,37 '3. 7- FORSCHUNGS museum KOENIG Bonn zoological Bulletin formerly: Bonner zoologische Beitrage Volume 63 Issue 2 2014 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 - Leibniz-Institut fur Biodiversitat der Tiere (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), excluding supplements, 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. © 2014 Zoologisches Forschungsmuseum Alexander Koenig - Leibniz-Institut fur Biodiversitat der Tiere (ZFMK). Bonn, Ger- many. ISSN 2190-7307. Produced by Eva-Maria Levermann, Kaiserstr. 129, 53 1 13 Bonn, Germany; emlevenriann@netcologne.de. Printed and bound by DCM, Wemer-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.de Managing Editor Supplement Series Thomas YVesener, ZFMK, tel. +49 228-9 122-125, fax: +49 228-9122-212; twesener@zfmk.de Editorial Board Dirk Ahrens, Insects: Coleoptera, ZFMK, tel. +49 228-9122-286; d.ahrens@zfmk.de Wolfgang Bohme, Amphibians and Reptiles, ZFMK, tel. +49 228-9122-250; w.boehme@zfmk.de Jan Decher, Mammals, ZFMK, tel. +49 228-9122-262; j .decher@zfmk.de Netta Dorchin. Insects: Diptera, Department of Zoology, Tel Aviv University, Tel Aviv 69978, Israel, tel. +972-3-6409808; ndorchin@post.tau.ac.il Bernhard Huber, Invertebrates except Insects, ZFMK, tel. +49 228-9122-294; b.huber@zfmk.de Rainer Hutterer, Mammals, ZFMK, tel. +49 228-9122-261; r.hutterer@zfmk.de Ximo Mengual, Insects: Diptera, ZFMK, tel. +49 228-9122-292; x.mengual@zfmk.de Gustav Peters, Mammals, ZFMK, tel. +49 228-9122-279; g.peters@zfmk.de Ralph Peters. Insects: Hymenoptera, ZFMK, tel +49 228-9122-290; r.peters@zfmk.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 K1A0C6, tel. + 1 613-759-1787; bradley.sinclair@inspection.gc.ca Dieter Stiining, Insects except Coleoptera and Diptera, ZFMK, tel. +49 228-9122-220; d.stuening.zfmk@uni-bonn.de Till Topfer, Birds, ZFMK, tel. +49 228-9122-246; t.toepfer@zfmk.de Philipp Wagner, Amphibians and Reptiles, Naturmuseum, Umweltschutz-Informationszentrum Lindenhof, Karolinen- reuther StraBe 58, 95448 Bayreuth, tel. +49 921-75942-0; philipp.wagner.zfmk@uni-bonn.dc Advisory Board Theo C. M. Bakker, Rheinische Friedrich-Wilhelms-Univer- sitat, Institut fur 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@vi 1 lanova .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 fur Naturkunde Berlin, Leib- niz-Institut fur 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.k17stufek@zrs.upr.si Wolfgang Schawaller, Staatliches Museum fur 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 89-8107-1 10; schliewen@zsm.mwn.de Michael Schmitt, Ernst-Moritz-Amdt-Universitat, Allge- meine & Systematische Zoologie, Anklamer Str. 20, 17489 Greifswald, Germany, tel. +49 3834-86^4242, fax: +49 3834-86-4098; 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, Spiclmannstr. 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 7071-2972616, fax +49 7071-295170; erich.weber@uni-tuebingen.de I , 3 7cZ 70* fj-tt FORSCHUNGS museum KOENIG Bonn zoological Bulletin formerly: Bonner zoologische Beitrage Index 2014 Volume 63 Issue 1-2 KOENIG Bonn zoological Bulletin Volume 63 Issue 1 2014 ■v^L KOENIG Bonn Volume 63 zoological lss2u0e,4 Bulletin Editor-in-Chief Fabian Herder Editorial Board Dirk Ahrens Wolfgang Bohme Jan Decher Netta Dorchin Bernhard A. Huber Rainer Hutterer Ximo Mengual Gustav Peters Ralph Peters Bradley Sinclair Dieter Stuning Till Topfer Philipp Wagner An open access journal of organismal zoology, published by Zoologisches Forschungsmuseum Alexander Koenig - Leibniz-lnstitut fur Biodiversitat derTiere, Bonn Contents Volume 63 Grebennikov, Vasily V.: 123-147 Morimotodes, a new genus for two minute wingless litter species from southwest China and Taiwan with an illustrated overview of Molityna and Plithina genera (Coleoptera: Curculionidae: Molityni) Liu, Wan-Gang, Ming Bai, Xing-Ke Yang & Dirk Ahrens: Towards an improved knowledge of Sericini of the Tibetan highland: new species and records (Coleoptera: Scarabaeidae) 157-172 Nguyen, Truong Quang, Trung My Phung, Nicole Schneider, Andreas Botov, Dao Thi Anh Tran & Thomas Ziegler: New records of amphibians and reptiles from Southern Vietnam 148-156 Peters, Gustav, Tanja Haus & Rainer Hutterer: Neotropical primates from the Cologne Zoo in the collections of the Zoologisches Forschungsmuseum Alexander Koenig: noteworthy specimens, taxonomic notes and general considerations 173-187 Peters, Ralph S.: First record of the parasitoid wasp Tachinaephagus zealandicus Ashmead, 1904 (FHymenoptera: Chalcidoidea: Encyrtidae) in Germany 115-118 Pimsai, Uraiporn, Malcolm J. Pearch, Chutamas Satasook, Sara Bumrungsri & Paul J.J. Bates: Murine rodents (Rodentia: Murinae) of the Myanmar-Thai-Malaysian peninsula and Singapore: taxonomy, distribution, ecology, conservation status, and illustrated identification keys 15-1 14 Stiels, Darius & Kathrin Schidelko: Erratum to Records of 'Indian' Baya Weaver Ploceus philippinus philippinus (Linnaeus, 1766) and Flooded Wheateater Oenanthe monacha (Temminck, 1825) from Afghanistan (Aves: Passeriformes) 119 Wagner, Philipp, Adam D. Leache & Matthew K. Fujita: Description of four new West African forest geckos of the Hemidactylus fasciatus Gray, 1842 complex, revealed by coalescent species delimitation 1-14 Book Reviews Grimmberger E (2014) Die Saugetiere Deutschlands: Beobachten und Bestimmen (Boris Krystufek) 120-121 Publication dates: Vol. 63 Issue 1 : June 2014 Vol. 63 Issue 2: December 2014 Bonn zoological Bulletin 63 (2): 123-147 December 2014 Morimotodes , a new genus for two minute wingless litter species from southwest China and Taiwan with an illustrated overview of Molytina and Plinthina genera (Coleoptera: Curculionidae: Molytini) Vasily V. Grebennikov Canadian Food Inspection Agency, K.W. Neatby Bldg., 960 Carling Ave., Ottawa, ON K1A 0Y9, Canada; E-mail: vasily. grebennikov@inspection.gc. ca Abstract. The new weevil genus Morimotodes g. n. is described for two new species: M. ismene sp. n. (Yunnan and Sichuan) and M. polynices sp. n. (Taiwan). Morimotodes ismene sp. n. is designated as the type species of Morimotodes g. n. Species are small (1. 6-2.1 mm), wingless inhabitants of leaf litter in montane deciduous forests with characteristically globular body and obliterate elytral striae and dorsal sculpture. Analysis of DN A barcodes and external similarity suggest that Mo- rimotodes g. n. is a sister to the West Palaearctic genus Leiosoma Stephens, 1829 with 32 similarly small-bodied and lit- ter-inhabiting species. Dated intraspecific phylogeographic analysis utilizing DNA barcodes of M. ismene sp. n. reveals seven strongly supported geographical clades, four of which are allopatric and three others are parapatric on Mt. Emei in Sichuan. Potentially allied genera belonging to two of the three Molytinini subtribes, Molytina and Plinthina, are il- lustrated and briefly discussed. Type specimens of five exceptionally poorly known genera ( Carbonomassula Heller, 1908, Clarkantlms Alonso-Zarazaga & Lyal, 1999, Gonotrachelus Champion, 1914, Vouauxia Hustache, 1920, Pterotomus Que- denfeldt, 1888) are re-examined and the original labels illustrated. Key words: Yunnan, Sichuan, Taiwan, COl, DNA barcoding, phylogeography INTRODUCTION In 2008, a first long series of conspicuous but unknown beetles was discovered by sifting forest leaf litter on the eastern slope of the Cang Shan Mountain Range in Yun- nan (China). The peculiarly small, lustrous and rounded body in combination with the rounded elytral shoulders strongly resembled other litter-inhabiting wingless Palaearctic Molytinae, i.e. Otibazo Morimoto, 1961 or Leiosoma Stephens (Fig. 7). The subsequent four years of intensive sampling in the same and nearby forests of Southwest China revealed additional similarly shaped specimens. A long series of another seemingly congener- ic species was detected among unidentified specimens col- lected two decades earlier in Taiwan by Ales Smetana (Fig. 6). The generic identity of these beetles, however, re- mained elusive. Since the original discovery in 2008, these beetles were repeatedly compared to many others, mainly belonging to the wingless members of Molytinae. Eventually it became evident that these finds represented a previously undetect- ed clade presumably deserving a formal status of a new genus. This thought intensified the search for its possible relatives. That pursuit was long inconclusive, since numer- ous unrelated wingless Molytinae were infamous for their convergent similarity linked to the loss of hind wings (Lyal 2014). DNA-based search for relatives of the new genus was similarly inconclusive, since the dataset did not Received: 07.08.2014 Accepted: 12.11.2014 contain many critical and important taxa. Moreover, due limitations in funding, the DNA work was done with the single marker of the DNA barcode (part of cytochrome oxidase I, Hebert et al. 2003), which had significant lim- itations in its applicability for detecting relatively old evo- lutionary events (Grebennikov & Heiss 2014). Eventual- ly, however, a representative of the genus Leiosoma be- came available for DNA analysis and consistently clus- tered with a clade representing the new genus. At that point the project’s results were judged mature enough to be for- mally presented, even though some shortcomings linked to a shortage of the material and data remained (see Dis- cussion). Another coincidental event further triggering the com- pletion of this project was the recent publication of the long-awaited “Phytophaga” volume of the Handbook of Zoology series, most specifically the chapter on the wee- vil subfamily Molytinae (Lyal 2014). This subfamily has long been and still is a phylogenetic challenge. In the aforementioned chapter the tribes of Molytinae were sig- nificantly reorganized, as compared to the earlier standard works of reference (Alonso-Zarazaga & Lyal 1999; Bouchard et al. 2011; Alonso-Zarazaga 2013). The most noticeable novelty, however, was the incorporation of the entire former “subfamily Cryptorhynchinae” into Molyti- nae, the latter presently containing 37 tribes plus ca. 28 Corresponding editor: D. Ahrens 124 Vasily V. Grebennikov incertae sedis genera (Lyal 2014: 530, 531). This decision is in line with the pivotal work of Kuschel (1987) amal- gamating a multitude of former “subfamilies” into a broad- ly defined Molytinae (then excluding “Cryptorhynchi- nae”). Both steps are seen as a much needed attempt to make the formal classification more in line with the cur- rent phylogenetic hypothesis (for example, see the remark- able mosaic of both former “subfamilies” in McKenna et al. 2009). Given an acute lack of the phylogenetic frame- work for the subfamily and for the great majority of its tribes, numerous further changes in Molytinae classifica- tions will undoubtedly follow. To move forward, a prac- tical approach was outlined (Lyal 2014: 549) as to “... pro- pose initial hypotheses for monophyletic groups within the Molytinae and then analyze them with both molecular and morphological characters...” This paper presents an at- tempt to answer this call for actions on Molytinae sub- groups and to shed light on a small subsection of this grand, if indeed monophyletic, radiation. This work has three main goals. First, to name formal- ly the newly discovered genus and its two species. Sec- ond, to utilize available mtDNA sequences to test the monophyly of the genus and to search for the possible sis- ter group. Third, to discuss and delimit species boundaries and to undertake a dated phylogeographical analysis. Forth, to advance understanding of Molytinae genera cur- rently attributed to the subtribes Molytina and Plinthina by providing an annotated list of taxa, illustrating a rep- resentative species for all but three genera and clarifying the identity of five obscure genera by re-examination of the type specimens of their type species. Overall, this pa- per continues the recently initiated efforts to assess phy- logeographical patterns detected from the low-dispersing Hightless weevils in southwestern China and adjacent lands and to clarify the taxonomy and diversity of wing- less Old World Molytinae (Grebennikov 2014). MATERIAL AND METHODS Museum abbreviations, followed by the name of the cu- rator: BMNH Natural History Museum, London, UK (M. Barclay); CMN Canadian Museum of Nature, Ottawa, Canada (R. Anderson, F. Genier); CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada (P. Bouchard); IZCAS Institute of Zoology, Chinese Academy of Science, Beijing, P.R. China (R. Zhang); MNHN Museum national d’Histoire naturelle, Paris, France (H. Perrin); MRAC Musee Royal de l’Afrique Centrale, Tervuren, Belgium (M. De Meyer); MTD Senckenberg Naturhistorische Sammlungen, Dresden, Germany (K.-D. Klass, O. Jager); NHMB Naturhistorisches Museum, Basel, Switzerland (E. Sprecher); NMW Naturhistorisches Museum, Wien, Austria (H. Schillhammer); ZIN Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia (B.A. Korotyaev). This study is based on examination of a total of 97 dry- mounted adult specimens of the new genus. Concepts of the Curculionoidea family-group names and assignment of weevil genera to tribes follow Bouchard et al. (201 1 ) and Alonso-Zarazaga & Lyal (1999), respectively, with subsequent significant modifications by Lyal (2014). The chronostratigraphic timing is that of Cohen et al. (2013) with the Pliocene-Pleistocene boundary set at 2.588 MY. Nomenclature of male genitalia follows that of Wanat (2007). The length of the body was measured in dorsal as- pect from the elytral apex to the front of the pronotum. Geographical labels of the newly described species are cit- ed verbatim in quotation marks. The hypothetical most re- cent common ancestor of a monophyletic group referred to in the discussion is abbreviated as MRCA. The distri- bution map was generated using the online SinrpleMap- pr tool (Shorthouse 2010). Species identification of the two sequenced specimens of Leiosoma reitteri (Bedel) was done based on the assumption that this is the only species of the genus known from Georgia (Alonso-Zarazaga 2013). The type species are cited using their presently valid names, which are not necesserely the same when the type species were designated; the latter can be found in Alonso-Zarazaga & Lyal ( 1999). DNA dataset. All specimens submitted to DNA barcod- ing have a unique identifier label with the code CNC- COLVG0000XXXX; this format is shortened to #XXXX when specimens are cited in the text. All details regard- ing sequencing, primers, PCR protocols as well as the list of specimen information including provenience and voucher number can be seen by accessing the online dataset at: dx.doi.org/10.5883/DS-MORIMO90. In a few cases a specimen failed to amplify DNA from the first at- tempt and was later resubmitted; in such cases a digit “1” after a dot follows the original four digit number (like #0251.1 on Fig. 7). The dataset used for the first DNA analysis contained 90 sequences and 658 aligned positions. The ingroup was formed by 46 newly generated se- quences, each representing a specimen of the continental new species of the new genus (GenBank accessions: HM4 1 7727-32, HQ986799, HQ986876-8, HQ987105, Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK Morimotodes, a new weevil genus from China 125 HQ9871 34-5, HQ987 1 59, HQ987 1 79-8 1 , KJ87 1 620-32, KJ87 1634-5, KJ87 1637-40, KJ87 1 642-5 1 ). All but one specimen representing another new species collected in Taiwan in 1992-1998 were submitted to DNA barcoding but failed to amplify, likely because of either their age, collecting methods or storage degraded DNA. No fresh specimens of the target genus were recovered in Taiwan during a month-long fieldwork in 2013. The majority of the ingroup sequences were 658 basepairs (bp) long and no one shorter than 407 bp. The outgroup consisted of 44 sequences and represented all Molytinae genera with avail- able DNA barcode data (Fig. 7), while three sequences of Graptus circassicus Solari, 1945 (Entiminae) were used to root the topology in the first analysis (see below). All outgroup sequences were of the full DNA barcode length (658 bp). Three of the outgroup sequences representing the genus Plinthus Germar, 1817 (KJ871633, KJ871636, KJ871641 ) were newly generated. The remaining 41 out- group sequences were taken from previous works (HM386450, HM4 17677-9, HQ986888-9, HQ987002-3, KJ427730-2, KJ427736-9, KJ427744, KJ427746, KJ427748, KJ445683-4, J445687, KJ445690-1, KJ445693-4, KJ445698-6, KJ445708-9, KJ445712-3, KJ445715 in Grebennikov 2014; KJ841729, KJ841731 in Grebennikov & Pham 2014). Specimen images (when present), geographical data, primers, original chro- matograms and other relevant data for all 90 sequenced specimens can be found online in the publicly accessible dataset “ Morimotodes ” on the Barcode of Life Database portal (doi: dx.doi.org/10.5883/DS-MORIMO90). DNA-based phylogenetic analyses. Two separate tree search analyses were run. The first analysis aimed at test- ing monophyly of the new genus represented by the type species, search for its sister-group and to examine its in- ternal relationships. The original DNA dataset containing 90 sequences was analysed using the Maximum Likeli- hood (ML) approach with statistical branch support test- ed by 1000 replications of bootstrapping in MEGA 5 (Tamura et al. 2013). The search for the optimal substi- tution model for both analyses was done in MEGA 5 and resulted in the same model (GTR+G+I). The second tree search analysis was performed to date the branching events in the continental species of the new genus and to re-test the interrelationships hypothesis, as already sug- gested in the first analysis. The original matrix of 90 se- quences was reduced to 33 to include only the full-length DNA barcodes (658 nt) of 3 1 ingroup specimens and two specimens of the sister group (the genus Leiosoma , as sug- gested in the first analysis). The second analysis was per- formed in BEAST vl.8.0 (Drummond et al. 2012) utiliz- ing the Bayesian inference (BI) approach with no a pri- ori internal grouping, all default priors and options, GRT+G+I nucleotide substitution model, strict linear mo- lecular clock and nucleotide substitution rate of 0.0 1 8 ( Pa- padopoulou et al. 2010). The “burn in’’ option was imple- mented eliminating the first 2500 of the 10000 obtained trees. Two resulting topologies, each representing either ML or BEAST analysis, were visualized in FigTree vl.4 (Rambaut 2013) and presented by Figs 7 and 8, respec- tively. Use of morphological data. A formal morphology-based phylogentic analysis of Molytinae genera to search for the sister group of the new genus was beyond the scope of the present paper. Even restricted to 8 1 genera of the tribe of Molytini (80 mentioned in Lyal 2014, and the new one), the task was judged as too daunting. An attempt was made to illustrate a representative of all genera currently as- signed to Molytina and Plinthina (13 and 17, respective- ly; see Lyal 2014: 530, 53 1 ; images of three genera were not obtainable). Five particularly poorly known genera were elucidated in more details, with their type specimens and original labels imaged. The third remaining Molyti- ni subtribe, Flylobiina, historically considered as a tribe of its own and incorporating 5 1 winged genera (Lyal 2014) was not included in this treatment. This decision is part- ly due to the large size of this tribe. The second reason for exclusion is that until very recently, Hylobiina were formally in a separate tribe and, therefore, not specifical- ly targeted since 2008, when this project was in prepara- tion. Morimotodes g. n. Type-species: Morimotodes ismene sp. n., by original des- ignation. Diagnosis. Adult specimens of Morimotodes g. n. can be immediately recognized from other Curculionoidea known both from the Eastern Palaearctic and the Oriental Region by their habitus. They are small (about 2 mm) brown or reddish beetles with lustrous dorsum, rounded body and nearly obliterated elytral shoulders and striae. In the area of their known distribution, namely in Yunnan, Sichuan and Taiwan, as well as in the adjacent regions, adults of Morimotodes externally resemble those of Otibazo Mo- rimoto. The latter, however, has not yet been discovered in China (although its presence is possible, since the first continental species has been recently detected in north- ern Vietnam, see Grebennikov & Pham, 2014). Adults of Otibazo are immediately distinguishable from nearly all weevils by their unique fields of short dense pubescence forming characteristic whitish tufts on flagella, femora and tibiae (Fig. 7), not found in Morimotodes. Field recogni- tion of Morimotodes can be facilitated by stressing their biological preferences. All known specimens have been found in wet leaf litter of primary deciduous forests in the altitudinal range of 1947^4072 m in southwestern China and 1110-2550 m in Taiwan. From the most closely re- Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK 126 Vasily V. Grebennikov Fig. I. Morimotodes ismene g. & sp. n., holotype, male, #0928 from Gaoligong Mt. Range, Yunnan. A-D: habitus; E-G: aedea- gus and tegment, dorsal (E), ventral (F), lateral; (G); H: stemites 8 and 9. lated genus Leiosoma, adults of Morimotodes g. n. can be easily distinguished by their brown or reddish colour (Leiosoma adults are black) and, additionally, by a wide gap in distribution ( Leiosoma is restricted to the West Palaearctic and, therefore, is widely allopatric with Mo- rimotodes g. n.). Description. Small (1.6-2. 1 mm) and uniformly brown or dark reddish, lustrous, wingless weevils. Dorsal surface of pronotum and elytra without longitudinal ridges, cari- nae or irregular sculpture; elytral striae weakly indicated by individual rows of fine setiferous punctures surround- ed by clearly restricted zone of darker pigmentation (Figs 1 A-C, 6A-C); setiferous punctures on pronotum shallow- er, not arranged in longitudinal rows and lacking pigment- ed surroundings (Figs 1 A-C, 6A-C). Length of dorsal se- tae variable, from almost indistinguishable under 50x mag- nification (Figs 6A-C, “polished” specimens) to relative- ly long and exceeding in length longitudinal distance be- tween setiferous punctures (Figs 4B, 5B, “hairy” speci- mens). Integument clean, not covered by incrustation or wax secretion. Elytra with shoulders entirely levelled, in dorsal view strongly and evenly rounded, 1 .5-1 .8x as wide as maximal prothoracic width. Body dorsal outline in lat- eral view evenly and relatively strongly curved (Fig. 5); elytra neither flattened dorsally, nor declivate apically. Head capsule moderately visible from above (Fig. 4). Eyes laterally, not reduced in size. Antennal attachments in api- cal third of rostrum; funicle with seven antennomeres; compact club consisting of three antennomeres. Pronotum not constricted laterally in anterior quarter; widest at mid- dle or at base; prosternum without rostral canal or ocular Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK. Morimotodes, a new weevil genus from China 127 Fig. 2. Morimotodes ismene g. & sp. n., paratype, female, #1020 from Gaoligong Mt. Range, Yunnan. A-D: habitus, E: genital stemites 8 & 9, ventral view. lobes; procoxae contiguous. All femora without tooth. Hind wings absent. Sclerolepidia absent (Fig. 3E). Aedea- gus weakly sclerotized, dorsally and ventrally membra- nous and translucent; endophallus without slerotization (Figs 1 E, F, 6E, F). Female hemistemites 9 each with sep- arate and clearly distinguishable basal and apical pieces (Fig. 2E; “coxite” and “stylus”, respectively). Species composition and distribution. As currently known, the genus Morimotodes g. n. comprises two species, each relatively widely distributed in Southwest China (Yunnan and Sichuan) or in Taiwan, respectively. Biology. All specimens of Morimotodes were collected by sifting deciduous leaf litter in montane primary forests with subsequent specimen extraction using Winkler fun- nels. These collecting circumstances were further corrob- orated by the loss of hind wings, which strongly suggest that Morimotodes species are habitat-dependant low-dis- persing forest litter inhabitants critically dependant on this habitat. Immature stages, host plants or any specific bio- logical preferences remain unknown. Etymology. Morimotodes , patronymic, derived by adding the Latin suffix -odes to the family name of Katsura Mo- rimoto (retired, formerly professor at Kyushu Elniversity, Fukuoka, Japan), in recognition of his outstanding con- tribution to the knowledge of weevils from the Asia-Pa- cific Region. Gender masculine. Morimotodes ismene sp. n. Figs 1-5. Diagnostic description. Holotype, male (Fig. 1). Gen- Bank accession: HQ987135. Length: 1.85 mm. Dorsal out- line of pronotum in lateral view evenly rounded; prono- tum and elytra dorsally with fine punctures and white pi- losity visible at magnification below 30x; each elytron with single longitudinal groove adjacent and parallel to edge. Acdeagus parallel-sided; sternite 9 not subdivided and not wider than 1.5x width of apodeme; sternal apodeme 9 with large paddle-shaped extension anterior- ly. Intraspecific variation. Length: 1.60-2.0 mm. Body outline in dorsal and lateral views variable (Figs 4, 5), pronotum in basal half either parallel-sided (Fig. 4D) or Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK 128 Vasily V. Grebennikov Fig. 3. Morimotodes ismene g. & sp. n., unsexed, #0427 from Cang Shan Mt. Range, Yunnan. SEM images. A: head, left later- al; B: head and pronotum, left lateral; C: head, frontal; D: rostrum, frontal; E: meso- and metathorax, left lateral; F: claws, later- al; G: pronotum and head, dorsal; H: meso- and metathorax, right ventro-lateral; I: elytra, left lateral. Mst: metepistemum, mtv: metaventrite, col-co3: pro-, meso- and metaxocae, respectively, vtl : ventrite 1 (=abdominal segment 3). Note absence of sclerole- pidia on metepisternal suture separating metepistemum and metaventrite; the presence of this suture in apterous weevils is unusu- al. weakly broadened posterad (Fig. 4E); dorsal outline of ros- trum-to-head junction even (Fig. 5B), evenly and weak- ly depressed (Figs 1 A, 5A) or marked more abruptly (Fig. 5D). Dorsal body puncture and pilosity variable, from mostly faint (Fig. 5D) to markedly developed (Fig. 5B). Material examined. Holotype male (IZCAS): #0928, “P.R. CHINA, Yunnan, E slope N Gaoligongshan, N27°47’22. 1 E098°32’17.7, 24.V.2010, 3027m, sifting20, V.Grebennikov”. Paratypes (CNC, IZCAS, MTD), 19 in total; 4 exx, #0114-6, #0503: “P.R. CHINA, Yunnan, E slope N Gaoligongshan, N27°46.212’ E098°36.221\ 12.vi.2009, 2312m, sifting02, V.Grebennikov”; 5 exx, #0144 and 4 not numbered: “P.R. CHINA, Yunnan, E slope N Gaoligongshan, N27°46.782’ E098°33.087\ 14. vi. 2009, 2956m, sifting04, V.Grebennikov”; 2 exx, not numbered: “P.R. CHINA, Yunnan, E slope N Gaoligong- shan, N27°46.665’ E098°33.32 1 ’, 15.vi.2009, 2982m, sift- ing05, V.Grebennikov”; I ex, not numbered: “P.R. CHI- NA, Yunnan, E slope N Gaoligongshan, N27°46.635’ E098°33.381’, 16.vi.2009, 2946m, sifting07, V.Greben- nikov”; 2 exx, #0927 and 1 not numbered: same data as holotype; 1 ex, #0977: 2010/21 “P.R. CHINA, Yunnan, E slope N Gaoligongshan, N27°47’22.1 E098°32’17.7, 24.V.2010, 3027m, sifting21, V.Grebennikov”; 4 exx, #1017-20: “P.R. CHIN A, Yunnan, E slope N Gaoligong- shan, N27°59’01 .0 E098°32’56.9, 27.V.2010, 3018m, sift- ing22, V.Grebennikov”. Additional non-type specimens: Cang Shan Mt. Range (15 in total, CNC): 14 exx, #0300-1, #0427, #0499-502, 7 not numbered: “P.R. CHINA, Yunnan, Cangshan, N25°40.381’ E100°08.016\ 30.V.2008, 2600m, sifting, V.Grebennikov”; 1 ex, #0867: “P.R. CHINA, Yunnan, E Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK Morimotodes, a new weevil genus from China 129 Fig. 4. Morimotodes ismene g. & sp. n., unsexed, dorsal view. A: #2409 from Mount Gongga, Sichuan; B: #4483 from Mount Haba, Yunnan; C: #6185 from Mount Emei, Sichuan; D: #1105 from Mount Emei, Sichuan; E: #0867 from Cang Shan Mt. Range, Yunnan; F: #0251 from Mount Emei, Sichuan. Scale: 1 mm. slope Cangshan at Dali, N25°40’15.5 E100°07’45.4, 18.V.2010, 2657m, sifting 1 7, V.Grebennikov”. Mount Ha- ba (6 in total, CNC): 4 exx, #4481-4: “CHINA, Yunnan, Haba Shan, N27°21’01“ El 00°05’44”, 21.vi.2012, 4072m, sift26, V. Grebennikov”; 2 exx, #4536-7: “CHI- NA, Yunnan, Haba Shan, N27°21'01“ E100°05’44”, 28.vi.20 12, 4072m, sift35, V. Grebennikov”. Mount Gongga (9 in total, CNC): 1 ex, #2409: “P.R. CHINA, Sichuan, NE slope Gongga Shan, N29°48’I5” E102°03’44”, 06.vi.201 1, 2765m, siftl 1, V.Grebennikov; 1 ex, #2456: “P.R. CHINA, Sichuan, NE slope Gongga Shan, N29°50’50” EI02°02’28”, 09.vi.2011, 3170m, sift 1 4, V.Grebennikov”; 3 exx, #2471-3: “P.R. CHINA, Sichuan, NE slope Gongga Shan, N29°50’05” E102°02’53”, 1 1 .vi.201 1, 3019m, siftl 5, V.Grebennikov”; 2 exx, #2559-60: “P.R. CHINA, Sichuan, NE slope Gong- ga Shan, N29°49’33” E102°03’06”, 21. vi.201 1, 2977m, sift24, V.Grebennikov”; 3 exx, #2562-3: “P.R. CHINA, Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK 130 Vasily V. Grebennikov Fig. 5. Morimotodes ismene g. & sp. n., unsexed, left lateral view. A: #2409 from Mount Gongga, Sichuan; B: #4483 from Mount Haba, Yunnan; C: #6185 from Mount Emei, Sichuan; D: #1 105 from Mount Emei, Sichuan; E: #0867 from Cang Shan Mt. Range, Yunnan; F: #0251 from Mount Emei, Sichuan. Scale: 1 mm. Sichuan, NE slope Gongga Shan, N29°49’29” E102°03’24”, 21 .vi.201 1, 2986m, sift25, V.Grebennikov”. Mount Emei, clade EM-A (10 in total, CNC): 10 exx, #1216-9: #6184-9: “P.R. CHINA, Sichuan, EmeiShan, N29°32’37.3 E103°19’57.5, 18.vi.2010, 2440m, sifting37, V.Grebennikov”; clade EM-B (6 in total, CNC): 4 exx, #0251-3, #0504: “P.R. CHINA, Sichuan, Emei Shan, N29°32.932’ EI03°20.466\ 01.vii.2009, 2310m, sift- ing 1 4, V.Grebennikov”; 1 ex, #0505: “P.R. CHINA, Sichuan, Emei Shan, N29°32.806’ E103°20.106\ 03.vii.2009, 2342m, sifting 1 5, V.Grebennikov”; 1 ex, #1172 and #4186 (two numbers on the same specimen): “PR. CHINA, Sichuan, EmeiShan, N29°32’37.3 E103°19’57.5, 18. vi. 2010, 2440m, sifting37, V.Greben- nikov”; clade EM-C ( 1 in total, CNC): 1 ex, #1105: “P.R. CHINA, Sichuan, EmeiShan, N29°33’36.3 E103°20’38.0, 15.vi.2010, 1947m, sifting33, V.Grebennikov”. Distribution. The species is known from five isolated high altitude forest regions in Yunnan and Sichuan provinces (Fig. 9). Etymology. The species epithet is the Latinized Greek mythical name of Ismene, a daughter and half-sister of Oedipus, daughter and granddaughter of Jocasta, and sis- ter of Polynices; noun in apposition. Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK Morimotodes , a new weevil genus from China 131 sternite 9 Fig. 6. Morimotodes polynices g. & sp. n., holotype, male, #5949 from Taiwan. A-D: habitus; E-G: aedeagus and tegment, dor- sal (E), ventral (F), lateral; (G); H: sternites 8 and 9. Morimotodes polynices sp. n. Fig. 4. Diagnostic description. Holotype, male (Fig. 6). DNAnot sequenced. Length: 2.00 mm. Dorsal outline of pronotum in lateral view with hump in basal third; pronotum and ely- tra dorsally with fine punctures and pilosity not visible at magnification below 30x; each elytron without longitu- dinal groove parallel to edge. Aedeagus rounded lateral- ly, sternite 9 Y-shaped and about 3-5x wider than apodeme; sternal apodeme 9 without extention anterior- ly. Intraspecific variation. Length: 1.98-2.12 mm. Material examined. Holotype male (CMN): #5949 “TAI- WAN, Kaohsiung Hs. Rd. Abv. Tona For. Sta. Km 16-17, 1700-1 800m 28. V. 1998, A. Smetana (T190)”. Paratypes (CNC, CMN), 26 in total: 2 exx, #5925-6: “TAIWAN, Kaohsiung Hsien, Kuanshan trail at Kaunshanchi Riv. 2400 m 24.VI. 1992 A. Smetana (T94)”; 9 exx, #5927-35: “TAIWAN, Kaohsiung Hsien, Peinantashan trail, 2000m 7. VII. 93 A. Smetana (T144)”; 1 ex, #0587: “TAIWAN, Kaohsiung Hsien, Kuanshan trail above Kaunshanchi Riv., 2550m, 22.VII.93 A. Smetana (T160)”; 1 ex, #5936: “TAI- WAN, Kaohsiung Hsien, Peinantashan trail 2450m, 2. V. 1995 A. Smetana (T170)”; 13 exx, #5940-8, #5950-3: Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK 132 Vasily V. Grebennikov Morimotodes ismene g. et sp. n. . Mt. Emei, Sichuan, clade EM-B #4186 #0252.1 #0253.1 #0505 #0251.1 Mt. Gongga, Sichuan 0 94 #2563 #2471 #2559 #2562 #2456 # 2409 #2473 #2472 #2560 . Mt. Haba, Yunnan 0.85 #4483 #4537 #4484 #4482 #4481 #4536 022 0.05 substitutions/site S.--14 0.03 0.01 0,57 0.07 0.24 0.33 0.23 , Mt. Emei, Sichuan, clade EM-A t Mt. Emei, Sichuan, clade EM-C #6185 #6186 . #6184 .#6188 ■#1216 - #1217 #6189 #1218 #1219 0.37 7 [ Cang Shan Mt. Range, Yunnan #1105 #0867 #0500 #0502 #0501 #0499 #1017 #1018 #1020 2^3 Gaoligong Shan Mt. Range, Yunnan 0.7 0.62 0.74 #0503 #0115.1 #0114 #0116.1 #0927 #0928 Holotype #0144 #0977 Leiosoma reitteri Lobosoma rausense Niphadomimus spp 0.88 Ectatorhinus adamsi Typoderus furcatus Acall/nus tuberculatus Hylobius sp. Adexius scrobipermis Plinlhus spp Anchonidium unguiculare Aparopion costatum Pissodes punctatus Thalasselephas spp Euthycus sp. Otibazo polyphemus Thromboslernus cuculatus Fig. 7. Maximum Likelihood inference phylogram positioning monophyletic Morimotodes ismene g. & sp. n. (in black) as a sis- ter to Leiosoma reitteri among other analysed Molytinae genera (in grey) using the 658 nt of the mtDNA barcoding COl gene fragment. Digits at intemodes are bootstrap values. The tree is rooted on Graptus circassicus (Entiminae; not shown). Seven ge- ographically linked lineages of M. ismene g. & sp. n. are marked in red; note that Mount Emei uniquely harbours three unrelated lineages. Red and italicized numbers are those of imaged specimens (Figs 1, 4, 5) illustrating respective clades. Two pink ovals with question marks denote two nodes not recovered in the alternative BI analysis (Fig. 8). Inserted image of the forest illustrates exact spot where the type specimen #0928 was found. same data as holotype. Additional non-type specimens: 3 exx, (CMN), #5937-9: “TAIWAN, Ilan Hsien Shen-Mi Lake 24°22’43”N 121°44’12”E, 1110m I0.V.1995 A. Smetana (T 1 77)”. Distribution. The species is known from two widely sep- arated regions in southern and northern Taiwan, respec- tively (Fig. 9). Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK Morimotodes , a new weevil genus from China 133 Pliocene Pleistocene Mt. Gongga, Sichuan Mt. Haba, Yunnan Mt. Emei, Sichuan, clade EM-A Cang Shan Mt. Range, Yunnan Mt, Emei, Sichuan, clade EM-C Mt. Emei, Sichuan, clade EM-B Gaoligong Shan Mt. Range, Yunnan Fig. 8. Ultrametic tree of Morimotodes ismene g. & sp. n. reconstructed using Baeysian inference method strict linear molecular clock approach to analyse 658 nt of the mtDNA barcoding COl gene fragment and calibrated at a rate of 0.018 substitutions/site/MY. Numbers at nodes and on the scale below are million years before present. Node bars represent 95% confi- dence intervals of the age estimate. Two pink ovals with question marks denote two nodes not recovered in the alternative ML analysis (Fig. 7). Fig. 9. Known distribution of Morimotodes g. n. Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK 134 Vasily V. Grebennikov Etymology. The species epithet is the Latinized Greek mythical name of Polynices, the son of Oedipus and Jo- casta, as well as a grandson of Jocasta; noun in apposi- tion. RESULTS The first tree search resulted in a Maximum Likelihood tree (Fig. 7) with the highest log likelihood of- 9190.09. All analyzed specimens of M. ismene g. & sp. n. formed a weakly supported clade (bootstrap value 25%). The genus Morimotodes g. n., represented by only its type- species, formed a moderately supported (52%) clade with the genus Leiosoma. Forty-six analyzed specimens of M. ismene sp. n. from five mountain regions formed seven geographical lineages (one monotypical, six others sup- ported with 100% bootstrap). Four of these lineages cor- responded to four mountain regions (Fig. 7), while all 15 analyzed specimens from Mount Emei formed three groups not most closely related to each other. Temporal (second) analysis in BEAST (Fig. 8) recovered all Mori- motodes specimens forming a clade with the same seven lineages as in the first analysis, although the grouping among themselves was not identical with the results of the first analysis. The inferred timing of the splits forming the seven geographically linked lineages within monophylet- ic M. ismene g. & sp. n. detected in the second analysis date between 10.34 MY to 3.46 MY (Fig. 8). DISCUSSION Monophyly and sister group of Morimotodes g. n. As presently defined, the genus Morimotodes g. n. is most likely monophyletic. This assertion is not tested by the cladistic methods, and is based entirely on circumstantial evidence. None of the 29 specimens of M. polynices g. & sp. n. responded positively to a DNA amplification effort and, therefore, the species was not represented in the DNA analyses. Continental populations of Morimotodes g. n. from Yunnan and Sichuan all assigned to a broadly defined M. ismene g. & sp. n. were consistently recovered as a monophyletic clade in the first tree analysis (Fig. 7). The assumption of both Morimotodes g. n. species forming a clade is based entirely on their adult external similarities, further corroborated by the analogous collecting circum- stances suggesting similar biology. The geographical gap of about 2000 km separating known distributional areas of both species is not large enough to shed significant doubts on their sister group relationships. Such are all presently available data to justify the implemented taxo- nomic decisions of grouping both species into a presum- ably monophyletic new genus. The most surprising aspect of the first analysis is the rel- atively weak statistical support for M. ismene g. & sp. n. (25%, Fig. 7), which can be at least partly attributed to the known phylogenetic limitations of COl (Maddison 2012), including the saturation effect. The sister-group relationships between Morimotodes g. n. and the West Palaearctic Leiosoma is moderately sup- ported (bootstrap 52% ( Fig. 7). Close affinities of both genera have been hypothesised ever since the first repre- sentatives of Morimotodes g. n. came to notice in 2008, and was grounded entirely on adult morphological simi- larities (small weevils less than 3 mm in body length with laterally rounded and not flattened bodies and with shiny dorsum) and biological preferences (inhabitants of the de- ciduous leaf litter). Analogous expectations for the simi- larly shaped genus Otibazo were, however, rejected by the ML tree, the latter clustered with the genus Euthycus (Fig. 7). At present, it is impossible to suggest phylogenetic affinities of the clade Morimotodes g. n. + Leiosoma , since the available data are highly inadequate. Pending further investigation, the genus Morimotodes g. n. is placed in the same higher taxonomic group containing its sister group Molytinae: Molytini: Molytina. If Morimotodes g. n. is indeed sister to Leiosoma , then attribution of both new species from China to the former genus could be a phylogenetically sound taxonomic alter- native. It was, however, rejected because establishing a new genus better reflects the significant external dissim- ilarity between their members and the wide gap in their known geographical distribution. Species delimitation of Morimotodes g. n. Both species of Morimotodes g. n. as presently defined are much dissimilar to each other. The species from Tai- wan, M. polynices sp. n., is known from a museum series of externally remarkably similar specimens, all but three of which were sampled from a geographically confined region (Kaohsiung, Fig. 9) not more than 50 km in its greatest detention and within the limited altitudinal range ( 1 700-2550 m). This evidence suggests that all type spec- imens most likely share the MRCA. Three additional non- type specimens from north of Taiwan ( llan. Fig. 9) are ex- ternally most similar to those from the type locality and, therefore, are thought to be conspecific, although with lesser certainty. These two hypotheses could not be test- ed using DNA data, since the sequences were not obtain- able. Judging from the morphological and geographical data, it seems unlikely that the Taiwanese specimens of Morimotodes g. n. could be grouped within the clade formed by their continental counterparts. This evidence, although not tested phylogenetically, strongly suggests separate evolutionary history for the Taiwanese lineage worthy, therefore, of establishing a formal species. Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK Morimotodes , a new weevil genus from China 135 Deciding what to all a species for the continental mem- bers of Morimotodes g. n. is ambiguous. The most inclu- sive and conservative approach is, therefore, employed. All specimens of Morimotodes g. n. from continental Chi- na are presently assigned to the broadly defined M. ismene sp. n. This uncertainty on species boundaries is not made any easier by the relatively large amount of available rel- evant data for the continental species. The implemented decision to assign all continental Morimotodes g. n. to a single species is a balanced temporary solution implement- ed to minimize the conflict among four independent lines of evidence (in the increasing order of their contribution): ( 1 ) biological characteristics, (2) morphological similar- ity/dissimilarity (3) geographical distribution, and (4) mtDNA sequences. The logic of their combined evalua- tion is described below. The condition determining the gene flow among Mori- motodes g. n. populations inhabiting five continental lo- calities (the Cang Shan and the Gaoligong Mountain Ranges, as well as Mounts Haba, Gongga and Emei; Fig. 9) can be considered analogous to those acting for terres- trial organisms confined to oceanic islands. This approx- imation is commonly used for the low-dispersing and flightless moisture- and habitat-dependent species found in relatively wet isolated forests widely separated by seem- ingly unsuitable lowlands. Such estimate might perhaps be even an understatement of the dispersal difficulties for the mountain species, since the salt water barriers are oc- casionally crossed by seemingly most unseaworthy organ- isms such as chameleons possibly rafting on dislodged floating trees, or blind subterranean Arthropods perhaps surviving the passage in soil attached to their roots (Rax- worthy et al. 2002 and Peck 1 990, respectively). The very first task with such allopatric mountainous low-dispers- ing organisms as Morimotodes g. n. was, therefore, to as- sess whether the specimens from each separate locality form distinct evolutionary clusters and whether any of the localities supports more than one such cluster. The four aforementioned lines of evidence contributed conflicting data. Biological data were the least informa- tive, since all sampled specimens of M. ismene sp. n. were sifted from seemingly similar forest litter and within com- parable altitude range. The only exception was shown by the Mount Haba specimens recovered from the upper for- est zone at altitude notably higher than known for other continental populations. Adult external morphology was the second least informative line of evidence (see below on the use of genital characters). Although the imaged specimens representing each of seven clades were some- what dissimilar to each other (note the hairy specimen #4484 from Mount Haba, or the large and non-hairy spec- imen #1105 representing the clade EM-C from Mount Emei; Figs 4, 5), it was impossible to consistently distin- guish the others. The most powerful signal came from the mtDNA sequences, which grouped specimens from four localities into four strongly supported corresponding clades (Fig. 7). The exception was Mount Emei, where 14 specimens clustered in two unrelated groups (EM- A and EM-B on Fig. 7) with an additional third equally unrelat- ed lineage (EM-C) represented by a single specimen #1 105. The emerging distributional pattern indicated that each locality supports at least one lineage of Morimotodes g. n., while Mount Emei has at least three such lineages. The main operational problem was that the majority of the seven detected radiations could neither be uniquely char- acterized morphologically, nor biologically. It should be noted that detecting seven evolutionary sep- arate lineages of continental Morimotodes g. n. inhabit- ing five discrete mountain regions is not prohibitive in it- self. There are no reasons to expect that no more than a single radiation of closely related low-dispersing organ- isms should exist in any suitable locality. Among all five continental localities. Mount Emei should be rightfully ex- pected to be the most diverse in Morimotodes g. n., be- cause it is overwhelmingly rich in other forms of life and by that exceeds many other comparable localities. For ex- ample, its relatively well documented flora condensed on just 154 km2 and lacking the entire alpine zone, encom- passes an astonishing 3703 species of higher plants (Wang et al. 2013). This diversity exceeds Canada (3,156; Mosquin et al. 1995) and is comparable with that of the British Isles (3842, Fitter & Peat 2014). Summing up, the phylogeographic pattern as represented on Fig. 7 did not pose significant biological inconsistencies. By far the ma- jor challenge was the necessity to implement the Linnaean taxonomy and decide on how many nominal species are there. At this stage the purely practical limitations of available material has to be mentioned. Mostly, specimens of Mo- rimotodes g. n. are relatively rarely encountered. For ex- ample, the radiation inhabiting the Cang Shan Mountain Range responded well to the original sifting in 2008, how- ever, DNA sequencing of the 2008 specimens resulted in only short fragments of 407 bp (four specimens #0499-0502). During the subsequent two visits to Cang Shan in 2010 and 2012, a significant greater sampling ef- fort resulted in the collection of only a single specimen (#0867), but provided a full-length DNA barcode. Mount Emei has also been visited three times with one month of intense fieldwork in total. Among the 27 sifting samples, only five contained representatives of Morimotodes g. n. Four focussed samples were taken in two years within 50 meters from where the unique specimen #1105 (lineage EM-C, Fig. 7) had been sifted, but 1 failed to discover ad- ditional specimens. The entire lineage EM-A was recov- ered from a single sample. Only the lineage EM-B was detected in three samples, two of them taken in exactly the same place in different years and the third one some 650 m away. Such sampling results strongly suggest that specimens of the continental Morimotodes g. n. are most Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK 136 Vasily V. Grebennikov unevenly distributed in time and across seemingly iden- tical landscape of mountainous forest. Species delimitation in insects often relies on the shape of male genitalia. However, due to the scarcity of Mori- motodes g. n. specimens coupled with possible structur- al damages during dissection of the minute genitalia, I re- frained to examine genitalia of all sequenced specimens. It was impossible to sex specimens externally and, there- fore, male genitalia dissection had to be attempted with a randomly chosen specimen, some of them females. Even more problematic were the difficulties linked to the me- chanical body properties of these highly compact minute beetles with globular elytra firmly fused to the hind body and forming together a most robust, inflexible and, accord- ingly, highly breakable capsule. After the muscle tissue was nearly completely dissolved and the remaining inter- nal structures much loosened, the genitalia were success- fully dissected for the two holotypes, only. It was, there- fore, judged inadvisable to perform extensive male gen- italia dissections. The practical conclusion reached after considering all was to define preliminarily a single species for all conti- nental representatives of Morimotodes g. n., even though this taxon might be later split into more separate species. The Haba Shan lineage having distinct external adult mor- phology further corroborated by the unique biological preferences of higher altitudes can perhaps merit its own species even at present. This action, however, will render the rest of the continental Morimotodes g. n. paraphylet- ic and, therefore, is not implemented. Considering a like- lihood of further taxonomic changes in the broadly defined M. ismene sp. n., the type series is purposely restricted to the specimens forming a separate lineage uniquely char- acterised by DNAand geographical data. Specimens from the Gaoligong Mountain Range forming the extreme west of the known generic distribution (Fig. 9) were chosen for this purpose, specifically because they are numerous in the DNA analysis. Additionally, this lineage is represented by specimens recovered in at least six separate samples tak- en from the area measuring 25 km in its greatest dimen- sion and, therefore, having the least chances of harbour- ing an undetected sympatric congeneric lineage (i.e., like Mount Emei with at least three such lineages). Conse- quently, all specimens of M. ismene sp. n. from localities other than the Gaoligong Mountain Range are not desig- nated as paratypes. Dated phylogeography of Morimotodes ismene g. & sp. n. Little can be presently said about the temporal aspect of Morimotodes g. n. evolution. The single and seemingly most interesting aspect of it, namely the time of the im- plied separation between continental M. ismene sp. n. and M. polynices sp. n. from Taiwan cannot be addressed at present because of the lack of mtDNA data to represent the latter species. Even though all seven lineages consti- tuting monophyletic M. ismene sp. n. were consistently re- covered in the first (ML) and the second (BI) analyses, their grouping was consistent in only three nodes (Fig. 8): (a.) the split between lineages inhabiting Mount Haba and Mount Gongga, (b.) their MCRA separating from the lin- eage EM-A inhabiting Mount Emei and (c.) the split be- tween Mount Emei clade EM-B and the lineage from the Gaoligong Mt. Range. All four consistently recovered evo- lutionary events (three aforementioned and the origin of the entire M. ismene sp. n.) date 4.45 MY, 8.08 MY, 3.46 MY and 10.34 MY, respectively. These estimations pro- vide a very approximate time frame for the origin and di- versification of the continental Morimotodes g. n., which is comparable to those of the sympartically distributed weevil genera Niphadomimus Zherikhin, 1987 (Greben- nikov 2014) and Notaris Gennar, 1817 (Grebennikov & Kolov, unpublished) likely linked to the advanced stages of the uplift of the Tibetan Plateau (Favre et al. 2014). Illustrated overview of genera of the subtribes Molyti- na and Plinthina The catalog of Alonso-Zarazaga & Lyal ( 1 999, 2002) had all genera currently assigned to Molytina and Plinthina al- ready listed as such (except for two proposed later) and all included in the broadly defined Molytini. Since then the following changes were proposed: ( 1 ) the monotypic Stenanchonus Voss, formerly of Anchonini, was syn- onymized with Euthycus Pascoe of Molytina (Meregalli 2003); (2) some species of the genus Minyops Schoenherr of Plinthina was assigned to the newly erected Paraminy- ops Osella & Bello (Osella & Bello 2010); (3) the tribe Sthereini was re-erected to incorporate nine genera, four of them formerly of Plinthina (Alonso-Zarazaga 2013; Lyal 2014); (4) the former Molytini subtribe Typoderini was elevated to tribe level; (5) the former Molytini sub- tribe Leiosomatina was synonymized with Molytina (Lyal 2014); (6) the former tribe Hylobiini was synonymized with Molytini by forming its third and the last subtribe (Lyal 2014); (7) one new genus, Morimotodes g. n., be- ing described herein. Moreover, the Palaearctic represen- tatives of both subtribes accounting for about 80% of their species were recently catalogued (Alonso-Zarazaga 20 1 3). All 30 genera presently assigned to both subtribes are listed below and 27 of them are illustrated. This summa- ry was done with the intention to bring them to the no- tice of the contemporary scientific community and hope- fully trigger their reinvestigation in a phylogenetic frame- work using modem data and analytical methods. Up to now very little of such work has been done for the entire subfamily Molytinae, including both Molytina and Plinthi- na. Most importantly, reciprocal monophyly of both sub- Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK Morimotodes, a new weevil genus from China 137 tribes has never been demonstrated and, therefore, their taxonomic existence as currently defined seems very pre- carious yet. Moreover, with the exception of Craw’s (1999) detailed analysis of the New Zealand’s Hadram- phus & Lyperobius , none of the non-monotypical genera shown in detail below has ever been tested for its mono- phyly. Some phylogenetically relevant statements were, however, made: Meregalli (1985) suggested a sister-group relation between Minyops (then containing Paraminyops) and Plinthus. Davidian (2008) proposed that the lineage presently consisting of Paraminyops and Minyops is most closely related to a subgroup of the genus Plinthus (ren- dering the former paraphyletic) and that Hoplopteridius forms a clade with all of them. Osella & Bello (2010), when separating their new genus Paraminyops from Miny- ops, suggested that both genera together with PIo- plopteridius form a group (“subtribe Minyopina”, a syn- onym of Plinthina, Bouchard et al. 2011 ). None of these hypotheses have been analytically tested. Subtribe Molytina The tribe includes 13 genera, which are numbered and list- ed alphabetically below. Nine of them, as noted by Lyal (2014), are widely scattered from New Zealand and Aus- tralia, through New Guinea, Borneo, Java, and Indochi- na, to Sri Lanka, eastern India, China and Japan. Three other genera ( Anisorhynchus , Leiosoma and Liparus) are distributed in the Western Palaearctic, with the former al- so known from Central Asia (Alonso-Zarazaga & Lyal 1999). One genus known only from the holotype (Ptero- tomus) is restricted to Angola. Anisorhynchus Sehoenherr, 1842 (Figs 10A, B). This is a genus of 13 large-bodied species and one non-nomina- tive subspecies restricted to the West Palaearctic and dis- tributed mainly around the Mediterranean Sea (Alonso- Zarazaga 2013). Apterylobius Marshall, 1942 (Figs 10C, D). Fewer than a dozen specimens of this genus are known, all exclusive- ly from Mount Kinabalu in northern Borneo. Since its de- scription (Marshall 1942), the genus contains two species and one non-nominative subspecies, and no newly collect- ed specimens have been since reported. Carbonomassula Heller, 1908 (Figs 10E, F, 16A). Spec- imens of this genus, apart from their much greater size, markedly resemble those of Morimotodes g. n. in having similarly evenly rounded and dorsally lustrous body of reddish or brownish color (compare Figs 6A and 1 0E). The genus contains the single species from Papua New Guinea, C. glaberrima Heller, 1908. The original description was based on an unknown number of syntypes and accompa- nied by a high quality habitus drawing (Heller 1908). The type locality is “Nova Guinea Britannica, ad flumen Aroa”. Two syntypes are in MTD, the smaller one best fit- ting the body length given in the original description is illustrated herein (Figs 10E, F). Both syntypes have a blue rectangular label “14523” (Fig. I6A), even though the original description mentions “14525”. One additional and seemingly conspecific specimen supplied to BMNH by M.E. Bacchus in 1965 has the locality label “NEW GINEA: Morobe Dist., Herzog Mts., Vagau, C.4,000 ft. 4-7.L1965”. No other specimens of this genus have been reported (Set! iff 2007). Clarkanthus Alonso-Zarazaga & Lyal, 1999 (Figs 10G, H, 1 6B). This genus consists of two poorly known species from Sri Lanka and the adjacent part of India. Its type species, C. taprobanus (Faust, 1895) was described from the holotype collected on “Ceylon". The genus has long been known under a preoccupied generic name Ischno- pus Faust, year (non Amyot). Second species, C. madrasensis Pajni & Sood, 1982, was described from a holotype collected in the South Indian state of Tamil Nadu just across the Palk Strait from Sri Lanka. Except for the two type specimens, representatives of the genus have not been reported in the literature. Two additional historical specimens are present in MTD: one seemingly conspecif- ic with the type species and labelled “Ceylon Plason” in Faust’s handwriting, and another at least congeneric and with a printed label “India or. Madura”. The locality might perhaps refer to the Madurai District in Tamil Nadu, which is the type locality of C. madrasensis. These four speci- mens are the sole representatives of this obscure genus. Euthycus Pascoe, 1885 (Figs 11 A, B). This genus em- braces 12 species distributed at the junction between the Palaearctic and Oriental Regions and recorded from Japan, southern China and Taiwan, southwards to Borneo and Ja- va, and westwards to Assam, Sikkim and Andaman Islands of India (Meregalli 2003). Little is known of their natu- ral history, aside from appearing to be associated with mid- altitude primary mountainous forests. Observation of two live specimens personally collected by sifting litter in Tai- wan and represented in the DNA analyses revealed re- markably strong adhesive power of their tarsi. This sug- gests that the beetles are climbers rather than true litter inhabitants (such as Morimotodes g. n.), where they might be encountered only accidently. Hadramphus Broun, 1911 (Figs 1 1C, D). This genus in- cludes four relatively large-bodied species restricted to New Zealand and some nearby islands and was revised by Craw (1999, including a summary on the immature stages and host plants). Together with Lyperobius , this genus can be considered as a weevd analogous to the moa or weta (i.e., flightless giant birds or crickets), that evolved Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK 138 Vasily V. Grebennikov * 1 Fig. 10. Representatives of the Molytini: Molytina genera. A-B: Anisorhynchus barbatus (Rossi), Italy, Abruzzo, no date, no col- lector, type species of Anisorhynchus , MTD; C-D: Apterylobius basalis (Heller, 1922), Malaysia, Borneo, no date, no collector, ?syntype, MTD; E-F: Carbonomassula glaberrima Heller, Papua New Guinea, syntype, see text, type species of Carbonomassu- Icy MTD; G- H: Clarkanthus taprobanus (Faust), Sri Lanka, holotype, see text, type species of Clarkanthus , MTD. in the unique New Zealand biota mostly deprived of ter- species quickly driven to extinction by invasive humans, restrial mammals and, therefore, highly vulnerable to in- all but one ( Tymbopiptus valeas Kuschel, 1987) known vasive predatory species. Unlike the moa with all nine large New Zealand weevils continue to survive, despite Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK Morimotodes, a new weevil genus from China 139 Fig. 11. Representatives of the Molytini: Molytina genera. A-B: Euthycus macilentus Pascoe, 1885, Indonesia, Java, Mt. Guntur, no date. Overbeck, type species of Euthycus, MTD; C-D: Hadramphus tuberculatus (Pascoe), New Zealand, Christchurch, 1910, no collector, BMNH. image: Vasily Grebennikov, © The Natural History Museum, London; E-F: Leiosoma deflexion (Panzer, 1 795), Poland, Muszkowice, 6.V.1986, M.Wanat, type species of Leiosoma , CNC; G-H: Lipams glabrirostris (Kiistcr, 1849), Poland, Sudetes, 1 5.vii. 1 975, M.Kupczynski, CNC. loosing most of their former range (Kuschel & Worthy 1996). Recently an extant population of H. tuberculatus (Pascoe, 1877), a species thought to be extinct, was dis- covered (Fountain et al. 2013). Kyliparus Moromoto, 1982 (not illustrated). This genus was proposed to accommodate a single species from Japan, K. lewisii (Roelofs), previously attributed to Li- parus. The original description of Kiliparus contains a habitus dorsal image of a single female studied by Mori- moto (1982, fig. 46E). Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK 140 Vasily V. Grebennikov Leiosoma Stephens, 1829 (Figs 11H, F). All 32 species and one non-nominative subspecies of this genus are re- stricted to the West Palaearctic, particularly the northern Mediterranean, with two species extended eastwards to the Caucasus (Alonso-Zarazaga 2013). Numerous species de- scriptions were made for localized allopatric populations and, therefore, a revision is desirable to assess their sta- tus as that of biologically valid entities. Liparus Olivier, 1807 (Figs 1 1 G, FI), the type genus of the subtribe Molytina, the tribe Molytini and the subfam- ily Molytinae. The genus includes 1 7 species and two non- nominative subspecies (Alonso-Zarazaga 2014) of large- bodied weevils. The genus is most diverse in the West Palaearctic, while two species of the subgenus Li- parorhynchus Reitter form the extreme eastward bound- ary of the generic distribution and are found in Central Asia (Uzbekistan, Tajikistan, Afghanistan, Iran; Bajtenov 1974; Alonso-Zarazaga 2013). Lyperobius Pascoe, 1876 (Figs 12A, B). Along with Hadramphus, Lyperobius is another genus of large-bod- ied and flightless weevils endemic to New Zealand. The majority of its 1 6 species have highly restricted distribu- tions due to human-induced habitat changes, including predation by introduced rodents (Craw 1999). Morimotodes g. n. (Figs 1-9). The genus includes two small-bodied litter species from Southwestern China and Taiwan, respectively, and is a sister to the West Palaeaerc- tic Leiosoma. See the taxonomic comments above. Pinacopus Marshall, 1932 (Figs 12C, D). This genus has 14 named species found in Malaysia (8 spp.), Thailand ( 1 sp. ), Laos (4 spp.) and northern India ( 1 sp.), with at least some specimens “collected from shrubs, which are more or less covered with lichens, by beating method” (Koji- ma 2010). Pterotomus Quedenfeldt, 1888 (Figs 12E, F, 16C). The genus includes a single species, P. moebiusi Quedenfeldt, 1888, known from the holotype collected in Angola’s province Malange and never reported since. The original description included a high quality habitus drawing (Que- denfeldt 1888). Subtribe Plinthina The subtribe Plinthina consists of 17 genera listed below. Eleven of them are restricted to the Mediterranean region and some, like Minyops, are also found in Central Europe, Ukraine, southern Russia and western Kazakhstan. Two genera, Kurilio and Metahylobius, with a total of three species are limited to the eastern Pacific (Ffonshu, Flokkaido and Kunashir). Four other inadequately known mono- and oligotypic genera are restricted to the Afrotrop- ical Region, namely Seychelles ( Gonotrachelus ), East Africa ( Molytophilus and Plinthinella ) and Madagascar ( Vouauxia). Adexius Schoenherr, 1834 (Figs 13A, B). This genus con- sists of three West Palaearctic species, two of them poor- ly known and narrowly distributed. The third one, A. scro- bipennis Gyllenhal, 1834 is found in forest leaf litter in most of Central and Southern Europe between Belgium and Ukraine (Alonso-Zarazaga 2013). Alloplinthus Solari, 1941 (Figs 13C, D). This genus con- sists of three species (originally described as Plinthus) from Tenerife and Gran Canaria (Alonso-Zarazaga 2013). Caulomorphus Faust, 1886 (Figs 13E, F). The genus con- tains 1 1 eyeless species from Turkey, Greece, Iran and Azerbaijan (Alonso-Zarazaga 2013). Osclla (1970) revised the genus taxonomically and later described three more species. Freyianus Zumpt, 1932 (Figs 13G, H). This is a monobasic genus for F liparoides Zumpt, 1932. It was de- scribed based on 10 syntypes (one of them adequately il- lustrated by a habitus photo) collected by “Kulzer” in “Ak- Schehir” (Zumpt 1932) and thought to be in Turkey (Alon- so-Zarazaga & Lyal 1999). Since its description, this genus has not been reported in the original literature and its iden- tity is inadequately known. The newly imaged specimen (Figs 13G, H), even if labelled as a “holotype”, is in fact a syntype and, together with two other syntypes, is stored in NHMB. Gonotrachelus Champion, 1914 (Figs 131, J, 16D). The genus includes a single species, G. quinquecarinatus Champion, 1914 described from two syntypes found on the Seychelles (Long Island, near Mahe). An additional specimen of this species was reported from Madagascar (Hustache 1925). The genus is characterized by the funi- cle consisting of fine antennomeres (Champion 1914, figs 11, lib) and was originally compared with Stypldoderes Wollaston (Typoderini). The type series (presently in BMNFI) was taken from seashores and was thought to be introduced via driftwood (Champion 1914). Hoplopteridius K. Daniel, 1908 (Figs 14A, B). This genus consists of five species and three non-nominative sub- species distributed in the northern and eastern Mediter- ranean (Alonso-Zarazaga 2013). Iberoplinthus Meregalli, 1986 (Figs 14C, D). This recent- ly erected genus (Meregalli 1986) consists of six species (Alonso-Zarazaga 2013) all found in northern Spain, while Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK Morimotodes, a new weevil genus from China 141 Fig. 12. Representatives of the Molytini: Molytina genera. A-B: Lyperobius huttoni Pascoe, 1876, New Zealand, Humboldt Mts., 30.xii. 1 944, C.E. Clarke, type species of Lyperobius , BMNH, image: Vasily Grebennikov, © The Natural History Museum, Lon- don; C-D: Pinacopus dentirostris Marshal, 1932, Malaysia, Perak, no date, no collector, MTD; E-F: Pterotomus moebiusi Que- denfeldt, Angola, holotype, see text, type species of Pterotomus , MNHN. the type species, I. imbricatus (Dufur, 1851), and is re- ported also from Andorra and southern France. Kurilio Zherikhin & Egorov, 1991 (Figs 14E, F). Its on- ly species, K. monachus Zherikhin & Egorov, 1 99 1 , is re- stricted to Hokkaido and Kunashir Islands (Zherikhin & Egorov 1990). Metahylobius Nakane, 1964 (Figs 14G, H). This genus consists of two species, both restricted to the alpine zone of Honshu: M.jonensis Nakane, 1964 and M. rubiginosus Morimoto, 1982. The adults of the former species “were captured by turning stones on the ground” (Morimoto 1982). Minyops Schoenherr, 1823 (Figs 141, J). This genus con- sists of 29 species and two non-nominative subspecies dis- tributed through most of southern Europe and also Turkey (Alonso-Zarazaga 2013). The genus was recently taxo- nomically revised with 22 of its 3 1 species-group taxa de- scribed as new and eight others re-validated from syn- onymy (Osella & Bello 2010). Mitoplinthus Reitter, 1897 (Figs 15 A, B). This genus con- sists of two species and one non-nominative subspecies of somewhat cylindrical medium-sized weevils distributed in southern parts of Europe between Spain and Ukraine. Meregalli (1986) provided the most recent treatment of the genus. Molytophilus Hartmann, 1904 (Figs 15C, D). The genus consists of three species, the type species Molytophilus carinatus Hartmann, 1904, described from Tanzania, and two more from Ethiopia: M. affinis Hustache, 1936, and M. puncticollis Hustache, 1936, whose affinities with the genus have not yet been demonstrated. Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK 142 Vasily V. Grebennikov Fig. 13. Representatives of the Molytini: Plinthinae genera. A-B: Adexius scrobipennis Gyllenhal, Poland, Holubla ad Przemysl, 4.vi,1997, M.Wanat, type species of Adexius, CNC; C-D: Alloplinthus musicus (Wollaston, 1860), Spain, Teneriffe, 1 7.V.1924, E.Ap- penhagen, type species of Alloplinthus, BMF1N, image: Vasily Grebennikov, ©The Natural History Museum, London; E-F: Caulo- morphus lederi (Chevrolat, 1880), Georgia, “Meskisch. Gb.”, no date, H.Leder, ?syntype, type species of Caulomorphus , MTD; G-H: Freyianus liparoides Zumpt, Turkey, “Ak-Schehir”, 22.V.1926, Kulzer, syntype, type species of Freyianus, NHMB; 1 -J: Gono- trachelus quinquecarinatus Champion, Seychelles, see text, syntype, type species of Gonotrachelus, BMNH, image: Vasily Greben- nikov, © The Natural History Museum, London. Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK Morimotodes, a new weevil genus from China 143 Fig. 14. Representatives of the Molytini: Plinthinae genera. A-B: Hoplopteridins lutosus (Frivaldszky, 1835) Italy, Rom, no date, no collector, type species of Hoplopteridius , MTD; C-D: Iberoplinthus curvirostris Meregalli, 1986 Spain, Asturias, 1 4.vii. 1 986, M.Meregalli, paratype, BMNH, image: Vasily Grebennikov, © The Natural History Museum, London; E-F: Kurilio monachus Zherikhin & Egorov, Russia, Kunashir Isl., 1 6.ix. 1 975, no collector, type species of Kurilio , CNC; G-H: Metahylobius jonensis Nakane, Japan, Honshu Isl., 1 .vii. 1 986, S.J. Martin, type species of Metahylobius , BMNH, image: Vasily Grebennikov, © The Nat- ural History Museum, London; 1 J: Minyops carinatus (Linnaeus, 1767), Austria, Wien, no date, no collector, type species of Miny- ops , MTD. Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK 144 Vasily V. Grebennikov Fig. 15. Representatives of the Molytini: Plinthinae genera. A-B: Mitoplinthus caliginosus (Fabricius, 1775), Germany, Amstadt, 16.x. 1949, W.Liebmann, MTD; C-D: Molytophilus carinatus Hartmann, Tanzania, Usambara, no date, no collector, syntype, type species of Molytophilus, MTD; E-F: Neoplinthus tigratus porculus (Fabricius, 1801 ), Austria, Graz, no date, no collector, MTD; G-H: Plinthus confiusus Meregalli, 1985, Russia, Caucasus, 21. vi. 1974, J.Vorisek, paratype, CNC; I-J: Vouauxia onopatroidea Hus- tache, Madagascar, see text, syntype, type species of Vouauxia , MNITN. Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK Morimotodes, a new weevil genus from China 145 Q’L.H'.'tyw.lA, j>. Vivjk* Stoatl. Museum fur Tierkunde. Dresden 'JU SMer TijJurV ANGOLA Mahe, 1908-9 Seychelles Exp. (JoH, J. Fausts Ankauf 1900- u u~i {afrtd> 'f*uu6 t f-titctl- Sf). figured. MUSfeUM PARIS IX ^ullSSevv&l it COU '« QBE fl T HUR I 4 vrvCo cr\\\A+,ti n .yvn j «4a<> c4 & Stoat!. M Tlerkunde.Drasden . \At(x£cl'X4C (SUi ur.eurn fur ' ' I! ?eyt helle Islands. P^rcy Sladen Trust Expedition. 1913.— 170. Tr.Linn.S.L 1914. Goifotrachelus ! D^CQniuituSj Cli. n frialat, MUSEUM PARIS MADAGASCAR Collection Uon Fairmaire 1906 COTYPE Yo U~iMAOC>VCL* ^o/usch-aiclta Hwt. MUSEUM PARIS c 1949 Col. A. HUSTACHE Fig. 16. Type specimen labels. A: Carbonomassida glaberrima Heller, ST; B: Clarkanthus taprobanus (Faust), holotype; C: Ptero- tomus moebiusi Quedenfeldt, holotype; D; Gonotrachelus quinquecarinatus Champion, syntype, image: Vasily Grebennikov, © The Natural History Museum, London; E: Vouauxia onopatroidea Hustache, syntype. Labels within each species are of propor- tional size; habitus images are not to scale. Neoplinthus Bedel, 1884 (Figs I5E, F). This genus in- cludes five species and two non-nominative subspecies of elongated and parallel-sided weevils distributed in Cen- tral, Eastern and Southern Europe; all of them recorded from Italy (Alonso-Zarazaga 2013). Paraminyops Osella & Bello, 2010 (not illustrated). This genus was recently separated from Minyops by Osella & Bello (2010) to accommodate nine species and two non- nominative subspecies from northern and eastern Mediter- ranean. Plinthinella Hofmann, 1965 (not illustrated). This genus includes its sole member, the type species by original des- ignation P. tenella Hoffmann, 1965. This is a small bee- tle of some 2. 0-2. 2 mm in body length and funicle with seven antennomeres described from the holotype and 12 paratypes collected in 1962 by Herbert Frantz. The type locality in “Tanganyika: Kilimandjaro” (= Mt. Kiliman- jaro in Tanzania) and the altitude is unknown. When de- scribed, this taxon was compared with Matxaldiella glacialis (Hustache, 1929) (cited as “ Mimus glacialis ”) from Mount Kenya in Kenya (depicted in Hustache 1929: fig. 15). Since its description, Plinthinella tenella has not been reported in the original literature and its identity is inadequately known. The curatorial search to locate the type series in NMW failed, while my extensive recent sampling in the type locality did not rediscover the species. Both the generic and species names should, therefore, be considered as nomina dubia. Bonn zoological Bulletin 63 (2): 123-147 Plinthus Gerinar, 1817 (Figs 15G, H). This genus in- cludes 68 species and 17 non-nominative subspecies of medium- to large-bodied weevils widely distributed in the West Palaearctic. The majority of species are found in mountains between the Pyrenees and Alborz in northern Iran (Alonso-Zarazaga 2013). The Caucasus appears to have disproportionally large number of species normally restricted to alpine mountaintops. This is one of a few gen- era receiving significant attention in recent years, includ- ing detailed (although mainly unpublished) studies on in- ternal classification, structure of male genitalia, species distribution and biology (Davidian 2008). Vouauxia Hustache, 1920 (Figs 151, J, I6E). The genus includes a single species, O. onopatroidea Hustache, 1920, described from Madagascar from unknown number of syntypes. The original brief description (Hustache 1920) was followed by a more detailed one (Hustache 1925), both relating this taxon to Gonotrachelus Champion. At least three syntype are present in MNHN. No other spec- imens of this genus have been reported in the literature. CONCLUDING REMARKS This review revealed, how little is actually known about wingless Molytinae genera, especially those, which are presently assigned to the subtribes Molytina and Plinthi- na. The discovery of a peculiar new genus from China strongly suggests that similar findings still remain to be made, especially using specialized collecting techniques ©ZFMK 146 Vasily V. Grebennikov and addressing specific habitats harbouring cryptic and small-sized organisms. Equally remarkable is the acute scarcity of data on the ma jority of other genera herein dis- cussed. In many cases species and even genera were men- tioned only once when formally described. Many mono- and oligobasic genera such as Apteiylobius, Carbonomas- sula, Clarkanthus, Kyliparus, Pterotomus , Freyianus , Gonotrachehis , Molytophilus , Plinthinella and Vouauxia are particularly poorly known and at least some of them appear only distantly related to the type genera of their respective subtribes. A detailed morphological and, par- ticularly, DNA analysis of Molytinae is indeed long over- due. Much remains to be done with wingless Molytinae weevils and hopefully this paper helps to reinforce this seemingly self-evident conclusion. Acknowledgement. Ales Smetana collected all specimens of Morimotodes polynices g. & sp. n. in Taiwan. Curators of the collections mentioned above variously helped in accessing spec- imens under their care. Genrikh Ed. Davidian (St. Petersburg, Russia) identified the sequenced specimens of Plinthus. Marek Wanat (Wroclaw, Poland) sent imaged specimens of Adexius, Li- parus and Leiosoma. Eva Sprecher contacted on my behalf Is- abelle Ziircher-Pfander (both Basel, Switzerland), who counted syntypes and took images of Freyianus liparoides (Figs 1 3G, H). Thierry Deuve (Paris, France) sent a reprint of Hustache (1920). Bradley J. Sinclair and Eduard Jendek (both Ottawa, Canada) read an early draft of this work. REFERENCES Alonso-Zarazaga MA(2013) Molytinae. Pp. 475^197 in: Lobl I & Smetana A (eds.) Catalogue of Palaearctic Coleoptera, Vol- ume 8, Curculionoidea II. Brill, Leiden Alonso-Zarazaga MA, Lyal C'HC (1999) A world catalogue of families and genera of Curculionoidea (Insecta: Coleoptera) (excepting Scolytidae and Platypodidae). Entomopraxis, Barcelona Alonso-Zarazaga MA, Lyal CHC (2002) Addenda and corrigen- da to “A World Catalogue of Families and Genera of Cur- culionoidea (Insecta: Coleoptera)”. Zootaxa 63: 1-37 Bajtenov MS (1974) Zhuki-dolgonosiki (Coleoptera: Attelabidae, Cucrulionidae) Srednej Azii i Kazakhstana Illjustrirovannyj OpredeliteF Rodov i Katalog Vidov. Izdatel’stvo Nauka Kaza- khskoy SSR, Alma-Ata Bouchard P, Bousquet Y, Davies AE, Alonso-Zarazaga MA, Lawrence JF, Lyal CHC, Newton AF, Reid CAM, Schmitt M, Slipinski SA, Smith ABT (2011) Family-group names in Coleoptera (Insecta). ZooKeys 88: 1-972 Champion GC (1914) Reports on the Percy Sladen Trust expe- dition to the Indian Ocean in 1905, under the leadership of Mr. J. Stanley Gardiner, M.A. No. XIX. Coleoptera, Curoulionidae. Transactions of the Linnean Society London 16: 393-497 Cohen KM, Finney S, Gibbard PL (201 3) International Chronos- tratigraphic Chart, International Commission on Stratigraphy. Online at http://www.stratigraphy.org/ICSchart/Chronos- tratChart2013-01.pdf last accessed March 12 2014 Craw RC (1999) Molytini (Insecta: Coleoptera: Curculionidae: Molytinae). Fauna of New Zealand, 39. Manaaki Whenua Press, Lincoln Davidian GE (2008) Weevils of the genus Plinthus Genu. (Coleoptera, Curculionidae) from the Caucasus. Autoreferat of unpublished PhD Thesis, 19 pp. [in Russian] Drummond AJ, Suchard MA, Xie D, Rambaut A (2012) Bayesian phylogenetics with BEAUti and the BEAST 1.7. Molecular Biology and Evolution 29: 1969-1973 Faust J (1895) Russelkafer der alten und neuen Welt. Stettiner Entomologische Zeitung 55: 140-160 Favre A, Packert M, Pauls SU, Jahnig SC, Uhl D, Michalak I, Muellner-Riehl AN (2014) The role of the uplift of the Qing- hai-Tibetan Plateau for the evolution of Tibetan biotas. Bio- logical Reviews doi: 10.1 1 1 1/brv. 12107 Fitter AH, Peat HJ (2014) Ecological Flora of the British Isles. Online at http://www.ecotlora.co.uk last accessed 2 July 2, 2014 Fountain ED, Wiseman BH, Cruickshank RH, Paterson AM (2013) The ecology and conservation of Hadramphus tuber- culatus (Pascoe 1877) (Coleoptera: Curculionidae: Molytinae). Journal of Insect Conservation 17: 737-745 Grebennikov VV (2014) DNA barcode and phylogeography of six new high altitude wingless Niphadomimus (Coleoptera: Curculionidae: Molytinae) from Southwest China. Zootaxa 3838: 151-173 Grebennikov VV, Heiss E (2014) DNA barcoding of flat bugs (Insecta: Hemiptera: Aradidae) with phylogenetic implications. Arthropod Systematics & Phylogeny 72: 213-219 Grebennikov VV, Pham H.T. (2014) First record of Otibazo (Coleoptera: Curculionidae: Molytinae) outside of Japan, with description of a new species from Vietnam. Zootaxa 3869: 597-600. Hebert PDN, Cywinska A, Ball SL, deWaard, JR (2003) Biolo- gical identifications through DNA barcodes. Proceedings of the Royal Society B, 270: 313-321 Heller KM (1908) Viertel Beitrag zur Papuanischen Kaferfau- na. Abhandlungen und Berichte des Koiiigl. Zoologischen und Anthropologisch-Etnographischen Museums zu Dresden 12: 1-33+pl Hoffmann A (1965) Curculionides nouveaux du Tanganyika [Col.]. Annales de la Societe entomologique de France, Nou- velle serie 1: 393-409 Hustache A ( 1 920) Diagnoses abregees de Curculionides de Ma- dagascar. L’Echange 36(398): 1M Hustache A (1925) Synopsis des Cuculionides de Madagascar. Bulletin de FAcademie Malagache, Nouvelle Serie 7 (1924): 1-582 Hustache A (1929) Insectes Coleopteres XIX Curculionidae. Pp. 366-560, plates X, XI in: Voyage de Ch. Alluaud et R. Jean- nel en Afrque Orientale (1911-1912). Resultats scientifique. Librarie P. Lechevallier, Paris Kojima H (2010) Weevils of the genus Pinacopus (Coleoptera, Curculionidae) from Laos. Elytra 38: 151-160 Kuschel G (1987) The subfamily Molytinae (Coleoptera: Cur- culionidae): general notes and descriptions of new taxa from New Zealand and Chile. New Zealand Entomologist 9: 1 1-29 Kuschel G, Worthy TH (1996) Past distribution of large wee- vils (Coleoptera: Curculionidae) in the South Island, New Zealand, based on Holocene fossil remains. New Zealand En- tomologist 19: 15-22 Lyal CH (2014) 3.7.7 Molytinae Schoenherr, 1823. Pp. 529-570 in: Leschen RAB & Beutel RG (eds.) Handbook of Zoology, Arthropoda: Insecta: Coleoptera. Volume 3: Morphology and Systematics (Phytophaga). Walter de Gruyter, Berlin Maddison DR (2012) Phylogeny of Bembidion and related ground beetles (Coleoptera: Carabidae: Trechinae: Bembidi- Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK Morimotodes , a new weevil genus from China 147 ini: Bembidiina). Molecular Phylogenetics and Evolution 63: 533-576 Marshall GAK ( 1942). New Malaysian Hylobiinae (Col., Cur- cul.). Annals & Magazine of Natural History, Series 11 9: 370-390 McKenna DD, Sequeira AS, Marvaldi AE, Farrell BD (2009) Temporal lags and overlap in the diversification of weevils and flowering plants. Proceedings of the National Academy of Sci- ences USA 106: 7083-7088 Meregalli M (1985) Revisione del genere Plinthus Germar (Coleoptere: Curculionidae). Memorie del Museo Civico di Storia Naturale di Verona, IIA Serie, Sezione Scienze della Vi- ta, Supplement 5, 1-133 Meregalli M ( 1986) Iberoplintlnis, n. gen. di Plinthini e note sul genere Mitoplinthus Reitter, 1897 (Coleoptera: Curculionidae: Hylobiinae) (5 contributo alia conoscenza dei Curculionidi della Penisola Iberica). Bollettino del Museo Regionale di Scienze Naturali (Torino) 4: 489-522 Meregalli M (2003) Stenanchonus Voss 1937, a junior synonym of Euthycus Pascoe 1885 (Insecta, Coleoptera, Curculionidae, Molytinae). Senckenbergiana Biologica 82: 1-7 Morimoto K (1982). The family Curculionidae of Japan. I. Sub- family Hylobiinae. Essakia 19: 51-121 Mosquin T, Whiting PG, McAllister DE ( 1995) Canada’s bio- diversity: the variety of life, its status, economic benefits, con- servation costs and unmet needs. Canadian Museum of nature, Ottawa Osella G ( 1 970) Revisione del genere Caulomorphus Faust e de- scrizione di cinque nuove specie delle regioni montuose del- la Turchia lungo il Mar Nero (Coleoptera, Curculionidae). Memorie del Museo Civico di Storia Naturale di Verona 17(1969): 359-395 Osella G, Bello C (2010) Revisione di Minyops Schoenherr e Paraminyops nov. gen. (Coleoptera, Curculionidae, Molyti- nae). Memorie del Museo Civico di Storia Naturale di Verona, 2. serie, Sezione Scienze della Vita 19: 3-133 Pajni HR, Sood S (1982) First record of genera Niphadonyx Dal- la Torre and Ischnopus Fst. and description of two new species from India (Coleoptera: Curculionidae: Hylobiinae). Journal of the Bombay Natural History Society 79: 363-369 Papadopoulou A, Anastasiou I, Vogler AP (2010) Revisiting the insect mitochondrial molecular clock: the mid-Aegean trench calibration. Molecular Biology and Evolution 27: 1659-1672 Peck SB (1990) Eyeless Arthropods of the Galapagos Island, Ecuador: composition and origin of the cryptozoic fauna of a young, tropical oceanic archipelago. Biotropica 22: 366-38 1 Quedenfcldt G (1888) Verzeichniss der von Herrn Major a.D. von Mechow in Angola und am Quango-Strom 1878-1881 gesammelten Curculioniden und Brenthiden. Berliner Ento- mologische Zeitschrift 32: 271-308+pl Rambaut A (2013) FigTree, version 1.4. Online at http://tree.bio.ed.ac.uk/software/figtree/ last accessed March 12, 2014 Raxworthy CJ, Forstner MRJ, Nussbaum RA (2002) Chameleon radiation by oceanic dispersal. Nature 415: 784-787 Setliff, G.P. (2007) Annotated checklist of weevils from the Papuan region (Coleoptera, Curculionoidea). Zootaxa 1536: 1-296 Shorthouse DP (2010) SimpleMappr, an online tool to produce publication-quality point maps. Online at http://www.sim- plemappr.net last accessed May 12, 2014 Tamura K, Peterson D, Peterson N, Steelier G, Nei M, Kumar S (201 1 ) MEGA5: molecular evolutionary genetics analysis using Maximum Likelihood, Evolutionary Distance, and Max- imum Parsimony Methods. Molecular Biology and Evolution 28: 2731-2739 Wanat M (2007) Alignment and homology of male terminalia in Curculionoidea and other Coleoptera. Invertebrate System- atics 21: 147-171 Wang L. Liao W, Chen C, Fan Q (2013) The seed plant flora of the Mount Jinggangshan Region, Southeastern China. PLoS ONE 8(9): e75834^ Zherikhin VV, Egorov AB (1990) Zhuki-dolgonosiki (Coleoptera, Curculionoidea) dal’nego vostoka SSSR (obzor podsemejstv s opisaniem novykh taksonov). Akademija Nauk SSSR, Dal’nevostochnoe Otdelenie, Biologo-Pochven- nyj Institut, Vladivostok. Zumpt F (1932) Curculioniden - Studien IV. (Col.) Drei neue Kaferarten aus Kleinasien. Mitteilungen der Deutschen En- tomologischen Gesellschaft 3: 87-94 Bonn zoological Bulletin 63 (2): 123-147 ©ZFMK Bonn zoological Bulletin 63 (2): 148-156 December 20 1 4 New records of amphibians and reptiles from southern Vietnam Truong Quang Nguyen12, Trung My Phung3, Nicole Schneider4, Andreas Botov4, Dao Thi Anh Tran5,6 & Thomas Ziegler4* ' Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Hanoi, Vietnam; E-mail: nqt2@yahoo.com ' Department of Terrestrial Ecology, Zoological Institute, University of Cologne, Ziilpicher Strasse 47b, D-50674 Cologne, Germany 3 Dong Khoi 9 A, Tam Hiep, Bien Hoa, Dong Nai Province, Vietnam; E-mail: pmytnmg@yahoo.com 4 AG Zoologischer Garten Koln, Riehler Strasse 173, D-50735 Cologne, Germany 3 Vietnam National University, Ho Chi Minh City, University of Science, Faculty of Biology, 227 Nguyen Van Cu, District 5, HCM city, Vietnam ; E-mail: ttadao.hcmuns@gmail.com 6 Zoologisches Forschungsmuseum Alexander Koenig, Adenaueralle 160, D-53113 Bonn, Germany ' corresponding author: E-mail: ziegler@koelnerzoo.de Abstract. We report six new records of poorly known species of amphibians and reptiles on the basis of a new herpeto- logical collection from southern Vietnam: Ophryophryne gerti and Raorchestes gryllus from Khanh Hoa Province, Acan- thosaura coronata from Ba Ria - Vung Tau Province, Gekko grossmanni from Ninh Thuan Province, as well as Cylin- drophis ruffus and Oligodon cinereus pallidocinctus from Binh Phuoc Province. Keywords. Ophryophiyne gerti , Raorchestes giyllus, Acanthosaura coronata, Gekko grossmanni, Cylindrophis ruffus, Oligodon cinereus pallidocinctus, new records, taxonomy. INTRODUCTION Geographic distribution records are crucial for assessing the conservation status and are also evidence to elucidate species’ evolution and biology. Based on newly collect- ed specimens of amphibians and reptiles from southern Vietnam, we herein report six new records of poorly known species, comprising two anuran species from Khanh Hoa Province, one agamid species from Ba Ria - Vung Tau, one geckonid species from Ninh Thuan Province as well as one cylindrophiid and one colubrid species from Binh Phuoc Province. MATERIAL AND METHODS Field work was conducted by Trung My Phung in Hon Ba Nature Reserve (Khanh Hoa Province) in July 2010, on Nui Dinh Mountain (Ba Ria -Vung Tau Province) in No- vember 2008, in Ca Na forest (Ninh Thuan Province) in March 2010 and by Trung My Phung and Thang Huu Khuong in Bu Gia Map National Park in September and October 201 1 (Fig. 1). After taking photographs, speci- mens were anaesthetized, fixed in 80-85% ethanol and subsequently stored in 70% ethanol. Specimens were de- posited in the collections of the Institute of Ecology and Biological Resources (1EBR) and Vietnam National Mu- seum of Nature (VNMN), Hanoi, Vietnam and the Zool- ogisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn, Germany. Received: 07.02.2013 Accepted: 25.1 1.2014 Measurements of specimens were taken with a digital caliper to the nearest 0. 1 mm. Abbreviation are as follows: SVL (snout-vent length): from tip of snout to anterior mar- gin of cloacal; TaL (Tail length): from posterior margin ofcloacal to tip of tail; TL (total length): SVL + TaL. Ter- minology of morphological characters followed Nguyen et al. (2012) for amphibians (except for forearm length, FAL: from axilla to elbow and hand length, HAL: from base of outer palmar tubercle to tip of finger III), Phung & Ziegler (2011) for lizards, and David et al. (2012) for snakes. RESULTS AND DISCUSSION Taxonomic accounts Megophryidae Ophryophryne gerti Ollier, 2003 Gerti’s Mountain Toad / Coc nui got (Fig. 2) Specimens examined: VNMN 983 and ZFMK 94220, males, collected by T. M. Phung on 4 July 2010 from Hon Ba Nature Reserve (ca. I2°10’N, 109°02’E), Dien Khanh District, Khanh Hoa Province. Corresponding editor: T. Ziegler New herpetofaunal records from Vietnam 149 Fig. 1. Map of sampling sites in southern Vietnam: 1 ) Khanh Hoa Province, 2) Ninh Thuan Province, 3) Binh Phuoc Province, 4) Ba Ria - Vung Tau Province. Morphological characters: Size small, SVL 28.2-29.7 mm (n = 2); head wider than long (HW 8. 5-8. 9 mm, HL 8. 1-8.7 mm); snout truncate, protruding, shorter than hor- izontal diameter of eye (SL 2. 6-2. 7 mm, ED 3. 2-3. 7 mm); canthus rostralis rounded, loreal region convex; interor- bital distance narrower than upper eyelid and internarial distance (IOD 4. 0-4. 3 mm, UEW 2. 6-2. 7 mm, IND 2. 5-2. 7 mm); nostrils closer to the eye than to the tip of snout (EN 1.1-1. 4 mm, SN 1.2-1. 5 mm); pupil rounded; tympanum oval, greater than half eye diameter and tym- panum-eye distance (TD 1 .8-2.3 mm, TYE 1 .5-1.9 mm); supratympanic fold distinct; vomerine teeth absent; tongue rounded posteriorly. Forelimbs: FAL 6. 5-7.4 mm, HAL 7. 5-7. 6 mm; fingers free of webbing, relative length of fingers: I16 years. Most of the photos published of Chiropotes albinasus from Cologne Zoo (with its hybrid offspring) show this indi- vidual. ZFMK_MAM 89.480: adult female, f 01. IX. 1988, received from Cologne Zoo IX. 1989; skull, postcranial skeleton, flat skin. This is the female with the Zoo house name ‘Bianca’ which arrived there 08.X.1970 (Windecker 1971 ) and is the individual with the maximal longevity (> 18 years) attained by this species in any zoo ( Weigl 2005). During the time this female lived at the zoo it gave birth to two offspring. Its estimated age of about 20 years noted by Weigl (2005) would imply that it was roughly 2 years old when it arrived at the zoo. The first documented individual of Chiropotes albina- sus in the care of Cologne Zoo, an adult female, arrived in 1965 (Hick 1965a; Pagel & SpieB 2011). In 1968 the zoo received a juvenile female (Windecker 1969, anoth- er female which arrived in 1966 only survived for a few months) and in 1970 another two juvenile females (Windecker 1971). Apart from popular print products of the Cologne Zoo with photos (black-and-white and colour) of several of these individuals (e.g. Hick 1968a: 35, 1973; Windecker 1969; front pages of the journal ‘Freunde des Kolner Zoo’ 9[3], 1966 and 1 1 [2], 1968), photos of them also appeared in technical publications (Hick 1968b: Fig. 31; Roosmalen et al. 1981: 420; Hershkovitz 1985: 26). Two of the white-nosed saki females gave birth success- fully to hybrid young: a male out of a mating of Chiropotes albinasus female x Chiropotes chiropotes male in 1968 (Hick 1968a) and five more hybrids (2 males, 3 females) out of matings of Chiropotes albinasus female x Chi- ropotes satanas male in 1973 (2), 1975, 1976, and 1979; in 1978 one female had a miscarriage after such a mat- ing. Like the first hybrid bom in 1968 (Hick 1968a), some of the others born later were figured in photos, usually to- gether with their mother (Kullmann 1976). The first hy- brid bom in 1968 is mentioned and also figured in tech- nical publications (Roosmalen et al. 1981: 420; Her- shkovitz 1985: 26). Hershkovitz’s statement is not correct; the sire of this hybrid offspring was a Chiropotes chi- ropotes male (Becker, pers. comm. 2014). Four of the hy- brid offspring were bom by the female ‘Bella’, two by the female ‘Bianca’, and the same Chiropotes satanas male was the sire of 4 of the 5 hybrids bom since 1973. A fe- male hybrid bom in 1975 lived at Cologne Zoo for more than 26 years (Weigl 2005). Occasionally one of these ju- venile hybrids was wrongly labelled as Chiropotes albi- nasus, resp. no mention was made that the juvenile is a hybrid, e.g. in the publications of Kullmann (1976) and Mittermeier & Coimbra-Filho (1981: Fig. 53, p. 86) or in a figure legend (Zeitschrift des Kolner Zoo 1 7 ( 1 ), p. 1 ). The same is probably tme for a colour photo of an adult ‘female white-nosed saki (Chiropotes albinasus)' in Pauly (20 1 0: 1 82, Abb. 1 ). If the date when the photo was taken (24.V.1990 as stated in the figure legend) is correct, this individual is a hybrid of Chiropotes albinasus female x Chiropotes satanas male because according to the zoo archives the last female white-nosed saki at Cologne Zoo died in 1988. In 1990 only two female hybrids were liv- ing at the zoo (SpieB, pers. comm. 2011; Becker, per. comm. 2012). Unfortunately, none of the altogether six hy- brids bom at the zoo was given to the ZFMK, another zo- ological museum or a scientific institute after their death. Chiropotes chiropotes (Humboldt, 1811) — Red-backed bearded saki Material: ZFMK MAM 66.001 ; adult male, f 04.01.1966, received from Cologne Zoo 1966; skull, taxidermy mount (Fig. 2). This individual arrived at the zoo as a juvenile in 1961 and was originally registered as a female which was lat- er corrected. It is mentioned in Anonymous (1962) and Bonn zoological Bulletin 63 (2): 173-187 ©ZFMK 180 Gustav Peters et al. shown at different ages in two black-and-white photo- graphs, the earlier taken in May 1962, the later in Sep- tember 1963 (Anonymous 1963: 87). Comparing both photographs the pronounced growth of the typical thick split tuft of hair on the head and that of the eponymous beard during the time span of about one year becomes ob- vious. Photos of Chiropotes chiropotes individuals kept at Cologne Zoo and published from 1964 onwards by zoo employees (Hick 1965a, 1966) very likely show other in- dividuals of this species living there, as two more indi- viduals arrived at the zoo in 1964 and 1965. Description of the specimen (Fig. 2): As the specimen was on exhibit in the museum for a few decades, a gen- eral fading of its pelage colouration is likely to have oc- curred, especially in respect of possible yellowish/reddish hues that may have been present in the living animal. Fol- lowing statements are made under this qualification. There is a marked contrast between the homogeneous light buffy ochre of the back and the dark brown to black coloura- tion of the fur of the head, extremities and the tail. The colouration of the back extends into the flanks and the arms and thighs. On the upper arms pelage colour grad- ually changes into blackish brown of the more distal por- tion of the anns, on the upper side interspersed with lighter hairs. This colour change is more abrupt on the proximal part of the thighs. The backs of hands and feet are cov- ered by brown hair. The venter is black. The fur colour on the head (with the beard and the split tuft of hair on top of it), nape and tail is basically black. The change in fur colour from the trunk to these body regions is abrupt. Taking into account the unsettled taxonomic issues ad- dressed earlier, the actual species identity of all relevant specimens that lived at Cologne Zoo cannot be verified resp. refuted retroactively, even more for those for which no photos are available and/or no material was given to the ZFMK or to another scientific institution after the an- imals’ death. In addition, it is difficult to determine the species identity of juvenile bearded saki individuals ret- rospectively on the basis of photographic portraits with adequate certainty. Therefore we use the scientific names here under which they are registered in the zoo archives. According to these (SpieB, pers. comm. 2011; Becker, pers. comm. 2011) Cologne Zoo received four specimens of Chiropotes chiropotes between 1961 and 1977, three males and one individual of unknown sex which died a few months after its arrival whereas the other individuals all lived at the zoo for several years. One of these males (house name ‘Ringo’), which arrived at the zoo as a ju- venile in 1964, was the father of the first hybrid saki (mother Chiropotes albinasus ) bom at Cologne Zoo in 1968 (Hick 1968a). This male, given to a private animal husbandry in 1 982, was figured in Hick ( 1 968a: 38 & 39), van Roosmalen et al. (1981: 421; fig. 2); photos in Napi- er and Napier (1967: 121, plate 33) and Hershkovitz (1985: 14, fig. 8) probably also show it. Unfortunately, no remains of this individual are conserved in a scientific in- stitution. Chiropotes sp. - Bearded saki Material: ZFMK_MAM_2008.244; adult male, f 1991, received from Cologne Zoo 5. IX. 1991; skull, postcranial skeleton, flat skin (Fig. 3). Animal received by Cologne Zoo as a present from the defunct Dierenpark (zoo) Wasse- naar, Netherlands, in 1978 and originally listed as Chi- ropotes satcmas chiropotes in the zoo archives (Kullmann 1979). A colour photo of this individual alive, taken 24. V. 1990 and referred to as Chiropotes chiropotes , was published by Pauly (20 1 0: 1 83, Abb. 2). However, the dor- i 0 02 H SSU53582 Saimiri sdureus AF289515 Alouatta belzebul FJ785423 Callicebus donacophilus ■ AF289988 Callicebus personalus AY226183 Pithecia irrorata AY226184 Cacajao melanocephalus Fig. 4. Cytochrome b phylogeny of the genus Chiropotes. A, Posterior probabilities of the Bayesian approach and bootstrap sup- port values of the maximum-likelihood analysis before and after the slash, respectively. B, Phylogenetic relationships and boot- strap support values based on the neighbor-joining approach. Sample IDs correspond to GenBank accession numbers. Bonn zoological Bulletin 63 (2): 173-187 ©ZFMK Neotropical primates from the Cologne Zoo in the collections of the Zoologisches Forschungsmuseum A. Koenig 181 sum of this specimen is light buffy to ochre contrasting the dark brown to blackish colour of the outer side of the upper and lower arms, legs, the fur of the head, and the outer side of the thighs. According to Hershkovitz ( 1985:13) characters of chiropotes and utahickae are de- scribed as ‘Dorsum dominantly orange or pale brown to dark brown; outer side of upper arms and proximal part of outer side of thighs orange to dark brown, not black- ish’ and as ‘Head, nape, lower arms, and legs blackish, sharply contrasted with orange of dorsum' in chiropotes vs. ‘Head, nape, lower arms, and legs pale brown to dark brown and not sharply contrasted with brown of dorsum’ in utahickae. Based on these external characters and the fact that no geographical origin is known, the species des- ignation of the ZFMK specimen remains questionable. The results of our genetic study of this specimen is shown in Fig. 4. Except of relationships among Cacajao , Pithecia and Chiropotes , which cannot be correctly resolved based on the short cyt b fragment using the ML and Bayesian approaches, we obtained the same pitheciine phylogeny as Bonvicino et al. (2003). Chiropotes albinasus forms a basal taxon to all remaining Chiropotes members and C. utahickae forms a sister clade to C. israelita. However, irrespective of the phylogenetic approach and of uncer- tainties among basal pitheciine relationships, our sample from the ZFMK clusters together with the C. chiropotes! sagulatus sample from east of the Rio Branco causing paraphyletic relationships between C. israelita (sensu Bonvicino et al. 2003; also Dare et al. 2011) and C. chiropotes! sagulatus west and east of the Rio Branco, respectively (Fig. 4). Chiropotes satanas (Hoffmannsegg, 1807) - Black bearded saki Material: ZFMK_MAM_60. 1 08; juvenile male, f 1 1. IV. 1960, received from Cologne Zoo 20.VII.1960; taxi- dermy mount. The animal died at the zoo shortly after its arrival. A photo each of a juvenile individual of Chiropotes satanas from Cologne Zoo published in Hill ( 1960) and in Anonymous (1963) may be this animal but it may al- so show one of the other two juveniles of this species which the zoo received in 1954 resp. 1959, each of each survived for a short time only. Description of the specimen: With the exception of the back which is covered with relatively long chestnut brown hair, the hair-coat of all other body parts is black. ZFMK_MAM_81.1819; adult male, | 04.VII.1981, re- ceived from Cologne Zoo VIII. 1 98 1 ; skull, skeleton, taxi- dermy mount. When this individual (house name ‘Nicky’) arrived at the zoo as a young juvenile on 26.VI.1970 it was originally registered as a female but this was correct- ed in the archives in 1971. Its black-and-white portrait tak- en soon after arrival was published (Windecker 1971 : 4). No other photo published later of a bearded saki from Cologne Zoo or one kept in the zoo archives can be re- lated to this individual with certainty; therefore its char- acters as an adult are not documented. It is the father of all hybrids of Chiropotes satanas and Chiropotes albina- sus females bom at Cologne Zoo between 1973 and 1978 (see above). Description of specimen ZFMK MAM 8I.1819 (Fig. 5): As the specimen was on exhibit in the Museum for nearly two decades, the same general qualifications as for the Chiropotes chiropotes specimen ZFMK_MAM_ 66.001 apply in respect of an assessment of the coat colour of this individual. Moreover, it lost hair in the tanning process, especially on the back. This is likely to result in a changed perception of the colour of the back coat as compared to that of a living animal. Generally, there is no sharp contrast in coat colouration between different parts of the body. The colour of the head (with beard and split tuft of hair on top of it), nape, tail, extremities and backs of hands and feet is black as is that of the venter. On the back, shoulders, upper arms and thighs a lighter dull brown shines through the blackish hue, dominating the general colour impression of the back. Whereas the last in-depth revision of the genus Chi- ropotes by Hershkovitz (1985) listed two subspecies of Chiropotes satanas in addition to the nominate form, the current notion is that Chiropotes satanas represents a dis- tinct species with no subspecies (Silva-Jr. & Figueiredo 2002; Bonvicino et al. 2003; Groves 2005; Figueiredo et al. 2006; Rylands & Mittermeier 2009; Ferrari et al. 2013; Fig. 5. Back view of mounted adult male Chiropotes satanas specimen ZFMK MAM 81.1819. The hair on the back is thin- ner than in the living animal (Photo: M. Weigt). Bonn zoological Bulletin 63 (2): 173-187 ©ZFMK 182 Gustav Peters et al. Silva-Jr. et al. 2013). As detailed above, the two remain- ing former subspecies and an additional one are accord- ed species rank each as well. We adopt this concept here. As in the case of Chiropotes chiropotes we have to rely on the species determinations in the Cologne Zoo archives for those individuals for which no detailed doc- umentation and/or collection material is available, appre- ciating the fact that these may have been wrong, especial- ly in young juveniles which are difficult to identify as to species in Chiropotes satanas sensu lato. Between 1954 and 1970 Cologne Zoo received 4 individuals of Chi- ropotes satanas , all as young juveniles: two males, one female and one individual the sex of which was not es- tablished with certainty and which died soon after its ar- rival. Only the male individual received by the zoo in 1970 lived to adulthood. Concluding taxonoinical remarks on Chiropotes specimens in the ZFMk collection Chiropotes chiropotes ZFMk MAM 66.0111: We con- sider this species identification to be well-founded. In con- sideration of the qualifications of such an appraisal the available evidence still documents an extensive similari- ty of this ZFMK specimen with the type specimen of Brachyurus israelita Spix, 1823 and the specimen referred to as Chiropotes cf. satanas in a colour photograph in Bou- bli (2002), both of which are synonyms of Chiropotes chi- ropotes. Chiropotes sp. ZFMKMAM 2008.244: Our cyt b phy- logeny shows a close genetic relationship of this speci- men to an individual from Rio Trombetas east of the Rio Branco, i.e. within the distribution of C. sagulatus accord- ing to Ferrari et al. (2013) and Silva-Jr. et al. (2013). How- ever, their description of sagulatus is ‘Dorsum and upper limbs ... are orange to reddish-brown. Head, nape, low- er arms and legs are blackish’ (Ferrari et al. 2013: 481), which does not fit to the general appearance of the ZFMK specimen. Despite the small amount of samples analysed, the unknown origin of the ZFMK specimen and of the low support values in reconstructed phylogenies, our data al- low us to draw some preliminary conclusions. Our results support the existence of different colour variants within groups of closely related individuals and possibly within the same geographic region, here within the Chiropotes form occurring east of Rio Branco. This finding corre- sponds to the statement by Hershkovitz (1985) that no clear geographic separation between different pelage vari- ants exists. Furthermore, our molecular results indicate that most probably both israelita and sagulatus do not de- serve species status and may represent synonyms of Chi- ropotes chiropotes. Therefore, we highly recommend fur- ther genetic studies on diverse Chiropotes samples from a variety of geographical areas, including Chiropotes sa- tanas to clarify the phylogeny of the genus, and most im- portantly, to allow final taxonomic conclusions. Chiropotes satanas ZFMk_MAM_60.108: There is very little reliable information on coat colour of juvenile indi- viduals of Chiropotes satanas sensu lato. To the best of our knowledge no such data have been published for ju- venile Chiropotes satanas sensu stricto; based on the rel- evant details provided for Chiropotes chiropotes (Her- shkovitz 1985: 17) it seems possible that this ZFMK in- dividual actually represents the latter species. Chiropotes satanas ZFMk_MAM_81.1819: In spite of the reservations with respect to the assessment of the fur colouration of the living animal, in light of the condition of this taxidermy mount we consider this species identi- fication to be well-founded. Pithecia Desmarest, 1804 The first revision of the genus Pithecia by Hershkovitz (1979) recognized four species, a concept also followed by Buchanan et al. ( 1981 ). The current taxonomy of this genus as adopted by Groves (2001, 2005), Rylands & Mit- tenneier (2009) and Ferrari et al. (2013) is based on the earlier and more detailed revision by Hershkovitz ( 1987b) in which he recognized five species. However, there is al- so evidence that a further revision of this genus is required (Marsh 2004, 2006, pers. comm. 2012). In her final revi- sion (Marsh 2014) 16 species are recognized. Pithecia mittermeieri Marsh, 2014 - Mittermeier’s Tapajos saki Hershkovitz ( 1987b: 427, Fig. 25) figured an adult male Pithecia irrorata irrorata (house name Sascha) from Cologne Zoo. Since its arrival at Cologne Zoo in 1 966 un- til its death on 20.1.1981 it was registered as Pithecia monachus in the Zoo archives. Hick ( 1 968b) published its black-and-white portrait and later on another photograph/portrait in colour (Hick 1973) under this name. The best photographic documentations of this individual referred to as Pithecia monachus are two black-and-white photographs and a colour photo in Tylinek and Berger (1984) and colour photos in several official guides of Cologne Zoo in the 1970es (Fig. 6). According to Weigl (2005), who listed this animal under the scientific name Pithecia irrorata irrorata , it reached the highest longevi- ty known of any individual of this species in human care (more than 14 years and 7 months). In the recent revision of the genus this individual is identified as Pithecia mit- termeieri sp. nov. (Marsh 2014: 5, Table 2). Regrettably no zoological material of it is preserved in the ZFMK, an- other zoological museum or a scientific institution. Bonn zoological Bulletin 63 (2): 173-187 ©ZFMK Neotropical primates from the Cologne Zoo in the collections of the Zoologisches Forschungsmuseum A. Koenig 183 Fig. 6-7. 6. (Left) Adult male Pithecia mittermeieri Marsh, 2014 in the Zoo Cologne; this animal may be the one shown in fig- ure 7 (Photo: P. De Prins, reproduced from 'Wegweiser durch den Zoo Koln’, c. 1976). 7. (Right) Adult male specimen of Pithe- cia aequatorialis (lying individual on the left) (ZFMK_MAM_81.1818) in the Zoo Cologne ( 1973 or later); the individual sitting on the right may represent Pithecia mittermeieri , possibly the same animal as shown in figure 6 (Photo: G. Peters). Pithecia aequatorialis Hershkovitz, 1987 - Equatorial saki Material: ZFMK_MAM_8 1.1 818; adult male, f 1981 (?), received from Cologne Zoo VI 11.1981; skull, postcranial skeleton, flat skin (Fig. 7, the living animal). This specimen has the diagnostic pelage characteristics of Pithecia aequatorialis. It was received from Cologne Zoo as Pithecia monachus and classified as such in the ZFMK collection then. After the publication of the second revi- sion of the genus Pithecia by Flershkovitz (1987b) the species determination of this specimen was reassessed and subsequently classified as Pithecia aequatorialis. This identification was confirmed by L. Marsh (pers. comm. 2011) on the basis of several colour photos of the ZFMK skin. Unfortunately, there is no photograph of the living animal or written record in the Cologne Zoo archives which can be unambiguously assigned to this individual (SpieB & Becker, pers. comm. 2011). However, a photo- graph by R. Mittermeier (in Marsh 2014: fig. 49) may show this individual. Several individuals classified as Pithecia monachus lived at Cologne Zoo during the second half of the 1970’s. Apart from the male individual of Pithecia mittermeieri mentioned above (and then wrongly classified as P. monachus at the zoo) none of the black-and-white pho- tographs of ‘ Pithecia monachus' individuals archived at the zoo shows an animal with the typical pelage charac- teristics of Pithecia aequatorialis males. In the five year period from 1977 until 1981 four individuals classified as P. monachus died at Cologne Zoo, one of which was the misidentified P. mittermeieri (Becker, pers. comm. 2012). The only photograph on file of a "Pithecia monachus' in- dividual at the zoo which died between 1977 and 1981 (other than the P.mittermeieri male mentioned) is one tak- en in 1969 of a "monachus' that died in 1979. So, there is definitely no ‘trace’ of the ZFMK Pithecia aequatori- alis adult male in the Cologne Zoo archives. A colour slide (Fig. 7) taken by one of us (G.R) shows this individual alive (or another male of this species if there were more than one in the zoo at that time) and therefore is addition- al proof of its species identification (confirmed by L. Marsh, pers. comm. 2013, based on this slide). The pho- to can be roughly dated to the time period 1973 to 1977 because it was taken in the Lemur House of Cologne Zoo which opened in 1973, and on written entries in the ani- mal records at the zoo. This species was described as new to science after all Pithecia individuals (other than P. pithecia ) at Cologne Zoo had been dead already for several years. In his ear- lier revision of the genus Hershkovitz (1979: 15, fig. 5) had figured a study skin of an adult male Pithecia as P. monachus , only to revise his taxonomic assessment of the same individual in the later revision and identify it as the new species Pithecia aequatorialis (Hershkovitz 1987b: 408, fig. 16). Diagnostic characteristics differentiating males of aequatorialis from male monachus are a ruff of orange hair and a horseshoe-like area of relatively short white dense hair around the sparsely haired facial disk in the former species (L. Marsh, pers. comm. 2011, Marsh 2014). Bonn zoological Bulletin 63 (2): 173-187 ©ZFMK 184 Gustav Peters et al. CONCLUDING REMARKS Our paper is based on only a few Neotropical primate specimens from Cologne Zoo in the ZFMK mammal col- lection. Nevertheless, it provides an example of the val- ue and of the conclusions that can be drawn from collec- tions obtained from zoos and housed at zoological muse- ums (or similar scientific institutions). The ZFMK re- ceived also primate specimens from other zoos, such as Duisburg and Frankfurt (see Appendix), which may be equally important. In the first place it shows that even to- day zoos may have scientifically important specimens in their care, sometimes unnoticed by the scientific commu- nity. To avoid such cases as far as possible it is desirable that zoos and zoological museums intensify their cooper- ation (cf. Gippoliti & Kitchener 2007). It is important that zoos are aware of the potential scientific importance of specimens in their care and try to gather and archive all pertinent information during the lifetime of such speci- mens: geographical origin (if known or if it can be traced back), colour photos at different ages, body mass, meas- urements, reproductive history, ISIS reports, CITES pa- pers (if applicable), references of publications dealing with the specimen (if the specimen is mentioned/figured in a publication), or other. This is, however, more than usual- ly recorded. It may be useful to agree on a uniform data- base, so that information from different zoos can be more easily combined. Taking into account relevant veterinary and other legal regulations the most reasonable procedure would be that zoos pass such specimens immediately after their demise to a zoological museum. The condition of the specimen should thereby allow all standard museum procedures, e.g., collection of tissue and hair samples, preparation of skin and skeleton or any other method of conservation. Ap- propriate storage of such samples should allow subsequent genetic analyses. Furthermore, copies of all documenta- tion of that specimen compiled at the zoo during the an- imal’s lifetime should be provided, including veterinary treatments. A scientifically important specimen in/from a zoo can only maintain its ‘voucher status’ and its signif- icance posthumously as a subject for later studies, if its remains are housed and curated properly in a zoological museum or another appropriate scientific institution in the long run. This ought to be in the genuine interest of the respective zoo and definitely is so for the scientific com- munity, especially if specimens have been referred to in the scientific literature like several Chiropotes individu- als from Cologne Zoo in the ZFMK mammal collection, which are listed here. The Pithecia mittermeieri individ- ual discussed above exemplifies the deplorable ultimate loss of information and research options if this procedure is not followed after an animal’s death. On the other hand, the specimen of Pithecia aequatorialis in the ZFMK col- lection detailed before illustrates that steady contact and information exchange between zoos and zoological mu- seums may have resulted in the timely awareness of the significance of this individual, especially in the context of a taxonomical revision of the genus Pithecia. Zoological museums with their taxonomic expertise ought to interact with (neighbouring) zoos regularly, in- cluding the offer to provide advice on matters of taxono- my if requested. Once a specimen that died in a zoo is giv- en to a museum, the museum has the responsibility of its proper long-term documentation and curation, preserving as much and diverse material of the animal as possible to enable diverse kinds of (future) studies. This includes the repeated verification of a specimen’s species identity on the basis of the most recent pertinent publications to en- sure its scientific significance. If a specimen is affected by a taxonomical change, either during its lifetime or posthumously, this information ought to be circulated among the institutions involved. Both zoos and zoological museums can only benefit from a closer cooperation as a means to increase taxonom- ic expertise and awareness of taxonomical problems as well as the proper long-term conservation of scientifical- ly interesting and important zoological specimens. Acknowledgements. We are grateful to the many colleagues who generously helped in gathering all the diverse information necessary to carry out this study. Our sincere thanks to Wilhelm SpieB and Ralf Becker (both Cologne Zoo archives) and Uta Ruempler (nee Hick) (formerly Cologne Zoo) for providing in- formation on the zoo’s relevant primate individuals and photos, Andrew Kitchener (National Museums Scotland, Edinburgh) and Ms. E.M. Lajtos (Library, Naturalis, Leiden) for help with lit- erature, Laura Marsh (Global Conservation Institute) for expert- ise in Pithecia identification, Richard Kraft and Michael Hier- meier (ZSM) for information on and photos of the type speci- men of Brachyurus israelita. Laura Parker and Wendy Lehkyi (both Colchester Zoo), Tremaine Gregory (National Zoo, Wash- ington, D.C.), Marilyn Norconk (Kent State University, Ohio), and Liza Veiga (deceased) kindly provided photos of living Chi- ropotes individuals and information. We are indebted to Melanie Weigt (ZFMK) for photography and to Jorg Adrian (ZFMK) for technical assistance, to Achim Winkler (Duisburg Zoo) for information on Pithecia individuals kept at that zoo, and to Carlos Ruiz Miranda (Universidade Estadual Norte Flu- minense) for help with contacts in Brazil. We also thank Michael Hofreiter for giving us the opportunity to conduct the genetic analysis of the Chiropotes sample in his ancient DNA laborato- ry at York University. We are also grateful for helpful comments and suggestions of the two reviewers. REFERENCES Anonymous (1962) Freunde des Kolner Zoo 5:112 Anonymous (1963) Rotruckensaki im Kolner Zoo. Freunde des Kolner Zoo 6: 87-88 Barnett AA (2005) Cacajcio melanocephalus. Mammalian Species 776: 1-6 Barnett AA & Brandon-Jones D (1997) The ecology, biogeog- raphy and conservation of uacaris, Cacajao (Pitheciinae). Fo- lia primatologica 68: 223-235 Bonn zoological Bulletin 63 (2): 173-187 ©ZFMK Neotropical primates from the Cologne Zoo in the collections of the Zoologisches Forschungsmuseum A. Koenig 185 Bonvicino CR, Lemos B & Seuanez HN (2001 ) Molecular phy- logenetics of howler monkeys (Alouatta, Platyrrhini). Chro- mosoma 110: 241-246 Bonvicino CR, Boubli .IP, Otazii IB, Almeida FC, Nascimento FF, Coura JR & Seuanez HN (2003) Morphologic, karyotyp- ic, and molecular evidence of a new form of Chiropotes (Pri- mates, Pitheciinae). American Journal of Primatology 61: 123-133 Boubli JP (2002) Western extension of the geographic distribu- tion of bearded sakis: a possible new taxon of Chiropotes sym- patric with Cacajao in Pico da Neblina National Park, Brazil. Neotropical Primates 10: 1-4 Boubli JP, da Silva-Jr MNF, Amado MV, Hrbek T, Pontual FB & Farias IP (2008) A taxonomic reassessment of Cacajao melanocephalus Humboldt (1811). with the description of two new species. International Journal of Primatology 29: 723-741 Boubli JP & de Lima MG (2009) Modeling the geographical dis- tribution and fundamental niches of Cacajao spp. and Chi- ropotes israelita in Northwestern Amazonia via maximum en- tropy algorithm. International Journal of Primatology 30: 217-228 Buchanan DB, Mittermeier RA & Roosmalen MGM van (1981) The saki monkeys, genus Pithecia. Pp. 391-441 in: Coimbra- Filho AF & Mittermeier RA (eds) Ecology and Behavior of Neotropical Primates Vol. 1 . Academia Brasileira de Ciencias: Rio de Janeiro Cabrera A ( 1961 ) Catalogo de los Mamiferos de America del Sur. Revisa del Museo Argentino de Ciencias Naturales ‘Bernar- dino Rivadavia’, Ciencias Zoologicas 4(2): v-xxii + 309-732 Chiarelli AB ( 1972) Taxonomic Atlas of Living Primates. Aca- demic Press: London Dare M, Hait SH, Soares EA, Cicala C, Seuanez HN, Macha- do ES, Arthos JA & Soares MA(201 1 ) Polymorphisms in the alpha4 Integrin of Neotropical Primates: Insights for binding of natural ligands and HIV-1 gpl20 to the Human alpha4be- ta7. PLoS ONE 6 (9), E24461 Ferrari SF, Guedes PG, Figueiredo WMB & Barnett AA (2010) Re-evaluation of the nomenclature of the blacked-faced uacaris (Cacajao melanocephalus group, sensu Hershkovitz, 1987). 23ri^ Congress of the International Primatological So- ciety, Kyoto, Japan: Abstract #350 Ferrari SF, Veiga LM, Pinto LP, Marsh LK, Mittermeier LK & Rylands AB (2013) Family Pitheciidae (Titis, Sakis and Uacaris). Pp. 432-483 in: Mittermeier RA, Rylands AB & Wil- son DE (eds) Handbook of the Mammals of the World, Vol. 3. Primates. Lynx Edicions: Barcelona Figueiredo WB, Silva-Jr, JM, Bates JM, Harada ML & Silva- Jr, JS (2006) Conservation genetics and biogeography of pitheciines. International Journal of Primatology 27 (Suppl 1): Abstract #5 1 0 Figueiredo-Ready WB, Schneider H, Ferrari SF, Harada ML, DaSilva JMC, Silva-Jr JS & Bates JM (2013) A molecular phylogeography of the uacaris (Cacajao). Pp. 23-30 in: Veiga LM, Barnett AA, Ferrari S & Norconk MA (eds) Evolution- ary Biology and Conservation of Titis, Sakis and Uacaris. Cambridge University Press, Cambridge Finstermeier K, Zinner D, Brameier M, Meyer M, Kreuz E et al (2013) A mitogenomic phylogeny of living Primates. PLoS ONE 8(7): e69504. doi:10.1371/joumal. pone. 0069504 Geoffrey I & Deville E (1848) Note sur huit especes nouvelles de Singes americains, faisant partie des collections de MM. de Castelnau et Emile Deville. Comptes Rendus Hebdoma- daires des Seances de FAcademie des Sciences 27: 497-499 Gippoliti S & Bruner E (2007) The role of historical research in the study of primatological collections: case-studies from Bonn zoological Bulletin 63 (2): 173-187 the Museo di Antropologica “G. Sergi”, Rome. Journal of An- thropological Sciences 85: 157-162 Gippoliti S & Kitchener AC (2007) The Italian zoological gar- dens and their role in mammal systematic studies. Conserva- tion biology and museum collections. Hystrix - Italian Jour- nal of Mammalogy (ns) 18: 173-184 Gregory T (2011) Socioecology of the Guianan bearded saki, Chiropotes sagulatus. PhD thesis, Kent State University, Col- lege of Arts and Sciences / School of Biomedical Sciences, downloaded at http://etd.ohiolink.edu/view.cgi7acc_num =kent 1300284081 Gregory T & Norconk M (2011) Behavioral responses to sea- sonal changes in Guianan bearded sakis (Chiropotes saga lo- tus): Brownsberg Nature Park, Suriname. American Journal of Physical Anthropology 144 Suppl. 52: 149 Groves C (2001) Primate Taxonomy. Smithsonian Institution Press; Washington, D.C. Groves C (2005) Order Primates. Pp. 111-184 in: Wilson DE & Reeder DM (eds) Mammal Species of the World - A Tax- onomic and Geographic Reference. 31C* edition. The Johns Hopkins University Press: Baltimore Guindon S & Gascuel O (2003) A simple, fast, and accurate al- gorithm to estimate large phylogenies by maximum likelihood. Systematic Biology 52: 696-704 Haus T, Akom E, Agwanda B, Hofreiter M, Roos C, & Zinner D (2013) Mitochondrial diversity and distribution of African green monkeys (Chlorocebus Gray, 1870). American Journal of Primatology 75: 350-360 doi: 10.1 002/ajp.22 113 Hershkovitz P (1979) The species of sakis, genus Pithecia, with notes on sexual dichromatism. Folia primatologia 31: 1-22 Hershkovitz P (1985) A preliminary taxonomic review of the South American bearded saki monkeys genus Chiropotes (Ce- bidae, Platyrrhini), with the description of a new subspecies. Fieldiana: Zoology (New Series) 27: 1-46 Hershkovitz P (1987a) Uacaris, New World monkeys of the genus Cacajao (Cebidae, Platyrrhini): A preliminary taxonom- ic review with the description of a new subspecies. American Journal of Primatology 12: 1-53 Hershkovitz P (1987b) The taxonomy of South American sakis, genus Pithecia (Cebidae, Platyrrhini): A preliminary report and critical review with the description of a new species and a new subspecies. American Journal of Primatology 12: 387-468 Hick U (1965a) Einige bemerkenswerte Neuanschaffungen im zweiten Halbjahr 1965. Freunde des Kolner Zoo 8: 141-143 Hick U (1965b) Red-backed saki Chiropotes chiropotes at Cologne Zoo. International Zoo Yearbook 5: 140 Hick U (1966) Aus der Kolner Saki-Sammlung. Freunde des Kolner Zoo 9: 75-82 Hick U ( 1 968a) Erstmalig gelungene Zucht eines Bartsakis ( Va- ter: Rotriickensaki, Chiropotes chiropotes (Humboldt, 1811), Mutter: WeiBnasensaki, Chiropotes albinasus (Geoffrey et De- ville, 1848) im Kolner Zoo. Freunde des Kolner Zoo 1 1 : 35 — 4 1 Hick, U. (1968b): The collection of saki monkeys at Cologne Zoo. International Zoo Yearbook 8: 192-194 Hick U ( 1 973) Wir sind umgezogen. Zeitschrift des Kolner Zoo 16: 127-145 Hick U ( 1 974) Das erste Jahr im neuen Lemurenhaus. Zeitschrift des Kolner Zoo 1 7: 123-135 Hill WCO (1960) Primates: Comparative Anatomy and Taxon- omy IV Cebidae Part A. Edinburgh University Press: Edin- burgh Huelsenbeck .1, Ronquist F, Nielsen R & Bollback .1 (2001) Bayesian inference of phylogeny and its impact on evolution- ary biology. Science 294: 2310-2314 ©ZFMK 186 Gustav Peters et al. Humboldt A von (1811) Sur les singes qui habitent les rives de FOrenoque, du Cassiquiare et du Rio Negro, pp. 305-335 in: Humboldt A von and Bonpland A (eds.) Receuil d’observa- tions de zoologie et d’anatomie comparee, faites dans l’Ocean Atlantique, dans Finterieur du nouveau continent et dans la Merdu Sud pendant les annees 1799, 1800, 1801, 1802 et 1803. Premier Volume. Schoell & Dufour: Paris Husson AM (1957) Notes on the primates of Suriname. Studies on the Fauna of Suriname and other Guayanas 2: 13-40 Husson AM (1978) The Mammals of Suriname (Zoologische Monographieen van het Rijksmuseum van Natuurlijke Histo- rie no. 2. EJ Brill: Leiden Hutterer R & Peters G (2010) Type specimens of mammals (Mammalia) in the collections of the Zoologisches Forschungsmuseum Alexander Koenig, Bonn. Bonn zoolog- ical Bulletin 59: 3-27 Hutterer R, Verhaagh M, Diller J, & Podloucky R ( 1995) An in- ventory of mammals observed at Panguana Biological Station, Amazonian Peru. Ecotropica 1: 3-20 1CZN (1999) International Code of Zoological Nomenclature, fourth edition. London, The International Trust for Zoologi- cal Nomenclature. Jahn 1 (1994) Zoologische Garten - Zoologische Museen. Pa- rallelen ihrer Entstehung. Bongo 24: 7-30 Kraft R (1983) Die von .1. B. v. Spix beschriebenen neotropi- schen Primaten und Chiropteren. Verzeichnis der in der Zoo- logischen Staatssammlung Miinchen aufbewahrten Typus- exemplare. Spixiana Suppl 9: 429 — 44 1 Kullmann. E. ( 1976): Jahresbericht 1975 der Aktiengesellschaft Zoologischer Garten Koln. - Zeitschrift des Kolner Zoo 19: 31-46 Kullmann E (1979) Jahresbericht 1978 der Aktiengesellschaft Zoologischer Garten Koln. Zeitschrift des Kolner Zoo 22: 75- 91 Landsberg H (1994) Das erste Zootier des Museums - ein Man- drill? Die Griindung des Zoologischen Gartens und dessen Be- deutung fur die Sammlungen des Zoologischen Museums der Berliner Universitiit. Bongo 24: 85-106 Marsh LK (2004) Primate species at the Tiputini Biodiversity Station, Ecuador. Neotropical Primates 12: 75-78 Marsh LK (2006) Identification and conservation of a new species of Pithecia in Amazonian Ecuador. International Jour- nal of Primatology 27 (suppl 1): Abstract #508 Marsh L (2014) A taxonomic revision of the Saki monkeys, Pithecia Desmarest, 1804. Neotropical Primates 21(1): 1-163 Mittermeier RA & Coimbra-Filho AF (1981) Systematics: species and subspecies. Pp. 29-109 in: Coimbra-Filho AF & Mittermeier RA (eds). Ecology and behavior of Neotropical primates. Volume 1 . Academia Brasileira de Ciencias: Rio de Janeiro Napier JR & Napier PH (1967) A Handbook of Living Prima- tes. Academic Press: London Napier PH (1976) Catalogue of Primates in the British Muse- um (Natural History). Part I: Families Callitrichidae and Ce- bidae. British Museum (Natural History): London Norconk MA (2007) Sakis, uacaris, and titi monkeys: Behav- ioral diversity in a radiation of primate seed predators. Pp. 123-138 in: Campbell CJ, FuentesA, MacKinnon KC, Panger M & Bearder SK (eds) Primates in perspective. Oxford Uni- versity Press: New York Oliveira LC, Loretto D, Viana LR, Silva-Jr JS & Fernandes W (2009) Primate community of the tropical rain forests of Sara- ca-Taquera National Forest, Para, Brazil. Brazilian Journal of Biology 69: 1091-1099 Oppennann J (1994) Tod und Wiedergeburt - Uber das Schick- sal einiger Berliner Zootiere. Bongo 24: 51-84 Pagel T & SpieB W (2011) Der Zoologische Garten in Coin er- offnet am 22. Juli 1860 - 150 Jahre Wildtierhaltung und - zucht. Der Zoologische Garten NF 80: 1 17-202 Pauly A (2010) Der Schwarzkopfuakari im Zoo Koln - die Neu- bestimmung einer Art. Milu 13: 181-186 Posada D (2008) jModelTest: phylogenetic model averaging. Molecular Biology and Evolution 25: 1253-1256 Rambaut A (2008) FigTree: Tree figure drawing tool, version 1.2.2. Institute of Evolutionary Biology, University of Edin- burgh http://tree.bio.ed.ac.uk/ software/figtree/ Rohland N, Siedel H, & Hofreiter M (2004) Nondestructive DNA extraction method for mitochondrial DNA analyses of museum specimens. BioTechniques 36(5): 814-821 Rohland N, & Hofreiter M (2007) Ancient DNA extraction from bones and teeth. Nature Protocols 2(7): 1756-1762 doi: 1 0. 1 038/nprot. 2007. 247 Rohland N, Siedel H, & Hofreiter M (2010) A rapid column- based ancient DNA extraction method for increased sample throughput. Molecular Ecology Resources 10: 677-683 doi: 10.1 1 1 1/j.l 755-0998. 2009. 02824.x Ronquist F & Huelsenbeck JP (2003) MrBayes 3: Bayesian phy- logenetic inference under mixed models. Bioinformatics 19(12): 1572-1574 doi : 1 0. 1 093/ bioinformatics/ btg 1 80 Roosmalen MGM van. Mittermeier RA & Milton K ( 198 1 ) The bearded sakis, genus Chiropotes. In Coimbra-Filho AF & Mit- termeier RA (eds) Ecology and behavior of Neotropical pri- mates. Rio de Janeiro: Academia Brasileira de Ciencias: 419-441 Rowe N (1996) The Pictorial Guide to the Living Primates. Pogonias Press: East Hampton, New York Rylands AB & Mittermeier RA (2009) The diversity of the New World primates (Platyrrhini): an annotated taxonomy. Pp. 23-54 in: Garber PA, Estrada A, Bicca-Marques JC, Heymann E & Strier KB (eds) South American Primates: Comparative Perspectives in the Study of Behavior, Ecology and Conser- vation. Springer Science and Business Media Press: New York Rylands AB, Mittermeier RA & Luna ER (1995) A species list for the New World primates (Platyrrhini): Distribution by country, endemism, and conservation status according to the Mace-Land system. Neotropical Primates 3 (suppl): 113-160 Rylands AB, Schneider H, Langguth A, Mittermeier RA, Groves CP & Rodriguez-Luna E (2000) An assessment of the diver- sity of New World primates. Neotropical Primates 8: 61-93 Silva-Jr, JS & Figueiredo WMB (2002) Revisao sistematica dos cuxius, genero Chiropotes Lesson, 1840 (Primates Pithecidae). Livro de Resumos do X. Congresso da Sociedade Brasileira de Primatologia, Amazonia, Belem, Para: 2 1 Silva-Jr, JS, Figueiredo-Ready WMB & Ferrari SF (2013) Tax- onomy and geographic distribution of the Pitheciidae. Pp. 31-42 in: Veiga LM, Barnett AA, Ferrari S & Norconk MA (eds) Evolutionary Biology and Conservation of Titis, Sakis and Uacaris. Cambridge University Press, Cambridge Spix JB Ritter von (1823) Simiarum et Vespertilionum Brasi- liensium species novae on Histoire Naturelle des especies nou- velles de singes et de chauves-souris observees et recueillies pendant le voyage dans Finterieur du Bresil execute par or- dre de S M Le Roi de Baviere dans les annees 1817, 1818, 1819, 1820. Typis Francisci Seraphi Hiibschmanni, Monachii: I— VIII, 1-72, 28 Taf. Swofford D (2003) PAUP'. Phylogenetic analysis using parsi- mony ('and other methods). Sinauer Associates: Sunderland, MA Bonn zoological Bulletin 63 (2): 173-187 ©ZFMK Neotropical primates from the Cologne Zoo in the collections of the Zoologisches Forschungsmuseum A. Koenig 187 Tamura K, Peterson D. Peterson N, Steelier G, Nei M, & Ku- mar S (2011 ) MEGA5: Molecular evolutionary genetics analy- sis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evo- lution 28: 2731-2739 Traill TS ( 1821 ) Description of the Simia sagulata , - or jacket- ed monkey. Memoirs of the Wernerian Society 3: 1 67— 1 69 Tylinek E & Berger G (1984) Das grofie Affenbuch. Landbuch Verlag: Hannover Veiga LM, Silva-Jr JS, Mittermeier RA & Boubli J-P (2008) Chi- ropotes chiropotes. In: IUCN 2011. IUCN Red List of Threat- ened Species. Version 201 1.2 . Down- loaded on 10 January 2012 Weigl R (2005) Longevity of mammals in captivity: from the living collections of the world. Kleine Senckenberg-Reihe 48: 1-214 Windecker W (1969) Jahresbericht des Zoologischen Gartens Koln 1968. Freunde des Kolner Zoo 12: 3-12 Windecker W (1971) Jahresbericht des Zoologischen Gartens Koln 1970. Freunde des Kolner Zoo 14: 3-12 Windecker W (1975) Jahresbericht des Zoologischen Gartens Koln 1974. Zeitschrift des Kolner Zoo 18: 3-12 Zwickl D (2006) Genetic algorithm approaches for the phylo- genetic analysis of large biological sequence datasets under the maximum likelihood criterion. The University of Texas: Austin, TX APPENDIX List of Pitheciidae in the collections of ZFMK Bonn. Cacajao calvus. ZFMK MAM 66.612 (Cologne Zoo 1966); ZFMKM AM_8 1.1816 (Cologne Zoo 1981); ZFMK MAM 200 1 147 (Cologne Zoo 1993); ZFMKM AM_20 1 2.427 (Cologne Zoo 2008). Cacajao hosomi. ZFMK MAM 2002.032 (Venezuela/Cologne Zoo 1992). Chiropotes albinasus. ZFMK MAM 7 1.097 (Cologne Zoo 1971); ZFMK_MAM_8 1.1815 (Cologne Zoo 1981); ZFMK MAM 89.480 (Cologne Zoo 1989). Chiropotes chiropotes. ZFMK MAM 66.001 (Brazil/ Cologne Zoo 1961). ZFMK_MAM_74.155 (Duisburg Zoo 1974). Chiropotes satanas. ZFMK MAM 60.108 (Cologne Zoo 1960); ZFMK_MAM_ 81.1819 (Cologne Zoo 1981). Chiropotes sp. ZFMK MAM 2008. 244 (Cologne Zoo). Pithecia aequatorialis. ZFMK MAM 8 1.1818, male (Peru/ Cologne Zoo 1981). Pithecia inusta. ZFMK_MAM_ 87.730, female (Peru, Panguana 1973; Hutterer et al. 1995). Pithecia pithecia. ZFMK MAM 71 .097, male (Cologne Zoo 1971); ZFMK_MAM_20 12.009, male (Frankfurt Zoo 2008). Pithecia sp. ZFMK MAM 75.108, female (Duisburg Zoo 1975). Bonn zoological Bulletin 63 (2): 173-187 ©ZFMK 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. Publication in BzB is free of charge, including colour illustrations or photographs contributing significantly to quality and / or read- ability of the manuscript. Authors retain full copyright of their pub- lished papers, may share them with colleagues, and are encouraged to post the original pdfs on their personal or institutional website for non-commercial use. All material must be original, unpublished work and not under consideration for publication elsewhere. Research articles: Manuscript preparation and submission Manuscripts should be written in English. For serving readers from the organism’s country of origin, bilingual abstracts are welcome. If you plan to submit a manuscript of more than 50 manuscript pages, please contact the editor-in-chief in advance. Please submit your manuscript as electronic file (see paragraph on electronic submis- sion below). Manuscripts should strictly follow the instructions spec- ified below. Manuscripts not compatible with these guidelines will not be taken into consideration. Format Typing should be double-spaced in 10 pt throughout the text, including tables, figure legends, and reference list. Pages and lines should be numbered consecutively. Short subheadings may struc- ture the text into sections. Do not number headings or subheadings. Structure Research articles should be organized as follows: Title, Author(s), Address(es) including email address and telephone. Ab- stract, Key words. Running Title, Introduction, Material and Meth- ods, Results, Discussion (or combined Results and Discussion), Con- clusions (optional). Acknowledgements, References, Tables, Figure Captions, Figures (separately numbered and identified), Appendices (if necessary). Footnotes should not be used except in Tables. Title The title should be brief (30 words limit, ideally shorter) and indicate clearly the field of study and group of animals investigat- ed. The systematic position of taxa listed in the title must be indi- cated (e.g. “Squamata: Colubridae”). A Running Title (maximum 45 characters inclusive of spaces) should describe the paper’s core top- ic. Abstract Each manuscript should contain a concise abstract as sin- gle paragraph (max. 200 words) summarizing the significant find- ings, followed by not more than six key words. References In the text, cited sources should be referred to as fol- lows: Ax (2001 ), Kim & Lumaret (1989), Holm et al. (1997) - for three or more authors, or, if cited consecutively: (Ax 2001, 2002; Holm et al. 1997; Kim & Lumaret 1989) - please follow exactly the use of commas and semicolons. Do not use commas between author and year (only exception: taxonomic names, in which a com- ma is placed between author and year). The list of references should be arranged alphabetically according to the surname of the first author; all authors should be included. Citations of two or more authors should be arranged alphabetical- ly according to the first- and then the following author’s surname. When more than one reference is given for a single author or the same combination of authors, these should be arranged chronolog- ically. If citing more than one reference by the same author(s) pub- lished in the same year, use a, b, etc. after the year in both, text and reference list (e.g. 2006a, b). The names of journals in the References should be given in full. References “in press” shall on- ly be cited when they have been accepted for publication. Exam- ples of reference style are: Kottelat M, Whitten T, Kartikasari SN, Wirjoatmodjo S (1993) Freshwater fishes of Western Indonesia and Sulawesi. Periplus Editions, Hong Kong Mayr E (2000) The biological species concept. Pp. 17-29 in: Wheel- er QD & Meier R (eds.) Species Concepts and Phylogenetic The- ory -A Debate. Columbia University Press, New York Parenti RP (2008) A phylogenetic analysis and taxonomic revision of ricefishes, Oryzias and relatives (Beloniformes, Adrianichthyi- dae). Zoological Journal of the Linnean Society 154: 494-610 Sullivan J (1994) Bufo boreas. In: Fire Effects Information System (U.S. Department of Agriculture, Forest Service, Rocky Moun- tain Research Station, Fire Sciences Laboratory). Online at http://www.fs.fed.us/database/feis/animals/amphibian/bubo/all.ht ml last accessed on December 28, 2009 Sztencel-Jablonka A, Jones G, Bogdanowicz W (2009) Skull mor- phology of two cryptic bat species: Pipistrellus pipistrellus and P. pygmaeus - a 3D geometric morphometries approach with land- mark reconstruction. Acta Chiropterologica 11:1 13-126 Authors are kindly requested to make sure that sources listed in the References match those cited in the text. Names of persons who pro- vided unpublished information should be cited by including the ini- tial(s) and city as follows: “(JG Frommen, Bonn, pers. comm. 2009)”. Tables and Figures Tables and Figures should not duplicate infor- mation provided in the text. Tables should be numbered consecu- tively in Arabian numbers in order of their mention in the text (Table 1, Tables 1-3; do not abbreviate “Table(s)”). A brief self-explana- tory title should be typed directly above each table, not on a sepa- rate page. Do not insert frames, vertical rules, dotted lines or foot- notes. Figures (maps, graphs, charts, drawings, photographs etc.) must be numbered consecutively using Arabian numbers in order of their mention in the text (abbreviate “Fig. 1”, “Figs 1-3” etc. in the text, but use the full term “Figure(s)” in the captions. If refer- ring to illustrations in cited sources, use “fig. / figs”. Grouped fig- ures should be lettered with a lower case block letter in the lower left comer. Scale should be indicated on figures by a scale bar. A fully descriptive caption of each figure should contain all relevant information; captions are to be typed together on a separate page. Colour figures will be published free of charge if contributing sig- nificantly to quality and / or readability of the manuscript. The de- cision to print in colour or black and white any figure submitted in colour will generally be based on the referee’s and handling editor’s recommendations, but remains with the editor-in-chief and publish- er. If appropriate, authors may be asked to group single colour fig- ures into plates. For review, tables and figures should be enclosed to text (preferable one electronic file per manuscript), but may al- ternatively also be submitted as separate files (jpg, pdf, tiff). At this stage, illustrations of low resolution allowing the referees to follow the contents are sufficient; high-resolution files (jpg, pdf, tiff of pho- tographs, maps etc. at > 300 dpi; line artwork etc. at 600 dpi or in eps format) will be requested after acceptance. Unless otherwise agreed with the editor, single electronic submissions should not ex- ceed 6 MB file size. Appendix Appendices should be numbered consecutively in Roman numbers, in order of their mention in the text (Appendix I, Appen- dices I VI ). Abbreviations Except of very common abbreviations such as mm, kg etc.; all abbreviations should be explained in the Methods sec- tion or figure legend if appropriate. Hyphenation or upper case let- ters for entire words are not permitted. All measurements must be metric units and given to the same decimal, i.e., 5. 3-6.0 mm (not: 5.3-6 mm). 9 Instructions to authors Taxonomy Names of animals and the description of new genera or species must follow the current version of the International Code of Zoological Nomenclature (ICZN, available at http://www.iczn.org/iczn/index.jsp). Type specimens should be de- posited in recognised institutions; deposition at ZFMK is highly ap- preciated. Italics are required for species names which are written in full the first time they appear in the text, e.g. Carassius auratus (Linnaeus, 1 758), but abbreviated at subsequent mention, e.g. C. au- ratus. List names in synonymies as follows: Attelabus asparagi Scopoli, 1763 (Scopoli 1763: 36, fig. 1 13.), and list the source un- der References. Dichotomous keys are desirable in taxonomic pa- pers. Statistics Statistics presented should include the name of the test, number of observations or degree of freedom, and probability lev- el (P > 0.05, P < 0.05, P > 0.001, P < 0.001). Values of test statis- tics are not required. Scientific Notes Notes should comply with the instructions given above, but have to be ( 1) confined to a single point or issue of progress, (2) must be short (typically no more than 3 printed pages including illustrations and references), and (3) do not include headings or subheadings. A brief abstract of no more than three lines is required. Review papers Reviews should be concise, critical and creative, seeking to sum- marize the state-of-the-art of complex topics, and stimulate debates and new research initiatives. Review papers should comply with the instructions for preparation of Research articles, with the exception that the main body of text may be structured as appropriate. Electronic submission Manuscripts should be submitted as text files (*.doc or * .rtf; *.pdf is welcome for the review process, but for production *.doc or *.rtf will be required) via e-mail (bzb.zfmk@uni-bonn.de) to the editor- in-chief. Alternatively, send a CD. If neither is possible for any rea- son, please contact the editor-in-chief. The file name must start with the submitting author’s last name. Review process Each manuscript will be peer-reviewed, generally by two or more referees. Manuscripts should be submitted to the editor-in-chief, and will be handled by the editor dealing with the respective taxon or topic (see list of editors). Authors are encouraged to suggest poten- tial referees for their manuscripts. Final acceptance for publication is in the responsibility of the editor-in-chief. This decision is main- ly based on the referees’ report and the handling editor’s recommen- dation. Revised manuscripts received more than 6 months after the reviewers’ comments had been sent will be treated as new submis- sions. After acceptance, authors are encouraged to submit sugges- tions for a cover image. 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 infonnation 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: 1 . Is the paper of interest to the readers of BZB? 2. How well does it meet the focus of BZB? 3. 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. 5. Has the welfare of any experimental animals been adequately tak- en into account? 6. Does the title aptly and correctly describe the paper? 7. Does the abstract summarize only the significant findings? 8. Is the length of the paper appropriate? 9. Are all (colour) figures and tables necessary and appropriate? Contents 3 9088 01798 0780 Grebennikov, Vasily V.: Morimotodes, a new genus for two minute wingless litter species from southwest China and Taiwan with an illustrated overview of Molityna and Plithina genera (Coleoptera: Curculiomdae: Molityni) Nguyen, Truong Quang, Trung My Phung, Nicole Schneider, Andreas Botov, Dao Thi Anh Tran & Thomas Ziegler: New records of amphibians and reptiles from Southern Vietnam Liu, Wan-Gang, Ming Bai, Xing-Ke Yang & Dirk Ahrens: Towards an improved knowledge of Sericim of the Tibetan highland: new species and records (Coleoptera: Scarabaeidae) Peters, Gustav, Tanja Haus & Rainer Hutterer: Neotropical primates from the Cologne Zoo in the collections of the Zoologisches Forschungsmuseum Alexander Koenig: noteworthy specimens, taxonomic notes and general considerations 123 148 157 173 Cover illustration: White-faced saki Pithecia pithecia (Photo: Rolf Schlosser) (this volume, pp. 173-187). Bundesministerium fur Bildung und Forschung Ministerium fur Innovation, Wissenschaft und Forschung des Landes Nordrhein-Westfalen