrophidion dendrophis (Schlegel, 1837) Material: ZSM 1616/0 [2 ex.] from “Ecuador” [no further locality specified] collected by “M. Wagner”.. Catalogue name: ,,/schnognathus occipitomaculatus” Diaphorolepis wagneri Jan, 1863 Material: ZSM 2708/0 [holotype] from “Andes de l’E- cuador” [= Andes in Ecuador, no further locality speci- fied] without further data. Catalogue name: “Diaphorolepis wagneri”’. Citation: As “Diaphorolepis wagneri” by Jan (1863, p. 98). Bonn zoological Bulletin 61 (2): 216-240 DOS Remarks: Even that no collector is indicated, the locality and the fact that the species is named after Moritz Wag- ner suggest he was the collector. Dipsas andiana (Boulenger, 1896) Material: ZSM 2278/0 from “Ecuador” [no further local- ity specified] collected by “M. Wagner”. Catalogue name: ,,Sibynomorphus variegatus”’. Eirenis modestus Martin, 1838 Material: ZFMK 31625 [ZMUG 234a] from “Tiflis” [=Tbilissi], Georgia collected by “M. Wagner” in 1846. Catalogue name: “Ablabes modestus”’ [ZMUG]. Citation: As “Coronella collaris Berth.” from “aus Grusien [Georgia] und von den Abhangen des Kaukasus” by Berthold (in M. Wagner 1850, p. 332). Remarks: Berthold (in M. Wagner 1850) was the first au- thor who assigned ,Coluber collaris’ as a species of the genus Coronella, but nevertheless, his identification of the specimen was wrong. He emended his own identification in the catalogue where the specimen is inventoried as Ablabes modestus. He described the pholidosis of the voucher as having 17 rows of rhomboidal smooth scales around midbody. Originally it was a series of three spec- imens but two of them are now lost. Farancia abacura (Holbrook, 1836) Material: ZMB 1778 from “Centralamer[ika]” [Central America, no further locality specified] collected by “Wa- gener” [sic!]. Catalogue name: “Calopisma abacurum”’. Remarks: According to the ZMB catalogue this specimen was collected by “Wagener”, but the locality and the time of the inventory suggests that this refers to Moritz Wag- ner, whose name was apparently misspelled. The locali- ty does not match the currently known distribution (east- ern USA) of the species but probably Wagner recognized eastern Texas rather as Central America than as North America and therefore the locality could be correct and can be restricted to this region. Farancia erythrogramma (Palissot de Beauvois, 1802) Material: ZSM 1905/0 from “Siid-Carolina [= South Car- olina],” USA collected by “M. Wagner”. Catalogue name: ,,Abastor erythrogrammus”. Hemorrhois hippocrepis (Linnaeus, 1758) Material: RMNH 471 [2 ex.] from “Algeria” [no further locality specified] collected by “M. Wagner” in 1839. Catalogue name: “Coluber hippocrepis ”’. Citation: As “Coluber hippocrepis” by Schlegel (in M. Wagner 1841, part III, 130). Remarks: According to Wagner (1841) the species is com- mon in southern Algeria where it is often found in hous- es. OZFMK Imantodes cenchoa (Linnaeus, 1758) Material: ZSM 30/0 from ‘“Mittelamerika” [=Central America, no further locality specified] collected by “M. Wagner”; ZSM 2000/0 from “Pastassathal [=Pataza val- ley],” Ecuador collected by “M. Wagner”. Catalogue name: “Elaps ancoralis” |ZSM 30/0]; ,,Jman- todes chenchoa” [sic!| [ZSM 2000/0]. Imantodes gemmistratus (Cope, 1861) Material: NM W 25528 from “Costa Rica” [no further lo- cality specified] collected by “M. Wagner”. Catalogue name: “/mantodes gemmistratus”’. Lampropeltis getula (Linnaeus, 1766) Material: ZSM 1839/0 from “Florida,” USA collected by “M. Wagner”. Catalogue name: ,,Lampropeltis getula var. sayi’’. Lampropeltis holbrooki Stejneger, 1902 Material: ZFMK 36445 [ZMUG 96c] from “Nord Ameri- ka” [North America, no further locality specified] collect- ed by “M. Wagner” in 1853. Catalogue name: “Coronella getula” |ZMUG]. Remarks: The species is only recognized from west of the Mississippi which restricts the locality to this area of North America.. Lampropeltis triangulum Lacépéde, 1789 Material: ZFMK 36837 [ZMUG 89b] from “Tennessee,” USA collected in 1853. Catalogue name: “Coronella triangulum” [ZMUG]. Leptodeira annulata (Linnaeus, 1758) Material: ZSM 2017/0, ZSM 2024/0 [4 ex.], ZSM 2025/0 from “Mittel-Amerika [= Central America, no further lo- cality specified]” collected by “M. Wagner”. Catalogue name: “Leptodeira annulata albofusca” |ZSM 2017/0]; ,.Leptodeira annulata personata” |ZSM 2024/0, ZSM 2025/0]. Macroprotodon cucullatus (Geoffroy de St-Hilaire, 1827) Material: RMNH 212 [2 ex.] from “Alges” [=Algeria, no further locality specified] collected by “M. Wagner”. Catalogue name: “Coronella mauritanica ”’. Citation: ? As “Coronella laevis” by Schlegel (in M. Wag- ner 1841, part III, 129). Remarks: The only Coronella which is mentioned by Schlegel (in M. Wagner 1841) is C. /aevis from Algier, Bona and Constantine. Therefore, these specimens could be identical with the cited ones. Mastigodryas boddaerti (Seetzen, 1796) Material: NMW 18744 from “Costa Rica” [no further lo- cality specified] collected by “M. Wagner”; ZSM 83/0 Bonn zoological Bulletin 61 (2): 216-240 228 Philipp Wagner et al. from ““Mittelamerika” [= Central America, no further lo- cality specified] collected by “M. Wagner”. Catalogue name: “Mastigodryas boddaerti” [NMW 18744]; “Drymobius boddaerti” [ZSM 83/0]. Nerodia erythrogaster Forster, 1771 Material: ZFMK 33472 [ZMUG 75d] from “Nord Ameri- ka [North America], Georgia”, USA collected in 1853. Catalogue name: “T7ropidonotus fasciatus var. erythro- gaster” [ZMUG]. Remarks: No collector specified but the year and locali- ty are consistent with Moritz Wagner being the collector. Nerodia fasciata (Linnaeus, 1766) Material: ZFMK 33474-478 [ZMUG 75a, 75b] from “Nord America” [=North America, no further locality specified] collected by “M. Wagner” in 1853; ZFMK 33479 [ZMUG 75c] from “Nord Amerika, Tennessee,” USA collected in 1853. Catalogue name: “Zropidonotus fasciatus” [ZMUG 75a, 75b]; “Tropidonotus fasciatus” [ZMUG 75c]. Remarks: No collector is specified with the specimen ZFMK 33479 but the year and locality are consistent with Moritz Wagner being the collector. Nerodia taxispilota (Holbrook, 1838) Material: ZSM 1475/0 from “Florida,” USA collected by “M. Wagner”. Catalogue name: ,,Natrix taxispilotus” Oxybelis aeneus (Wagler, 1824) Material: NMW 27266 from “Costa Rica” [no further lo- cality specified] collected by “M. Wagner”. Catalogue name: “Oxybelis aeneus”. Opheodrys aestivus (Linnaeus, 1766) Material: ZSM 1919/0 from “Siid-Carolina [= South Car- olina],” USA collected by “M. Wagner”. Catalogue name: ,,Contia aestiva’, ,,Opheodrys aestiva’. Pituophis catenifer sayi (Schlegel, 1837) Material: ZMB 2087 from “Nordamerika” [North Amer- ica, no further locality specified] collected by “Wagener” [sic! ]. Catalogue name: “Coronella sayi”’. Remarks: According to the ZMB catalogue this specimen was collected by “Wagener”, but the locality and the time of the inventory suggests that this refers to Moritz Wag- ner, whose name was apparently misspelled. Platyceps sp. Material: ZFMK 31706 [ZMUG 104e] from “Tiflis” [=Tbilissi] collected by “M. Wagner” in 1846. Catalogue name:“Zamenis gemonensis* |[ZMUG]. Citation: As “Coluber viridiflavus Lacépéde” from “aus ©ZFMK Amphibians and reptiles collected by Moritz Wagner 229 Grusien [Georgia] und von den Abhangen des Kaukasus* by Berthold (in M. Wagner 1850, p. 334). Remarks: Berthold (in M. Wagner 1850) mentioned a ju- venile ,Coluber viridiflavus’, which is most probably iden- tical with the specimen Zamenis gemonensis in the cata- logue of the Zoologisches Museum (ZMUG). He de- scribed his voucher as having 15 rows of smooth scales around midbody, 171 ventralia and 102 subcaudalia. It has so far not been possible to identify this specimen at the species level, but it is most similar and probably identi- cal with P. najadum. Regina regida (Say, 1825) Material: ZSM 1471/0 from “Florida, Amerika [=USA]” collected by “M. Wagner”. Catalogue name: ,,Natrix rigidus” Regina septemvittata (Say, 1825) Material: ZFMK 33468 [ZMUG 77] from “Nord Ameri- ka [=North America], Georgia”, USA collected in 1853; ZSM 1470/0 from “Florida, Amerika [=USA]” collected by “M. Wagner”. Catalogue name: “7ropidonotus septemvittatus” [ZMUG 77), ,.Natrix septemvittatus” [ZSM 1470/0]. Remarks: In the ZFMK specimen no collector is speci- fied but the year and locality are consistent with Moritz Wagner being the collector. Storeria dekayi (Holbrook, 1836) Material: ZSM 1530/0 from “Florida,” USA collected by “M. Wagner”; ZFMK 33518 [ZMUG 76] from “Nord Amerika” [=North America, no further locality specified] collected by “M. Wagner” in 1853. Catalogue name: ,,/schnognathus dekayi” [ZSM]; “Tropi- donotus cyclopium {[sic!]|” [ZMUG]. Telescopus fallax iberus Eichwald, 1831 Material: ZFMK 31731 [ZMUG 47a] from “Tiflis” [=Tbilissi], Georgia collected by “M. Wagner” in 1846. Catalogue name: “Zarbophis savignyi” [ZMUG]. Citation: As “Dipsas fallax Schlegel” from “Tiflis” by Berthold (in M. Wagner 1850, p. 334); as “7rigonophis iberus Eichwald” from “Tiflis” by Berthold (in M. Wag- ner 1850, p. 334). Remarks: Berthold (in M. Wagner 1850) specified the voucher as ‘Dipsas fallax Schlegel’ but also mentioned ,Trigonophis iberus’ as synonym. It can be assumed that this voucher is identical with the one described by Berthold. As pholidosis he mentioned 13 rows of smooth scales around midbody, 235 ventral- and 66 subcaudal scales. Additionally he described the voucher as: body de- pressed; head broader than neck; pupil vertical; head scales short; occiput scales rounded and short; two postocular and no praeocular scale; body scales lanceolate, small and smooth. The coloration was cinereous with dark oblique Bonn zoological Bulletin 61 (2): 216-240 markings, bordered upwards by pale dots, belly marbled blue. Thamnophis sauritus (Linnaeus, 1758) Material: ZSM 1416/0 [3 ex.], ZSM 2596/0 from “Flori- da,” USA collected by “M. Wagner”. Catalogue name: ,,7ropidonotus saurita” |ZSM 1416/0]; »Lhamnophis sauritus sackenii” [ZSM 2596/0]. Tropidodipsas fasciata Giinther, 1858 Material: NUW 19872 from “Nicaragua” [no further lo- cality specified] collected by “M. Wagner”. Catalogue name: “Sibon fasciata”’. Xenodon rabdocephalus (Wied, 1824) Material: ZSM 1818/0 from “Mittel-Amerika” [=Central America, no further locality specified] collected by “M. Wagner”. Catalogue name: ,,Xenodon colubrinus”’. [Elapidae] Micrurus latifasciatus Schmidt, 1933 Material: ZSM 2263/0 [paratype] from ‘““Nordamerika” [=North America, no further locality specified] collected by “M. Wagner”. Catalogue name: “Micrurus latifasciatus”’. Citation: As “Micrurus latifasciatus””’ by Schmidt (1933). Remarks: The locality is mentioned as erroneous in Franzen & Glaw 2007. Micrurus multifasciatus hertwigi (Werner, 1897) Material: ZSM 2268/0 [holotype] from “Mittelamerika” [=Central America, no further locality specified] collect- ed by “M. Wagner”. Catalogue name: “Elaps hertwigii.” Citation: As “Elaps hertwigii” by Werner (1897b). Pelamis platura (Linnaeus, 1766) Material: ZSM 2191/0 from “Panama” [no further local- ity specified] collected by “M. Wagner”. Catalogue name: ,,Hydrus platurus” {[Psammophiidae| Malpolon monspessulanus (Herrman, 1804) Material: RMNH 730 from “Algeria” [no further locali- ty specified] collected by “M. Wagner”. Catalogue name: “Psammophis lacertina”. Citation: As “Psammophis lacertina” by Schlegel (in M. Wagner 1841, part III, 131). [Viperidae] Agkistrodon contortrix (Linnaeus, 1766) Material: ZFMK 32527-528 [ZMUG 219c] from “Nord Amerika” [=North America, no further locality specified] collected by “M. Wagner” in 1853. Catalogue name: “Ancistrodon lebetina” [ZMUG]. ©OZFMK 230 Philipp Wagner et al. Fig. 4. Holotype of Vipera wagneri Nilson & Andrén, 1984 collected by Moritz Wagner. Bonn zoological Bulletin 61 (2): 216-240 ©ZFMK Amphibians and reptiles collected by Moritz Wagner 23] Agkistrodon piscivorus (Lacépéde, 1789) Material: ZSM 2327/0 from “Florida,” USA collected by “M. Wagner”. Catalogue name: ,,Agkistrodon piscivorus” Bothriechis schlegelii (Berthold, 1846) Material: NMW 28002 from “Costa Rica” [no further lo- cality specified] collected by “M. Wagner”; ZSM 2363/0 from “Panama” [no further locality specified] collected by “M. Wagner”. Catalogue name: “Bothriechis schlegelii”” |NMW 28002}; Bothrops schegeli” [ZMS 2363/0]. Bothrops lanceolatus (Bonnaterre, 1790) Material: ZSM 2339/0 from “Mittel-Amerika [= Central America, no further locality specified],”collected by “M. Wagner”. Catalogue name: ,,Bothrops lanceolatus”’. Montivipera wagneri (Nilson & Andrén, 1984) Material: ZFMK 32495 [ZMUG 233a] [holotype], “Ur- mia,’ Iran collected by “M. Wagner” in 1846. Catalogue name: “Vipera lebetina” [ZMUG]. Citation: As “Vipera aspis Merr[em] (Varietas: V. occel- lata, Latr[eile])” from “von der armenisch-persischen Grenze” [= from the border between Armenia and Iran] by Berthold (in M. Wagner 1850, p. 337); as “Vipera xan- thina” by Strauch (1869); as “Vipera wagneri” by Nilson & Andrén (1984). Remarks: As mentioned above, Berthold (in M. Wagner 1850: 337) mentioned this specimen as variety ocellata of Vipera aspis, but later, he inventoried it as V. /ebetina. It was later seen by Strauch (1869) and who cited it as if Berthold was the author of the taxonomic name ocellata, and identified it as synonym of V. xanthina. Nilson & An- dren (1984) were therefore correct to describe it as a new species, Vipera wagneri, closely related to V. xanthina. Later, this group was accommodated in its own genus Montivipera (Nilson et al. 1999). Nevertheless, the de- scription of the pholidosis by Berthold (in M. Wagner 1850) is not identical with the description of the holotype (Fig. 4), but it must be assumed that the mentioned spec- imens are identical. From the recent distribution of the species it seems obvious that “Urmia” as the locality men- tioned in the catalogue is incorrect and the locality pub- lished by Berthold (in M. Wagner 1850) “von der ar- menisch-persischen Grenze” seems to be more likely. Porthidium lansbergi (Schlegel, 1841) Material: NMW 28392 from “Costa Rica” [no further lo- cality specified] collected by “M. Wagner”. Catalogue name: “Porthidium lansbergi”’. Citation: None. Bonn zoological Bulletin 61 (2): 216-240 Porthidium nasutum (Bocourt, 1868) Material: NVW 28398 from “Costa Rica,” [no further lo- cality specified] collected by “M. Wagner”. Catalogue name: “Porthidium nasutum””’. Material lost or apparently lost at the ZSM & ZMUG col- lections Specimens are listed under their current name based on the last catalogue entry. In many cases the mentioned ma- terial must be recognized as lost, but some material, par- ticularly from the ZSM collection, may still be extant and this should be clarified in the future. The reason for the loss of specimens in the ZMUG collection is unknown but was documented during two inventories of the collection in the 1960s and 1970s. The material in the ZSM collec- tion was lost because of bombing during the Second World War, but it is possible that some of the specimens are still available. Amphibia Amphibia, Caudata [Ambystomidae] Ambystoma jeffersonianum (Green, 1827) Material: ZMUG 4la [lost] from “Nord Amerika” [=North America, no further locality specified] collected in 1856. Catalogue name: “Ambystoma jeffersonianum”. Remarks: At the second inventory of the ZMUG collec- tion in 1970 the specimen was present but it seems to have been lost subsequently. Ambystoma tigrinum (Green, 1825) Material: ZMUG 11b, 11g [2 ex.] [lost] from ‘Nord Amerika” [=North America, no further locality specified] collected in 1856. Catalogue name: “Ambystoma tigrinum”’. Remarks: For neither voucher is a collector mentioned but the date and locality are consistent with M. Wagner. At the second inventory of the ZMUG collection in 1970 the specimen was still present but it is now apparently lost. [Plethodontidae] Bolitoglossa palmata (Werner, 1897) Material: ZSM 1272/0 [2 ex, paralectotypes] [lost] from “Ecuador” [no further locality specified] collected by “M. Wagner”. Catalogue name: “Oedipus palmatus” Citation: As “Spelerpes palmatus” by Werner (1897b). Remarks: NMW 22862 was designated as lectotype by Brame & Wake (1962: 173). ©ZFMK tO Ww No Eurycea longicauda (Green, 1818) Material: ZMUG 4a [lost] from “Tennessee, North Amer- ica” collected in 1853. Catalogue name: “Spelerpes longicauda’’. Remarks: In the catalogue determined without a collec- tor but date and locality are consistent with other materi- al collected by M. Wagner. Plethodon cinereus (Green, 1818) Material: ZSM 1276/0 [8 ex., apparently lost] from “Wis- consin,” USA collected by “M. Wagner”. Catalogue name: “Plethodon erythronotus”; “Plethodon cinereus’. Pseudotriton montanus Baird, 1850 Material: ZMUG 7d [lost] from “Tennessee,” USA col- lected in 1853. Catalogue name: “Spelerpes ruber var. montana”. Remarks: In the catalogue no collector is added but date and place refer to M. Wagner. At the time of the second inventory of the collection in 1970 the voucher was still present but it has been lost subsequently. Pseudotriton ruber (Sonnini de Manoncourt and La- treille, 1801) Material: ZSM 1264/0 [apparently lost] from ‘“Nord- Amerika” [= North America, no further locality specified] collected by “M. Wagner”. Catalogue name: “Pseudotriton ruber”. {[Salamandridae| Notophthalmus viridescens (Rafinesque, 1820) Material: ZSM 1239/0 [4 ex, lost], from “Wisconsin,” USA collected by “M. Wagner”. Catalogue name: “T7riturus viridescens”’. Triturus karelini Strauch, 1870 Material: ZMUG 16d [lost] from “Constantinopel” [=Is- tanbul], Turkey collected by “M. Wagner”. Catalogue name: “7riturus cristatus”’. Remarks: Berthold (in Wagner 1850) reported ‘7riton tae- niatus’ and ,Triton nycthemerus’ from the ‘tiirkisch-pon- tischen Kiiste’ (southern coastal parts of the Black Sea). However, no vouchers are preserved and it is not possi- ble to determine if this was additional material or a cor- rect identified voucher. But ‘cf. Woltersdorff [sic!] 1907’ is noted handwritten in the catalogue, so it is most prob- ably a correctly identified specimen. At the time of the sec- ond inventory of the ZMUG collection in 1970 the vouch- er was still present but it has since been lost. Amphibia, Anura Hylodes spec. Material: ZSM 1048/0 [status unclear] from “Pastassathal [Pastaza valley],” Ecuador” collected by ““M. Wagner”. Bonn zoological Bulletin 61 (2): 216-240 Philipp Wagner et al. Catalogue name: “Hylodes spec.”. Citation: None. Remarks: Today the genus Hylodes Fitzinger 1826 in- cludes 24 species, but non of them occur in Ecuador. Therefore, the specimen identity remain unclear. [Bufonidae] Atelopus ignescens (Cornalia, 1849) Material: ZSM 1014/0 [7 ex., apparently lost] from “Pich- incha,” Ecuador collected by “M. Wagner”. Catalogue name: “Atelopus laevis”. Atelopus longirostris (Cope, 1868) Material: ZSM 1018/0 [apparently lost] from “Ecuador” [no further locality specified] collected by “M. Wagner”. Catalogue name: “Atelopus longirostris”. Atelopus varius (Lichtenstein & Martens, 1856) Material: ZSM 1016/0 [apparently lost] from “Ecuador” [no further locality specified] collected by “M. Wagner”. Catalogue name: “Atelopus varius”. Bufo variabilis Pallas, 1769 Material: ZMUG 65ba [lost] from “Tiflis” [=Tbilissi], Georgia collected by “M. Wagner” in 1846. Catalogue name: “Bufo virdis”. Remarks: Berthold (in M. Wagner 1850) indeed mentioned one juvenile voucher of this taxon but with the locality ‘ttirkisch-pontischer Kiiste’ (= southern coastal parts of the Black Sea), but as recognized from present vouchers, the catalogue locality is often more precise than the locality given by Berthold (in M. Wagner 1850). Rhinella crucifer (Wied-Neuwied, 1821) Material: ZSM 2653/0 [apparently lost] from “Mittel- Amerika [=Central America, no further locality speci- fied],” collected by “M. Wagner”. Catalogue name: “Bufo crucifer’’. [Dendrobatidae] Dendrobates tinctorius (Cuvier, 1797) Material: ZSM 1010/0 [apparently lost] from “Chiriqui, Central Am[erika] [=Panama]” collected by “M. Wagn- Clan Catalogue name: “Dendrobates tinctorius”’. {[Hemiphractidae] Gastrotheca marsupiata (Duméril & Bibron, 1841) Material: ZSM 1187/0 [3 ex., apparently lost] from “Pich- incha,” Ecuador collected by “M. Wagner”. Catalogue name: “Gastrotheca marsupiata”. [Hylidae] Hyla arborea (Linnaeus, 1758) Material: ZMUG 88a [lost] from “Algier,“ [=Algeria, no further locality specified] collected by “M. Wagner”. ©ZFMK Amphibians and reptiles collected by Moritz Wagner 233 Catalogue name: “Hyla arborea”. Citation: As “Hyla arborea™ by Schlegel (in M. Wagner 1841, p. III, 133). Hyla cinerea (Schneider, 1799) Material: ZSM 1165/0 [apparently lost] from “Louisiana,” USA collected by “M. Wagner”. Catalogue name: “Hyla carolinensis”. Hypsiboas albomarginatus (Spix, 1824) Material: ZSM 2500/0 [apparently lost] from ‘“‘Pastassathal [Pastaza valley], Ecuador collected by “M. Wagner”. Catalogue name: “Hyla albomarginata”’. Hypsiboas geographicus (Spix, 1824) Material: ZSM 1154/0 [apparently lost] from “Ecuador” [no further locality specified] collected by “M. Wagner”. Catalogue name: “Hy/a aff. cryptomelas”. Trachycephalus typhonius (Linnaeus, 1758) Material: ZSM 1148/0 [apparently lost] from “Ecuador” [no further locality specified] collected by “M. Wagner”; ZSM 2492/0 [3 ex., ? apparently lost] from “Panama” col- lected by “M. Wagner”. Catalogue name: “Bufo typhonius”’. [Leptodactylidae] Leptodactylus wagneri (Peters, 1862) Material: ZSM 1080/0 [holotype] [lost] from “Pastas- sathal” [=Pastaza valley], Ecuador collected by “M. Wag- netee Catalogue name: “Plectromantis wagneri” Citation: As “Plectromantis wagneri” by Peters (1862). Remarks: According to the original description, the type locality is “an der Westseite der Anden in Ecuador” [= on the western side of the Andes in Ecuador]. [Microhylidae] Gastrophryne carolinensis (Holbrook, 1835) Material: ZSM 1024/0 [apparently lost] from “Wisconsin,” USA collected by “M. Wagner”. Catalogue name: “Engystoma carolinense’’. Remarks: This species does not occur in Wisconsin. [Ranidae] Lithobates pipiens (Schreber, 1782) Material: ZSM 984/0 [2 ex., apparently lost] from “Wis- consin,” USA collected by “M. Wagner”. Catalogue name: “Rana halecina brachycephala’; “Rana pipiens” Pelophylax k\. esculentus (Linnaeus, 1758) Material: ZMUG 13c [lost] from “Urmia,” Iran collected by “M. Wagner” in 1846. Catalogue name: “Rana esculenta”. Bonn zoological Bulletin 61 (2): 216-240 Citation: As “Rana esculenta Linnaeus” by Berthold (in M. Wagner 1850, p. 337); as “Rana tigrina Eichwald non R. tigrina Daudin” by Berthold (in M. Wagner 1850, p. 337). Remarks: Berthold (in M. Wagner 1850) gives a detailed description of the voucher and mentioned that, apart from the coloration, the specimen was typical of Rana esculen- ta. At the same time he synoymized Rana caucasica Pal- las, 1814 [currently a synonym of Pelophylax ridibundus (Pallas, 1771)] with Rana esculenta and noted that this tax- on is a Rana s. str. and not a taxon within the genus Bu- fo as mentioned by Menéetriés (1832) and Eichwald (1831). At the second inventory of the ZMUG collection in 1970 the voucher was still present but it was subsequently lost. [Scaphiopodidae] Scaphiopus holbrookii Harlan, 1835 Material: ZMUG 39a [lost] from “Georgia,” USA collect- ed in 1853. Catalogue name: “Scaphiopus solitarius”. Remarks: In the ZMUG catalogue no collector is listed, but the date and locality are consistent with other mate- rial collected by M. Wagner. [Strabomantidae] Pristimantis chalceus (Peters, 1873) Material: ZSM 1045/0 [2 ex., syntypes] [lost] from “Pas- tassathal [=Pastaza valley, Ecuador]” collected by “M. Wagner”. Catalogue name: “Phyllobatus chalceus”. Citation: As “Phyllobates chalceus” by Peters (1873); as “Eleutherodactylus chalceus” by Glaw & Franzen (2006). Remarks: In the original description three (type) speci- mens are mentioned, but according to Glaw & Franzen (2006) only two are listed in the ZSM catalogue. The third specimen is extant in Berlin as ZMB 7814 (Bauer et al. 1995: 46). Reptilia, Testudinae [Emydidae] Malaclemys terrapin pileata (Wied, 1865) Material: ZSM 2508/0 [apparently lost] from “Nord- Amerika” [= North America, no further locality specified] collected by “M. Wagner”. Catalogue name: ,,Malaclemys pileata” [Kinosternidae] Kinosternon subrubrum (Bonnaterre, 1789) Material: ZMUG 29c [lost] from “Nordamer[ika]” [=North America, no further locality specified] collected in 1853; ZSM 2376/0 [? apparently lost] from “Nord- Amerika” [= North America, no further locality specified] collected by “M. Wagner”. Catalogue name: “Cinosternum pennsylvanicum” [ZMUG 29c]; ,,Nicoria punctularia” [ZSM 2376/0]. ©OZFMK 234 Philipp Wagner et al. Remarks: The ZMUG specimen is an apparently lost stuffed voucher. The fluid collection only was transferred from Gottingen to Bonn and therefore this specimen could still be present in the public exhibition of the Zoological Museum in Gottingen, but this needs to be clarified. [Geoemydidae] Mauremys caspica (Gmelin, 1774) Material: ZMUG 16a [lost] from “Tiflis” [=Tbilissi], Geor- gia collected by “M. Wagner” in 1846. Catalogue name: ,, Clemmys caspia”. Remarks: Berthold (in M. Wagner 1850) referred to two individuals with ‘sehr sch6nen Flammenlinien’. Howevy- er, the catalogue of the museum lists only one voucher. Rhinoclemmys punctularia (Daudin, 1801) Material: ZSM 88/0 [apparently lost] from ““Centralameri- ka” [= Central America, without further locality] collect- ed by “M. Wagner”. Catalogue name: ,,Nicoria punctularia”. Reptilia, Sauria [Agamidae] Trapelus sanguinolentus (Pallas, 1814) Material: ZMUG 151a [lost] from “Tiflis” [=Tbilissi], Georgia collected by “M. Wagner” in 1846. Catalogue name: ,, Agama sanguinolenta”. Citation: As “Trapelus sanguinolutus Eichwald” from “aus Grusien und von den Abhangen des Kaukasus* by Berthold (in Wagner 1850, p. 330). Remarks: Berthold (in Wagner 1850) indicated two indi- viduals of which only one has the characteristic striped throat of the species. At the first inventory of the ZMUG collection in 1968 the voucher was still present but it has apparently been lost. [Dactyloidae] Anolis carolinensis Voigt, 1832 Material: ZMUG 99a-—c [3 ex., lost] from “Nord Ameri- ka” [=North America, no further locality specified] col- lected by “M. Wagner” in 1853; ZSM 470/0 [many ex.; ? apparently lost] from “Louisiana” collected by “M. Wag- ner”. Catalogue name: “Anolis carolinensis”. Remarks: At the time of the inventory of the collection in 1968 all vouchers were still present but they were subse- quently lost. [Lacertidae] Mesalina guttulata (Lichtenstein, 1823) Material: ZMUG 29°, ZMUG 29b [2 ex., lost] from ,,Al- gerien™ [= Algeria, no further locality specified] donated by the ,,Zool[ogisches] Inst[itut]”. Catalogue name: “Evemias guttata”’. Citation: As “Lacerta guttulata” by Schlegel (in M. Wag- ner 1841, part II, 113). Bonn zoological Bulletin 61 (2): 216-240 Remarks: No collector or date is specified with this spec- imen, but the locality and the circumstance that it was a donation of the Zoological Institute (where Rudolf Wag- ner was the head) points to Moritz Wagner. [Scincidae] Plestiodon fasciatus (Linnaeus, 1758) Material: ZMUG 66c [lost] from “Nord Amerika” [=North America, no further locality specified] collected by “M. Wagner” in 1853. Catalogue name: “Eumeces quinquelineatus”. Scincella lateralis (Say, 1823) Material: ZSM 806/0 [4 ex., apparently lost] from “Wis- consin,” USA collected by “Wagner”. Catalogue name: “Lygosoma (Leiolepisma) laterale”’. Remarks: This species does not occur in Wisconsin. [Teiidae] Ameiva praesignis (Baird & Girard, 1852) Material: ZSM 1930/0 [apparently lost] from “Mittel- Amerika” [=Central America, no further locality specified] collected by “M. Wagner”. Catalogue name: ,,Ameiva ameiva praesignis” Aspidoscelis sexlineata (Linnaeus, 1766) Material: ZMUG 17a [lost] from “Nord Amerika” [=North America, no further locality specified] collected by “M. Wagner” in 1853. Catalogue name: ,, Cnemidophorus sexlineatus”’. Remarks: At the time of the inventory of the collection in 1968 the voucher was still present but it is considered to have been lost since that time. Reptilia, Serpentes [Colubridae| Atractus occipitoalbus (Jan, 1863) Material: ZSM 1375/0 [holotype] [lost] from “Ecuador” [no further locality specified] collected by “M. Wagner”. Catalogue name: ,, Atractus occipitoalbus”’. Remarks: According to the original description the type locality is “Andes de |’Ecuador, 4000 ft.” [=Andes of Ecuador, 4000 ft.]. Clelia clelia (Daudin, 1803) Material: ZSM 2664/0 [2 ex., apparently lost] from “Ecuador” [no further locality specified] collected by “M. Wagner’’.. Catalogue name: ,,Pseudoboa clelia” Coluber constrictor Linnaeus, 1758 Material: ZMUG 110a [lost] from “Nord Amerika” [=North America, no further locality specified] collected by “M. Wagner” in 1854. Catalogue name: ,,Zamenis constrictor’. OZFMK Amphibians and reptiles collected by Moritz Wagner 235 Remarks: In contrast to the other North American mate- rial of the Wagner collection this voucher was collected in 1854. Coroneila austriaca Laurenti, 1768 Material: ZMUG 50d [lost] from “Tiflis” [=Tbilissi], Georgia collected by “M. Wagner” in 1846, and later “ded. Strauch”. Catalogue name: ,,Coronella austriaca”. Citation: As “Coronella laevis Boie” from “aus Grusien und von den Abhangen des Kaukasus” by Berthold (in M. Wagner 1850, p. 332). Remarks: Berthold (in M. Wagner 1850) refers to two specimens of ‘Coronella laevis’ but mentioned that they rather should be determined as ,Coronella cupreus’. He specified 13 scale rows around midbody, 171 ventral body and 50 ventral paired tail scales. The coloration was green- ish-gray to copper, with a metallic glossiness. He men- tioned a small stripe between eye and neck and a dark blotch, followed by dark sports behind every occiput scale. Dipsas catesbyi (Sentzen, 1796) Material: ZSM 2279/0 [apparently lost] from “Ecuador” [no further locality specified] collected by “M. Wagner”. Catalogue name: ,,Sibynomorphus catesbyi”. Elaphe guttata (Linnaeus, 1766) Material: ZMUG 88b [lost] from “Nord Amerika” [=North America, no further locality specified] collected by “M. Wagner” in 1853. Catalogue name: ,, Coluber guttatus”’. Imantodes gemmistratus (Cope, 1861) Material: ZSM 30/0 [holotype] [lost] from “Muittel-Ameri- ka” [=Central America, no further locality specified] col- lected by “M. Wagner”. Catalogue name: ,,Himantodes cenchoa var. elegans” Citation: As ,,Himantodes cenchoa var. elegans” from “Amérique central” by Jan and Sordelli (1871). Remarks: This is the holotype of Himantodes cenchoa vat. elegans Jan in Jan & Sordelli, 1871, which is today rec- ognized as a synonym of /. gemmistratus. Leptophis ahaetulla occidentalis (Giinther, 1859) Material: ZSM 1795/0 [apparently lost] from “Mittel- Amerika” [=Central America, no further locality specified] collected by “M. Wagner”. Catalogue name: ,,Leptophis occidentalis” Liophis epinephelus (Cope, 1862) Material: ZSM 1783/0 [2 ex., apparently lost] from “Ecuador” [no further locality specified] collected by “M. Wagner”. Catalogue name: ,,Leimadophis albiventris” Bonn zoological Bulletin 61 (2): 216-240 Mastigodryas boddaerti (Sentzen, 1796) Material: ZSM 83/0 [apparently lost] from ““Mittelameri- ka” [=Central America, no further locality specified] col- lected by “M. Wagner”. Catalogue name: ,,Drymobius boddaerti (Sentz)” Natrix natrix persa Pallas, 1814 Material: ZMUG 67k [lost] from “Urmia,” Iran collect- ed by “M. Wagner” in 1846. Catalogue name: ,, Tropidonotus natrix var. persa”. Citation: As ,,7ropidonotus natrix Kuhl” from “von der ar- menisch-persischen Grenze” by Berthold (in M. Wagner L8SOspsssm)): Remarks: Berthold (an M. Wagner 1850) described this specimen in detail and used this voucher to downgrade ,[ropidonotus persa’, viewed as a full species by Eich- wald (1831), to a subspecies of Natrix natrix. Natrix tessellata (Laurenti, 1768) Material: ZMUG 69c [lost] from “Tiflis” [=Tbilissi], Geor- gia collected by “M. Wagner” in 1846. Catalogue name: ,, Zropidonotus tessellates”. Citation: As “Tropidonotus tessellatus Boje” from “aus Grusien [Georgia] und von den Abhangen des Kaukasus [and the mounatin slopes of the Caucasus]” by Berthold (in M. Wagner 1850, p. 335). Remarks: Berthold (in Wagner 1850) mentioned one voucher of this taxon with 17 keeled scale rows around midbody, 175 ventral body and 60 ventral tail scales. The coloration was described as leaf green on the upper side and yellow and black on the ventral side. This voucher is most probably the specimen mentioned by Berthold. Nev- ertheless, it has apparently been lost since 1968. Nerodia cyclopion (Duméril, Bibron & Duméril, 1854) Material: ZMUG 76 [lost] from “Nord Amerika” [=North America, no further locality specified] collected by “M. Wagner” in 1853. Catalogue name: ,, Thamnophis cyclopium”. Opheodrys vernalis (Harlan, 1827) Material: ZSM 1913/0 [apparently lost] from “Florida,” USA collected by “M. Wagner”. Catalogue name: ,,Contia vernalis”. Oxyrhopus petolarius (Linnaeus, 1758) Material: ZSM 2060/0 [apparently lost] from “Ecuador” [no further locality specified] collected by “M. Wagner”; ZSM 2063/0 [apparently lost]; ZSM 2065/0 [apparently lost] from “Mittel-Amerika [Central America, no further locality specified]” collected by “M. Wagner”; ZSM 2673/0 [apparently lost] from “Ecuador” [no further lo- cality specified] collected by “M. Wagner”. Catalogue name: ,,Clelia petola” [ZSM 2060/0, ZSM 2063/0, ZSM 2065/0]; ,,Pseudoboa petola” [ZSM 2063/0, ZSM 2065/0]; ,,Pseudoboa petola” [ZSM 2673/0]. ©ZFMK 236 Philipp Wagner et al. Sibon nebulata (Linnaeus, 1758) Material: ZSM 2606/0 [2 ex., apparently lost] from “Mit- tel-Amerika [= Central America, no further locality spec- ified]” collected by “M. Wagner”. Catalogue name: ,,Petalognathus nebulosus” Sistrurus miliarius Linnaeus, 1766 Material: ZMUG 216a, ZMUG 216b [2 ex., lost] from “Nord Amerika” [=North America, no further locality specified] collected in 1854. Catalogue name: ,,Sistrurus miliarius” Remarks: In the old catalogue the collector is not named, but locality and date agree with Moritz Wagner. Storeria occipitomaculata (Storer, 1839) Material: ZSM 1538/0 [apparently lost] from “Florida,” USA collected by “M. Wagner”. Catalogue name: ,,/schnognathus occipitomaculatus” Tantilla melanocephala (Linnaeus, 1758) Material: ZSM 2172/0 [apparently lost] from “Mittel- Amerika [= Central America, no further locality speci- fied]” collected by “M. Wagner”. Catalogue name: ,,7antilla melanocephala” Thamnophis sauritus (Linnaeus, 1758) Material: ZSM 81/0 [apparently lost] from “Florida,” USA collected by “M. Wagner”. Catalogue name: ,,7ropidonotus saurita”. Thamnophis sirtalis (Linnaeus, 1758) Material: ZMUG 82c-—e [3 ex., lost] from “Nord Ameri- ka” [=North America, no further locality specified] col- lected by ““M. Wagner [see remarks]” in 1853; ZSM 86/0 [apparently lost] from “Stidcarolina” [= South Carolina], USA collected by “M. Wagner”. Catalogue name: “Thamnophis ordinatus” |ZMUG 82c—e]; ,,7ropidonotus ordinates var. sirtalis” [ZSM 86/0] Remarks: In the ZMUG catalogue the collector’s name for voucher 82c is lacking but date and locality indicate M. Wagner as collector. Specimen ZMUG 82e was recog- nized by a student in Gottingen in 1968, but it has been missing since that time. Thamnophis ordinatus was de- scribed by Linnaeus in 1766 as Coluber ordinatus, but lat- er recognized as subspecies of 7: sirtalis by Cope (1900) and by subsequent authors as a color phase (e.g., Ross- man 1965) Xenodon rabdocephalus (Wied-Neuwied, 1824) Material: ZSM 1824/0 [apparently lost] from ‘“‘Pastassa [=Pastaza valley], Ecuador collected by ““M. Wagner”. Catalogue name: ,,Xenodon rabdocephalus”’. Zamenis longissimus (Laurenti, 1768) Material: ZMUG 86c [lost] from “Tiflis” [=Tbilissi], Geor- gia collected by “M. Wagner” in 1846. Bonn zoological Bulletin 61 (2): 216-240 Catalogue name: ,,Coluber longissimus ”. Citation: As “Coluber aesculapii Shaw” from “aus Grisien und von den Abhangen des Kaukasus” by Berthold (in M. Wagner 1850, p. 333). Remarks: Berthold (in M. Wagner 1850) mentioned ‘Col- uber aesculapii’ with 21 scale rows around midbody, 206 ventral body and 78 ventral tail scales. This voucher has apparently been lost since 1968. [Elapidae] Micrurus ancoralis (Jan in Jan & Sordelli, 1872) Material: ZSM 210/0 [holotype] [lost] from “Ecuador” [no further locality specified] collected by “M. Wagner”. Catalogue name: “(Elaps) ancoralis”’. Citation: As “Elaps Marcgravii var. ancoralis” by Jan and Sordelli (1872). Micrurus fulvius (Linnaeus, 1766) Material: ZSM 2263/0 [apparently lost] from “Nord Amerika [= North America, no further locality specified]”, ZSM 2266/0 [apparently lost] from “Florida,” USA col- lected by “M. Wagner”. Catalogue name: ,,Micrurus fulvius nigrocinctus” [ZSM 2263/0]; “Micrurus fulvius” [ZSM 2266/0] Questionable Material There is some doubtful material available from the ZSM collection. In most of the material a collector is not spec- ified and locality is not mentioned in other catalogue en- tries together with Moritz Wagner as collector, but the cat- alogue entry is together within a series of entries of ma- terial collected by M. Wagner or immediately following such a series. Furthermore, this material was collected at localities which usually refer to Wagner material, or it is from localities Wagner visited during his travels. If the ma- terial is not lost (as indicated in the material section), fur- ther examination is needed to clarify the status of this ma- terial. [Plethodontidae| Plethodon cinereus (Green, 1818) Material: ZSM 1277/0 from “Philadelphia,” Pennsylva- nia, USA; ZSM 1278/0 [8 ex., apparently lost] from “New York,” USA; ZSM 1279/0 [apparently lost] from “Nord- Amerika” [= North America, without further locality]. Catalogue name: “Plethodon erythronotus”; “Plethodon cinereus’. [Plethodontidae] Pseudotriton ruber (Sonnini de Manoncourt and Latreille, 1801) Material: ZSM 1266/0 [Status unclear] from “Nord- Amerika” [= North America, without further locality]; ZSM 1265/0 [4 ex., apparently lost], ZSM 1267/0 [2 ex.] from “Philadelphia,” Pennsylvania, USA. Catalogue name: “Pseudotriton ruber”. ©ZFMK Amphibians and reptiles collected by Moritz Wagner 237 Remarks: Both series, ZSM 1265 and ZSM 1266, have been exchanged with “Dr. [Goreslan?]”. [Salamandridae] viridescens (Rafinesque, 1820) Material: ZSM 1040/0 from ‘“Nord-Amerika” [=North America]; ZSM 1042/0 [2 ex, apparently lost] from “Philadelphia,” Pennsylvania, USA. Catalogue name: “Triturus viridescens”. Notophthalmus [Bufonidae] Atelopus ignescens (Cornalia, 1849) Material: ZSM 1015/0 [2 ex., apparently lost] from “Siid Amerika” [= South America]”. Catalogue name: “Atelopus laevis”. [Lacertidae] Mesalina rubropunctata (Cope, 1861) Material: ZSM 753 [2 ex.] from “Suez,” Egypt collected by “Wagner”. Catalogue name: ,,Eremias rubropunctata” Remarks: The collector Moritz Wagner is usually indicat- ed as “M. Wagner” in the ZSM catalogue, while here the entry only refers to “Wagner”. Even that only the family name is used in some other entries which clearly refer to Moritz Wagner, here the locality “Suez” was, according to our knowledge today, never visited by Wagner. There- fore, it remains questionable if he probably visited Egypt during one of his many shorter travels, or got these spec- imens in exchange. [Scincidae] Plestiodon fasciatus (Linnaeus, 1758) Material: ZSM 832/0 [2 ex.] from “Philadelphia,” Penn- sylvania; ZSM 833/0 [2 ex.] from “Nordamerika” [= North America, without further locality]. Catalogue name: “Ewmeces fasciatus”. [Boidae] Boa constrictor (Linnaeus, 1758) Material: ZSM 1376/0, ZSM 1377/0 from “Siid-Ameri- ka” [=South America, no further locality specified]. Catalogue name: ,,Boa constrictor” [Colubridae] Cubophis cantherigerus (Bibron, 1843) Material: ZSM 1786/0 [? Status unclear] from “Siid- Amerika” [=South America, without further locality]. Catalogue name: ,,Leimadophis angulifer (Bibr.)” [Colubridae] Lampropeltis nigra (Yarrow, 1882) Material: ZSM 1840/0 from “Tenesee [Tennesee], USA”. Catalogue name: ,,Lampropeltis getula nigra’.\\ [Colubridae] Lampropeltis triangulum (Lacépéde, 1789) Material: ZSM 1841/0 [4 ex.] from “New York,” USA. Catalogue name: ,,Lampropeltis getula triangulum’’. Bonn zoological Bulletin 61 (2): 216-240 [Colubridae] Lygophis lineatus (Linnaeus, 1758) Material: ZSM 1784/0 [status unclear], ZSM 1785/0 [2 ex., status unclear] from ‘“Siid-Amerika” [= South Amer- ica, without further locality]. Catalogue name: ,,Leimadophis triscalis”, ,,.Lygophis lin- eatus” [ZSM 1784/0]; “Dromicus lineatus” [ZSM 1785/0] [Colubridae] Nerodia sipedon (Linnaeus, 1758) Material: ZSM 1472/0 [status unclear], ZSM 1473/0 [sta- tus unclear] from “New York,’ USA; ZSM 1474/0 [sta- tus unclear] from “Florida,” USA. Catalogue name: ,,Natrix sipiton [sic!]” DISCUSSION This “checklist” encompasses 321 (207 extant, 114 lost or apparently lost; see also Table 1 for details) herpeto- logical specimens collected by Moritz Wagner and pres- ent in the collections of the NUW, RMNH, ZFMK, ZMB, or ZSM. However, it is possible that more material will be discovered in other collections, e.g., the Paris collec- tion to which Wagner probably donated some of the Al- gerian material. In particular, the Zoological Museum of the Friedrich-Alexander-University Erlangen-Nurnberg [=Nuremberg] possibly also holds material collected from Algeria (especially from his short stay in 1834) or short- er travels in Europe, e.g. southern France. His brother, Rudolf Wagner, being appointed as professor of Zoolo- gy, became the head of the zoological collection when the museum was separated into distinct collections on Janu- ary 18 1833. Up to 1840, when Rudolf Wagner left the university upon accepting a position in G6ttingen, he en- larged the collection which became one of the most im- portant zoological collections in Germany at this time. During this time, Moritz Wagner often visited Erlangen and the university awarded him with a doctoral degree. Therefore, it can be presumed that Wagner also donated material to the collection, but nevertheless, as in Gottin- gen, space for collections is limited at the university and the collection has not been accessible since 1985 and parts remain inaccessible today. The Museum Koenig in Bonn probably houses the largest number of herpetological vouchers collected by M. Wagner (see Table 1) from this time. He donated a large amount of specimens from North America to both the ZMUG and the ZSM collection, but no material from Central America was donated to the Got- tingen collection, probably due to his emerging affiliation to Munich and the Bavarian king after the voyage to North and Central America. Nearly all material from his last voy- age to Central- and South America was donated to the Mu- nich collection (ZSM). However, as collection dates are lacking in the ZSM catalogue, it is often not possible to reconstruct if the material was collected during his first or the second expedition to Central America. ©OZFMK 238 Philipp Wagner et al. Table 1. Number of specimens collected by Moritz Wagner, referring to his five general collecting areas and the investigated col- lections. Collection Algeria Caucasus North America Central America South America NMW 18 1 RMNH 20 ZMB l 5 1 ZMUG (3) (9) (20) ZFMK 4 15 19, ZSM 75 (39) 24 (12) 24 (31) Total 27 24 154 59 57 Table 2. Currently identified type material collected by Moritz Wagner. Original name Current name Type(s) Bufo mauritanicus Schlegel, 1841 Chalcides heteropus Wiegmann, 1856 Diaphorolepis wagneri Jan, 1863 Ecpleopus affinis Peters, 1863 Elaps hertwigii Werner, 1897 Elaps macgravii var. ancoralis Jan, 1872 Hemiphractus fasciatus Peters, 1862 Himantodes cenchoa var. elegans Jan, 1871 Hylaemorphus Pluto Schmidt, 1858 Micrurus latifasciatus Schmidt, 1933 Micrurus latifasciatus Schmidt, 1933 Phyllobates chalceus Peters, 1873 Plectromantis wagneri Peters, 1862 Rhabdosoma occipitoalbum Jan, 1862 Spelerpes palmatus Werner, 1897 Spelerpes palmatus Werner, 1897° Triton ophryticus Berthold, 1846 Vipera wagneri Nilson & Andrén, 1984 ‘= for discussion of this specimen see species account. Bufo mauritanicus Bachia heteropa Diaphorolepis wagneri Pholidobolus affinis Micrurus multifasciatus hertwigi Micrurus ancoralis Hemiphractus fasciatus Imantodes gemmistratus Atelopus varius Micrurus latifasciatus Micrurus latifasciatus Pristimantis chalceus Leptodactylus wagneri Atractus occipitoalbus Bolitoglossa palmata Bolitoglossa palmata Ommatotriton ophryticus Montivipera wagneri Holotype RMNH 2122 Paralectotype ZMB 52679 Holotype ZSM 2708/0 Holotype ZSM 644/0/1 Holotype ZSM 2268/0 Holotype ZSM 210/0 [lost] Holotype ZSM 36/0 Holotype ZSM 30/0 [lost] Syntypes NMW 3875.14 Paralectotype ZSM 2263/0 Paratype ZSM 2263/0 Syntypes ZSM 1045/0 [2.ex] Holotype ZSM 1080/0 Holotype ZSM 1395/0 [lost] Paralectotypes ZSM 1272/0 [2 ex, lost] Lectotype NMW 22862 Lectotype ZFMK 27793 Holotype ZFMK 32495 Moritz Wagner never worked as a herpetologist and on- ly his first books about his travels have a separate chap- ter about amphibians and reptiles (Wagner 1841, 1850). However, he collected some important material, e.g., the type material of Ommatotriton ophryticus (Berthold, 1846), Montivipera wagneri (Nilson & Andrén, 1984), Di- aphorolepis wagneri Jan, 1863 and Leptodactylus wag- neri (Peters, 1862) (see also Table 2 for details). Altogeth- er, 23 of the 149 herein recognized specimens were later used as the basis to describe new species. However, in his Bonn zoological Bulletin 61 (2): 216-240 several publications about zoogeography and speciation (e.g. M. Wagner 1889), Wagner often referred to reptiles. One dubious taxon is mentioned several times. Wagner (1868, 1880) mentioned a pygmy Crotalus which he col- lected on the plateau of Costa Rica and which was, ac- cording to Wagner (1868, 1880) described as a new species by Leopold Fitzinger. However, Wagner never mentioned the name of the new species or the citation of the description or the collection of the type specimen. Moreover, there is no Crotalus taxon described by OZFMK Amphibians and reptiles collected by Moritz Wagner 239 Fitzinger and no Crotalus specimen was found within the Wagner collection. Only two Crotalinae collected by Wag- ner in Costa Rica were found. Bothriechis schlegelii (NMW 28002) and Porthidium lansbergi (NMW 28392), but both were never recognized in the genus Crotalus nor as closely related to C. horridus as mentioned by Wagn- er (1868, 1880). The same problem exists with a putative Alligator species. Wagner (1880) mentioned that after an examina- tion of his material by Carl Theodor von Siebold and Leopold Fitzinger one specimens from western Panama was recognized as a new species. Again, Wagner (1880) did not provide any further data and no corresponding specimen was found. Even though type material of several species was col- lected by Moritz Wagner, his material is mainly of historic interest. However, especially the case of Montivipera wag- neri shows that even historical material can provide new insights, especially if it was collected in poorly known ge- ographic regions. Acknowledgements. We are thankful to the following curators: Frank Tillack and Mark-Oliver Rédel (both Museum fiir Naturkunde, Berlin) Frank Glaw, Michael Franzen and Dirk Neu- mann (all Zoologische Staatssammlung, Munich), Franz Tiede- mann (Naturhistorisches Museum, Vienna) and Ronald de Ruiter (Naturalis, Leiden). Without their important help an identifica- tion of Wagner’ specimens in their respective collections would not have been possible. We are especially very grateful to Michaela Appel from the Vélkerkundemuseum Miinchen and Wolfgang Smolka from the University of Munich for their con- tinuous help and encouragement to get information and docu- ments about Moritz Wagner. We are additionally thankful to the two reviewers, one anonymous and Kraig Adler, for their help- ful comments on the manuscript. Finally it is gratefully acknowledged that the academic staff of the Géttingen museum ultimately agreed with one of the authors (WB) in the mid-1970s to transfer the entire ethanol-preserved herpetology collection, including the historically important ma- terial collected by Moritz Wagner, to the Museum Koenig in Bonn. REFERENCES Anonymus (1856) Vereine, Gesellschaften und Anstalten. Oster- reichische botanische Zeitschrift 6: 116-119 Bauer AM, Giinther R (1994) An annotated type catalogue of the teiid and microteiid lizards in the Zoological Museum, Berlin (Reptilia: Squamata: Teiidae and Gymnophthalmidae). Mitteilungen des Zoologischen Museums in Berlin 70: 267— 280 Bauer AM, Giinther R, Klipfel M (1995) Synopsis of taxa. p. 39-81. In: Bauer AM, Giinther R Klipfel M (eds.) Herpeto- logical Contributions of W.C.H. Peters (1815-1883). 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Macmillan & Co., London 1876 [two volumes] Wagner M (1841) Reisen in die Regentschaft Algier in den Jah- ren 1836, 1837 und 1838. 3 Bande, Verlag von Leopold Voss, Leipzig Wagner M (1848) Reise nach dem Ararat und dem Hochland Ar- menien, mit einem Anhange: Beitrage zur Naturgeschichte des Hochlandes Armenien. J.G. Cotta Verlag, Stuttgart, Germa- ny, x1, 331 pp. Wagner M (1850) Reise nach Kolchis und nach den deutschen Colonien jenseits des Kaukasus. Mit Beitragen zur V6lkerkun- de und Naturgeschichte Transkaukasiens. Arnoldische Buch- handlung, Leipzig. viii, 341 pp. Bonn zoological Bulletin 61 (2): 216-240 Wagner M (1861) Beitrage zu einer physisch-geographischen Skizze des Isthmus von Panama. Mittheilungen aus Justus Per- thes’ Geographischer Anstalt, Erganzungsheft 5: 1-25 Wagner M (1862) Eine Reise in das Innere der Landenge von San Blas und der Cordillere von Chepo in der Provinz Panama, mit besonderer Beriicksichtigung der hypsometrischen Ver- haltnissen und der Kanalfrage. 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Arnoldische Buchhandlung, Leip- zig Wagner M, Scherzer K von (1856) Die Republik Costa Rica in Central Amerika mit besonderer Beriicksichtigung der Natur- verhaltnisse und der Frage der deutschen Auswanderung und Colonisation. Reisestudien und Skizzen aus den Jahren 1853 und 1854. Armoldische Buchhandlung, Leipzig Wagner P (2008) Moritz Wagner und sein Werk — Kurzbiogra- phie eines wegweisenden deutschen Wissenschaftlers. Miinch- ner Beitrage zur V6lkerkunde 12: 25-53 Wagner R (1852) Physiologische Briefe. XII. Beilage zur All- gemeinen Zeitung 61: 969-971 Werner F (1897a) Uber einige noch unbeschriebene Reptilien und Batrachier. Zoologischer Anzeiger 20: 261-267 Werner F (1897b) Ueber einige neue und seltene Reptilien und Frésche der zoologischen Sammlung des bayrischen Staates in Miinchen. Sitzungsberichte der Bayerischen Akademie der Wissenschaften zu Mtinchen, Mathematisch-Physikalische Klasse 27 (2): 203-220 ©ZFMK Bonn zoological Bulletin 61 (2): 241-254 December 2012 Studies on the taxonomy of the Gekko vittatus Houttuyn, 1782 complex (Squamata: Gekkonidae) I. On the variability of G. vittatus Houttuyn, 1782 sensu lato, with the description of a new species from Palau Islands, Micronesia Herbert Rosler , Ivan Ineich , Thomas M. Wilms & Wolfgang Béhme. ‘Senckenberg Naturhistorische Sammlungen Dresden, Museum fiir Tierkunde, Sektion Herpetologie, Kénigsbriicker Landstr. : 159, D-01109 Dresden, Germany. ‘Département Systématique et Evolution, Muséum national d'Histoire naturelle, UMR 7205 (OSEB), CP30, 25, rue Cuvier, F- 75005 Paris, France. ‘Zoologischer Garten Frankfurt, Bernhard-Grzimek-Allee 1, D-60316 Frankfurt am Main, Germany. ‘Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany. Abstract. The present study focuses on morphological variation among conspicuous geographic colour morphs of Gekko vittatus (sensu lato). Meristic data revealed four distinct, allopatric groups of phenotypes, whereas the morphometric char- acters examined do not differ among colour morphs. One of these, endemic to the Palau Islands in the Pacific, is also ge- netically distinct and is here described as a new species. Key words. Gekko vittatus phenotypes; taxonomy; Palau Islands; Gekko remotus sp.n. INTRODUCTION Gekko vittatus Houttuyn, 1782 is one of the few gecko species which is known to science already since the end of the 18" century. Its conspicuous white vertebral stripe on a light brown background, bifurcating in the neck, and its banded tail give this gecko a characteristic appearance. Subsequently, several new taxa, regarded as closely relat- ed to G. vittatus, were described, as well as some colour or pattern morphs which did not receive taxonomic recog- nition so far (e.g. Dumeéril & Bibron 1836, Peters 1872, Mertens 1934, McCoy 1980, 2006). Generally, the taxonomy of the nominal species G. vit- tatus, including the known morphs, synonyms and puta- tive taxa, is far from being satisfactorily solved. Here, we investigate whether forms differing in colouration pattern from typical G. vittatus can also be characterized by meris- tic and morphometric characters. We identify several ax- es of variation among the populations analysed, and for- mally describe a new species from the Pacific Palau Is- lands. MATERIAL AND METHODS A total of 48 specimens tentative assigned to G. vittatus (sensu lato) were examined (listed in the Appendix). The following characters (morphometric, meristic and quali- tative) were recorded for each complete specimen (dam- Received: 02.05.2012 Accepted: 29.05.2012 aged specimen resulted in partially reduced data sets): SVL-— snout vent length, TL — tail length, HL —- maximum head length (from tip to snout to posterior margin of ear), HW — maximum head width, HH — maximum head height, SE — length from snout tip to anterior margin of eye, EE — length between posterior margin of eye to anterior mar- gin of ear, RW — maximum rostral width, RH — maximum rostral height, MW — maximum mental width, ML — max- imum mental length, DTL — dorsal tubercle length (in one of the two median dorsal rows, DTW — dorsal tubercle width (in one of the two median vertebral rows), SPL — supralabials, IF — infralabials, N — nasals (in direction from rostral to labial: nasorostrals, supranasals, postnasals), NP — nostril contact rostral, I — internasals, S6S — scales be- tween the 6th supralabials across the snout, IO — interor- bitals, SC — spiny ciliaries, PM — postmentals, GP — gu- lars bordering postmentals, ESRM — enlarged scale row behind mental, DTR — dorsal tubercle rows, GSDT — gran- ules surrounding dorsal tubercles, GTL — granules between two dorsal tubercles in longitudinal direction, GTC — gran- ules crosswise between two dorsal tubercles , SMC — gu- lar and ventral scales between mental and cloacal slit, V — ventrals, TVF — tubercles on ventrolateral fold, SR — scales around mid-body, LF 1 — subdigital lamellae under Ist finger, LF4 — subdigital lamellae under 4th finger, LT] — subdigital lamellae under Ist toe, LT4 — subdigital lamel- lae under 4" toe, FTW — webbing present between finger Corresponding editor: F. Herder 242 Herbert Résler et al. and toes, TFL — tubercles fore limbs, THL — tubercles hind limbs, PP — precloacal and femoral pores (in males only), PS — precloacal and femoral scales with minute pores or shallow depressions (in females only), PCT — postcloa- cal tubercles, T1 W — tubercle rows in the Ist caudal whorl, T3W — tubercle rows in the 3" caudal whorl, S1 W — dor- sal scale rows in the middle of the first caudal whorl, S3W — dorsal scale rows in the middle of the third caudal whorl, SC1W — subcaudal rows in the Ist whorl, SC3W — sub- caudal rows in the 3" whorl, and SC5W — subcaudal rows in the 5th whorl. Bilateral scale counts were given as right/left. Measurements were recorded with a dial calliper to the nearest 0.5 mm (except the values for rostral, men- tal and dorsal tubercle size which were measured to the nearest 0.02 mm). Collection acronyms. CPHR — Herbert Résler private col- lection, Thale; MNHN — Muséum national d’ Histoire na- turelle, Paris; SMF — Senckenbergmuseum, Frankfurt am Main, now Forschungsinstitut Senckenberg, Frankfurt; ZMB — Zoologisches Museum Berlin, now Museum fir Naturkunde; ZFMK — Zoologisches Forschungsmuseum Alexander Koenig, Bonn; ZSM — Zoologische Staats- sammlung, Miinchen. Specimens were assigned to four different phenotype groups according to their coloration pattern, the shape of their nuchal tubercles, and their ge- ographic origin. Subsequently differences in respect to mensurable and meristic characters were assessed using non-parametric statistics (one-way NPMANOVA) and Principal Coordinates Analysis (PCO) using PAST (Ham- mer et al. 2001). RESULTS Phenotypes of Gekko vittatus sensu lato G. vittatus phenotype | (Fig. 1). A distinct light vertebral stripe, several millimetres wide and bordered with dark brown, bifurcating in the neck re- gion and usually reaching the posterior margin of the eyes is present. This stripe can be interrupted, shortened (Mc- Coy 2006: Fig. 14) or even missing (Ro6sler et al. 2011: Fig. 7A) between neck and eye. The vertebral stripe slight- ly widens more or less saddle-like in the anterior caudal region. The tail is annulated with sharply defined light and dark rings, the dark rings are usually twice as broad as the light ones. Juveniles are similar to adults, but with more intensive caudal colouration. Roundish to oval pointed tu- bercles present only within the white rami of the verte- bral band, all remaining nuchal tubercles blunt and slight- ly convex. Origin of the specimens examined: Wokan (Aru Islands, Indonesia); Asmat, Nabire, (Western New Guinea, Indone- sia), Kordo, Mysore (Schouten Group, Western New Bonn zoological Bulletin 61 (2): 241-254 Guinea, Indonesia), Pulau Ambon (Maluku, Indonesia), Didessa, Airdhills (Papua New Guinea), Ralum, Ratavul (New Britain, Papua New Guinea), Mioko (Duke of York Group, Papua New Guinea). G. vittatus s.1. phenotype 2 (Fig. 2). Irregularly flecked pattern on head, body and limbs. Some- times a light vertebral stripe densely interspersed with darker flecks is discernible. Two dark parallel paraverte- bral sacral stripes framing a lighter sacral are character- istic. Tail above with short irregular dark stripes and flecks, below monochromatic, whitish, medially with narrow, dark streaks. Subadults with all characters of the adults. All nuchal tubercles round to oval, conical, pointed and irregularly arranged. Origin of the specimens examined: Palau Islands (Re- public of Palau). G. vittatus s.1. phenotype 3 (Fig. 3). Colour pattern as in phenotype 1, but nuchal tubercles larg- er and more distinctly conical, pointed. Origin of the specimens examined: Kei Islands (Indone- sia). G. vittatus s.1. phenotype 4 (Fig. 4). Upper side variegated with light and dark. Vertebral stripe entirely lacking or rarely slightly indicated (McCoy 2006: Fig. 115). Two light sacral stripes sometimes present. Tail banding only indicated, predominantly flecked or varie- gated. All nuchal tubercles conical, pointed, larger than in phenotype 1. Origin of the specimens examined: Nissan Atoll (Green Islands, Papua New Guinea). ANALYSIS TERS OF MORPHOLOGICAL CHARAC- Besides of the conspicuous differences in colour pattern, the present sample of G. vittatus s.1. showed considerable variation in several scalation characters while size and body proportions appeared to be less distinct. Mean val- ues as well as minimal and maximal values are provided in Tables | and 2. The four pre-defined groups of pheno- types differ not significantly in the morphometric charac- ters examined (p < 0.2424; F: 1.254; one-way NPMANO- VA; 50000 permutations; distance measure: Gower), while differences in meristic characters are highly significant (p < 0.00004; F: 3.707, one-way NPMANOVA; 50000 per- mutations; distance measure: Gower). A pairwise compar- ison of the four groups revealed that only phenotypes 1 and 2 show significant differences in meristic characters, while the other phenotypes differ mainly in respect to colouration pattern (Table 3, Figs 1-4). A Principal Co- ordinates Analysis (PCO; Fig. 5a) on the morphological ©ZFMK Studies on the taxonomy of the Gekko vittatus Houttuyn, 1782 complex 243 Figs 1-4. 1. Gekko vittatus phenotype 1, ZFMK 20612, male, Pulau Ambon, Indonesia; 2. Gekko vittatus s.l. phenotype 2 (sp.n.), ZMB 5698, female, Republic of Palau; 3. Gekko vittatus s.1. phenotype 3, ZMB 48737, male, Kei Islands, Indonesia; 4. Gekko vit- tatus s.1. phenotype 4, SMF 9159, male, Nissan Atoll, Green Islands, Papua New Guinea. data obtained from 21 specimens [28 characters; includ- ing specimens from all four phenotypes identified in the present study as well as one of the syntypes of G. bivit- tatus (MNHN 6714) and the holotype of G. trachylaemus (ZMB 7511)] revealed that the two respective type spec- imens fall far outside of the remaining clusters. Hence, and supported by the finding that more differences in coloura- tion are present between those two nominal taxa and the four phenotypes of G. vittatus s.l. (see next paragraph), we consider the evaluation of the taxonomic status of G. bivittatus and G. trachylaemus as not within the scope of the present study, because we shall deal with these nom- inal taxa in detail in a forthcoming paper. The replication of this PCO excluding G. bivittatus and G. trachylaemus revealed two major clusters — one encom- passing all specimens of phenotype 2 while the second cluster includes roughly all remaining specimens with ex- ception of a single specimen related to phenotype 3 (Fig. 5b). The dataset used in the previous analysis was select- ed to maximize the number of specimens included in the analysis (because of the preservation state of most spec- imens, datasets of single specimens are mostly incomplete, missing one or more characters). Therefore the resulting datasets for analysis are either selected to maximize the Bonn zoological Bulletin 61 (2): 241-254 number of specimens (which means having fewer char- acters) or to maximize the number of characters (which means encompassing fewer specimens). To check if the use of these two different datasets results in significantly different PCO analyses, a PCO was carried out on a dataset maximizing the number of characters (44 characters; 14 specimens; Fig. 5c). The results of the two PCO analyses are in principal accordance, showing a single cluster for the specimens assigned to phenotype 2. Differences oc- cur in the positions of the specimens assigned to pheno- types 3 and 4, which cluster either together with the phe- notype | specimens (Fig. 5b) or completely outside of the clusters of phenotype | respectively two specimens (Fig. Sc.). On the basis of the PCO results presented here, phe- notypes | and 2 can be regarded as morphologically dif- ferent entities, while there is no evidence regarding the morphological affiliation of phenotypes 3 and 4. Available names in the synonymy of G. vittatus s.1. Gekko vittatus has been described by Houttuyn (1782) é” (certainly from India) where- on the type locality was founded. Subsequently, the nomen ©OZFMK 244 Herbert Rosler et al. Table 1. Mensural data of the Gekko vittatus phenotypes 1-4, the syntypes of Gekko bivittatus and the holoype of Gekko trachy- laemus (mean + SD; max; min; Abbreviations as in material and methods). phenotype 1 _— phenotype 2 phenotype 3 phenotype 4 G. bivittatus G. trachylaemus no 2 a 2 A a Si _. 2 1 SVL 76.35+25.20 81.63+30.56 105.3—125.4 110.5742.25 109.0—120.0 103.7 38.2-108.3 39.0-117.5 108.0-112.2 TL 70.4744 1.17 70.81423.18 116.0—123.0 — 115.0-125.0 106.6 39.4-114.5 53.0-114.8 HL 21.04+6.34 21.23+7.16 30.0-32.5 30.57+1.56 28.0—-30.0 28.3 11.1-29.1 11.1-29.4 29.1-32.2 HW 14.73+4.71 14.88+45.52 21.7-22.0 21.53+1.50 19.8-18.9 19.6 7.6—20.9 7.8-21.6 19.8-22.5 HH 9.0742.83 10.53+7.25 13.1-13.9 12.77+1.06 11.4-11.8 10.6 5.0-12.6 4.9-31.5 11.8=13.9 SE 10.9141.54 10.16+3.50 15.0-16.6 14.63+0.75 11.5—-12.8 13.5 5.2-13.9 4.9-14.5 13.9-15.4 EE 8.99+1.54 8.48+2.94 11.7-12.7 11.83+0.32 10.6-12.8 9.9 4.3-11.9 3.9-12.6 11.6-12.2 RW 3.92+0.48 4.28+0.47 3.90-4.50 4.50+0.34 3.9-5.5 4.00 3.20—-4.60 3.40-4.80 4.10-4.71 RH 2.06+0.37 2.09+0.27 2.10—2.90 2.45+0.38 1.82.3 2.50 1.40—2.50 1.72.3 2.14-2.88 MW 2.32+0.52 2.02+0.35 2.40—2.50 2.58+0.22 —= 2.60 1.90—3.60 1.70—2.30 2:35—2.18 ML 1.72+0.27 1.82+0.32 1.90—2.30 2.02+0.27 1.53—1.65 1.80 1.10-1.92 1.30—2.20 1.71-2.18 DTL 0.67+0.11 0.85+0.16 0.72—0.92 0.81+0.08 0.63—0.81 1.19 0.49-1.91 0.52-0.94 0.73-0.89 DTW 0.56+0.13 0.73+0.20 0.56—0.72 0.63+0.09 0.75—0.67 0.80 0.43—0.89 0.35—0.93 0.53-0.71 SVL/TL 0.95+0.02 0.97+0.03 0.91—1.02 == 0.95—0.96 0.97 0.89—1.03 0.93—1.02 SVL/HL 3.62+0.18 3.80+0.19 3.51-3.86 3.62+0.12 3.89-4.00 3.66 3.32-3.94 3.51-4.14 3.48-3.71 HL/HW 1.43+40.05 1.44+0.07 1.38-1.48 1.42+0.05 1.48—1.52 1.44 1.34-1.49 31S 1.57 1.36—1.47 HL/HH 2.32+0.09 2.27+0.44 2.29-2.34 2.40+0.19 2.46-2.54 2.67 2.22—2.46 2.18—2.60 2.19-2.56 SE/EE 1.21+0.07 1.2040.07 1.18—1.42 1.24+0.04 1.00—1.08 1.36 1.13-1.31 1.12-1.34 1.19—-1.26 RW/RH 1.93+0.34 2.07+40.29 1.34-2.14 1.85+0.19 DANG 239) 1.60 1.68—2.56 1.91—2.67 1.64—2.00 MW/ML 1.34+0.26 1.13+0.22 1.09-1.26 1.29+0.09 —= 1.44 1.11—1.89 0.86—1.47 1.20=1.37 RW/MW 1.74+0.25 2.16+0.37 1.63—1.80 1.74+0.06 = 1.54 1.14—2.00 1.68—2.53 1.68—1.80 DTL/DTW 1.24+0.35 1.20+0.18 1.28-1.29 1.29+0.07 — 1.49 0.55—1.82 0.97-1.49 1.24—1.38 DTLx100/SVL 0.71+0.08 0.77+0.08 0.68—0.73 0.73+0.06 — 1.15 0.55—0.86 0.62—0.85 0.68—0.79 DTWx100/SVL 0.62+0.26 0.66+0.13 0.53—0.57 0.57+0.07 — 0.77 0.44-1.26 0.42—0.79 0.49—0.63 Bonn zoological Bulletin 61 (2): 241-254 ©ZFMK Studies on the taxonomy of the Gekko vittatus Houttuyn, 1782 complex 245 Table 2. Meristic data of the Gekko vittatus phenotypes |—4, the syntypes of Gekko bivittatus and the holotype of Gekko trachy- /aemus (mean + SD; max; min; Abbreviations as in material and methods). 6.SPL PCT TILW T3W S1W S3W SC1W SC3W SC5W phenotype I chia -13.4540.94 1-16 13.5141.13 11-16 3.02+40.15 3-4 1.15+0.76 0-3 40.55+4.74 32-50 27.46+1.76 24-31 3.9541.52 1-6 5 3.69+0.70 3-5 5.08+0.71 4-6 26.27+2.28 2923) 8.74+1.11 6-10 1.96+0.57 1-3 1.7240.57 13 191,05+20.77 161-222 134.89+11.58 115-155 32.26+2.82 27-39 13.08+1.27 10-15 18.08+1,94 13-22 14.28+1,36 iy 21.22+2.07 17-26 51.5547.55 39-65 16.20+4.76 11-26 2.04+0.74 125 3.0840.74 2-4 3.08+0.00 3-4 10.58+1.27 8-12 11.25+0.82 10-13 4.42+0.52 4-5 4.08+0.32 4-5 4.00+0.00 4 phenotype 2 Pee phenotype 3 13.00+0.93 11-15 12.38+40,82 10-14 3.13+0.45 23 1.1740.58 13 40.55+2.81 6-45 27,4242.15 25-31 4.50+1.25 4) 1.92+0.29 1-2 4.00+1.28 2-6 5,250.62 4-6 24.75+1,36 2G 10.67+0.64 10-12 3.00+40.98 15 2.33+0.70 1-4 189.29+10.63 175-205 137.2943.73 131-142 30.63+2.88 25-34 15.80+40.77 14-17 21.50+0.95 202 16.89+1.08 15-1 24.15+1.23 22-26 39.57+5.68 31-45 2 1.55+0.51 1-2 3.42+0.51 3-4 3.33+0.65 3=5 11.91+0.83 10-13 12.40+1.17 1-14 4.17+0.39 4-5 4.00+0.00 4 4.00+0.00 4 194-208 139-142 31-38 [S=17 18-22 16-17 23-25 51-55 12-13 11-13 phenotype 4 0.3340.58 0-1 44.00+1.73 43-46 32.50+0.71 32-33 5.00+0.71 4-6 5 4.00+1.41 3-5 6.00+1.00 5-7 27.67+0.58 OB 9.17+0.98 8-11 1.6740.82 123 2.00+0.63 1-3 193.3343.51 190-197 130.6743.21 127-133 34.6744.73 31-40 16.67+1.21 15 —18 19.67+0.82 18—20 16.50+0.55 16-17 i) Bonn zoological Bulletin 61 (2): 241-254 G. bivittatus 2 10-11 11-14 53 35 G. trachylaemus 1 12/12 10/10 3 0 246 Herbert Rosler et al. | Gekko bivittatus (MNHN 6714) eae ae ee eae 4 5a Stan ne henotype 3“ Gekko ipacnvlecns (ZMB 7511) Phenotype 2 5b Phenotype 4 Fig. 5a. Principal Coordinates Analysis (PCO) on 28 morpho- logical characters [8 metric (ratios of original biometric data), 18 meristic and 2 qualitative] of 21 specimens within the Gekko vittatus complex (OTUs including the nominal taxa G. bivitta- tus and G. trachylaemus). Raw data log), transformed (after adding x=1 to all values); Similarity Index: Gower. Axis Eigenvalue Percent 1 0.17682 35.08 2 0.09013 17.88 3 0.05113 10.16 Sum 63.11 Fig. 5b. Principal Coordinates Analysis (PCO) on 28 morpho- logical characters [8 metric (ratios of original biometric data), 18 meristic and 2 qualitative] of 19 specimens within the Gekko vittatus — complex (OTUs excluding the nominal taxa G. bivit- tatus and G. trachylaemus). Raw data logy, transformed (after adding x=1 to all values); Similarity Index: Gower. oe Axis Eigenvalue Percent 1 0.21369 32,68 2) 0.08660 13,24 3 0.06508 9,95 Sum 55,87 Fig. 5c. Principal Coordinates Analysis (PCO) on 44 morpho- logical characters [10 metric (ratios of original biometric data), 31 meristic and 3 qualitative] of 14 specimens within the Gekko vittatus Was used in combination with various generic names: Lacerta (e.g. Gmelin 1789), Platydactylus (e.g. Fitzinger 1826) and Lomatodactylus (e.g. Van der Hoeven 1833). Currently, four names are considered to be synony- mous with Gekko vittatus Houttuyn, 1782 (see Wermuth 1965, Bauer 1994, Kluge 2001): 1. Lacerta unistrigata Shaw, 1792. Independently from Houttuyn (1782), Shaw (1792) described G. vittatus again, as Lacerta unistrigata, but without data on any type ma- terial. Later, he changed this name into Lacerta vittata (see Shaw 1802). Despite the lack of type material, the origi- nal description is sufficiently conclusive to place Lacer- ta unistrigata in the synonymy of G. vittatus sensu lato. 2. Stellio bifurcifer Schneider, 1792. In the same year, Schneider (1792) published a substitute name for G. vit- tatus Houttuyn, 1782, viz. Stellio bifurcifer, and comment- ed on it again (Schneider 1812) after having compared it with the description by Daudin (1802). Again, the origi- nal description allows synonymization of Ste/lio bifurcifer with G. vittatus sensu lato. Bonn zoological Bulletin 61 (2): 241-254 eens Ds vittatus — complex. Raw data log, transformed (after adding x=1 to all values); Similarity Index: Gower. pheneaee , Axis Eigenvalue Percent Phenovire 1 0.17682 35,08 2 0.09013 17,88 3 0.05113 10,15 Sum 63,11 3. Platydactylus bivittatus Duméril & Bibron, 1836. While Dumeéril & Bibron (1836) ignored the priority of Gekko Laurenti, 1768 over Platydactylus Goldfuss, 1820 (Gold- fuss 1820, Gray 1825), they generally accepted Cuvier’s (1817) classification of geckos who had placed G. vitta- tus as member of Platydactylus under the unavailable name “Le Platydactyle a bande”. Duméril & Buibron (1836) compared Platydactylus vittatus (= Gekko vittatus) with their newly described Platydactylus Bivittatus (sic) but admitted that both species are hardly distinguished by morphology and might be only colour morphs rather than species; “Il pourrait se faire que ce Platydactyle a deux bandes ne soit qu’une variété du Platydactylus vittatus, dont il ne différe bien réellement que par la systeme de coloration”. Boulenger (1885) downgraded Platydactylus bivittatus Duméril & Bibron, 1836 to a “variety” (current- ly mostly seen as a subspecies) of Gekko vittatus Hout- tuyn, 1782 (see also Strauch 1887, Boettger 1893, Wern- er 1900), before Loveridge (1948) finally synonymized this name with the nominotypic form (see also Guibé 1954, Brygoo 1990). ©OZFMK Studies on the taxonomy of the Gekko vittatus Houttuyn, 1782 complex 247 Table 3. Significance levels of morphological differences between four OTU’s within the Gekko vittatus -complex (p < 0.00004; F: 3.707, one-way NPMANOVA; 50000 permutations; distance measure: Gower); pairwise uncorrected significances shown. a ERODE LE Phenotype 1 0.00156" Phenotype 2 0.00156" Phenotype 3 0.6259 0.8558 Phenotype 4 0.5576 0.8675 We re-examined both syntypes of Platydactylus bivit- tatus Duméril & Bibron, 1836 (Tabs 1, 2). MNHN 6714, an adult from “Nouvelle Guinée”, has a relatively strong tuberculation and a high number of precloacal and femoral pores (53), and the unforked vertebral stripe is without flecks (Fig. 6). This character combination distinguishes it from both G. cf. vittatus from the Palau and the Kei Is- lands. The second syntype, MNHN 2285, an adult female from “Waigiuo” (= Pulau Waigeo) is badly preserved. Nei- ther a vertebral stripe nor a light, dark-bordered sacral stripe is discernible within the variegated dorsal pattern igs 7): Its dorsal pattern resembles that of G. cf. vittatus from Nissan Atoll, but further comparisons and a larger sam- ple size are necessary to see whether also the numbers of precloacal and femoral scales fit. 4. Gekko trachylaemus Peters, 1872. Described by Peters (1872) from “Nordaustralien” (North Australia) who di- agnosed it by larger dorsal tubercles, less supralabials and infralabials, a larger mental and more extended webbing between fingers and toes. Peters & Doria (1878), howev- er, synonymized G. trachylaemus with G. vittatus, and Cogger et al. (1983) pointed out that the alleged type lo- cality should be wrong, as no Australian records of this gecko exist. We re-examined the data given by Peters (1872) for G. trachylaemus on the basis of its holotype ZMB 7511 (Tab. 1 and 2): the mental relative to the rostral is broader than that of the Palau form and the phenotypes 3 and 4, but is within the range of G. vittatus s.str. (see Tab. 1). The scales behind the mental are relatively larger than in all other specimens studied, but the number of rows of these scales is the same (Tab. 2).The flat and smooth to weakly keeled nuchal, dorsal and lateral tubercles of G. trachylaemus are distinctly larger than in G. vittatus s.str. but less so in the Palau form and the phenotypes 3 and 4. The colour pat- tern in ZMB 7511 is strongly faded; only the short, sacral streaks as described by Peters (1872) are still a bit better discernible (Fig. 8). The holotype of Gekko trachylaemus is distinguished from the Palau form and from the G. vittatus (s.1.) phe- notypes 3 and 4 by a different ventral tail scalation: ex- cept the first three whorls, there are always three rows of subcaudals present in each whorl of the original, unregen- Bonn zoological Bulletin 61 (2): 241-254 _ Phenotype 2 Phenotype 3 ; _ Phenotype 4 0.6259 0.5576 0.8558 0.8675 erated tail while there are constantly four to five rows per whorl in G. vittatus s.str. as is also the case in the Palau form and in the phenotypes 3 and 4. Moreover, G. trachy- laemus differs from the Palau form by larger and flatter nuchal and dorsal tubercles, larger tail tubercles and a more banded colour pattern. The assessment the taxonom- ic status of Gekko trachylaemus requires more material and is therefore beyond the scope of the present study. CONCLUSIONS Among the four phenotypes of Gekko vittatus s.l., only phenotype | has rather small, flat tubercles on head, body and limbs, und only the nuchal region bears sparsely dis- tributed conical, pointed tubercles, while all other three phenotypes have larger, relatively high and pointed tuber- cles, particularly in the nape, so that they have a more spiny appearance. Together with its characteristic colour pattern, phenotype | corresponds closely with the origi- nal description and can thus be attributed to Gekko vitta- tus sensu stricto. The greatest affinity to this form in dor- sal and tail colour pattern as well as regarding their posi- tion in the PCO plot is shown by the two Kei Islands males (ZMB 48737-48738; phenotype 3) which are, however, distinguished by their distinctly larger and more pointed nuchal tubercles as well as a possible different tail scala- tion. Pending further morphological studies and molecu- lar genetic data, this phenotype should be termed Gekko cf. vittatus (“Kei Islands”). Despite the overall similarity of the specimens assigned to phenotype 4 to phenotype | which was shown in the PCO analysis certain differences in morphology and colouration [lower number of precloa- cal and cloacal pores (28-37) and the overall dark coloura- tion including the lack of a vertebral stripe] suggests, that the three males from Nissan Atoll (SMF 9157-9159; phe- notype 4) are also distinguishable from G. vittatus s.str. This phenotype should respectively be termed as Gekko cf. vittatus (“Nissan Atoll’). Based on the considerable ge- netic distance (see Rosler et al. 2011) and the significant differences in scalation and colour pattern as compared with the other three phenotypes outlined above, and as there is no older available name for the Palau population, we describe phenotype 2 below as a new species. ©ZFMK 248 Herbert Rosler et al. It TU 4219 14 49) ANU Sine Figs 6-10. 6. Syntype of Platydactylus bivittatus, MNHN 6714, male, New Guinea; 7. Syntype of Platydactylus bivittatus, MNHN 2285, female, Pulau Waigeo, Indonesia; 8. Holotype of Gekko trachylaemus, ZMB 7511, male, Australia; 9. Holotype of Gekko remotus sp.n., ZFMK 20611, male, Republic of Palau; 10. Gekko remotus sp.n., ZMB 76979, juvenile, Republic of Palau. Gekko remotus sp.n. Diagnosis. A species of the Gekko vittatus s.1. complex with which it is connected by its tubercles on throat and lateral folds as well as by colour pattern. From Gekko vit- tatus s.str. (phenotype 1) it is distinguished by pointed (versus blunt) nuchal tubercles and by the lack of a white, Bonn zoological Bulletin 61 (2): 241-254 anteriorly forked vertebral stripe and of an annulated tail pattern. From Gekko cf. vittatus (phenotype 3, “Kei Is- lands”) Gekko remotus sp.n. is distinguished by less nu- merous precloacal and femoral pores (31—45 vs. 51—55) and from Gekko cf. vittatus (phenotype 4, “Nissan Atoll”) by a distinct flecked dorsal (vs. dark and light variegat- ed) colour pattern. ©ZFMK Studies on the taxonomy of the Gekko vittatus Houttuyn, 1782 complex 249 Holotype. ZFMK 20611, male, Palau Islands, coll. by Brock, before 1874, received from Godeffroy Museum, Hamburg (via Zoological Museum, University of Géttin- gen, 1977). Description of holotype. Head moderately depressed, dis- tinct from neck. Body not depressed, roundish, belly flat. Tail round, not swollen at base; SVL 117.5 mm, TL 118.5 mm (incomplete); HL 29.4 mm; HW 21.3 mm; HH 13.5 mm; SE 14.2 mm, EE 11.5 mm; ED 5.65 mm; EAD 4.26 mm. Proportions: SVL/HL 4.00; HL/HW 1.38; HL/HH PRRSSE/ EE sle23 52 D/EE) 133: Rostral triangular, wider than high (RW 4.39 mm, RH 2.23 mm, RW/RH 1.97), wider than mental (RW/MW 1.82), on the upper margin a converse Y-shape suture; supralabials 14/13, 11/10 to the center of eye; nostril in contact with rostral and Ist supralabial; nasals 3/3, na- sorostrals squarish, twice as large as supranasals and post- nasals; 1 rectangular internasal, half as big than the na- sorostrals. Snout with a longitudinal shallow groove; lat- eral scales on snout polygonal, smooth, flat to weakly domed, juxtaposed, one and a half times as large as on the mid-snout; 17 scales between postnasals and orbit, 44 scales between the two 6th supralabials; median snout scales polygonal, smooth, flat to weakly domed, juxta- posed; dorsal ciliaries as large as median snout scales, 5/5 spiny tubercles posteriorly. Ear opening vertical, oval; in- terorbitals 27, polygonal, smooth, flat to weakly domed, juxtaposed; orbital scales weakly conical, and one a half times as large as those in the middle of head. Occipital and nuchal scales polygonal, flat, juxtaposed, half as big as the median interorbitals; temporal, occipital and nuchal tubercles round, conical, five times as big as the surrounding scales. Mental triangular, wider than long (MW 2.41 mm, ML 2.10 mm, MW/ML 1.15); infralabi- als 12/13, larger than supralabials; posterior of mental five rows of enlarged roundish, oval scales, two of them bor- dering the mental; no enlarged postmentals; gulars as big as nuchals, round, flat, juxtaposed, interspersed with dou- ble-sized conical tubercles. Dorsal and lateral scales sim- ilar to nuchals; Dorsal tubercles 2—3 times as big as the bordering dorsal scales (DTL 0.94 mm, DTW 0.79 mm, DTL/DTW 1.19), oval, conical, weakly keeled, surround- ed by 10-11 dorsals, in 22 more or less regularly arranged longitudinal rows. Lateral tubercles as large as dorsal tu- bercles, mostly round, conical to pointed. Tubercles on lat- eral fold large, blunt, conical. separated by 1—2 small flank scales. Ventrals flat, smooth, imbricate, the median ones being 3-4 times larger than dorsals, 31 between the lat- eral folds. Midbody scale count 136; 184 scales between mental and cloaca. Scales on upper side of forelegs as large as dorsals, flat, smooth and juxtaposed, passing in small, granular scales below; tubercles on forelegs roundish to oval, blunt, conical, 2—3 times larger than the surround- ing scales. Upper and lower thigh scales flat, imbricate Bonn zoological Bulletin 61 (2): 241-254 passing posteriorly into smaller granules. Tubercles on hind limbs resemble those on forelimbs. Digits connect- ed by narrow webbing, the claws of digits 2-5 encircled by 3 scales each, the median claw of the sheath 2-3 times bigger than the two lateral sheath scales. Subdigital lamel- lae 16/16 under the Ist finger, 22/23 under the 4th finger, and 17/17 under the Ist toe, and 24/26 under 4" toe, in- terdigital webbing present but weakly developed. 31 pre- coacal and femoral pores in one angular, continuous row, 17/17 smooth femoral scales towards the knees behind them small, round tubercles. Original tail distinctly whorled, not thickened at the base. Dorsal tail scales as large as on the back, flat or weakly domed, juxtaposed to weakly subimbricate, arranged in irregular oblique rows. The Ist whorl has 12 scale rows middorsally, the 3rd whorl 13; down to the 11" whorl 3 rows of tubercles per whorl. Tail tubercles similar to dorsal tubercles in size, conical and submucronate, subcaudals not widened, mostly 2, rarely 3 per oblique row, in the Ist whorl 5, in all remain- ing whorls 4 oblique scale rows per whorl, the last pair of scales a bit enlarged. The colour in preservative is lilac-grey on head and dor- sum, pale grey on tail. Head, dorsum and limbs densely flecked with brown, the flecks being partly confluent. Sacral area with two short blackish brown stripes, border- ing a light median zone between them. Tail dorsally var- iegated with brown. Underside of head, body and limbs greyish brown. Tail medially with a narrow, interrupted dark gray streak (Fig. 9). Variation. The infraspecific variability of measurements, scalation characters and colour pattern was evaluated in a series of 11 topotypic specimens from the ZMB collec- tion which are, however, regarded as further material rather than paratypes, because neither the holotype nor these specimens have specific data as to their particular island origin within the Palau archipelago. Measurements, proportion indices and scalation values are shown in Tabs 4 and 5. As compared with the predominantly greyish-coloured holotype, the ground colour of five adult males from Palau varies between pale brown and dark brown. A light ver- tebral strip, stretching from the nape to the tail base is on- ly shadowy indicated in four males while it is more dis- tinct with a lateral bordering in one. Four specimens have only a weakly discernible flecked pattern, and only one specimen is comparable to the holo- type in this respect (Fig. 2). Of six juveniles of various sizes, only the biggest three (SVL 57—65 mm) have a pat- tern similar to that of the holotype (Fig. 10), in two small- er ones (SVL 51-54 mm) the light vertebral stripe is forked and reaches the orbital hind margin; the smallest juvenile (SVL 39.0 mm) is uniformly pale brown. 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(see above). Distribution. Known only from the type locality (i.e. Palau Islands). Gekko remotus sp. n. has been recorded from the Palau Islands Babeldaob, Bablomekang, Beliliou, Kmekumer Island, Malakal, Ngeaur, Ngercheu, Ngeruais, Ngerekebesang, Ngerikeuid Island, Oreor, Ulebsechel and Ulong (Crombie & Pregill 1999). Natural History. Gekko remotus sp. n. is active at night and mostly arboricolous, but has occasionally also been found on house walls. It feeds on insects as well as on fruit. Crombie & Pregill (1999) observed not only strong ag- gressive (territorial) behaviour among single adult males, but also attacks on non-receptive females. Moreover they reported cannibalism. Comparisons. Like Gekko vittatus (s.1.) Gekko remotus sp.n. shows tubercles on the throat and lateral folds which makes both species easily distinguishable from all other species of the genus Gekko (Rosler et al. 2011). Moreover, G. remotus sp.n. differs from G. albofasciolatus Gunther, 1867, G. gecko (Linnaeus, 1758), G. reevesi (Gray, 1831), G. siamensis Grossmann & Ulber, 1990, G. smithii Gray, 1862, and G. verreauxi Tytler, 1864 (a species endemic for the Andaman Islands) by its smaller SVL (117.5 mm vs. >150 mm), and also — except G. verreauxi, by a con- tact of nostril and rostral; it differs from G. verreauxi by more dorsal tubercle rows (22 vs. 11). Gekko athymus Brown & Alcala, 1962, G. melli Vogt, 1922, G. scientiadventura Rosler, Ziegler, Vu, Herrmann & Boéhme, 2004, G. subpalmatus Gunther, 1864, and G. tawaensis Okada, 1956 lack the dorsal tubercles present in G. remotus sp.n. The latter differs from G. auriverrucosus Zhou & Liu, 1982, G. badenii Szczerabk & Nekrasova, 1994, G. can- hi Rosler, Ngyuyen, Doan, Ho, Nguyen & Ziegler, 2010, G. grossmanni Ginther, 1994, G. hokouensis Pope, 1928, G. japonicus (Schlegel, 1836), G. petricolus Taylor, 1962, G. russeltraini Ngo, Bauer, Wood & Grismer, 2009, G. scabridus Liu & Zhou, 1982, G. sibatai Toda, Senhoku, Hikida & Ota, 2008, G. similignum Smith, 1923, G. swin- honis Gunther, 1864, G. taibaiensis Song, 1985, G. tak- ouensis Ngo & Gamble, 2010, G. vertebralis Toda, Sen- goku, Hikida & Ota, 2008, G. vietnamensis Nguyen, 2010, G. wenxianensis Zhou & Wang, 2008, and from G. yvakuensis Matsui & Okada, 1968 by a higher number of cloacal and femoral pores (31 vs. 0-15), and moreover, except from G. taibaiensis, G. wenxianensis and G. yvakuensis, by non-dilated subcaudals. Gekko remotus sp.n. differs from G. crombota Brown, Oliveros, Siler & Diesmos, 2008, G. gigante Brown & Al- cala, 1978, G. mindorensis Taylor, 1919, G. palawanen- Bonn zoological Bulletin 61 (2): 241-254 Herbert Rosler et al. sis Taylor, 1925, G. porosus Taylor, 1922, G. romblon Brown & Alcala, 1978, and G. rossi Brown, Oliveros, Siler & Diesmos, 2009 by less numerous precloacal and femoral pores (31 vs. 50-88) and a lower midbody count (136 vs. 145-211). The new species is distinguished from G. ernstkelleri Rosler, Siler, Brown, Demegillo & Gaulke, 2006, G. /i- boensis Zhao & Liu, 1982, G. palmatus Boulenger, 1907, and G. similignum Smith, 1923 by its numerous, densely placed tubercles on the forelimb, and by a bigger SVL (117.5 vs. 82.5—92.1 mm). G. canaensis Ngo & Gamble, 2011 lacks tubercles on the limbs (present in G. remotus sp.n.), has fewer precloacal pores (14-18 vs. 31) and widened subcaudals. G. carusadensis Linkem, Siler, Dies- mos, Sy & Brown, 2010 is smaller than G. remotus sp.n. and has a smaller SVL (97.2 versus 117.5 mm), more in- terorbitals (35-48 vs. 27) and more precoacal and femoral pores (46—S0 vs. 31), moreover widened subcau- dals. G. remotus sp. n. differs from G. chinensis Gray, 1942 by only weakly developed webbing between fingers and toes (distinct and well developed in G. chinensis) and by its larger SVL (117.5 vs. 72 mm), more subdigital lamel- lae under the Ist and 4th toe (17 vs. 8-10 and 24—26 vs. 12-19 respectively), and non-widened subcaudals. G. kikuchii Oshima, 1912 has a smaller SVL than G. re- motus sp.n. (80 vs. 117.5 mm) and has more precloacal and femoral pores (48 vs. 31) and has moreover widened subcaudals. G. monarchus (Schlegel, 1836) has more in- terorbitals than G. remotus sp.n. (34-35 vs. 27) a higher midbody count (148-177 vs. 136) and widened subcau- dals. Finally, the endemic Thai species G. nutaphandi Bauer, Sumontha & Pauwels, 2008 has, in contrast to G. remotus sp.n., a nostril without contact to the rostral, and both species are moreover distinguished by different num- bers of precloacal and femoral pores, viz. 17—22 vs. 31. Etymology. The species name is derived from the Latin adjective remotus, -a, -um, meaning far away, and refers to the remote, isolated distribution range of the new species. DISCUSSION The genus Gekko contains several phylogenetic lines or clades with different geographic centres. Molecular diver- gence within the Gekko vittatus group (mitochondrial and nuclear genes: ND2, tRNA, RAG-1, PDC) revealed the distinctness of G. remotus sp.n. from all other members of this group (Résler et al. 2011). In addition to the dif- ferences in scalation and colour pattern, G. remotus sp.n. differs from G. vittatus s.str. in some behavioural traits: G. remotus have a strictly solitary lifestyle including ag- gressive behaviour against conspecifics and even canni- ©OZFMK Studies on the taxonomy of the Gekko vittatus Houttuyn, 1782 complex 253 balism, whereas G. vittatus live in familial groups and per- form brood care (Treu 2001). The herpetofauna of the Palau Islands is dominated by faunal elements of New Guinean, Solomonian or Moluccan origin (Crombie & Pregill 1999). The northernmost representative of the Gekko vittatus group so far known is from Pulau Moro- tai, and also the records from Halmahera and Pulau Waigeo are geographically closer to the Palau islands as for instance New Guinea or the Admiralty Islands (De Roo 1915). Further studies are, however, necessary to elucidate, from where the ancestors of Gekko remotus sp.n. may have reached the Palau Islands. Acknowledgements. 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Annali del Museo Civico di Storia Naturale, Genova, Seria. 1, 13: 323-450 Rosler H, Bauer AM, Heinicke MP, Greenbaum E, Jackmann T, Nguyen QT, Ziegler T (2011) Phylogeny, taxonomy, and zoo- geography of the genus Gekko Laurenti, 1768 with the reva- lidation of G. reevesii Gray, 1831 (Sauria: Gekkonidae). Zoo- taxa 2989: 1-50 Schneider JG (1792) Amphibiorum physiologiae alternum his- toriam et species generis stellionum seu geckonum sistens. Chr. Ludov. Frid Apitzil: Traiecti ad Viadrum Schneider JG (1812) Kritische Ubersicht der einzelnen Arten aus der Gattung von Eidechsen, welche ich Wandkletterer nenne, Linné aber und andere, Geckonen. Denkschrift der Akademie der Wissenschaften, Mtinchen, 1812: 31-70. [Reprinted 1994, Fauna Exotica Reproduktionen, Mainz-Kostheim] Shaw G (1792): The single-striped lizard. Naturalist’s Miscel- lany 3: [1-2], pl. 89 Shaw G (1802) General Zoology, or Systematic Natural Histo- ry 3 (1) Amphibia. Davison, London, pp. 313-615 Strauch A (1887) Bemerkungen tiber die Geckoniden-Sammlung im Zoologischen Museum der Kaiserlichen Akademie der ©ZFMK 254 Herbert Rosler et al. Wissenschaften zu St. Petersburg. Memoires de |’ Académie Impériale des Sciences, St. Pétersbourg (7) 35: I-II, 1-72 Treu B (2001) Haltung und Zucht des Streifengeckos, Gekko vit- tatus Houttuyn, 1782. Sauria, Berlin, 23 (4): 31-35 Van der Hoeven J (1833): Handboek der dierkunde, of Grond- beginsels der natuurlijke geschiedenis van het dierenrijk. Tweede en Laatste Deel. Sulpke, Amsterdam, 698 pp. APPENDIX Gekko remotus sp. n. ZFMK 20611 (holotype) — male, Palau islands (= Republic of Palau); ZMB 5698 — female, Palau islands (= Republic of Palau); ZMB 5889 — male, Palau islands (= Republic of Palau); ZMB 6239 — juvenile, Palau islands (= Republic of Palau); ZMB 7922 — male, Palau islands (= Republic of Palau); ZMB 7982 — male, Palau islands (= Republic of Palau); ZMB 76976-76977 — male, female, Palau islands (= Republic of Palau); ZMB 76978-76981 — 4 juvenile, Palau islands (= Re- public of Palau). Gekko vittatus s. str. CPHR 393 — female, Dutch New Guinea; ZFMK 20612 — male,, Amboina (= Pulau Ambon), Indonesia; ZMB 7940 — ma- le, Wokan, Aru islands, Indonesia; ZMB 8788 — female, Mysore, Kordo (Schouten Group), Indonesia; ZMB 9457 — ma- le, Ratavul, New Britain, Bismark archipelago ago Papua New Guinea; ZMB 14628-14632 — | male, 2 females, 2 juveniles, Ralum, Gazelle peninsula, New Britain, Bismark archipelago ago Papua New Guinea; ZMB 14650 — female, Mioko, Duke or York Group, Papua New Guinea; ZMB 14651-14652 — 2 males, Ralum, Gazelle peninsula, New Britain, Bismark archi- pelago ago Papua New Guinea; ZMB 76982-76983 — 2 juve- niles, Ralum, Gazelle Halbinsel, Neubrittanien, Bismark Archi- pel, Papua New Guinea; ZSM 285/0/1 — male, Amboina Bonn zoological Bulletin 61 (2): 241-254 Wermuth H (1965) Liste der rezenten Amphibien und Reptilien. Gekkonidae, Pygopodidae, Xantusiidae. Das Tierreich, Ber- lin, 80: I-XXII, 1-246 Werner F (1900) Die Reptilien- und Batrachierfauna des Bis- marck-Archipels. Mitteilungen aus der Zoologischen Samm- lung des Museums ftir Naturkunde in Berlin 1 (4): 1-132 (=Pulau Ambon), Indonesia; ZSM 30/1972/1—30/1972/9 — 4 males, 5 females, Asmat, near Agats, Irian Jaya (Western New Guinea), Indonesia; ZSM 105/1979 — female, Mt. Bosavi, Didessa, southern highland, Papua New Guinea; ZSM 106/1979 — female, Airdhills, Papua New Guinea; ZSM 507/1998 — male, 54 km south from Nabire (800 m a.s.1.), Irian Jaya (= West- ern New Guinea), Indonesia. Gecko trachylaemus PETERS, 1872 ZMB 7511 (holotype) — male, Northern Australia. Platydactylus bivittatus (DUMERIL & BIBRON, 1836) MNHN 6714 (syntype) — male, Nouvelle-Guinée (= New Guinea), Indonesia, MNHN 2285 (syntype) — female, “le Wai- giou” (= Pulau W), Indonesia. Gekko vittatus- phenotype 3 ZMB 48737-48738 — 2 males, Kei islands, Moluccas, Indone- sia. Gekko vittatus- phenotype 4 SMF 9157-9159 — 3 males, Nissan atoll, Green Islands, Papua New Guinea. ©ZFMK Bonn zoological Bulletin 61 (2): 255-281 December 2012 The herpetofauna of the Bijagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist Mark Auliya, Philipp Wagner & Wolfgang Béhme ‘Helmholtz Centre for Environmental Research — UFZ, Department of Conservation Biology, Permoserstr. 15, D-04318 Leipzig, d Germany. ~ Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA. * Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn. Abstract. An annotated checklist of amphibians and reptiles from the Bijagos archipelago (Guinea-Bissau) with com- ments on the species’ distribution, systematics and natural history traits is presented here for the first time. During two field surveys 13 anurans and 17 reptile species were recorded from the archipelago of which several species represent either first records for the islands, i.e., Silurana tropicalis, Hemisus g. guineensis, Leptopelis viridis, Hemidactylus an- gulatus, Chamaeleo gracilis, Trachylepis perrotetii, Philothamnus heterodermus, Toxicodryas blandingii, Naja melanoleuca and Thelotornis kirtlandii or first country records, i.e., Amietophrynus maculatus, Ptychadena pumilio, P. bibroni, Phrynobatrachus calcaratus, P. francisci, Leptopelis bufonides, Hyperolius occidentalis, H. nitidulus, H. spatzi, Kassina senegalensis and Thrasops occidentalis. Species diversity reflects savanna and forest elements and a complete herpetofaunal checklist of the country is provided. Key words. West Africa, Guinea-Bissau, Bijagos archipelago, herpetofauna, first country records. INTRODUCTION The former Portugese colony Guinea-Bissau is an au- tonomous country since 1974 and is bordered by Senegal in the north, Guinea in the east and south, and by the At- lantic Ocean in the west (Fig. 1). It covers an area of ap- proximately 36,125 km? between 10°52' and 12°40' N and 13°38' and 16°43' W. According to recent data, its human population is estimated at 1.596.677 (July 2011, The World Factbook 2012). The country can be divided into three characteristic types of landscape (1) scattered plain islands together with the flooded valleys describe the coastal zone, (2) coastal estuaries or "rias" outlined with mangrove swamps extend deep into the continent on the main rivers Rio Cacheu, Rio Mansoa, Rio Geba, Rio Grande de Bu- ba and Rio Cacine, and (3) this second zone borders a moist savannah on a very low elevated shelf with the high- est peak of 310 ma.s.] in the southeast of Guinea-Bissau, the foothills of the Fouta Djallon. The Bijagos archipelago consists of 83 islands and islets with a surface of about 1,500 km7?, distributed across an area of about 11,000 km? (Fortes et al. 1998). According to IUCN (1991) 20 islands are permanently inhabited, 26 seasonal inhabited and 37 uninhabited. The islands soils are ferralitic, hydromorphic and halomorphic (Said & Fon- seca 1990). The four islands surveyed within the study are Orango (158 km?), Bubaque (43 km/?), Imbone (19 km’), and Soga Island (13 km*) (Fig. 1). Received: 03.04.2012 Accepted: 23.11.2012 Guinea-Bissau's tropical climate is characterised by a dry season (November to May), and a wet season from June to October with average annual rainfall between 1,500—2,000 mm (IUCN 1992), whereas on the islands and the southeast of the mainland annual precipitation increas- es to 2,250—3,000 mm (Joop 1968). The Sudan-Zone in the north, the Guinea-Congolian zone and the West African Forest Block, in the south, frame Guinea-Bissau. A Guinean forest-savanna mosaic is the characteristic ecoregion of the country. The moist tropical Guinea sa- vannas are of climatic origin, which are defined accord- ing to the length of the dry season, lasting two to six months. Of the edaphic-based vegetation, mangrove forests reflect the major formation. Originally mangrove forests represented 11 % of all vegetation of Guinea-Bis- sau, with 30 % of mangroves distributed on the Bijagos Islands. Dominant taxa are Rhizophora racemosa, R. man- gle, and Avicennia africana. On the islands of Orango and Bubaque, the inland veg- etation, behind the mangroves, is characterized by a belt of littoral bush (Limoges & Robillard 1991a) and floral elements have xeromorphic leaves and > 5 m tall e.g., Hi- biscus tiliaceus (Malvaceae). Chrysobalanus orbicularis (Chrysobalanaceae) and /pomoea pes-caprae (Convolvu- laceae) are interspersed with Baobab, Adansonia digita- ta (Bombacaceae) and dense undergrowth e.g., Strophan- tus spp. (Apocynaceae). Also of edaphic origin are palm savannas dominated by the oil palm (Elaéis guineeensis), Corresponding editor: F. Herder Go P = 2 w . iL, at a eu —_ Bs —= BISSA \ yy, we t om KS ar? MEG: ee cs vad Carache ne? das es ae? y, Soga Bio < Ta AOS fe Roxa Un G bi pd Bijagos- ag Bubaque Archipelago KR ° of Imbone V 50km Mark Auliya et al. oe N Eee. ee me cee ee ck ee em ey ee es Fig. 1. Map of Guinea-Bissau; Capital Bissau (black square); major Bijagos islands are named. the next characteristic vegetation belt when moving to- wards inland. These are bordered by "depressions" of moist grass savanna (Hyparrhenia, Andropogon). Also Raphia exica (Arecaceae) is a characteristic element of these "valleys" (Limoges & Robillard 1991a). Evergreen bush-forests (up to 8 m tall) follow the partly swampy grass savannas inland e.g., Mussaenda spp. (Rubiaceae) or Zrema guineensis (Ulmaceae) (Knapp 1973). Rich rain- fall indicates patches of primary forest, representing the natural vegetation cover. According to Limoges & Robil- lard (199 1a) these consist of half-moist and half-dry three- storied primary forests. In half-moist forests the upper sto- ry (canopy height around 30 m) is characterized by Chlorophora regia (Moraceae), and Afzelia africana (Cae- salpiniaceae). Elaiés guineensis is the dominant species in the story below, whereas the undergrowth comprises shrubs and palm-shoots. Apart from these moist forests, which are distributed in the southern archipelago, the northern islands also support half-dry primary forests char- acterized by Khaya senegalensis (Meliaceae) and Parinari excelsa (Chrysobalanaceae). There are close phylogenet- ic relationships to the coastal regions of Guinea, Sierra Leone and Liberia (belonging to the western Guinean low- land forests), thus Guinea-Bissau's flora is assigned to the Bonn zoological Bulletin 61 (2): 255-281 West African tropical Flora (IUCN 1986). Therefore, the flora of Guinea-Bissau consists of typical Sudano-ele- ments e.g., Acacia senegal, Dicherostachys cinerea (Mi- mosaceae) together with elements of the tropical lowland evergreen rain forests of West Africa e.g., Trichilia prieureana (Meliaceae). J. V. Barboza du Bocage published first data of Guinea-Bissau's herpetofauna in 1867. The majority of specimens originated from Bolama Isl. and Bissau (Bocage 1867, 1873, 1896a). Costa Martins, the Assistant Director of the Public Health Office of the Cabo Verde Ar- chipelago made one of these collections (Bocage 1896b). Later, between 1898 and 1908, Francisco Newton who collected for the Lisbon museum, conducted inventories of the mainland herpetofauna (e.g. Antula [Bissau], Bam- badinca, Buba, Geba, Farim, Sambel n'anta, including the nearby island Bolama Isl.). Boulenger (1905) identified and published an amphibian and reptile collection made by Leonardo Fea in 1898-1900. These specimens origi- nated from Bissau, Bolama Isl., Farim, Cacheu, Cassine and Cambec. J. Bethencourt Ferreira (1902) published a checklist of amphibians and reptiles collected by Francis- co Newton in 1900/1901. Some findings were recorded from Bolama Isl., the major data though originate from ©ZFMK The herpetofauna of the Bijagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist 57 Table 1. Habitat composition of the Biyagés islands. Source: Limoges & Robillard (1991). Habitat % of the archipelago Moist forest 0.6 Closed Palm forest 25.8 Open Palm forest 5.8 Bush land 3.6 Forest savannah 5.6 Grass savannah 1.4 Moist savannah 6.9 Cultivated savannah 0.9 Mangroves Sod the mainland. Almost 40 years later in 1937-1938, Dr. Al- bert Monard, perpetuated herpetological research of Guinea-Bissau publishing many new records (Monard 1940a, b). The most detailed herpetofaunal surveys, as a program of the Portuguese inventory missions "Junta de Investigacoes Colonais" were conducted in 1945-1946, supervised by Fernando Frade. These collections were ex- amined and published by Managas (1947, 1949, 1950), in particular with reference to lizards, while Frade (1950) published the research results of the Varanidae, Typhlop- idae, Leptyphlopidae and Colubridae. A small reptile col- lection was provided by Naurois (1969), who conducted a survey on the reproductive cycles of birds in the West African coastal zone from the Western Sahara to Guinea. He mentioned taxa from the islands Bubaque and Uno (Fig. 1). This collection was identified by Jean Guibé from the "Muséum National d'Histoire Naturelle" in Paris. Due to the isolation of the Biyagos islands, not includ- ing Bolama Island (close to the mainland), no detailed field studies document the herpetofauna of the archipel- ago so far. In May 1991, Benoit Limoges and Marie-Josée Robillard carried out the most recent inventory of the in- sular vertebrate fauna (excluding amphibians) with the fo- cus to establish protected areas (Limoges & Robillard 1991b). The present paper provides a first overview of the herpetofauna of the Bijagos archipelago including a check- list of all amphibians and reptiles recorded in Guinea-Bis- sau. MATERIAL AND METHODS Guinea-Bissau was visited twice; the initial survey was conducted in April and May 1993, while the second field survey was carried out mid May until mid August 1994, the transition from the dry to the wet season. The base- camp was on Bubaque Island, while three islands (Soga, Orango and Imbone) were visisted by motorboat. Due to Bonn zoological Bulletin 61 (2): 255-281 this fact most specimens originate from Bubaque. The is- land of Orango was visited for eleven, while Imbone on- ly for four days. All voucher specimens were sampled by visual en- counter, noosing and and setting nets during day and night excursions. Occasionally villagers captured specimens, or reported on taxa observed. Several taxa were collected dead after being killed by locals. Voucher specimens were preserved in 70 % ethanol and deposited in the Zoologis- ches Forschungsmuseum Alexander Koenig (ZFMK) in Bonn. For additional taxonomic analysis aforementioned herpetological collection of R. de Naurois (1969) in 1961/62, from the Muséum National d'Histoire Naturelle in Paris/ France, was taken into account. Comparatively, collections from the localities Macenta and Sérédou, both in Guinea, were analysed. RESULTS Annotated Herpetofaunal Checklist of the Bijagos ar- chipelago AMPHIBIA Arthroleptidae Leptopelis viridis (Gunther, 1869) Common name.— Savanna Tree Frog Specimens examined.— Three, ZFMK 58265: Orango Isl., 7. VII.1994. 2 uncatalogued live specimens, Orango Isl. & Bubaque Isl., 2. VII. 1994. Distribution— Schietz (1967) re-identified Boulenger's holotype of Leptopelis hyloides as a specimen of L. viridis. Thus, the type locality “Bolama” previously assigned to “L. hyloides” represents the first record for Guinea-Bis- sau. This study’s findings provide the first record from the archipelago and the second record for the country after 88 years (Fig. 2). Remarks.— The specimens were found in a dense second- ary forest patch, surrounded by water basins (presumably used for livestock), with diverse herbaceous layers and aquatic plants. All specimens were located on branches of shrubs (around 1m) adjacent to the water sites. Advertise- ment calls were produced between 20.00h and 0.00h. L. viridis was associated with Hyperolius occidentalis and Kassina senegalensis, and represented the only arboreal species within this anuran community. At the time of the study this species was tentatively identified as Leptopelis cf. hyloides (Boes| 1995). Leptopelis bufonides Schiotz, 1967 Common name. Ground Tree Frog. Specimens examined.— Five, ZFMK 58263—64, 58266—67: Orango Isl., 7. VH.1994. One uncatalogued live specimen, Bubaque Isl., 28. VII. 1994. ©OZFMK 258 Mark Aultya et al. Figs 2-9. 2. Leptopelis viridis, Bubaque Isl.; 3. Leptopelis bufonides, Orango Isl.; 4. Hemisus g. guineensis, Bubaque Isl.; 5. Hy- perolius occidentalis, Bubaque Isl.; 6. Hyperolius spatzi, Bubaque Isl.; 7. Hyperolius spec., Bubaque Isl.; 8. Kassina senegalensis, Bubaque Isl.; 9. Ptychadena bibroni, Bubaque Isl. Bonn zoological Bulletin 61 (2): 255-281 OZFMK The herpetofauna of the Bijagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist 259 Distribution— L. bufonides is restricted to West African savanna, and so far has been recorded from Senegal (Bohme 1978), the Gambia (Barnett et al. 2001), Ghana (Schiotz 1967) Benin (Gilles et al. 2006), Nigeria (Walk- er 1968) and northern Cameroon (Amiet 2004). The pres- ent records are the first from the archipelago and Guinea- Bissau (Fig. 3). Remarks— This second Leptopelis savanna species occurs in syntopy with L. viridis and interspecific competition can be ruled out as this species appears to display strictly ter- restrial behaviour, even calling from land (Gilles et al. 2006). Both localities of L. bufonides are characterised as savanna habitats. In one case (Orango Isl.) the habitat was arid-adapted vegetation dominated by grasses, no tree canopy, with some isolated Acacia trees. Three specimens were captured in a well. One frog, producing "advertise- ment calls", was located in a palm leaf about 2.5 m above the ground, and 5 m away from the coast (Imbone Isl.). On 8 July 1994 no fresh water could be located nearby. The species depends on temporary pools during the wet season. Bufonidae Amietophrynus maculatus (Hallowell, 1854) Common name— Hallowell's Toad Specimens examined.— Two, ZFMK 58268-—69: Eticoga (Orango Isl.); 12./13.VI.1994. Distribution— This study documents first records from Guinea-Bissau. However, a collection of 197 specimens of Amietophrynus regularis, made by Frade 1945/1946 (Managas 1949), could include A. maculatus. This toad was given species level rank by Hulselmans (1970), and isolated from the A. regularis-complex by Laurent (1972a). However, none of Frade's specimens originated from the archipelago. Remarks.— Both specimens were collected in the village Eticoga, Orango Isl., characterised by diverse plantations (Anacardium occidentale), and paddy fields. A depression adjacent to the village cottages represents a large pond in the wet season. Large parts of this island are character- ized by savanna-like vegetation, with many plants adapt- ed to dry periods. Hemisotidae Hemisus guineensis guineensis Cope, 1865 Common name.— Guinea Shovelsnout Frog Specimens examined. Two, ZFMK 58259: female, Eticoga (Orango Isl.), 7.VII.1994. ZFMK 58260: male, Bubaque Isl., 30. VII.1994. Distribution A first record from Guinea-Bissau was pro- vided by Boulenger in 1905, who referred to the species Hemisus marmoratus from Bolama Isl. close to the main- land (Laurent 1972b). The specimens in this study are the second record after 88 years, and represent the first record from the archipelago. Bonn zoological Bulletin 61 (2): 255-281 Remarks — ZFMK 58259 was captured by locals on 19.V1I.1994 (Fig. 4). For habitat descriptions refer to Ami- etophrynus maculatus. The remaining three specimens were collected in the morning on a moist shady slope. The habitat was at the edge of a depression, which most like- ly is filled with water during the wet season. Small pools were observed amid the clearing bordered by densely veg- etated slopes. The loose soil was a mixture of clays and sands, and was covered by a layer of rotten leaves inter- spersed with seedlings. Andersson (1937) records termites as stomach contents in the taxon H. marmoratus sudanen- sis. The specimens collected in this study were feeding on termites at daylight. When grasped the frogs inflated their lungs and pressed their bodies to the forest floor. This an- ti-predator adaptation could disguise the frog, making it difficult for a predator to identify it as a tetrapod. Spiel- er (1997) observed this behaviour in nest guarding fe- males. Hyperoliidae Hyperolius nitidulus Peters, 1875 Common name. Peter’s Reed Frog. Specimens examined.— One uncatalogued live specimen, Bubaque Isl., 10.VII.1994. Distribution— This is the first record from the archipela- go and Guinea-Bissau. Remarks.— R6del et al. (2010) have shown that this species is morphologically and genetically distinct from H. spatzi, and both species are here recorded from Guinea-Bissau for the first time. No advertisement calls were heard dur- ing the entire field survey, most likely indicating that the reproductive period had not begun. This assumed time shift might reduce interspecific competition with H. oc- cidentalis, with both species utilizing the same microhab- itats. Hyperolius occidentalis Schietz, 1967 Common name.— Western Reed Frog. Specimens examined. Five, ZFMK 58278—80 males, (12.-27.VII.1994); ZFMK 58281 (male), ZFMK 58282 (female), 10.VIII.1994; all Bubaque Isl. (also see Table 2). Distribution— This species record is the first from the ar- chipelago and from Guinea-Bissau (Fig. 5). Remarks.— According to villagers the secondary forest patch, where the species was recorded, was unaffected by fires for the previous 3—4 years. Slash-and-burn activities for rice cultivation, create clearings (“bolanhas”’) bordered by secondary forest. During the wet season they resem- ble shallow lakes, providing favourable living conditions for H. occidentalis, during both the dry and wet seasons. This species does not occur in primary forest with closed canopy or in Sudan-savanna formations, where second- ary or gallery forests are not present. For further habitat descriptions see Leptopelis viridis. The species was ob- ©ZFMK 260 served in syntopy with L. viridis and Kassina senegalen- sis. H. occidentalis occurs sympatrically with H. pictura- tus, the latter not having exposed calling sites (Schietz 1967). All specimens were localised in vegetation approx- imately 0.5 m above the ground when advertisement calls were recorded. The majority of specimens were observed on leaf surfaces or twigs of shrubs. During the dry sea- son and heat of the day, one light yellow specimen was observed on a palm leaf in the characteristic resting po- sition reducing evaporative water loss (Spieler 1997). Hyperolius spatzi Ahl, 1931 Common name. Ahl’s Reed Frog. Specimens examined.— One, ZFMK 58291: Bubaque Isl., 28.VI.—12.VII.1994. Distribution. The record of H. spatzi from Bubaque Isl. represents the first record for the country and thus the most southern distribution of the species range (Fig. 6). H. spatzi has been recorded in Senegal and in the Gambia (BOhme 1978; Rédel et al. 2010; Schiotz 1971). Remarks .— The live specimen was observed amid a sec- ondary forest patch on a white wall of a small abandoned house at approximately 2.5 m height. Bohme (1993) re- ported a similar habitat for the form nitidulus. According to Schigtz (1971), H. spatzi, as a typical savanna species, is associated with habitats that resemble those of H. oc- cidentalis (see above). Hyperolius spec. Specimens examined. One, ZFMK 58277: Bubaque Isl., 28.V1.1994. Distribution. This conspicuous phenotype does not re- semble any morph of the region (Senegal to Guinea) (Fig. 7). It has been recognized as a new colour morph, and is recorded for the first time from Guinea-Bissau, and may represent a new taxon. Remarks.— The specimen’s snout-vent length is 26 mm. Compared to the other three species, there are some strik- ing differences in colouration: more than half of the throat region is dotted and the belly is orange/red; dorsum dis- tinctly marbled with black markings on a white back- ground; interior part of the front limbs and femora, as well as the inside and underside of tibia and tarsus are coloured bright orange/red; iris is blackish. A villager collected the specimen from a secondary habitat, characterised by stands of E. guineensis and 3—5S m shrubs. The distinct pat- tern may be a form of mimesis, resembling bird faeces. Kassina senegalensis (Duméril & Bibron, 1841) Common name.— Running Frog. Specimens examined. — Two, ZFMK 58261—62: male & fe- male, Bubaque Isl., 13. VII.1994. Distribution. Locality records on the archipelago pres- ent the species’ northwesternmost distribution and are the first from Guinea-Bissau (Fig. 8). —_— Bonn zoological Bulletin 61 (2): 255-28 Mark Auliya et al. Remarks .— For habitat descriptions also refer to Leptopelis bufonides. The water basins (resembling troughs) meas- ured 3 m in length and 1.5 m in width; water depth was approximately 30 cm. The first heavy rains of the wet sea- son filled these basins. Initially this terrestrial species was acoustically recorded under dense creeping herbaceous plants (Convolvulaceae), providing excellent shelter. Ap- proximately 2 m high shrubs surrounded the troughs. Ad- vertisement calls were first recorded in early June, always after sunset, around 20h00. The characteristic “plopping sounds” were first introduced by a solitary male acting as the initiator of the population, as described by Herrmann (1993). Consequently, all other males commenced calling, answering previous calls, and producing a distinctive call overlap as is described by Grafe (1999). These calls drowned out calls of other species, i.e., Leptopelis bu- fonides and Hyperolius occidentalis. Calling males were well camouflaged in their microhabitat. Even though call- ing sites were located approximately 30 cm away, it was not possible to make out most of the individuals. During several capture attempts it was observed that the frogs se- cretively crept to other sites sheltered by the dense herb layers. Pipidae Silurana tropicalis Gray, 1864 Common name.— Tropical Clawed Frog Specimens examined.— 21, ZFMK 58253—254: two suba- dults, Bubaque Isl., 27.V1I.1994. ZFMK 58255—258: four adults, Imbone Isl., 08.VII.1994. ZFMK 58292: series of 15 larvae, Bubaque Isl., 29. V1.1994. Distribution. The island records within this study (from Bubaque and Imbone) represent the first records from the archipelago. Close to the mainland the species was record- ed from Bolama Isl. (Boulenger 1905), Bissau, Margo and Calequisse (Managas 1947, 1951a). Remarks.— The two subadult specimens and tadpoles were captured in a 1m? well amid dense secondary forest. The water in the well was clear, and depth was about 2 m. The site was about 10 m from a formation of Avicennia africana. The adult specimens were collected in deeper puddles in the only village on Imbone Isl. Both locations were almost closed by the canopy. Several behavioural ob- servations were made at the well and under captive con- ditions. The breathing procedure of the frogs lasted about 10 seconds. The frogs emerged almost vertically from the dark bottom of the well, and submerged in the same man- ner. Besides the occurrence of adult frogs tadpoles of dif ferent metamorphic stages were also observed. Nieuwkoop & Farber (in Arnoult & Lamotte 1968) de- scribed stage "45" without barbles, which was observed together with "tentacled stages" resembling fish schools, about 10-25 cm below the water surface. The tentacled tadpoles never had contact with the base of the well or with its steep edges. It was difficult to make out distinct swim- ©ZFMK The herpetofauna of the Bijagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist 261 ming movements of the tadpoles. Only slight undulations of the tail filaments were observed, and breath-taking was practised in the same way as the adult frogs. The translu- cent appearance of all larval stages may represent a pro- tection against predators from water and land. Predators from under water probably have difficulties in approach- ing an individual tadpole, due to their transparency and back-lit conditions. Besides, light reflection on the water surface also handicaps potential predators approaching from the land to follow the slow moving. Tadpoles and their post-metamorphic stages were kept to- gether in water tank. It was observed that subadult frogs fed on their tadpoles. The tank did not provide natural con- ditions, thus schooling behaviour was absent, and space was distinctly restricted. If stress behaviour was not re- sponsible for cannibalism under captive conditions, it may also be a natural feeding behaviour of the metamorphic frogs to feed on their larvae, thus to minimise cannibal- ism, schooling behaviour and transparency may play a cru- cial role. Cannibalism enables the parental population to "exploit the nutrient resource, the energy of primary pro- duction by algae and micro-organisms" (Tinsley et al. 1996). Also, taking into consideration that during the dry season permanent water bodies are rare and overland mi- gration probably is negligible, however in other Pipidae spp. i.e. Xenopus muelleri overland migration over long distances of has been observed (Loveridge 1953; in Tins- ley et al. 1996). Ptychadenidae Ptychadena bibroni (Hallowell, 1845) Common name. Broad-banded Grass Frog. Specimens examined.— Seven, ZFMK 58288 (female) and 58289 (male), Orango Isl., 15./16.VI.1994. ZFMK 58283-—87: one female (58283), all others males, Bubaque Isl., 19. VII.1994. Distribution. These island records also represent the first for the country (Fig. 9). Remarks — For habitat descriptions of the two specimens from Orango Isl. see Leptopelis bufonides. The remain- ing individuals were captured in a well in Bubaque vil- lage (approximately 30 m from the sea) between houses and secondary vegetation. The ZFMK specimens were collected between June and November similar to records of Gruschwitz et al. (1991). Active Ptychadena bibroni have been recorded during the first rains of the wet sea- son. Phrynobatrachus calcaratus (Peters, 1863) Common name. Boutry River Frog Specimens examined.— Two, ZFMK 58273, 58275: males, Bubaque Isl., 12. VII.1994. Distribution. This study records P. calcaratus from the archipelago and Guinea-Bissau for the first time based on findings provided by Perret (1988) (Fig. 10). Bonn zoological Bulletin 61 (2): 255-281 Remarks — Irregular but intense rainfalls brought out this species after the dry season in mid June. Unidentified froglets of this genus (ZFMK 58276) were also observed in early August within the anuran community described below (see Leptopelis viridis). For habitat descriptions on Bubaque Isl. see Ptychadena bibroni. Phrynobatrachus francisci Boulenger, 1912 Common name.— Warty River Frog. Specimens examined.— Six, ZFMK 58270-72: males, Orango Isl., 19.VI.1994. ZFMK 58274: male, Bubaque Isl., 12.VIL.1994. ZFMK 58276: male, Bubaque, 27.VII.1994. Distribution.— This study records P. francisci from the ar- chipelago and also represents the first record from Guinea- Bissau (Fig. 11). Remarks.— The collecting locality on Orango Isl. is de- scribed below (see Leptopelis bufonides). The specimens from Bubaque Isl. were found during the construction of a well in moist soil and in a well associated with Ptychade- na bibroni (see above). Shrubby vegetation with a few sheltering coconut palms amid some houses characteris- es the habitat. These findings support the fact that P. fran- cisci is found near both artificial and natural water bod- ies during the dry season. Compared to the nocturnal P. natalensis, P. francisci 1s explicitly diurnal and even he- liophilic, as characterized by Lamotte & Xavier (1966: 361). Gruschwitz et al. (1991) observed P. francisci at day- time under bark, litter and logs. Ptychadena pumilio (Boulenger, 1920) Common name.— Little Rocket Frog. Specimens examined.— One, ZFMK 58290: male, Oran- go Isl., 14.V1.1994. Distribution— The single specimen collected represents the first record from Guinea-Bissau. Remarks.— The voucher specimen was trapped in a well together with Leptopelis bufonides, Ptychadena mac- carthyensis, and Phrynobatrachus francisci. The well was surrounded by grass savanna, and according to Rédel (1995) and Schiotz (1999) all four species inhabit savan- na. The onset of the wet season initiated the first activi- ties of P pumilio. For detailed habitat descriptions see Lep- topelis bufonides. Descriptions of 31 specimens collect- ed on the mainland, published by Managas (1949), do not allow unequivocal species identification. Referring to the description of the lateral position of the vocal sac by Man- acas (1949), Ptychadena [bibroni| arnei is probably the species involved, however future examinations of that col- lection may prove to be helpful. OZFMK N lon N Mark Auliya et al. Figs 10-14. 10. Phrynobatrachus calcaratus, Bubaque Isl.; 11. Phrynobatrachus francisci, Orango Isl.; 12. Pelusios castaneus, Bubaque Isl.; 13. Hemidactylus angulatus, Orango Isl., 14. Lygodactylus gutturalis (above), Soga Isl. (below, throat pattern of male individuals). Bonn zoological Bulletin 61 (2): 255—281 ©OZFMK The herpetofauna of the Bijagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist 263 TESTUDINES Cheloniidae Caretta caretta (Linnaeus, 1758) Common names.— Loggerhead Turtle; P: Tartaruga caretta. Distribution— The species was observed throughout the entire archipelago (Limoges 1989) Remarks .— To date, however, nesting sites have never been recorded from the archipelago; the species 1s very rare and only one more record in 2003 provides evidence of the species roaming around the islands (Catry et al. 2009). Chelonia mydas (Linnaeus, 1758) Common names.— Green Turtle; P: Tartaruga verde; C: Tartaruga preto; B: Entchunko (Orango Isl.). Distribution— This is the most common marine turtle species of the archipelago (Limoges 1989, Limoges & Ro- billard 1991b). Remarks.— Major nesting sites are located on the islands of Poilao, Amegue, Meio, Cavalos, Jodo Vieira, Adonga, Orango and Porcos (Fortes et al. 1998). Eretmochelys imbricata (Linnaeus, 1766) Common names.— Hawksbill; P: Tartaruga verdadeira; C: Tartaruga vermelho; B: Djassaka (Orango Isl.). Distribution— The species was observed in the entire ar- chipelago (Limoges 1989). Remarks.— Nesting sites were found on Adonga Island (Limoges 1989), and the species has been observed on the main beaches of Poilao Isl., Unhocom and Porcos Isl. Lepidochelys olivacea (Eschscholtz, 1829) Common names .— Olive Ridley Turtle; P: Tartaruga de ri- dley; B: Emvara (Orango Isl.). Distribution— The species was observed throughout the entire archipelago (Limoges & Robillard 1991b). Remarks .— Major nesting sites are located on Orango Isl., lesser nesting activity occurs on Adonga and Poilao Island (Limoges & Robillard 1991b). Dermochelyidae Dermochelys coriacea (Vandelli, 1761) Common names.— Leatherback Turtle; P: Tartaruga de cou- ro; C: Tartaruga gigante; B: Djunumémeée. Distribution.— The species occurs throughout the archipe- lago (Limoges & Robillard 1991b). Remarks — Egg deposition was observed on the islands of Oran- go, Joao Vieira and Unhocomo (Limoges & Robillard 1991b). Pelomedusidae Pelusios castaneus (Schweigger, 1812) Common names.— East African Black Mud Turtle; P: Tar- taruga aquatica, Cagado preto; C: Tartaruga de agua do- ce; B: Epototo (Bubaque, Canhabaque), Nun-é (Galinhas, Formosa, Caravela), Iun-¢ (Orango). Bonn zoological Bulletin 61 (2): 255-281 Specimens examined.— Seven, ZFMK 60762-63, plus fi- ve uncatalogued live specimens (vouchers lost): Bubaque Isl., Orango Isl., 21.V1.1994, 11./13.VIL1994, 6. VIIL1994. Distribution.— Monard (1940b) recorded the species first from the mainland. Limoges (1989) and Limoges & Ro- billard (1991b) first report P. castaneus (as P. subniger) from the islands Enu and Joao Vieira Isl. and larger islands. Remarks — Specimens were collected in secondary forests with temporary water bodies ”lagunas”’. One specimen was found inside a watering tank. During the wet season (mid July to August), adult specimens could be observed in larger, permanent ponds. Next to the water line more individuals were found hidden beneath the grass. Hatch- lings and juveniles were examined and released on 6.VI- 11.1994 (Fig. 12). Carcasses of P. castaneus were found near water bodies. Commonly observed Palm-nut Vultures (Gyphohierax angolensis) may represent one potential predator. SAURIA Amphisbaenidae Cynisca feae (Boulenger, 1906) Distribution— The species was recorded from Bubaque Isl. (Gans 1987). On the mainland recorded from Rio Caci- ne, Bissau, Ponta de Marques, Bissalanca (Boulenger 1905, Managcas 1955). Gekkonidae Hemidactylus angulatus Hallowell, 1854 Common names.— West African House Gecko; P: Osga de brook; C: Oziga. Specimens examined.— Twelve, ZFMK 58299300: juve- niles, Bubaque Isl., 17. VII.— 9. VII.1994. ZFMK 58301: Eticoga (Orango Isl.), 18.VI.1994. ZFMK 58302: Soga Isl., 10. VII.1994. Distribution— Bocage (1896a) reported this gecko first from Bissau and Bolama Isl. A collection made by L. Fea published by Boulenger (1905) again included material from Bolama Isl., Ferreira (1902) mentions Geba, Monard (1940b) from Ponte Robalo, Mansoa, Sama, Pitche, Ma- dina Boé and Catio and Managas (1951b) additionally from Ponta de Marques, Bissalanca and Cacine. This study records the species from the Bijagos islands, Orango, Bubaque, and Soga for the first time. Remarks.— All specimens were collected on house walls and juveniles were commonly observed isolated from adults (Fig. 13). Under captive conditions on Bubaque Isl. cannibalism was observed. Lygodactylus gutturalis (Bocage, 1873) Common names.— West African Dwarf Gecko; P: Oziga; C: Lagarticha; B: Dongat. Specimens examined— Six, ZFMK 5829394: Eticoga ©ZFMK 264 Mark Aultya et al. (Orango Isl.), 18.VI.1994. ZFMK 58295—96: Imbone Isl., 8.VI.1994. ZFMK 58297: Soga Isl., 19.VII.1994. ZFMK 58298: Bubaque Isl., 16.V1.1994. Distribution.— Loveridge (1947) recorded the species first from Bolama Isl. and therefore the series reported here presents the second record from the archipelago. L. gut- turalis was described from the mainland (Nogueira) and was later also recorded from Contubo-el, Madina Boé and Pitche (Monard 1940b). Remarks.— Specimens were found in a variety of arbore- al habitats e.g., Acacia, Anacardium and Elaiés as well as in Ficus spp. in gardens, but also in human settlements on roofs and fences. Specimens can be individually iden- tified by their throat patterns (Fig. 14). The species was usually observed active from afternoon to sunset, occa- sionally also at noon. The Grey-headed Kingfisher (Hal- cyon leucocephala) preyed on this gecko. Other potential predators among the kingfishers include Halycon sene- galensis and Alcedo cristata. In addition, a captive held Chamaeleo gracilis on Bubaque Isl. was observed prey- ing on L. gutturalis. Agamidae Agama picticauda Peters, 1877 Common names.— West African agama; C: Lagarticha. Specimens examined.— Two, ZFMK 58314—15: Bubaque Isl., 20. VII.1994. Distribution Bocage (1896a) first recorded this species from Guinea-Bissau. Later, Manacas (1950) recorded A. agama from Formosa Isl., whereas Naurois (1969) found the species on Bubaque Isl. During his study, 4. agama was recorded from the islands Uno, Bubaque and Soga. Remarks.— We follow the taxonomic concept for Agama agama published by Wagner et al (2009) who restricted A. agama to Central Africa. Therefore the western popu- lations bear the nomen Agama picticauda. This species was more abundant in coastal areas, in ruines, but also on Ficus or Mangifera trees (Fig. 15). On one occasion the species was observed in a Avicennia tree, which at high tide was at the waterline. The species was never record- ed in treeless areas. Gravid and egg-laying females were observed at the end of July. Chamaeleonidae Chamaeleo gracilis gracilis Hallowell, 1842 Common names.— Graceful Chameleon; P: Camaleao; B: Nhanguti (Balanta). Specimens examined.— Two, ZFMK 58312—13: females, Eticoga (Orango Isl.), 17./21.VI.1994. Distribution— The species was first recorded from Guinea-Bissau by Bocage (1896a). Boulenger (1905) recorded the species from Bolama Isl., the first record from the island group. Limoges & Robillard (1991b) only men- tioned C. senegalensis from the archipelago, however; pre- sumably both species are represented in their collection. Bonn zoological Bulletin 61 (2): 255-281 During this study this species was recorded from Orango Isl. Remarks.— Specimens were only recorded in savannah- like habitats with single trees providing shelter. One spec- imen was collected on the beach, 3 m from coastal shrubs (Fig. 16). On 24.VI.1994 copulation was observed in cap- tivity on Bubaque Isl. Chamaeleo senegalensis Daudin, 1802 Common names.— Senegal Chameleon; P: Camaleaéo do Senegal; C: Camaleao. Distribution— On the mainland this chameleon was recor- ded from Rio Cacine (Boulenger 1905), Ponte Robalo, Mansoa, Contubo-el, Pitché, Catio, Cacheu (Monard 1940b); Bijimita, Bissoram, Mansoa (Managas 1951b); there is one record from Caravela Island (Limoges & Ro- billard 1991b). Remarks.— The species is considered to be rare, however it also occurs on other islands (Limoges & Robillard 1991b). Scincidae Trachylepis affinis (Gray, 1838) Common names.— Senegal Mabuya; C: Lagarticha; B: Dongat. Specimens examined.— Three, ZFMK 58305—06: Bubaque Isl., 25./29.V1.1994. ZFMK 58307: Eticoga (Orango Isl.), 18.VI.1994. Distribution.— Bocage (1896a) recorded this species first from Guinea-Bissau, Boulenger (1905) recorded it from Bolama Isl. and Naurois (1969) from Bubaque Isl. Remarks.— The species was observed in shaded and dense- ly vegetated habitats with shrubs and mango trees. 7. affi- nis was also observed foraging in the litoral zone (Fig. 17). Trachylepis perrotetii (Duméril & Bibron, 1839) Common names .— Teita Mabuya; C: Lagarticha; B: Don- gat. Specimens examined. Two, ZFMK 58303: Soga Isl., 14. VIL.1994. ZFMK 58304: Bubaque Isl., 8. VII.1994. Distribution.— Initially, Bocage (1872) recorded this scin- cid from Guinea-Bissau and the same author (Bocage 1896a) provided the Bolama record. In this study 7! per- rotetii was first recorded from the islands Bubaque and Soga. Remarks.— The species utilizes exposed structures e.g., ter- mite mounds or logs in various forested habitats such as remnants of virgin humid and drier forests, forest edge habitats, secondary forests aswell as gardens and vegetat- ed roadsides. Varanidae Varanus exanthematicus (Bosc, 1792) Common names .— Savannah Monitor; P: Linguana de ma- to; C: Cutd; B: Cagueda (Bubaque, Canhabaque, Formo- ©ZFMK The herpetofauna of the Bijagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist sa, Orango), Cagada (Caravela), Caguedje (Galinhas). Distribution— On the mainland this monitor lizard was recorded from Bissau and nearby Bolama Isl. (Bocage 1896a; Boulenger 1905), on Buba (Ferreira 1902) and Bissalanca, Canchungo, Pecixe and Pitche (Managas 1955). The species occurs on several islands of the archi- pelago (Limoges & Robillard 1991b). Although the au- thors do not mention specific islands, several local island names of this species are available (see above). Remarks.— This species 1s less common than V. niloticus (Limoges & Robillard 1991b). Varanus niloticus (Linnaeus, 1758) Common names.— Nile Monitor; P: Linguana de agua; C: Linguana; B: Cagueda, Caguedje, Cagada. Specimens examined.— Five, ZFMK 58308—09: Eticoga (Orango Isl.), 12.VI.1994; ZFMK 58310: Soga Isl., 14. VII.1994. One uncatalogued live specimen, Bubaque Isl., 2. VIII.1994; 1 uncatalogued live specimen Bissau, 10.VHI. 1994. Distribution— V. niloticus was first recorded from Guinea- Bissau by Bocage (1896a). Manacas (1955) noted the species from Formosa Isl., the first record from the archi- pelago. Naurois (1969) reported V. niloticus from Bubaque Isl. Limoges (1989) also mentioned Varela Isl. and Limo- ges & Robillard (1991b) indicated that the species inhab- its the entire archipelago including smaller islands. Remarks.— The species was only found in coastal areas. Adult specimens were observed in densely vegetated steeper slopes near the coast. Juveniles were observed next to a freshwater pond (Fig. 18). In 1993, one juvenile was observed in a saltwater pond. Locals consume both, the eggs and the animals. SERPENTES Colubridae Philothamnus heterodermus (Hallowell, 1857) Common name.— Variable Green Snake Specimens examined. One, ZFMK 58332: Orango Isl., 13.VI.1994 Distribution— Monard (1940b) reported the species first from Guinea-Bissau. This voucher is the first record from the archipelago and represents the westernmost range of the species. Remarks The specimen was found dead next to a path in a dense stretch of coastal forest. Thelothornis kirtlandii (Hallowell, 1844) Common name.— Forest Vine Snake. Specimens examined.— One, ZFMK 60764: Bubaque Isl., 27.VII.1994. Distribution. This voucher documents the first record of the species from the archipelago (cf. Riquier & Bohme Bonn zoological Bulletin 61 (2): 255-281 265 1996) (Fig. 19); Loveridge (1944) indicated only ‘Por- tuguese Guinea”. . Remarks .— The specimen was detected at a height of 34 m in a Mangifera tree surrounded by secondary forest. An alerted sunbird (Cinnyris cf. cupreus) indicated the pres- ence of the snake, which was approaching the sunbird’s nest. Thrasops occidentalis Parker, 1940 Common name.— Western Black Tree Snake Specimens examined.— One, ZFMK 58331: Bubaque Isl., 12. VIII.1994. Distribution — This record is the first from the archipela- go and Guinea-Bissau and presents the northwesternmost locality within the species range (Fig. 20). Remarks— The voucher was killed by locals in a tradition- al village with mud huts interspersed with shrubs. The first third of specimen shows light brown lateral bands on both sides, and the anal scale is entire. Toxicodryas blandingii (Hallowell, 1844) Common names.— Blanding’s Tree Snake Specimens examined. One, ZFMK 61093: Bubaque Isl., 4. VIII.1994. Distribution— This snake has been documented from Bo- lama Isl. (Bocage 1896a, b) and Bissau (Managas 1955). The Bubaque Isl. voucher specimen represents the first record from the archipelago (Fig. 21). In addition, one specimen was observed on Soga Isl. Remarks.— The voucher specimen was found during a night survey on the roadside amid dense shrubs of sec- ondary bush and palm vegetation. The specimen from So- ga Isl. was observed in an oil palm at a height of approx- imately 8 m. The taxonomic status of the species remains unresolved; the species is also treated in the genus Boiga (e.g. Pauwels & Vande weghe 2008). Elapidae Dendroaspis viridis (Hallowell, 1844) Common names.— Western Green Mamba; P: Cobra ver- de de palmeira; C: Cacdba; B: Ianne (Orango Isl.), Ed- janro (Caravela Isl.). Distribution— Naurois (1969) recorded this species from Bubaque Isl., Managas (1981) further mentions Bolama Isl., whereas Limoges & Robillard (1991b) indicate that the species occurred on all larger islands. Remarks.— According to Limoges & Robillard (1991b) the species is particularly associated with raffia palms (Raphia exica) commonly dominant in swamp forests. Elapsoidea semiannulata moebiusi (Werner, 1897) Common names.— Angolan Garter Snake Distribution— On the islands this species has only been reported from Bubaque Isl. (Naurois 1969). Manacas (1981) provided a second record from Bubaque Isl. OZFMK 266 Mark Auliya et al. Figs 15-22. 15. Agama picticauda, left male, right female; 16. Chamaeleo gracilis, Orango Isl.; 17. Trachylepis affinis, Bubaque Isl.; 18. juv. Varanus niloticus, Bubaque Isl.; 19. Thelotornis kirtlandii, Bubaque Isl.; 20. Thrasops occidentalis, Bubaque Isl.; 21. Toxicodryas blandingii, Bubaque Isl.; 22. Naja melanoleuca, Bubaque Isl. Bonn zoological Bulletin 61 (2): 255-281 ©OZFMK The herpetofauna of the Bijagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist 267 Loveridge (1944) indicates mainland locations in Bissau and Rio Cacine. Remarks.— Voucher specimens are deposited in the col- lections of the Muséum nationale d’ Histoire Naturelle de Paris (MNHN 1965-463). Loveridge (1944) lists the species as Elapsoidea sundevalli gtintherii. Naja melanoleuca Hallowell, 1857 Common names.— Forest Cobra; C: Bida; B: Cadjipon. Specimens examined.— One, ZFMK 58329: Bubaque Isl., 1. VIL.1994. Distribution The first record from Guinea-Bissau was provided by Monard (1940b), and this voucher is the first record from the archipelago (Fig. 22). A second specimen was observed on Orango Isl. Remarks.— The voucher specimen was found at midday in a depression surrounded by slopes with dense vegeta- tion. A second specimen was observed in a palm forest with dense scrub. Naja nigricollis nigricollis Reinhardt, 1843 Common names.— Black-necked Spitting Cobra; P: Cus- pideira; C: Vida; B: Cadjipon Specimens examined.— One, ZFMK 58330: Bubaque Isl., 2.VIII.1994. Distribution— Bocage (1896a) documented the first record from Bolama Isl., and Manacas (1981) added Bis- sau and Buba. Limoges & Robillard (1991b) first record the species far off the mainland on the archipelago. Remarks.— The voucher was killed around midday by lo- cals in secondary bushland on a densely vegetated slope near the coast (Fig. 23). Lamprophiidae Boaedon fuliginosus (Boie, 1827) Common name.— Brown House Snake Distribution— Boulenger (1905) reports the species on the mainland from Cacine, Manacas (1955) from Bissau, Bissalanca and Bijimita. The only record from the archi- pelago is provided by Naurois (1969) who reports the species was collected 1962 from Bubaque Isl. Remarks.— Hallermann & Schmitz (2007) indicated that the “Lamprophis fuliginosus” complex includes several cryptic species. Most recent taxonomic research suggests the resurrection of the generic name Boaedon Duméril, Bibron & Duméril, 1854 for the widely distributed fulig- inosus complex (Kelly et al. 2011), a view which is fol- lowed herein. Psammophis phillipsi (Hallowell, 1844) Common names .— Phillip’s Sand Snake; P: Cobra amon- ton. Specimens examined.— Five, ZFMK 58316—17: Bubaque Isl., 26./27. VI. 1994. ZFMK 58318: Bubaque Isl. 20. VII. Bonn zoological Bulletin 61 (2): 255-281 1994. ZFMK 58319: Bubaque Isl., 15. VII. 1994. ZFMK 58320: Bubaque Isl., 2. VIII. 1994. Distribution— Naurois (1969) reported the species from Uno Island. This study recorded the species based on vouchers from Bubaque Isl. (Fig. 24) and one specimen was observed on Soga Isl. Remarks.— The species was recorded during daytime in secondary bushland interspersed with E. guineensis, in agroecosystems and home gardens. Two specimens were observed preying on a female Agama picticauda and Tra- chylepis affinis. Both species have been found to repre- sent the most common prey items of P. phillipsi (Akani et al. 2003). Pythonidae Python sebae (Gmelin, 1789) Common names .— Northern African Python; P: Giboia; C: Ira Cego; B: Emeno, Emeni (Orango Isl.). Specimens examined.— Four, ZFMK 58333: Soga Isl., 14.VII.1994. Three uncatalogued live specimens, Buba- que Isl., 13./14./23.VI.1994 (Fig 25). Distribution— Bocage (1896a) was the first to record P. sebae from Guinea-Bissau while Naurois (1969) first re- ported the species from the archipelago, without provid- ing any specific locality. Limoges & Robillard (199 1b) in- dicated that this species ranges on all larger and some smaller islands. In this study the species is documented by vouchers from Soga Isl. and Bubaque Isl., one skin owned by locals on Orango Isl. was observed and a skele- ton including a jawbone was found on Imbone Island. Remarks. Limoges & Robillard (1991b) recorded the snake from agroecosystems, salt marshes and mangrove swamps, usually near water bodies. Voucher specimens were collected in secondary bushland, rice fields and near mangrove swamps. Limoges & Robillard (1991b) report- ed P. sebae as a predator of bird colonies, also on very small islands. This may prove that the species is capable of crossing greater marine passages between the islands. According to locals, domestic fowl and livestock also rep- resent prey items of P. sebae. In one event, a python de- voured a sleeping child in a hut (IUCN staff, pers. comm.). P. sebae is always killed by locals if it enters a village. The meat is consumed, and it is believed that it heals lep- rosy. Skins are sold in local markets. Viperidae Bitis arietans arietans Merrem, 1820 Common names.— Puff Adder; P: Surucucu; C: Tutu; B: Evé-ve. Specimens examined— Two, ZFMK 58327: Soga Isl., 15.VII.1994. ZFMK 58328: Bubaque Isl., 17.VII.1994. Distribution— Bocage (1896a) documented the species first from Bolama Island. Limoges & Robillard (1991b) reported the species from the archipelago, but did not pro- OZFMK 268 Mark Auliya et al. Figs 23-31. 23. Naja n. nigricollis, Bubaque Isl.; 24. Psammophis phillipsi, Bubaque Is].; 25. Python sebae, Bubaque Isl.; 26. Bitis a. arietans, Soga Isl. ; 27. juv. Crocodylus suchus, Bissau Zoo; 28. Latastia ornata, Bafata, mainland ; 29. Tarentola ephip- piata senegambiae, Mansoa, mainland; 30. Lycophidion albomaculatum, Bissau; 31. Python regius, Bissau. Bonn zoological Bulletin 61 (2): 255-281 ©OZFMK The herpetofauna of the Bijagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist Table 2. Colour morphs of six live Hyperolius occidentalis. Specimens No. 1 No. 2 No.3 No.4 No.5 No.6 Locality Orango Bubaque Bubaque Bubaque Bubaque Sex Female? Female? Male Male Male Male? SVL (mm) 33 30-32 28-29 30 28-29 25 Throat Yellow Yellow Intense yellow Yellow Dark yellow ~ Belly Yellow White with Light yellow Yellow Yellow Yellow small yellow with white mark mark in the centre Fore limbs Interior - = Only pigmented Joints without = light yellow above, interior pigmentation yellow Fingers 1 and 2 yellow — 1 and 2 yellow 1 and 2 - 1 and 2 yellow less pigmented, including 269 Discs / fingers = - - Hind limbs Backside of thighs Underside of (concerning thighs also of thighs, orange/red parts) with scarce underside yellowish of tarsus pigmentation Toes 1, 2 and 3 are red - — including the corresponding webbing Discs / toes red, 4 and 5 ~ 4 reddish more pigmented, dark red Dorsal-lateral — = ss line/stripe Inside and outside Posterior, interior light yellow the web = 1 not pigmented, — yellow, 2 slightly pigmented dark yellow, 3 and 4 pigmented Upper side of thigh Upper side of framed with red and underside of thigh slightly thigs. femur, tibia, spot and dark pigmented, and tarsus yellow pigmented, underside of outside of thigh with 3x larger inside, towards tarsus (1, 2, and 3 anus yellowish toe), underside mottled red mark of thighs with compared to larger upper side yellowish mottling 1, 2 and 3 less in front of 3 toe 1, 2 and 3 pigmented, red mark including the little reddish and webbing red, whitish 4 and 5 pigmented uf all dark, 1, 2, all red and 3 lighter red Distinctly Distinctly Scarcely pronounced pronounced pronounced vide explicit locality records. Garcia (1991) provided a first record from Bubaque Island. In this study, B. arietans was recorded from the islands Soga and Bubaque. Remarks. Specimens from both islands Orango and Bubaque were detected in savannah-like habitats period- ically altered through slash-and-burn farming practises. On Soga Isl. a dead juvenile specimen was found in dense sec- ondary bush and palm forest (Fig. 26). Bonn zoological Bulletin 61 (2): 255-281 Bitis nasicornis (Shaw, 1802) Common names.— Rhinoceros Viper; P: Vibora cornuda Distribution Apparently only known from Bubaque Isl., where a juvenile and adult have been recorded (Naurois 1969) Remarks.—Manacas (1981) indicates that Schmidt (1933) provides a quote by Boulenger, who states that the species occurs in Guinea-Bissau. This viper occurs in diverse ©OZFMK 270 Mark Aultya et al. ecosystems from lowland savannah to high altitude forest- ed ecosystems (Angel et al. 1954). Crocodylidae Crocodylus suchus Geoffroy, 1807 Specimens examined.— One, ZFMK 58334: dead juv., Bis- sau Zoo (Fig 27). Distribution — A first record from Bolama Isl. is provid- ed by Bocage (1896b). Populations are recorded from the islands Orango, Formosa, Imbone and Uno (Limoges & Robillard 199 1a, b). Remarks.— Limoges & Robillard (1991b) reported an in- crease of populations from the islands mentioned above. West and central African populations proved to have high genetic divergence from eastern populations of the Nile Crocodile (Schmitz et al. 2003). Osteolaemus tetraspis tetraspis Cope, 1861 Common names.— Dwarf Crocodile; P: Crocodilo anao; C: Lagarto preto; B: Eneguebe (Bubaque, Canhabaque), Etchega (Galinhas, Formosa), Eod (Orango), Equetch (Caravela). Distribution.— Bocage (1867) reports one specimen from Bissau. Waitkuwait (1985) could not provide any locali- ty records from Guinea-Bissau, however, according to Methot (1989, in litt. Limoges 1989) remnant populations probably occur on the islands Formosa and Orango. Remarks.— Similarly as in C. suchus Limoges & Robil- lard (1991b) reported an increase of populations from Orango Island. DISCUSSION Available reports on amphibians or reptiles recorded in Guinea-Bissau are scarce. Preliminary studies go back to Bocage (1866, 1867, 1872, 1873, 1888, 1896a, b), Monard (1940a, b) and Manacas (1947, 1949, 1950, 1951a). Oth- er reports provide more general data of the entire verte- brate fauna as well as essays on the herpetofauna (e.g., Frade 1950; Naurois 1969; Limoges 1989; Limoges & Ro- billard 199 1a, b). Latest herpetological published records mainly refer to sea turtles, in particular to Chelonia my- das (Catry et al. 2002; Fortes et al. 1998; Godley et al. 2003). The present study demonstrates that at least 13 amphib- ian species occur on four of the main Bijagos islands, ten of these represent first records for the country. Therefore, a total of 25 amphibian species are now known from Guinea-Bissau (App. 1). Five marine turtle species and 26 reptile species (including two crocodilians) occur on the archipelago. Alike amphibians, reptile diversity is clearly lower on the islands compared to the mainland (App. 1). However, the present study has clear limits and most likely covers only parts of the herpetofauna of the islands. Bonn zoological Bulletin 61 (2): 255-281 The phenomen of lower species richness on the islands was also recognized by other authors (e.g., Mertens 1964) and may relate to the islands distinctly smaller land sur- faces, isolation from the mainland, lacking elevated ecosystems and freshwater wetlands or it is correlated with forest cover, habitat diversity and rainfall (Leaché et al. 2006, Rodel et al. 2008). The differences in species rich- ness are more distinct in amphibians, and naturally dis- persed colonization is more restricted although possible via rafting and potentially with birds (egg clusters stick to feathers) (cf. Measey et al. 2007; Queiroz 2005), and with few species 1.e. Ptychadena mascareniensis indicat- ing a certain tolerance to salinity (Vences et al. 2004). However, the translocation of amphibians through humans appears to be the most common mechanism of dispersal for species roaming oceanic islands (Vences et al. 2003). The vegetation of the Bijagos archipelago comprises both, Guinean/Congolian and Sudano-Sahel elements, which is reflected in its herpetofauna. Typical savannah species (e.g., Hemisus guineensis, Trachylepis perroteti and Bitis arietans) are recorded from the islands as well as typical rainforest dwellers (Si/urana tropicalis, Bitis na- sicornis and Thelotornis kirtlandii). In some species, e.g., Kassina senegalensis and Thrasops occidentalis differ- ences in morphology between island and mainland pop- ulations have been recognized and further research is nec- essary to clarify the taxonomic status of these island pop- ulations. Also, the Hyperolius sp. may represent an endem- ic species. This is important as the islands are impacted by various threats, in particular by illegal logging for com- mercial timber and slash-and-burn shifting agriculture and therefore many species could be threatened before they become known to science. Acknowledgements. We are grateful to the following people: U. Bott prepared the map to illustrate the archipelago; E. Fis- cher provided relevant references and identified herbarized plant material. The “Muséum National d'Histoire Naturelle" in Paris provided voucher specimens. Many thanks to P. Campredon, C. Cassama, G. da Costa, L. Giindling, P. K. Mendy, R. Miranda, B. Paris, J.-Y. Pirot A. Rachid, for the logistical and bureaucra- tic support. The following people provided valuable or rare li- terature: P. Catry, E. Crespo, F. Reiner and S. Trape. During both trips N. Riquier provided enormous stamina, patience and indis- pensable assistance — a very special thanks to her. REFERENCES Adalsteinsson SA, Branch WR, Trape S, Vitt LJ, Hedges SB (2009) Molecular phylogeny, classification, and biogeography of snakes of the Family Leptotyphlopidae (Reptilia, Squama- ta). Zootaxa 2244: 1-50 Akani GC, Eniang EA, Ekpo IJ, Angeleci FM, Luiselli L (2003) Food Habits of the Snake Psammophis phillipsi from the Con- tinuous Rain-Forest Region of Southern Nigeria (West Africa). Journal of Herpetology 37: 208-211 ©ZFMK The herpetofauna of the Byagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist 27] Amiet JL (2004) A propos de deux Leptopelis nouveaux pour la faune du Cameroun (Anura, Hyperoliidae). 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Copeia: 164-167 Wanger TC (2005) The Amphibians of Kiang West National Park, The Gambia. Salamandra 41: 27—33 Wermuth H (1967) Liste der rezenten Amphibien und Reptilien: Agamidae. Das Tierreich 86: 1-127 The World Factbook (2012) Washington, DC: Central Intelli- gence Agency, 2012. https://www.cia.gov/library/publica- tions/the-world-factbook/index.html ©OZFMK The herpetofauna of the Bijagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist 275 APPENDIX | Checklist of the herpetofauna of Guinea-Bissau. AMPHIBIA Artholeptidae Arthroleptis poecilonotus Peters, 1863 Localities — Guinea-Bissau. References — Managas (1949, 1950). Remarks — The species is recognized as a member of a spe- cies complex (Rédel & Bangoura 2004); the taxonomic status of A. poecilonotus in Guinea-Bissau requires veri- fication. Leptopelis bufonides (Gunther, 1868) Localities — Orango Isl., Bubaque Isl. References.— Present study. Leptopelis viridis (Boulenger, 1906) Localities.— Bolama Isl., Orango Isl., Bubaque Isl. References. Bocage (1896a), Boulenger (1905), Mana- cas (1949), present study. Remarks.— Managas (1949, 1950) published a record from Bolama Isl. as Leptopelis bocagei (Gunther, 1865), but the occurrence of this species in western Africa is questiona- ble as the westernmost confirmed record is from Central Africa (eastern Cameroon) (Rédel 2000). Therefore we suggest that this specimen might refer to L. viridis. Bufonidae Amietophrynus maculatus (Hallowell, 1854) Localities — Orango Isl. References.— Present study. Amietophrynus regularis (Reuss, 1833) Localities.— Bissau, Ponta do Marques Mano, Bissalan- ca, Tor, Bissoram; Enxalé, Contubo-el, Catio, Cacheu, Bo- lama Isl., Pecixe, Mansoa, Cacine, Pitche, Orango Isl. References Monard (1940a), Manacas (1949, 1950), Naurois (1969). Hemisotidae Hemisus guineensis Cope, 1865 Localities.— Bolama Isl., Bubaque Isl., Orango Isl. References. Boulenger (1905), Managas (1949, 1950), present study. Hyperoliidae Hyperolius concolor (Hallowell, 1844) Localities. — Ponta de Marques Mano, Cutia, Pitche, Bo- lama Isl., Ponta Robalo, Bissalanca, Bissau, Sansanto (Mansoa). References Boulenger (1905), Managas (1949, 1950), Monard (1940a), Naurois (1969). Remarks.— Hyperolius Bonn zoological Bulletin 61 (2): 255-281 guineensis Ahl, 1931 is considered as a synonym of H. concolor and was listed next to H. concolor by Monard (1940a), Managas (1949, 1950), Naurois (1969). Hyperolius nitidulus Peters, 1875 Localities.— Bubaque Isl. References.— Present study. Hyperolius occidentalis Schiotz, 1967 Localities — Bubaque Isl. References.— Present study. Hyperolius spatzi Ahl, 1931 Localities. — Bubaque Isl. References.— Present study. Kassina senegalensis (Duméril & Bibron, 1841) Localities.— Bubaque Isl. References.— Present study. Ranidae Hoplobatrachus occipitalis (Giinther, 1858) Localities.— Ponta de Marques Mano, Bissalanca, Tor, Pra- bis, S. Domingos, Gabu, Bafata, Canchungo, Calequisse, Pecixe, Bissoram, Mansoa, Catid, Bolama Isl., Antula, Bambadinca, Sambel n’anta, Farim, Buba, References — Ferreira (1902), Boulenger (1905), Monard (1940a), Manacas (1949, 1950). Hylarana galamensis (Duméril & Bibron, 1841) Localities.— Bissau, Catid, Cacheu References. Monard (1940a), Manacas (1949, 1950). Pipidae Pseudhymenochirus merlini Chabanaud, 1920 Localities. Pitche (Gabu), Contubo-el References— Monard (1940a), Manacas (1947, 1950, 195 1a) Silurana tropicalis (Gray, 1864) Localities.— Wha de Bissau, Bolama Isl., Calequisse (Can- chungo), Bubaque Isl., Imbone Isl. References. — Managas (1947, 1950, 1951a), present study Phrynobatrachidae Phrynobatrachus calcaratus (Peters, 1863) Localities.— Bubaque Isl. References.— Present study. ©OZFMK 276 Mark Auliya et al. Phrynobatrachus francisci Boulenger, 1912 Localities — Orango Isl., Bubaque Isl. References.— Present study. Phrynobatrachus minutus (Boulenger, 1895) Localities — Bolama Isl. References.— Managas (1949, 1950). Remarks.— listed as Arthroleptis minutus; Phrynobatra- chus minutus is endemic to Ethiopia, congeners of Arthroleptis recorded from Guinea-Bissau remain valid species (see below). Phrynobatrachus natalensis (Smith, 1849) Localities — Pitche. References.— Monard (1940a), Managas (1949, 1950). Phrynobatrachus tokba (Chabanaud, 1921) Localities — Dandum (Boé). References.— Monard (1940a), Managas (1949, 1950). Remarks — Listed as Arthroleptis “tobka”. Ptychadenidae Ptychadena ansorgii (Boulenger, 1905) Localities.— Cacine. References.— Managas (1949, 1950). Ptychadena bibroni (Hallowell, 1845) Localities — Ponte de Marques Mano, Bissoram, Mansoa, Madina-Boé, Caira, Catiod, Orango Isl., Bubaque Isl. References Monard (1940a), Manacas (1949, 1950), Naurois (1969), present study. Ptychadena mascareniensis (Duméril & Bibron, 1841) Localities. — Bissau, Ponta Robalo, Ponta de Marques Ma- no, Bissoram, Prabis; Catio, Enxalé, Contubo-el, Pitche, Bolama Isl., Gabu, Madina do Boé, Caira, Cacine, Man- soa, Cutia. References.— Bocage (1896a), Boulenger (1905), Mana- cas (1949, 1950), Monard (1940a), Naurois (1969). Ptychadena oxyrhynchus (Smith, 1849) Localities — Bolama Isl. References.— Managas (1950). Ptychadena pumilio (Boulenger, 1920) Localities — Orango Isl. References — Present study. Reptilia Testudines Cheloniidae Caretta caretta (Linnaeus, 1758) Localities — Bijagos archipelago. References.— Limoges (1989), Catry et al. (2009). Bonn zoological Bulletin 61 (2): 255-281 Chelonia mydas Linnaeus, 1758 Localities — Bijagos archipelago. References.— Bocage (1896a), Limoges (1989), Limoges & Robillard (1991b). Eretmochelys imbricata (Linnaeus, 1766) Localities. — Bijagos archipelago. References— Limoges (1989), Limoges & Robillard (199 1b). Lepidochelys olivacea (Eschscholtz, 1829) Localities — Bijagos archipelago. References. Limoges & Robillard (1991b). Dermochelyidae Dermochelys coriacea (Vandelli, 1761) Localities.- Bijagos archipelago, mainland References.- Limoges & Robillard (1991b), Catry et al. (2009) Pelomedusidae Pelomedusa subrufa (Lacepéde, 1788) Localities. Cufada, Cufar, Catio, Cacine. References.— Garcia (1991). Pelusios castaneus (Schweigger, 1812) Localities.— Bissau, Ponte Robalo, Catio, Cacheu, Cata- nhez, Enu, Joao Vieira, Bubaque Isl., Orango Isl. References. Bocage (1867), Broadley (1981), Garcia (1991), Monard (1940b), Limoges (1989), Limoges & Ro- billard (1991b), present study [party mentioned as Pelu- sios subniger]. Testudinidae Kinixys belliana nogueyi (Lataste, 1886) Localities — Bolama Isl. References.— Bocage (1896a), Garcia (1991). Kinixys homeana Bell, 1827 Localities — Cajadude, Boé. References.— Limoges (1989), Garcia (1991). Trionychidae Cyclanorbis senegalensis (Duméril & Bibron, 1835) Localities — Cufada. References.— Present study. Trionyx triunguis (Forskal, 1775) Localities. — Bissau, Corubal, Cufada References. Bocage (1867), Limoges (1989), Garcia (1991). Remarks .— Listed by Bocage (1867) as Gymnopus aegyp- tiacus. ©OZFMK The herpetofauna of the Bijagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist 277 SAURIA Amphisbaenidae Cynisca feae (Boulenger, 1906) Localities. — Rio Cacine, Ponta de Marques Mano, Bissa- lanca, Bubaque. References.— Gans (1987), Managas (1955). Cynisca leonina (Miller, 1885) Localities. Rio Pongo. References.— Loveridge (1941), Gans (1987). Cynisca oligopholis (Boulenger, 1906) Localities.— Rio Cacine. References — Loveridge (1941), Gans (1987). Agamidae Agama boensis Monard, 1940 | Agama sankaranica Chabanaud, 1918 Localities — Madina Boé, Pitche. References— Managas (1951b), Grandison (1968), Joger (1979). Remarks— According to Grandison (1968), Monard (1940b) described Agama boensis on the basis of a series intermingled with adult specimens referring to A. sankaranica and juveniles referring to A. weidholzi. Lat- er, A. boensis is often recognized as subspecies of Aga- ma agama (e.g. Wermuth 1967), which is in fact very doubtful. Managas (1951b) collected three specimens as- signed to to A. boensis while Wagner (2010) recognized A. boensis as valid species. In this context A. boensis should be added to the herpetofauna of Guinea-Bissau, but further research is needed to clarify the presence of A. sankaranica within the country. Agama picticauda Peters, 1877 Localities — Bissau, Ponta Robalo, Ponta de Marques Ma- no, Tor, Biombo, Sama, Pitche, Enxalé, Bolama Isl., Fa- rim, Mansoa, Contubo-el, Madina do Boé, Gabu, Ilha For- mosa, Bambadinca, Pecixe, Canchungo, Bissalanca, Ba- fata, Bissora, Chitole, Rio Cacine, Capé, Bor, Uno, Bu- baque, Soga References — Bocage (1896a), Ferreira (1902), Boulenger (1905), Monard (1940b), Managcas (1950, 1951b), Frade (1950), Garcia (1991), Naurois (1969), present study Remarks. Listed as Agama colonorum by Bocage (1896a), Ferreira (1902), Monard (1940b) and Manacas (195 1b). Agama weidholzi Wettstein, 1932 Localities. Madina do Boé, Pitche. References.— Joger (1979), Monard (1940b), Grandison (1968). Remarks. Monard (1940b) described A. boensis (see above) on the basis of a single voucher of A. weidholzi Bonn zoological Bulletin 61 (2): 255-281 therefore this was the first record of the species from Guinea-Bissau. Chamaeleonidae Chamaeleo gracilis Hallowell, 1842 Localities. — Bissau, Antula, Ponta de Marques Mano, Brene, Tor, Prabis, Bissalanca, Bissoram, Canchungo, Bis- soram, Cacheu, Bilama, Chitole, Rio Cacine, Farim, Can- chungo, Mansoa, Catio, Nhampurbane (Gabu), Farim, Rio Cacine, Ponta Robalo, Cantubo-el, Pitche, Cacheu, Bola- ma Isl., Contabani, Orango Isl. References — Bocage (1896a), Ferreira (1902), Boulenger (1905), Manacas (1951b), , Gardete-Correira (1971-73), Garcia (1991), present study. Chamaeleo senegalensis Daudin, 1802 Localities Bissoram, Pitche, Cacheu, Mansoa, Catio, Contubo-el, Bijimita, Rio Cacine, Caravela. References. Monard (1940b), Managas (1951b), Limo- ges & Robillard (1991b). Gekkonidae Hemidactylus angulatus Hallowell, 1854 Localities. Bissau, Ponta Robalo, Ponta de Marques Mano, Bissalanca, Sama, Pitche, Madina do Boé, Bola- ma Isl., Geba, Mansoa, Pitche, Chitole, Catid, Cacine, Rio Cacine, Bubaque, Orango, Soga. References.— Bocage (1896a), Ferreira (1902), Boulenger (1905), Monard (1940b), Manacas (1951b), Garcia (1991), present study. Lygodactylus gutturalis (Bocage, 1873) Localities. Bissau, Geba, Contubo-el, Madina do Boé, Pitche, Bolama Isl., Orango Isl., Imbone Isl., Soga Isl., Bubaque Isl. References Bocage (1873), Garcia (1991), Monard (1940b), Loveridge (1947), present study. Lacertidae Latastia ornata Monard, 1940 Localities.— Bafata (Fig. 28). References. Monard (1940b). Remarks.— Additional records remain outstanding; the on- ly West African congener recorded in coastal wetlands of Mauritania (Ineich 1997) and coastal Senegal (B6hme 1978) is L. longicaudata (Reuss, 1839). Phyllodactylidae Tarentola ephippiata senegambiae Joger, 1984 Localities.— Ponta de Marques Mano, Tor, Antula, Biom- bo, Bissalanca, Enxalé, Mansoa, Contubo-el, Bafata, Pit- che, Canchungo, Farim, and Mansoa present study (Fig. 29): References.— Boulenger (1905), Monard (1940b), Mana- cas (1951b), Joger (1981, 1984). OZFMK 278 Mark Aultya et al. Remarks.— Considered as a full species by Trape et al. (2012). Scincidae Chalcides armitagei Boulenger, 1922 Localities — Sucujaque, coastal Guinea-Bissau References.— Trape (2007) Remarks — Previously considered endemic to coastal dunes in The Gambia (Gruschwitz et al. 1991). Chalcides pulchellus Mocquard, 1906 Localities — Madina do Boé. References.— Monard (1940b). Remarks.— C. pulchellus 1s restricted to western Guinea, southeastern Senegal and Mali, whereas C. thierryi occurs further south extending into northern Ghana and the east of Nigeria (Joger & Lambert 2002, Greenbaum et al. 2006). Mochlus guineensis Peters, 1879 Localities. — Rio Corubal e Cacine. References.— Boulenger (1905), Managas (1951b). Panaspis tristaoi Monard, 1940 Localities — Corubal. References Monard (1940b), Naurois (1969). Trachylepis affinis (Gray, 1838) Localities. — Bissau, Ponta de Marques Mano, Bissalanca, Bissoram, Geba, Antula, Bolama Isl. Isl., Cacheu, Rio Cas- sine, Farim, Canchungo, Calequisse, Pecixe, Mansoa, Ma- dina do Boé, Enxalé, Orango Isl., Bubaque Isl. References.— Bocage (1896a), Ferreira (1902), Boulenger (1905), Monard (1940b), Manacas (1951b), Naurois (1969), Garcia (1991), present study. Trachylepis perrotetii (Duméril & Bibron, 1839) Localities — Bissau, Bolama Isl. Isl., Farim, Cacheu, Man- soa, Madina do Boé, Canchungo, Calequisse, Cacine, Bis- soram, Farim, Soga Isl., Bubaque Isl. References. — Bocage (1896a), Boulenger (1905), Monard (1940b), Manacas (1951b), Gardete-Correira (1971-73), Garcia (1991), present study. Varanidae Varanus exanthematicus (Bosc, 1792) Localities.— Bissau (present study, ZFMK 58311), Buba, Bissalanca, Tor, Brene, Pitche, Pecixe, Canchungo, Bo- lama Isl., and Bijagos archipelago. References — Bocage (1896a), Boulenger (1905), Limo- ges & Robillard (1991b), Garcia (1991), Varanus niloticus (Linnaeus, 1758) Localities — Bissau, Brene, Bigimita, Formosa, Bambadin- ca, Ponta Robalo, Bolama Isl., Dungal, Corubal, Varala, Bonn zoological Bulletin 61 (2): 255-281 Orango Isl., Soga Isl., Bubaque Isl., Caravela Isl., Cara- che Isl., Uno Isl., Canhabaque, Ca-Balanta. References.— Bocage (1866), Ferreira (1902), Boulenger (1905), Monard (1940b), Managas (1951b, 1955), Nau- rois (1969), Limoges (1989), Limoges & Robillard (1991b), Garcia (1991), present study. SERPENTES Colubridae Crotaphopeltis hotamboeia (Laurenti, 1768) Localities — Bissau, Contubo-el, Cacheu, Marques Mano, Biimita, Bissalanca, Ponta de Machado, Bafata. References.— Monard (1940b), Managas (1955), Chippaux (2006). Dasypeltis confusa Trape & Mané, 2006 Localities. Bissau, Bissalanca, Rio Cacine. References. — Bocage (1867, 1896a), Managas (1955), Gar- dete-Correira (1971—73), Naurois (1969), Trape & Mané (2006), Chippaux (2006). Remarks. — Dasypeltis scaber scaber was recorded from Guinea-Bissau. Scalation data matches with that of D. con- fusa described by Trape & Mané (2006), thus assigning D. s. scaber as a synonym of D. confusa. Trape & Mane (2006) further record this species from Senegal, Guinea and Benin. Dispholidus typus (Smith, 1829) Localities — Bolama Isl., Cacheu, Rio Cacine. References.— Bocage (1896b), Managas (1955). Grayia smithii (Leach, 1818) Localities — Madina Boé, Bissau. References.— Monard (1940b), Managas (1955), Gardete- Correira (1971-73), Chippaux (2006). Remarks.— The sympatric G. tholloni was mistaken with a Senegal record of G. smithii by Villiers (1950), as was determined by Trape & Mané (2000). However the record here provided by the authors matches with G. smithii with respect to scalation data. Hapsidophrys smaragdina (Schlegel, 1837) Localities — Bolama Isl. References.— Bocage (1896b). Natriciteres olivacea (Peters, 1854) Localities.— Guinea-Bissau. References — Frade et al. (1946), Naurois (1969). Remarks. The species has been recorded in Guinea (Broadley 1966) and Burkina Faso (Trape 2005), in Chip- paux (2006), the species’ most northern distribution along the West African forest belt is Sierra Leone. ©ZFMK The herpetofauna of the Bijagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist 279 Philothamnus heterodermus (Hallowell, 1857) Localities — Catio, Madina do Boe, Orango Isl. References — Monard (1940b), present study. Philothamnus heterolepidotus (Giinther, 1863) Localities — Ponte Robalo, Caita. References — Monard (1940b). Remarks.— Sierra Leone is the most northwestern distri- bution of the species (Chippaux 2006). Philothamnus irregularis (Leach, 1819) Localities — Bissau, Cacheu, Bolama Isl., Pimenta, Pitche, Barahona, Ponte Robalo. References.— Bocage (1896a), Monard (1940b), Gardete- Correira (1971-73). Philothamnus semivariegatus (Smith, 1847) Localities — Bissau. References.— Gardete-Correira (1971—73). Thelotornis kirtlandii (Hallowell, 1844) Localities — Bubaque Isl., “Portuguese Guinea” References.— Riquier & Bohme (1996), Loveridge (1944), present study. Toxycodryas blandingii (Hallowell, 1844) Localities — Bissau, Bissalanca, Soga Isl., Bubaque Isl. References.— Managas (1955), present study. Elapidae Elapsoidea semiannulata moebiusi Werner, 1897 Localities. Bubaque Isl., Bissau, Rio Cacine References — Loveridge (1944), Naurois (1969), Manacas (1981). Dendroaspis jamesoni (Traill, 1843) Localities. Bissau, Bolama Isl. References.— Bocage (1896a, b, 1888), Naurois (1969), Manacas (1981). Dendroaspis viridis (Hallowell, 1844) Localities — Bolama Isl.. Bubaque Isl. References.— Naurois (1969), Managas (1981). Naja haje (Linnaeus, 1758) Localities — Bolama Isl. References.— Garcia (1991). Naja melanoleuca Hallowell, 1857 Localities — Contubo-el, Bolama Isl., Farim, Bubaque Isl. References — Monard (1940b), Managas (1981), present study. Naja nigricollis Reinhardt, 1843 Localities — Bolama Isl., Bambadinca, Aldeia de Cuor, Bonn zoological Bulletin 61 (2): 255-281 Bissau, Buba, Bijagos archipelago, Bubaque Isl. References.— Bocage (1896a), Frade et al. (1946), Man- agas (1981), Limoges & Robillard (1991b), present study. Lamprophiidae Atractaspis aterrima Ginther, 1863 Localities. — Bolama Isl. References.— Managas (1981), Chippaux (2006). Amblyodipsas unicolor (Reinhardt, 1843) Localities.— Rio Cacine. References.— Villiers (1963), Broadley (1971), Chippaux (2006). Boaedon fuliginosus (Boie, 1827) Localities.— Bubaque Isl., Rio Cacine, Bissalanca, Bigi- mita. References.— Boulenger (1905), Naurois (1969), Managas (1955), Gardete-Correira (1971-73). Boaedon lineatus (Dumeéril, Bibron & Duméril, 1854) Localities —Cacheu, Bissau, Madina Boé, Rio Cacine, Bo- lama Isl. Isl., Ponta de Machado. References — Bocage (1896a), Boulenger (1905), Monard (1940b), Manacas (1955). Gonionotophis grantii (Gunther, 1863) Localities — Guinea-Bissau. References. Loveridge (1939), Villiers (1963, 1975), Chippaux (2006). Gonionotophis poensis (Smith, 1847) Localities — Bissau. References — Bocage (1873). Remarks. Based on current taxonomic revisions all Mehelya spp. are included in the genus Gonionotophis (Kelly et al. 2011). Bocage received one specimen (Simo- cephalus poensis) from M. Ferreira Borges, and empha- sizes its similarity with Gonionotophis grantii; however G. poensis’ most northwestern distribution is southeast- ern Guinea (Chippaux 2006), thus Bocage’s provisional identification may confirm G. poensis with a distinct range extension. Gonionotophis stenophthalmus (Mocquard, 1887) Localities.— Bissau. References — Bocage (1896a), Chippaux (2006). Lycophidion albomaculatum Steindachner, 1870 Localities — Bissau, Bolama Isl. References.— Bocage (1896a), Boulenger (1905), Ferrei- ra (1902), Managas (1955), Villiers (1975), present study (Fig. 30). Remarks.— Aforementioned authors assigned the species to L. semicinctum. ©ZFMK 280 Mark Aultya et al. Lycophidion irroratum (Leach, 1819) Localities — Rio Cacine, Bissau, Ponta de Machado. References— Boulenger (1905), Managas (1955), Chip- paux (2006). Prosymna meleagris (Reinhardt, 1843) Localities — Mansoa. References.— Monard (1940b). Remarks.— Broadley (1980) distinguished two populations, P. m. melagris and P. m. greigerti and intergrades from Senegal. Trape & Mané (2002) list both populations from the Senegal, however later Trape & Mané (2006) ressur- rect P. meleagris (Reinhardt, 1843) and P. greigerti (Moc- quard, 1906), and latter species into, P. g. greigerti and P. g. collaris. On the basis of Monard’s description it is not possible to distinguish between the two, hence P. melea- gris remains listed here. P. greigerti must therefore be con- sidered as a potential species of Guinea-Bissau (see be- low). Psammophis elegans (Shaw, 1802) Localities — Bissau, Bissalanca, Bigimita, Bolama Isl., Farim, Cacine, Catio. References.— Bocage (1867, 1896a), Ferreira (1902), Ma- nacas (1955), Gardete-Correira (1971-73). Psammophis lineatus (Duméril, Bibron & Dumeéril, 1854) Localities — Contubo-el, Rio Cacine, Catio, Bafata, Pon- ta de Machado, Bissau References.— Monard (1940b), Loveridge (1940), Mana- cas (1955), Gardete-Correira (1971-73), Garcia (1991). Remarks. The species was previously included in the genus Dromophis (also see Chippaux 2006), and relegat- ed to the synonymy of Psammophis by Kelly et al. (2008). Psammophis phillipsi (Hallowell, 1844) Localities.— Uno Isl., Bubaque Isl. References.— Naurois (1969), present study. Remarks.— See Psammophis sibilans. Psammophis sibilans (Linnaeus, 1758) Localities — Bissau, Cacheu, Bolama Isl. Isl., Antula, Rio Cacine, Madina Boé, Catio. References — Bocage (1866, 1896b), Ferreira (1902), Mo- nard (1940b), Manacas (1955). Remarks.— According to Kelly et al. (2008) the Psam- mophis sibilans “species complex” remains unresolved, however the authors provisionally distinguish two lineag- es, the “phillipsii” and “‘subtaeniatus” complex. Leptotyphlopidae Myriopholis narirostris (Peters, 1867) Localities — Ponta de Marques Mano, Bissalanca, Bissau. Bonn zoological Bulletin 61 (2): 255-281 References— Manacas (1955), Gardete-Correira (1971-73). Remarks. The originally described Glauconia debilis Chabanaud 1918 has been synonymized with Leptoty- phlops narirostris boueti (Hahn & Wallach 1998). The species was renamed by Adalsteinsson et al. (2009). Pythonidae Python regius (Shaw, 1802) Localities — Bissau. References.— Monard (1940b), present study (Fig. 31). Python sebae (Gmelin, 1789) Localities — Ponte Machado, Bissau, Enchalé, Bor, Biyagos archipelago, Cufada, Catid, Bucaré, Imbone Isl., Orango Isl., Soga Isl., Bubaque Isl. References.— Bocage (1896a), Frade et al. (1946), Garcia (1991), Naurois (1969), Broadley (1984), Limoges & Ro- billard (1991b), Chippaux (2006), present study. Typhlopidae Afrotyphlops punctatus (Leach, 1819) Localities — Ponta de Machado, Bissau, Bolama Isl. Isl., Rio Cacine, Cacheu. References.— Bocage (1896a), Ferreira (1902), Boulenger (1905), Managas (1955). Remarks .— Typhlops punctatus was renamed by Broadley & Wallach (2009). Viperidae Bitis arietans (Merrem, 1820) Localities — Bolama Isl., Antula, Bissau, Bubaque Isl., Orango Isl., Soga Isl. References.— Bocage (1896a), Ferreira (1902), Manacas (1981), present study. Bitis nasicornis (Shaw, 1802) Localities — Bubaque Isl. References.— Naurois (1969). Causus maculatus (Hallowell, 1842) Localities. — Cacheu, Contubo-el, Rio Cacine. References Manacas (1981), present study (ZFMK 88323). Crocodylidae Crocodylus suchus Geoffroy, 1807 Localities — Rio Geba, Corubal e Cacheu; Contubo-el, Bo- lama Isl., Nhacra, Orango Isl., Formosa Isl., Imbone Isl., Uno Isl. References— Limoges (1989), Limoges & Robillard (1991b), present study. ©ZFMK The herpetofauna of the Biyagos archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist 28] Mecistops cataphractus (Cuvier, 1825) Localities — Rio Cacheu?, Rio Corubal, Cacine. References.— Garcia (1991), de Buffrenil (1993). Osteolaemus tetraspis Cope, 1861 Localities — Antula, Madina do Boé, Formosa, Orango. References — Monard (1940b), de Buffrenil (1993), Li- moges (1989). APPENDIX 2 Previously recognized, but now doubtful records from Guinea-Bissau Arthroleptidae Arthroleptis variabilis Matschie, 1893 Localities. Guinea-Bissau. References.— Managas (1949, 1950). Remarks.- According to Rédel & Bangoura (2004) the spe- cies 1s confined to central Africa. Leptopelis cynnamomeus (Bocage, 1893) Localities — Bolama Isl. References.— Bocage (1896a). Remarks.— The species’ northern range limit is in the Dem- ocratic Republic Congo; according to Perret (1976) the paratypes of L. cynnamomeus from Guinea-Bissau are referable to L. viridis. Hyperoliidae Hyperolius argus Peters, 1854 Localities.— Ponta de Marques Mano, Bolama Isl. References.— Monard (1938), Managas (1949, 1950). Remarks.— Listed as H. cinctiventris Cope; H. argus is ge- ographically confined to eastern Africa (Frost 2011), hence the orginal H. cinctiventris reflects a synonym (at this stage unknown) of a congener native to Guinea-Bissau. HAyperolius ferreirai Noble, 1924 Localities — Ponta de Marques Mano, Bissalanca, Caio- nete, Caio, Contubo-el. References — Monard (1940a), Managas (1949, 1950). Remarks .— The species appears to be confined to Angola (Frost 2011), hence H. ferreirai reflects a synonym (at this stage unknown) of a congener native to Guinea-Bissau. Hyperolius marmoratus Rapp, 1842 Localities. — Bolama Isl., Geba. References.— Bocage (1896a), Monard (1940a), Manacas (1949, 1950). Remarks.— The species appears to be confined to eastern and southern Africa (Frost 2011), hence H. ferreirai re- flects a synonym (at this stage unknown) of a congener native to Guinea-Bissau. However the taxonomic status Bonn zoological Bulletin 61 (2): 255-281 of this species complex remains highly distorted and con- troversial. Gerrhosauridae Gerrhosaurus nigrolineatus Hallowell, 1857 Localities — Bolama Isl. References — Ferreira (1902). Remarks.— Hallowell (1857) describes the species from Gabon, and Pauwels & David (2008) confirm the species for Gabon. The record from Bolama Isl. remains doubt- ful. Scincidae Chioninia delalandii (Duméril & Bibron, 1839) Localities — Garcia (1991) mentions ““S6 se encontra men- cionada nas obras de Bocage, vinda de Bissau, no litoral da Guinée”, which indicates that “the species was only re- corded, as was reported in the studies of Bocage, upon ar- rival in Bissau in coastal Guinea-Bissau” (Garcia 1991). However, Bocage (1867) lists the species “Euprepres de- lalandii” from the Cape Verde Islands. References.— Bocage (1867), Garcia (1991). Lamprophiidae Gonionotophis brussauxi (Mocquard, 1889) Localities. — Guinea-Bissau. References.— Villiers (1963, 1975). Remarks.— The species is geographically confined to Cameroon, Republic of the Congo and Gabon (Chippaux 2006, Pauwels et al. 2006). A doubtful record for G. brus- sauxi in Togo 1s indicated by Chippaux (2006), which may in fact refer to G. Alingi. A record for Guinea-Bissau at this stage must be considered doubtful. Lycophidion meleagris Boulenger, 1893 Localities.— Guinea-Bissau. References — Gardete-Correira (1971-73), Garcia (1991). Remarks.— Garcia (1991) mentions L. horstocki and L. gambensis as synonyms of L. meleagris. Former is rec- ognized as a synonym of L. albomaculatum and L. capense while latter synonyms could not be traced. The most northern record of a West African congener, L. ni- gromaculatum (Peters, 1863) is southeastern Guinea (Chippaux 2006). At this stage it is not possible to ascer- tain the validity of this species. Viperidae Causus rhombeatus (Lichtenstein, 1823) Localities — Cacheu, Contubo-el, Bissau, Rio Cacine. References.— Bocage (1896a), Monard (1940b). Remarks .— This record very likely reflects C. maculatus, based on the number of ventrals (137) as indicated by Monard (1940b); in C. rhombeatus ventrals count more than 150 (Chippaux, 2006). OZFMK Bonn zoological Bulletin (BzB) Instructions to authors Scope The Bonn zoological Bulletin (BzB), formerly “Bonner zoologi- sche Beitrage”, is an international, peer-reviewed, open access jour- nal publishing original research articles, reviews, and scientific notes dealing with organismal zoology. Focus of the BzB are (1) taxon- omy, (2) systematics and evolution, and (3) biodiversity and bio- geography, all with respect to terrestrial animals. Terrestrial animals as understood here include those inhabiting fresh or brackish wa- ters. Contributions from related fields like ecology, morphology, anatomy, physiology or behaviour are welcome when of clear rel- evance to the focus topics. 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Ethical and legal aspects Authors are obliged to state clearly that they followed the legal reg- ulations and laws for the collection and ethical treatment of animals. Publication and open access policy Manuscripts will be published online at www.zoologicalbulletin.de after acceptance as preprint version, which will be replaced by the final pdf after publication of the printed issue of BzB. The corre- sponding author must clearly state so if preprint online publication is not desired. In that case, only title, authors and the information that this manuscript has been accepted for publication will be made available at that stage. Immediately after publication, the correspon- ding author will receive an e-mail containing the final pdf. Reprints can only be purchased on the author’s cost. Reprint orders must be submitted to the editor at latest with the approval of the final proofs. All published content is freely available without charge to the user or his/her institution. Authors are invited to place the final pdf on their personal or institutional website. Questions we ask our reviewers Authors preparing manuscripts for BZB should keep in mind the fol- lowing questions which referees for the journal are asked: . Is the paper of interest to the readers of BZB? . How well does it meet the focus of BZB? . Is it acceptable for publication, or should it be reconsidered af- ter revision? 3. Please grade originality, quality of data, quality of statistical analy- ses, and quality of interpretation on a scale from 1 = low to 5 = high. 4. Please grade clarity and quality of text, figures and tables from 1 = low to 5 = high. . Has the welfare of any experimental animals been adequately tak- en into account? . Does the title aptly and correctly describe the paper? . Does the abstract summarize only the significant findings? . Is the length of the paper appropriate? . Are all (colour) figures and tables necessary and appropriate? WN re Nn Oo COND iy Tyee ar mt j 7 Oe ye: Teh oa OL ee iT, ite Me tert ; , i hy e olnee | t r MITHSONIA Contents il HM Assing, Volker: 142 A revision of the Lathrobium species of the Himalaya (Coleoptera: Staphylinidae: Paederinae) _ 210 Assing, Volker: The genus ASR in the ee Wagner, Philipp, Aaron M. Bauer & Wolfgang Bohme: 216 Amphibians and reptiles collected by Moritz Wagner, with a focus on the ZFMK collection 241 Résler, Herbert, lvan Ineich, Thomas M. Wilms & Wolfgang BOhme: Studies on the taxonomy of the Gekko vittatus Houttuyn, 1782 complex (Squamata: Gekkonidae) |. On the variability of G. vittatus Houttuyn, 1782 sensu lato, with the description of a new species from Palau Islands, Micronesia Auliya, Mark, Philipp Wagner & Wolfgang Bohme: 255 The herpetofauna of the Bijagds archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist Cover illustration: Holotype of Vipera wagneri Nilson & Andrén, 1984, ZFMK 32495 (this volume, pp. 216-240) Bundesministerium ,,..,_. i : Uy, fiir Bildung Ministerium fir Innovation, und Forschung Wissenschaft und Forschung G3 des Landes Nordrhein-Westfalen ft Leibniz-Gemeinscha